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Barley Genetics Newsletter<br />
Volume 37<br />
Editorial Committee<br />
P. Bregitzer<br />
U. Lundqvist<br />
V. Carollo Blake
Table of Contents<br />
Information about the Barley Genetics Newsletter<br />
The Barley Genetics Newsletter is published electronically at<br />
http://wheat.pw.usda/gov/ggpages/bgn<br />
Contributions for publication in the Barley Genetics Newsletter should be sent to the<br />
Technical Editor, Dr. Phil Bregitzer (see Instructions to contributors below for details on<br />
submission). Reports on research activities (including ongoing projects, descriptions of<br />
new genetic and cytological techniques, and current linkage maps) are invited for the<br />
Research Notes section. Researchers are encouraged to submit descriptions of genetic<br />
stocks for the Barley Genetic Stocks section. Letters to the editors are welcome and will<br />
be published. Please inform the Technical Editor of any errors you notice in this volume<br />
or in previous volumes; corrections will be published in future volumes, and electronic<br />
copy will be appropriately corrected.<br />
All correspondence and contributions to the Barley Genetics Newsletter should be in<br />
English. Please include your complete mailing address, including your country, and an email<br />
address (if you have one) with all correspondence and contributions.<br />
Acknowledgements<br />
The contributors of research reports and the diligence of the many Coordinators make<br />
this publication possible. Victoria Carollo Blake and the <strong>GrainGenes</strong> team at the <strong>US</strong>DA-<br />
ARS make possible the electronic version of BGN.
Instructions for contributors<br />
Volume 37 (2007) of the Barley Genetics Newsletter<br />
Submissions will be published via the internet (http://wheat.pw.usda.gov/ggpages/bgn/).<br />
Approximately quarterly, new submissions will be appended to existing submissions; the<br />
page numbers of existing submissions will not change and citation information will<br />
remain constant. BGN 37 will consist of all submissions received prior to the end of the<br />
calendar year of 2007, and will be compiled into a printable version that will be available<br />
via the Graingenes website. Send submissions to:<br />
Dr. Phil Bregitzer, BGN Technical Editor,<br />
<strong>US</strong>DA-ARS, 1691 S. 2700 W.,<br />
Aberdeen, ID, 83210, <strong>US</strong>A.<br />
Telephone: (voice) 208-397-4162 ext. 116; (fax) 208-397-4165<br />
e-mail: pbregit@uidaho.edu<br />
Coordinators assigned at the International Barley Genetics Symposium should submit<br />
their reports to:<br />
Dr. Udda Lundqvist<br />
Coordinator, Barley Genetics Newsletter<br />
Nordic Genetic Resource Center<br />
P.O. Box 41<br />
SE-230 53 Alnarp, Sweden<br />
Phone: +46 40 536640<br />
FAX: +46 40 536650<br />
Cell phone: +46 70 624 1502<br />
E-mail: udda@nordgen.org<br />
Manuscripts should be written in English and be brief. Each paper should be prepared<br />
separately, with the title and the authors' names and addresses at the top of the first page.<br />
For each paper to be included in Barley Genetics Newsletter Volume 37, submit a file to<br />
the Technical Editor in one of the following ways (in order of preference):<br />
1. As an attachment to an e-mail message<br />
2. A 3.5" diskette or CD; include also one high quality printed copy<br />
3. High-quality paper copy (acceptable only in unusual circumstances)<br />
Instructions for file preparation<br />
Text and tables for each article can be in one file, but place each figure in a separate file.<br />
Name these files with your initials, a manuscript number, and "txt" or "fig" (in the case of<br />
separate files for text and figures), and a number for each figure (in the case of separate<br />
files). For example, the files for a manuscript plus two figures from Phil Bregitzer would<br />
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File Formats for Text and Tables: Preferred: Word or WordPerfect, PC-compatible<br />
formatting. Macintosh formats are not acceptable.<br />
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information and tables, which should be centered. Please use only tabs and hard returns in<br />
the preparation of text files. Tables must be prepared using the table feature of your word<br />
processing program, and not prepared using tabs. Please do not add additional lines to<br />
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that is used for text). Figures can be submitted in any common format but avoid the<br />
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white without losing information contained in the figures.
Table of Contents<br />
Barley Genetics Newsletter, Volume 37, 2007<br />
[Other volumes of the Barley Genetics Newsletter]<br />
Click here for a complete printable version of Volume 37 (Please Note: 2.12 MB)<br />
Information about the Barley Genetics Newsletter, p. i<br />
Instructions for contributors, p. ii<br />
Research Notes<br />
Dominance and recessiveness of parameters of Aluminum-resistance of<br />
barley F2 hybrids at different concentrations of stress factor (.doc, .pdf)<br />
F. M. Lisitsyn, I. I. Lisitsyna<br />
Frequency distributions and composite interval mapping for QTL<br />
analysis in ‘Steptoe’ x ‘Morex’ barley mapping population (.doc, pdf)<br />
H. S. Rao, O. P. Basha, N. K. Singh, K. Sato, H. S. Dhaliqal<br />
PCR-based markers targeting barley putative grain yield and quality<br />
QTLs regions (.doc, pdf)<br />
I. A. del Blanco, D. A. Schmierer, A. Kleinhofs , S. E. Ullrich<br />
Genetic architecture for yield and quality component traits over two<br />
environments in barley (Hordeum vulgare L.) (.doc, .pdf)<br />
A. K. Verma, S.R. Vishwakarma and P.K. Singh<br />
Line x tester analysis in barley (Hordeum vulgare L.) across<br />
environments (.doc, .pdf)<br />
A. K. Verma, S.R. Vishwakarma and P.K. Singh<br />
Four new barley mutants (.doc, .pdf)<br />
S.A.I .Wright, M. Azarang, A. B. Falk<br />
The Scandinavian barley chlorophyll mutant collection (.doc, .pdf)<br />
M. Hansson<br />
CAPS markers targeting barley Rpr1 region (.doc, .pdf)<br />
L. Zhang, A. Kleinhofs<br />
1 – 4<br />
5 - 20<br />
21 - 23<br />
24 - 28<br />
29 - 33<br />
34 - 36<br />
37 - 43<br />
44 - 46<br />
Tolerance to high copper ions concentration in the nutrient medium of 47 - 49
some Bulgarian cultivars (.doc, .pdf)<br />
J. Stoinova, S. Phileva, M. Merakchyiska, S. Paunova<br />
Coordinator's Reports<br />
Rules for Nomenclature and Gene Symbolization in Barley (.doc, .pdf)<br />
J. D. Franckowiak , U. Lundqvist<br />
Overal Coordinator's Report (.doc, .pdf)<br />
U. Lundqvist<br />
Barley Genetic Stocks for 2007 (.doc, .pdf)<br />
L. Dahleen, J.D. Franckowiak, U. Lundqvist<br />
Descriptions of Barley Genetic Stocks for 2007 (.doc, .pdf)<br />
L. Dahleen, J.D. Franckowiak, U. Lundqvist<br />
100 - 104<br />
105 - 153<br />
153 - 187<br />
188 - 301
Barley Genetics Newsletter 37:1-4 (2007)<br />
Dominance and recessiveness of parameters of Aluminum-resistance of barley<br />
F2 hybrids at different concentrations of stress factor<br />
E. M. Lisitsyn and I. I. Lisitsyna<br />
North-East Agricultural Research Institute, 166-а Lenin street, Kirov, 610007, Russia<br />
e-mail: edaphic@mail.ru<br />
Till now there is no uniform opinion in the scientific literature about number and type of<br />
action of genes coding barley aluminum resistance. For example, Rigin, Yakovleva (2001)<br />
considers, that it is controlled by two oligogenes, as a minimum, with possible action of genes<br />
with weaker effect. Other authors assume the control of the parameter by one dominant gene Pht<br />
[Stolen, Andersen, 1978], or gene Alp [Reid, 1971], both located on chromosome 4 [Minella,<br />
Sorrells, 1997].<br />
Gourley et al [1990] concluded, that type of action of genes coding aluminum resistance in<br />
sorghum (additive, partially or completely dominant), depends not only on a researched genotype,<br />
but also on used Al concentration. Aniol [1995, 1996] has established, that when the<br />
concentration of aluminum in test solution is low (30-40 µМ), cereal cultures (wheat, rye) use<br />
mechanisms that block accumulation of aluminum in roots (for example, chelation of aluminum<br />
at the expense of exudation of organic acids by root system), at higher concentration of<br />
aluminum in root growth environment (200-300 µМ) the basic role is played by other<br />
physiological mechanisms of Al-resistance. Thus the author remarks [Aniol, 1997] that the<br />
aluminum resistance of wheat plants at a concentration of 296 µМ is controlled by 2 genes and<br />
at a concentration of 592 µМ aluminum resistance is controlled by three genes. He wrote, that<br />
genes located in D-genome of wheat, are expressed only at high Al concentration, and genes<br />
located on chromosome 5А are expressed at all studied concentrations.<br />
The aim of our research was to determine the influence of Al concentration on a direction<br />
and character of dominance of parameters of roots growth of barley F2 hybrid seedlings.<br />
Material and Methods<br />
The direct and reciprocal F2 hybrids of four selection numbers of barley (№№ 565-98,<br />
889-93, 999-93 and 1030-93), bred in North-East Agricultural Research Institute (Kirov, Russia)<br />
were taken for the analysis. By results of the preliminary laboratory analyses the parental forms<br />
of these hybrids differed significantly on a level of Al-resistance that corresponded to the<br />
research aim. A level of Al-resistance (relative root length - RRL) was estimated under<br />
conditions of rolled culture on five-day barley seedlings according to the technique described<br />
earlier [Lisitsyn, 2000] by division of value of root length of each individual seedling in test<br />
treatment variant (0.5 and 1.0 mM of aluminum as sulphate salt, рН 4.3) on value of average root<br />
length of control variant (without the stress factor, рН 6.0). Each sample volume consists of 99-<br />
105 seedlings in each treatment variant.<br />
Character of dominance for parameters of root growth of F2 hybrid plants was estimated by<br />
equation [Petr, Frey, 1966]:<br />
d = F2 - MP<br />
HP-MP<br />
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Barley Genetics Newsletter 37:1-4 (2007)<br />
,<br />
where d = degree of dominance; F2, НР, МР = means of F2 hybrids, resistant parent value, and<br />
mid parent value, respectively.<br />
Results and Discussion<br />
Expression of Al-resistance genes appreciably depend on a concentration of aluminum in<br />
test solution and with its increase the resistance of all hybrids was reduced without exception<br />
(table 1).<br />
Table 1. Parameters of root growth of barley F2 hybrids under laboratory condition<br />
Hybrid<br />
0 mM Al<br />
Root length, mm<br />
0.5 mM Al 1.0 mM Al<br />
RRL, %<br />
0.5 mM 1.0 mM<br />
565-98 x 889-93 109.2±1.5 71.2±1.2 56.3±1.0 65.2±0.6 51.6±0.5<br />
889-93 x 565-93 103.0±1.1 84.7±1.3 71.3±1.2 82.3±0.7 69.2±0.7<br />
565-98 x 999-93 113.6±1.6 71.8±1.8 48.3±1.2 63.2±0.9 42.6±0.6<br />
999-93 x 565-98 103.3±2.2 65.8±0.9 55.3±1.2 63.7±0.5 53.5±0.7<br />
565-98 x 1030-93 112.7±1.1 74.2±1.1 57.0±1.3 65.8±0.6 50.6±0.7<br />
1030-93 x 565-98 110.8±1.4 79.3±1.3 65.2±1.0 71.6±0.7 58.8±0.5<br />
889-93 x 999-93 107.7±2.2 76.0±1.7 61.0±1.5 70.5±0.7 56.6±0.8<br />
999-93 x 889-93 106.9±1.1 70.7±1.5 58.2±0.8 66.1±0.8 54.5±0.4<br />
889-93 x 1030-93 107.6±1.4 75.0±2.0 64.5±1.5 69.7±1.1 59.9±0.8<br />
1030-93 x 889-93 102.1±1.6 79.0±1.0 62.9±0.8 77.4±0.6 61.6±0.5<br />
999-93 x 1030-93 104.9±1.6 75.6±1.6 52.2±1.7 72.0±0.9 49.8±0.9<br />
1030-93 x 999-93 111.2±1.2 77.4±1.0 59.6±1.2 69.5±0.5 53.6±0.6<br />
As it is visible from data, submitted in table 2, depending on the cross and aluminum<br />
concentration used, for some hybrids the large value of root length was dominated, for others<br />
hybrids – the smaller value, but for the third part of hybrids dominance of root length was absent<br />
practically. It is possible to note the same character of dominance for RRL parameter. The<br />
similar phenomenon was earlier marked in the literature for other cereals. So, [Camargo, 1981,<br />
1984] pointed out, that Al-resistance of wheat F2 population was coded by dominant genes at<br />
concentration of aluminum 3 mg/l, but became recessive at increase of concentration of the<br />
stressful factor up to 10 mg/l. Similar results were described in the researches with wheat [Bona<br />
et al., 1994].<br />
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Barley Genetics Newsletter 37:1-4 (2007)<br />
Table 2. Influence of direction of crossing on character of dominance of parameters of root<br />
growth of barley F2 hybrids<br />
Degree of dominance of a parameter<br />
Hybrid<br />
Root length RRL<br />
0 mM Al 0.5 mM Al 1.0 mM Al 0.5 mM 1.0 mM<br />
565-98 x 889-93 0.33 -0.78 -1.94 -1.21 -5.35<br />
889-93 x 565-93 -0.94 0.34 1.26 0.99 5.00<br />
565-98 x 999-93 1.19 -1.42 -1.51 -4.65 -3.13<br />
999-93 x 565-98 -0.44 -2.12 -0.70 -4.50 -0.85<br />
565-98 x 1030-93 -0.81 -0.45 -0.51 -0.37 -0.40<br />
1030-93 x 565-98 -2.00 -0.05 0.44 0.11 0.13<br />
889-93 x 999-93 4.33 1.18 1.10 -0.16 0.23<br />
999-93 x 889-93 3.80 -0.38 0.40 -1.16 -0.47<br />
889-93 x 1030-93 -0.25 10.14 1.98 1.67 1.15<br />
1030-93 x 889-93 -1.11 15.96 1.58 3.47 1.40<br />
999-93 x 1030-93 -0.35 1.05 -4.00 0.59 0.17<br />
1030-93 x 999-93 0.44 1.49 7.00 0.30 1.22<br />
As it is follows from data, submitted in table 2, depending on a concrete combination of<br />
crossing domination of root length under control conditions, under both Al treatments and of<br />
RRL parameter can have positive or negative meanings, changing from negative superdomination<br />
till positive super-domination. Character and direction of domination can coincide<br />
for parameters of roots length under control conditions and under aluminum treatment, but<br />
sometimes can have an opposite direction.<br />
Directions of crossing caused opposite character of dominance of researched parameters of<br />
Al-resistance for hybrids 565-98 х 889-93 and 889-93 х 565-98. This tendency is some less expressed<br />
at hybrids received from crossing of breeding numbers 565-98 and 1030-93. At the same<br />
time direct and reciprocal hybrids between breeding number 565-98 and breeding number 999-<br />
93 for main part of researched parameters have shown only different degree of dominance, but<br />
not its different direction.<br />
Direct and reciprocal hybrids between selection numbers 889-93 and 1030-93 had least<br />
differences on a direction and character of dominance.<br />
References:<br />
Aniol A.M. 1995. Physiological aspects of aluminum tolerance associated with the long arm of<br />
chromosome 2D of the wheat (Triticum aestivum L.) genome. Theor Appl Genet. 91: 510-<br />
516<br />
Aniol A. 1996. Aluminum uptake by roots of rye seedlings of differing tolerance to aluminum<br />
toxicity. Euphytica 92: 155-162.<br />
3
Barley Genetics Newsletter 37:1-4 (2007)<br />
Aniol A. 1997. the aluminum tolerance in wheat. plant Breeding: Theories, achievements and<br />
problems. Proc. Int. Conf., Dotnuva - Akademija, Lithuania: 14-22<br />
Ригин Б.В., Яковлева О.В. 2001. Генетические аспекты толерантности ячменя к токсичным<br />
ионам алюминия [Genetic aspects of barley tolerance against toxic Al ions] Генетические<br />
ресурсы культурных растений. Межд. науч.-практ. конф., 13-16 ноября, С-Пб: 397 [In<br />
Russian]<br />
Minella E., Sorrells M.E. 1997. Inheritance and chromosom locationof Alp. A gene controlling<br />
aluminum tolerance in 'Dayton' barley. Plant Breeding, V.116: 465-469<br />
Reid D.A. 1971. Genetic control of reaction to aluminum in winter barley. Barley Genetics II –<br />
Proc. 2 nd Int. Barley Genetics Symp., Pullman, Wash.: 409-413<br />
Stolen O., Andersen S. 1978. Inheritance of tolerance to low soil pH in barley. Hereditas, V.88.<br />
101-105<br />
Gourley L.M., Rogers S.A., Ruiz-Gomez C., Clark R.B. 1990. Genetic aspects of aluminum<br />
tolerance in sorghum. Plant Soil, V.123: 211-216.<br />
Petr F.C., Frey K.J. 1966. Genotype correlations, dominance and heritability of quantitative<br />
characters in oats. Crop Sci., V.6. 259-262<br />
Bona L., Carver B.F., Wright R.J., Baligar V.C. 1994. Aluminum tolerance of segregating wheat<br />
populations in acidic soil and nutrient solutions. Communic. Soil Sci. Plant Anal., V.25.<br />
327-339<br />
Camargo C.E.O. 1981. Wheat improvement. 1. The heritability of tolerance to aluminum<br />
toxicity. Bragantia, V.40. 33-45<br />
Camargo C.E.O. 1984. Wheat improvement. IV. Heritability studies on aluminum tolerance<br />
using three concentrations of aluminum in nutrient solutions. Bragantia, V.44. 49-64<br />
Lisitsyn E. M. 2000. Intravarietal Level of Aluminum Resistance in Cereal Crops. J Plant Nutrit.,<br />
V.23(6): 793-804<br />
4
Barley Genetics Newsletter 37:5-20 (2007)<br />
Frequency distributions and composite interval mapping for QTL analysis in<br />
‘Steptoe’ x ‘Morex’ barley mapping population<br />
ABSTRACT<br />
H. S. Rao 1, 2† , O. P. Basha 1 , N. K. Singh 1 , K. Sato 3 , H. S. Dhaliwal 1 *<br />
Indian Institute of Technology Roorkee, UA, India<br />
1 <strong>Department</strong> of Biotechnology, Indian Institute of Technology Roorkee,<br />
Roorkee 247667, Uttaranchal, INDIA<br />
2 Present address: 101, Maize Genetics Unit, Division of Genetics, LBS Building,<br />
Indian Agricultural Research Institute, New Delhi 110012, INDIA<br />
3 Barley Germplasm Center, Okayama University, Kurashiki 710-1146, JAPAN<br />
*corresponding author Email: hsdhafbs@iitr.ernet.in<br />
FAX: 91-1332-273560 Tel.: 01332-285259<br />
† Additional author Email: hsr_f12@rediffmail.com<br />
With the advancement of QTL mapping strategies, the traditional approaches for the<br />
identification of genes and their effects responsible for trait expression are gradually losing<br />
significance. The phenotypic data for heading days, plant height, peduncle length, number of<br />
tillers and stripe rust was recorded on 150 recombinant inbred line (RIL) population developed<br />
from a barley cross of ‘Steptoe’ and ‘Morex’. Preliminary examinations of frequency distribution<br />
plots were fairly useful in the prediction of number of genes governing traits expression in barley<br />
(Hordeum vulgare L.). The predictions were then systematically confirmed through QTL<br />
mapping. Molecular marker data for the population available at the public domain (<strong>GrainGenes</strong><br />
website) was used for the construction of linkage map and QTL analysis. Clear colinearity was<br />
observed between the number of QTLs identified and the number of genes predicted based upon<br />
frequency distribution study alone. A total of 17 QTLs were detected for the five traits evaluated.<br />
Several major QTLs were detected on chromosome 2H, which could serve as candidate for mapbased<br />
studies of phenomena such as pleiotropism, recombination hot spots, gene-rich regions and<br />
QTL clustering.<br />
Keywords. Barley, frequency distribution, quantitative trait loci, recombinant inbred line,<br />
peduncle length, stripe rust, gene rich region.<br />
INTRODUCTION<br />
Cultivated barley (Hordeum vulgare L.) is a diploid (2n=14) which ranks fourth among<br />
the most important cereal crops in the world after rice, wheat and maize. Barley belongs to the<br />
same tribe Triticeae as that of wheat and rye and it resembles wheat in many respects. Barley is<br />
however, more tolerant to soil salinity and drought than wheat. Many important traits of<br />
economic and agronomic importance in barley are quantitative in nature displaying continuous<br />
5
variation. The efficiency of breeding programs depends on our knowledge of the genetic control<br />
and genomic location of QTLs governing the trait(s) of interest. With the advent of molecular<br />
markers (Botstein et al. 1980) and the user-friendly statistical software it has become possible to<br />
resolve and map the QTLs for complex traits on chromosomes.<br />
QTL analysis in barley is significant not only for the crop itself but also for comparative<br />
mapping with other cereal crops. The identification of diverse taxa sharing segments of similar<br />
gene orders throughout their genomes has been the major outcome of comparative mapping<br />
where chromosome alignment has hastened identification of new genes for their ultimate<br />
introgression into suitable cultivars. The genetic maps of diploid wheat, Triticum monococcum<br />
and barley, Hordeum vulgare L. are remarkably conserved except for few regions where<br />
translocation and inversion of chromosome segments have taken place (Dubcovsky et al. 1996).<br />
There is also a high level of genomic conservation at specific regions with rice, maize and oat<br />
(Deynze et al. 1995, Han et al. 1998).<br />
The results of several QTL mapping studies have indicated that there are only few major<br />
genes, which interact with numerous minor genes and environment to give continuous trait<br />
phenotype characteristic of quantitative traits. Thus examination of frequency distribution plots<br />
at single location is useful in assessing the probable number of major genes controlling a trait.<br />
This is particularly useful when resources are limiting and a decision has to be taken on priority<br />
basis before starting a breeding program for which trait QTL mapping experiment should be<br />
undertaken. The present article deals with the mapping of QTLs for a number of traits in a barley<br />
‘Steptoe’ x ‘Morex’ RIL mapping population and its relationship with the predictive value of<br />
frequency distribution for the traits.<br />
MATERIALS AND METHODS<br />
Plant Material<br />
150 recombinant inbred lines (RILs) derived from a cross between two barley (Hordeum<br />
vulgare L.) genotypes ‘Steptoe’ and ‘Morex’, obtained from Dr. Kazuhiro Sato, Okayama<br />
University, Japan was used as a mapping population and it was phenotyped for five traits. Each<br />
of the RILs and the parents were sown as a single one meter row with row to row spacing of<br />
30cm in the experimental field at the Indian Institute of Technology Roorkee, India in the winter<br />
season of 2005. Normal fertilizers, irrigation and other agronomic practices were followed for<br />
growing the population. Both ‘Steptoe’ and ‘Morex’ are hulled, six-rowed spring barley<br />
varieties. ‘Steptoe’ is a late flowering and dwarf line with reduced peduncle, few tillers per plant<br />
and moderate resistance to stripe rust. On the other hand, ‘Morex’ is early flowering and tall<br />
genotype with long peduncle, more tillers per plant and high susceptibility to stripe rust.<br />
Phenotypic Data<br />
The data on five quantitative traits were recorded as the average of five competitive<br />
plants per RIL. The number of days to heading was recorded as the number of days from sowing<br />
till half of the tillers in a RIL had flowered. Plant height (cm) was recorded as the length of plant<br />
from the base at the soil surface to the tip of spike of the tallest tiller excluding awns. Peduncle<br />
6
length was measured as the length of peduncle from the base of flag leaf to the base of basal<br />
spikelet of a spike. The number of effective tillers bearing spikes was taken as the number of<br />
tillers per plant. The data on natural incidence of stripe rust at the adult plant stage under field<br />
conditions was recorded as percent severity of the leaf area covered by stripes of uredia and the<br />
type of pustules, where the symbol S for susceptible was used for large pustules without any<br />
necrotic area; MS, MR for moderately susceptible/resistance for small pustules with or without<br />
necrotic areas around them and R for hypersensitive immune reaction without any pustule.<br />
Data Analysis<br />
The basic statistical analysis was performed for all the traits recorded. Mean, range and<br />
standard deviations were estimated. Correlation coefficient among various traits was calculated<br />
to infer probable inter-relationships between the traits studied. Frequency distribution plots for<br />
traits were presented as about ten phenotypic classes in the RIL population.<br />
Genotypic data and Linkage mapping<br />
Genotypic data for 343 molecular markers available at the public domain (<strong>GrainGenes</strong><br />
website: http://www.gene.pbi.nrc.ca) for 150 RILs of ‘Steptoe’ x ‘Morex’ mapping population<br />
was utilized for the construction of linkage map. Standard χ 2 test was used to test the segregation<br />
pattern of each marker. Linkage map was constructed by using the software package<br />
MAPMAKER/EXP version 3.0 (Lincoln et al. 1992). A LOD score of 3.0 and a maximum<br />
recombination frequency of 0.40 were used to declare linkage between two markers.<br />
QTL mapping<br />
QTL analysis was performed using the method of Composite Interval Mapping (CIM)<br />
(Zeng 1994) as in QTL Cartographer version 2.5 (Wang et al. 2005). Composite interval<br />
mapping combines the approaches of interval mapping (IM) and Single Marker Analysis in a<br />
multiple regression framework. Initially it builds cofactors by selecting most significant markers<br />
through Single Marker Analysis methodology. Once the model containing cofactors is built, the<br />
entire genome is rescanned using interval mapping. We used model 6 with window size 10 cM<br />
where forward and backward regression method was utilized. Walk speed was set at 2 cM to<br />
scan the entire genome. We performed 1000 permutations at 0.05 significance level to balance<br />
type 1 and type 2 errors and declare appropriate threshold levels for QTL (Churchill and Doerge<br />
1994). The best estimate of QTL location was assumed to correspond to the position having the<br />
peak significance level and the confidence interval was drawn according to 1-LOD support<br />
interval (Lander and Botstein 1989).<br />
RESULTS<br />
Frequency distribution for various traits<br />
The frequency distributions for the five traits evaluated are given in Figure 1. Only plant<br />
height showed normal distribution while all other traits displayed various levels of skewedness.<br />
The days to heading trait was roughly partitioned into two phenotypic classes, one with early<br />
7
heading habit while the other showing late heading trait. Peduncle length also showed two<br />
phenotypic classes but the distribution tended to be strongly influenced by embedded peduncle<br />
genotype rather than emerged peduncle. Tiller numbers showed broad range with transgressive<br />
segregation towards both ends. The histogram for stripe rust showed continuous disease<br />
distribution pattern and the phenotypic classes were clearly partitioned into five clusters.<br />
Phenotypic data Analysis<br />
Except for the heading days all traits showed transgressive segregation and their phenotypic<br />
values exceeded beyond both of the mean parental values (Table 1). The correlation coefficients<br />
between most of the trait combinations were found to be significant ( Table 2). As expected<br />
peduncle length had extraordinarily high correlation with heading days and plant height.<br />
Peduncle length and heading habit were negatively correlated, i.e. the emerged peduncle inbred<br />
lines had more probability of early heading habit whereas the embedded peduncle lines had late<br />
heading. The significant positive correlation between plant height and peduncle extrusion could<br />
explain the fact that in taller plants, peduncle grew faster to emerge out of the boot leaf before<br />
anthesis.<br />
TABLE 1. Trait means in parent and trait means, standard deviations (SD) and range in RILs of<br />
Steptoe x Morex population<br />
Parents RIL population<br />
Trait Steptoe Morex Range Mean ± SD<br />
Heading days 126.0 100.0 101.0 – 126.0 112.05 ± 7.35<br />
Plant height (cm) 66.4 100.1 61.0 – 126.6 95.57 ± 13.27<br />
Peduncle length (cm) 5.7 24.2 0 – 28.6 10.21 ± 6.61<br />
Tiller number 2.6 3.1 1.2 – 6.4 3.49 ± 1.08<br />
Stripe rust (%) 12.5 75 0 – 100 43.11 ± 22.83<br />
TABLE 2. Correlation coefficient (r) among various traits in RIL population<br />
Plant height Peduncle length Tiller number Stripe rust<br />
Heading days -0.5491* -0.7392* -0.4987* -0.2056<br />
Plant height 0.6360* 0.5112* 0.1595<br />
Panicle exertion 0.3880* 0.1334<br />
Tiller number 0.2111<br />
* P < 0.0001<br />
Linkage map<br />
Out of the 434 polymorphic molecular markers data available at <strong>GrainGenes</strong> website, we<br />
selected 343 by rejecting markers with more than 40% missing genotypes and those showing<br />
segregation distortion at 0.05 significance level (χ 2 =3.841). Except for the two regions in<br />
chromosomes 5 (1H) and 6 (6H), the whole genome was adequately covered with markers and<br />
the centromeres were placed on consensus positions based on marker orders along the<br />
chromosomes. The order of markers and centromere positions did not vary much from their<br />
8
established map positions. The combined length of the linkage map was 824.1 cM with average<br />
spacing of 2.40 cM between adjacent markers.<br />
Genomic distribution of QTLs<br />
QTLs detected by CIM analysis as implemented in QTL Cartographer version 2.5 are<br />
presented in Table 3. The parent contributing respective alleles for increasing trait value for<br />
heading days (HD), plant height (PH), peduncle length (PL), number of tillers (TN) and the allele<br />
conferring resistance to stripe rust (SR) were indicated in the table. In the present study, five<br />
QTLs were identified for heading days exceeding the threshold LOD score. ‘Morex’ parent<br />
contributed early heading alleles for all the QTLs identified. Most of the phenotypic variance<br />
was explained by the QTLs on 2H and 1H while those on 7H and 3H had only a minor effect.<br />
Identification of several plant height QTLs spread throughout genome (7H, 2H, 2H, 3H, 6H, 6H<br />
and 5H) supported the results of frequency distribution. The two QTLs on 2H and 3H together<br />
explained about 40% of the phenotypic variance. The alleles responsible for increase or decrease<br />
in height had come from both of the parents and thus supporting transgressive segregation<br />
observed for some of the RILs. Two major QTLs, one each on chromosomes 2H and 3H were<br />
identified for peduncle length while a third putative QTL detected on 1H had only a minor effect.<br />
Only one QTL was identified for tillering ability on chromosome 2H. For stripe rust, although<br />
four resistance QTLs spread across chromosomes 2H, 3H, 4H and 5H were identified, none of<br />
them explained significant phenotypic variance.<br />
TABLE 3. Chromosome mapping of various QTLs with nearest linked molecular marker for<br />
different traits<br />
Trait Chrom Marker interval Position LOD Additiv R 2 x Allele<br />
Heading<br />
date<br />
Plant<br />
height<br />
Peduncle<br />
length<br />
osome<br />
(cM)<br />
1 (7H) ABC156d - ABG022A 41.61 4.35 1.8964 6.60 S<br />
2 (2H) ABG005 - Pox 33.41 18.78 4.1443 31.42 S<br />
2 (2H) Adh8 – CDO537 47.00 6.39 2.47 8.00 S<br />
3 (3H) ABG471 - ABG399 38.11 3.22 1.6616 4.96 S<br />
5 (1H) ABC307A-cMWG706A 31.31 11.69 3.2381 19.35 S<br />
1 (7H) Pgk2B - PSR129 67.91 2.49 2.8861 4.72 S<br />
2 (2H) ABG005 - Pox 33.41 8.83 -5.7128 18.43 M<br />
2 (2H) Adh8 - CDO537 43.31 8.41 -6.1379 20.57 M<br />
3 (3H) ABC156c - AtpbB 46.81 12.37 -5.7998 18.77 M<br />
6 (6H) CDO497 - ABR335 40.91 3.07 3.2705 6.00 S<br />
6 (6H) BCD340E - ksuD17 45.91 3.74 3.5796 7.24 S<br />
7 (5H) ABG708 - Dor5 30.31 4.93 -4.2142 9.72 M<br />
2 (2H) ABG358 - ABG459 28.71 19.30 -3.9943 36.24 M<br />
3 (3H) ABG471 - ABG399 38.11 4.63 -2.6111 15.38 M<br />
5 (1H) ABC307A 92.31 2.61 -1.7200 6.76 M<br />
Tiller # 2 (2H) MWG858 - ABG358 28.61 4.89 -0.3615 11.02 M<br />
Stripe rust<br />
resistance<br />
2 (2H) MWG858 - ABG358 26.61 2.95 -4.9528 6.22 S<br />
3 (3H) ABG377 - MWG555b 61.31 5.10 7.0202 11.30 M<br />
4 (4H) ABC321 - ABR315 30.21 3.24 -5.1908 6.88 S<br />
7 (5H) ABG391 124.81 2.64 -4.6547 5.56 S<br />
9<br />
e<br />
100
DISC<strong>US</strong>SIONS<br />
Frequency distribution of various traits<br />
RILs are inbred lines derived from a cross of two diverse parents in which the individual<br />
genes are resolved into homozygous progenies. If we construct a histogram on such a population,<br />
the number and size of phenotypic classes obtained is directly related to the number of genes<br />
influencing the trait. For example, if a single gene controls the trait, there will be two phenotypic<br />
classes of equal sizes and if two genes control the trait there will be three phenotypic classes in<br />
1:2:1 size proportion. If we assume the genes are additive and explaining equal variance, their<br />
should be n + 1 number of phenotypic classes observed for n number of genes in the population<br />
for as many number of additive gene combinations. On the other hand if epistatic interactions<br />
were significant and the individual genes were contributing unequally towards the overall<br />
phenotype, the distribution of phenotypic classes becomes skewed. From the preliminary<br />
observation of histograms (Figure 1) the approximate number of genes responsible for each trait<br />
could be predicted. For example, there should be one gene explaining most of the phenotypic<br />
variance for heading days as we observed two broad phenotypic classes in the histogram. There<br />
should also be one more gene with lesser but significant effect, which is responsible for minor<br />
third phenotypic class in between the two major classes. Height is the perfect example of<br />
quantitative inheritance where we expect large number of alleles acting additively to give normal<br />
distribution. The presence of two phenotypic classes is the indicative of single gene inheritance<br />
for peduncle length, but their unequal proportion could be explained by presence of one more<br />
parallel allele contributing towards short peduncle length phenotype. Tillering trait showed<br />
normal distribution and large transgressive segregation beyond both parental types. Thus, in<br />
barley we do not expect tiller number to be governed by single gene. Although we observed only<br />
one QTL exceeding the threshold LOD score, there is an indication of two more putative QTLs<br />
on chromosomes 7H and 1H (Figure 3). Also, some of the QTLs on chromosomes 3H and 6H<br />
identified by Franckowiak and Lundqvist, 2002, Buck-Sorlin, 2002 and Babb and Muehlbauer,<br />
2003, remained undetected in the present study. The most probable explanation could be that<br />
‘Steptoe’ and ‘M’ are not diverse enough with respect to tillering ability and it actually limited<br />
the QTL mapping approach to detect all genes of the trait. Frequency distribution of stripe rust<br />
has shown overall normal disease distribution pattern clustered in three major segregative and<br />
two minor transgressive groups. Both parents were susceptible to stripe rust but the rust<br />
progressed slowly in ‘Steptoe’ with very low terminal severity, usually called slow rusting. The<br />
five classes for rust severity could be explained by the presence of four genes, each of which was<br />
contributing additively towards resistance to stripe rust. When the resistance alleles for all four<br />
genes were present in a single genotype (4R), maximum resistance is expected which was<br />
observed with zero percent rust severity in the first transgressive group. Similarly, genotypes<br />
with 3R+1S, 2R+2S, 1R+3S and 4S allele combinations explained second, third, fourth and fifth<br />
groups, respectively in the frequency distribution plot.<br />
QTL Analysis<br />
Heading Days: Days to Flower or heading days is considered to be an important trait for<br />
planning of a breeding program. Early-heading genotypes are preferred when the objective is to<br />
10
grow a cultivar late or early in the growing season. Heading time in barley and wheat is governed<br />
by three major genetic systems: vernalization requirement (response to low temperature at the<br />
initial stages of plant development), photoperiod sensitivity (day length) and narrow-sense<br />
earliness (response to sum of temperature over a long period). Vernalization is the requirement<br />
of low temperature period to plants for transition from a vegetative to a reproductive phase.<br />
Among the vrn loci, Vrn1 on the group 5 chromosomes of Triticeae (A, B, D and H genomes)<br />
are the most extensively characterized in terms of its effects and inheritance (Law et al. 1976,<br />
Galiba et al. 1993, Kato et al. 1999). The vernalization responsive phenotype is often found in<br />
conjunction with photoperiod sensitivity, a delay in flowering when plants are grown under<br />
short-day conditions (Karsai et al. 2001). Low vernalization requirement of barley was probably<br />
met with partially but longer photoperiod could be available only in the last week of March and<br />
hence delayed flowering in RILs with Ppd. The Ppd – H1 photosensitivity locus was first<br />
described by Laurie et al. (1994) on the short arm of chromosome 2H. In barley, eps 2S, located<br />
near the centromere region of the 2H chromosome has been reported to be a QTL for the<br />
environment independent narrow-sense earliness (Laurie et al. 1995). The RILs in the present<br />
investigation were sown without vernalization treatment. The QTL analysis carried out without<br />
partitioning of heading trait into three categories identified two significant QTL on 2H which<br />
probably represent Ppd and eps loci. Thus major effect on heading days under field conditions<br />
was due to the photoperiod and environment independent narrow sense earliness genes. Major<br />
role of 2H in determining heading days in barley is also inferred from the investigations carried<br />
out by Marquez-Cedillo et al. (2001), Kicherer et al. (2000), Karsai et al. (2005) and Qi et al.<br />
(1998) who detected heading days QTL on chromosomes 2H; 2H; 2H, 4H, 5H; and 2H, 7H,<br />
respectively.<br />
Plant Height: Before the green revolution, one of the major causes of yield loss was lodging of<br />
tall cultivars during rain and strong winds. Plant height and culm stiffness are reported to be the<br />
two most important traits determining lodging resistance in cereal plants (Keller et al. 1999).<br />
Murthy and Rao (1980) and Stanca et al. (1979) have worked out significant correlation between<br />
lodging resistance and dwarfness in barley. Recently Chloupek et al. (2006) demonstrated the<br />
significance of different sets of semi-dwarf genes in the determination of different root system<br />
size that was further implicated with biotic and abiotic stresses. With the development of semidwarf<br />
varieties, yield loss was largely overcome and the plant utilized its resources in increasing<br />
harvest index rather than its biomass. If the objective of the breeding program is to enhance the<br />
grain production as in most instances, the breeder prefers dwarf variety but if the objective is to<br />
utilize the plant for dual purpose for fodder as well, then the breeder obviously goes with taller<br />
varieties. Several studies in the recent past had identified QTL for plant height in barley<br />
distributed throughout genome but the major QTLs identified in the present study on<br />
chromosome 2H and 3H were found to occur more often than others. For example, Thomas et al.<br />
(1995) detected plant height QTLs on 1H, 3H and 7H; Marquez-Cedillo et al. (2001) on 2H, 3H,<br />
4H and 5H; Kicherer et al. (2000) identified on 2H and 3H; Qi et al. (1998) on 2H, 3H and 7H;<br />
Teulat et al. (2001) on 2H, 3H, 4H, 5H, 6H and 7H; Zhu et al. (1999) on 1H, 3H, 4H and 6H; and<br />
Chloupek et al. (2006) on 3H, 4H, 5H and 7H.<br />
Peduncle length: Peduncle length is an important character in barley as the inability of spikes to<br />
emerge out of boot leaf not only eliminates outcrossing but also adversely affects the use of its<br />
photosynthetic contribution to seed-setting and seed development. From the preliminary<br />
11
observation on correlation coefficient, we can predict that heading date and plant height jointly<br />
influences the ability of peduncle to emerge out since early heading and tall plants had emerged<br />
peduncle. We can infer that photoperiod (2H) had major effect on peduncle length while narrow<br />
sense earliness (2H and 1H) had relatively minor effect on peduncle length. Chromosome 3H<br />
also had significant influence on the trait which might be coming from major plant height QTL<br />
on the same chromosome. Hence, we can conclude that peduncle length is a composite trait<br />
whose component traits include heading days and plant height.<br />
Number of Tillers: Tiller number is regarded as an important yield component in wheat, barley<br />
and rice as the number of effective tillers is equal to the number of spikes. The plants with lesser<br />
tiller number generally have long spikes with increased grain weight and overall sturdy plant<br />
architecture (Vasu at al. 2006). But the advantage of high tiller numbered plant is that the overall<br />
harvest index from single plant is much higher and it saves the unit area of land required to grow<br />
plants for similar yields. In the present study, the tillering ability of plants behaved as a single<br />
gene inherited trait and only one QTL on chromosome 2H could be detected. Although earlier<br />
studies have also indicated single gene inheritance pattern for the trait, chromosomal locations of<br />
QTLs were not consistent with the present study. Franckowiak and Lundqvist, 2002, Buck-Sorlin<br />
2002 and Babb and Muehlbauer 2003 have identified major QTL lnt1 on chromosome 3HL and<br />
a second QTL cul2 on 6HL.<br />
Stripe Rust: Stripe rust is a major disease in Triticeae and its causal organism in barley is a<br />
biotrophic fungus, Puccinia striiformis f. sp. hordei. The characteristic feature of the disease is<br />
the appearance of pale stripes on leaves followed by emergence of orange brown uredosori that<br />
contain fungal spores. Yield loss of upto 30 % may occur, as photosynthetic and metabolic<br />
capabilities of the plant are severely impaired. Rust resistance in Triticeae is mainly of two types<br />
viz., vertical and horizontal. Vertical resistance is race-specific conditioned by gene-for-gene<br />
interaction between host and pathogen (Flor 1946). Horizontal resistance, on the other hand is<br />
race-non-specific governed by multiple genes with small effects. The results of the present<br />
analysis show that the resistance QTLs identified were acting in a race non-specific manner<br />
where they prevent pathogen(s) to form a basic compatibility reaction. The ‘Steptoe’ is a slow<br />
stripe rusting genotype. The rust infection is of susceptible type but the disease progresses very<br />
slowly without causing an appreciable loss. Some of the QTLs for resistance mapped in the<br />
present study were colinear with the previously mapped QTL using different barley populations<br />
on chromosomes 4H, 5H (Chen et al. 1994); 2H, 3H, 1H, 6H (Toojinda et al. 2000); 1H, 2H, 4H,<br />
6H, 7H (Berloo et al. 2001); 2H, 4H (Kicherer et al. 2000); and on 3H, 4H, 7H (Toojinda et al.<br />
1998) using local rust pathotypes further confirming their race-non-specific resistance. The<br />
QTLs with their low individual phenotypic variances when pyramided in a single genotype could<br />
confer high overall resistance. The approach for pyramiding of resistance QTLs was well<br />
demonstrated for 1H, 4H and 7H conferring resistance to stripe rust at seedling stage in barley by<br />
Castro et al., (2003).<br />
QTL clusters<br />
Although the large genome size (4.9 × 10 9 bp) of barley makes the genetic manipulations<br />
difficult, some of the recent advances in comparative genomics have suggested the presence of<br />
recombination hot spots and gene-rich regions in Triticeae where targeted approaches could be<br />
utilized for genome analysis (Gill et al. 1996). Identification of multiple QTLs on chromosomes<br />
12
2H and 3H were also indicative of gene-rich regions in barley. 2H is especially important<br />
because it contained QTLs for all of the traits analyzed in the present study. Aissani and Bernardi<br />
(1991) suggested the distribution of less conserved genes in euchromatic regions while the<br />
distribution of conserved genes in the heterochromatin region. This could be of evolutionary<br />
significance as the genes in euchromatin region exposed to manipulations were providing<br />
diversity for the plants to adjust to different environmental conditions while the genes of<br />
heterochromatin region might be regulatory in function, which must be conserved for their<br />
essential role in the very existence of the organism.<br />
Although the QTL analysis is of tremendous use in the identification of genomic regions<br />
pertaining to the traits of economic importance, it fails to identify the complete set of genes of a<br />
biochemical pathway leading to the expression of trait. Many of the conserved genes for which<br />
there are no allelic variants cannot be detected by this approach. Such genes have been conserved<br />
through evolutionary forces and any attempt to change them could have deleterious effect on the<br />
survival ability of the organism.<br />
CONCL<strong>US</strong>ION<br />
It has been shown here that QTL mapping is an accurate approach for the identification<br />
of genes underlying a trait but it is also cost intensive to carry out such exercise for all traits. On<br />
the other hand, construction of frequency distribution plots in routinely developed mapping<br />
populations could be used as diagnostic assessment for the prediction of number and effects of<br />
genes before planning QTL mapping experiment. Although such an approach is very crude, it<br />
would be useful in making appropriate allocation of resources.<br />
In the present study, it has been found that proximal region of chromosome 2H is of<br />
special interest as it contains QTLs for all of the traits studied explaining large phenotypic<br />
variances and thus the region could serve as candidates for further investigation. Significant<br />
correlation among various traits was because of QTL linked in coupling phase and in some of the<br />
instances pleiotropism cannot be ruled out. The other genomic regions of interest include<br />
chromosome 3(3H) and chromosome 5(1H), which harbors QTL for heading, peduncle length<br />
and plant height. Further study needs to be carried out to fine map and ultimately dissect out the<br />
individual genes in the region. Map-based cloning is gradually becoming a feasible approach for<br />
dissection of the QTL genes in Triticeae (Li et al. 2003). The individual QTL effects could be<br />
studied efficiently by generating near isogenic lines (NILs) in an organized backcross program,<br />
which specifically nullifies the background noise. In the recent past bioinformatics tools have<br />
gained significant importance to aid in silico comparative genomic studies in species with large<br />
unmanipulable genomes like barley and wheat with the help of well studied genomes such as<br />
Arabidopsis thaliana, rice and maize.<br />
ACKNOWLEDGEMENT<br />
The authors extend thanks to Dr. Firoz Hossain, Indian Agricultural Research Institute, New<br />
Delhi for his valuable suggestions and discussion while preparing the manuscript.<br />
13
Figure 1. Frequency distribution plots for heading days (HD), plant height (PH), peduncle length<br />
(PL), tiller number (TN), and stripe rust (SR) in RILs of Steptoe x M population.<br />
14
Figure 2. Genetic map of SM barley RIL population with 343 molecular markers indicating QTLs<br />
for heading days (HD), plant height (PH), peduncle length (PL), number of tillers (TN) and stripe<br />
rust (SR). Vertical bar and the symbols represent confidence interval and peak LOD scores<br />
respectively for various QTLs<br />
15
Figure 2. (continued) Genetic map of SM barley RIL population with 343 molecular markers<br />
indicating QTLs for heading days (HD), plant height (PH), peduncle length (PL), number of<br />
tillers (TN) and stripe rust (SR). Vertical bar and the symbols represent confidence interval and<br />
peak LOD scores respectively for various QTLs.<br />
16
Figure 3. QTL likelihood maps for various traits obtained from composite interval mapping<br />
(CIM) analysis indicating LOD score along the ordinate while genetic map (all chromosomes<br />
together) along the abscissa. The respective threshold LOD estimated by 1000 permutations at<br />
0.05 significance, are represented as horizontal line.<br />
17
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Lincoln, S. E., M. Daly, and E. S. Lander, 1992: Constructing genetic maps with<br />
MAPMAKER/EXP 3.0. Whitehead Institute technical report, 3 rd edn. Cambridge<br />
Marquez-Cedillo, L. A., P. M. Hayes, A. Kleinhofs, W. G. Legge, B. G. Rossnagel, K. Sato, S.<br />
E. Ullrich, and D. M. Wesenberg, The North American Barley Genome Mapping Project,<br />
2001: QTL analysis of agronomic traits in barley based on the doubled haploid progeny of<br />
two elite North American varieties representing different germplasm groups. Theor. Appl.<br />
Genet. 103, 625-637<br />
Murthy, B. N., and M. V. Rao, 1980: Evolving suitable index for lodging resistance in barley.<br />
Indian J. Genet. Plant. Breed. 40, 253-261<br />
Qi, X., R. E. Niks, P. Stam, and P. Lindhout, 1998: Identification of QTLs for partial resistance<br />
to leaf rust ( Puccinia hordei) in barley. Theor. Appl. Genet. 96, 1205–1215<br />
Stanca, A. M., G. Jenkins, and P. R. Hanson, 1979: Varietal responses in spring barley to natural<br />
and artificial lodging and to a growth regulator. J. Agric. Sci. Camb. 93, 449-456<br />
Teulat, B., O. Merah, I. Souyris, and D. This, 2001: QTLs for agronomic traits from a<br />
Mediterranean barley progeny grown in several environments. Theor. Appl. Genet. 103,<br />
774–787<br />
Thomas, W. T. B., W. Powell, R. Waugh, K. J. Chalmers, U. M. Barua, P. Jack, V. Lea, B. P.<br />
Forster, J. S. Swanston, R. P. Ellis, P. R. Hanson, and R. C. M. Lance, 1995: Detection of<br />
quantitative trait loci for agronomic, yield, grain and disease characters in spring barley<br />
(Hordeum vulgare L.). Theor. Appl. Genet. 91, 1037-1047<br />
Toojinda, T., E. Baird, A. Booth, L. Broers, P. Hayes, W. Powell, W. Thomas, H. Vivar, and G.<br />
Young, 1998: Introgression of quantitative trait loci (QTLs) determining stripe rust<br />
resistance in barley: an example of marker-assisted line development. Theor. Appl. Genet.<br />
96, 123–131<br />
Toojinda, T., L. H. Broers, X. M. Chen, P. M. Hayes, A. Kleinhofs, J. Korte, D. Kudrna, H.<br />
Leung, R. F. Line, W. Powell, L. Ramsay, H. E. Vivar, and R. Waugh, 2000: Mapping<br />
19
quantitative and qualitative disease resistance genes in a doubled haploid population of<br />
barley (Hordeum vulgare). Theor. Appl. Genet. 101, 580–589<br />
van Berloo, R., H. Aalbers, A. Werkman, and R. E. Niks, 2001: Resistance QTL confirmed<br />
through development of QTL-NILs for barley leaf rust resistance. Mol. Breed. 8, 187-195<br />
Vasu K., S. Sood, H. S. Dhaliwal, P. Chhuneja, B. S. Gill, 2006: Identification and mapping of a<br />
tiller inhibition gene (tin3) in wheat. Theor. Appl. Genet. (Published online)<br />
Wang, S., C. J. Basten, Z. B. Zeng, 2005: Windows QTL Cartographer 2.5. <strong>Department</strong> of<br />
Statistics, North Carolina State University, Raleigh<br />
http://www.statgen.ncsu.edu/qtl/crt/WQTL.htm<br />
Zang, Z. B., 1994: Precision mapping of quantitative trait loci. Genetics 136, 1457-1468<br />
Zhu, H., L. Gilchrist, P. Hayes, A. Kleinhofs, D. Kudrna, Z. Liu, L. Prom, B. Steffenson, T.<br />
Toojinda, and H. Vivar, 1999: Does function follow form? Principal QTLs for Fusarium<br />
head blight (FHB) resistance are coincident with QTLs for inflorescence traits and plant<br />
height in a doubled-haploid population of barley. Theor. Appl. Genet. 99, 1221–123<br />
20
Barley Genetics Newsletter 37:21-23 (2007)<br />
PCR-based markers<br />
targeting barley putative grain yield and quality QTLs regions<br />
Isabel A. del Blanco*, Deric A. Schmierer, Andris Kleinhofs and Steven E. Ullrich<br />
<strong>Department</strong> of Crop and Soil Sciences, Washington State University, Pullman, WA, 99164-6420<br />
*e-mail: mailto:blanco@wsu.edu<br />
Abstract<br />
Eight restriction fragment length polymorphism (RFLP) and two genes delimiting, or included<br />
in, chromosome fragments containing putative QTLs for grain yield and quality were sequenced<br />
and converted to PCR-markers. Eight markers were co-dominant between two-rowed barley<br />
cultivars Harrington and Baronesse after digestion with restriction enzymes. Three were<br />
dominant-recessive after designing specific primers exploiting single nucleotides polymorphisms<br />
(SNPs) between those two cultivars.<br />
Introduction<br />
Two chromosomal regions from Baronesse have been reported as containing putative grain yield<br />
QTLs, one on chromosome 2(2HL) (between markers ABG461C and MWG699) and the other<br />
on chromosome 3(3HL) (between markers MWG571A and MWG961) (Schmierer et al., 2004).<br />
Subsequent analysis of Harrington/Baronesse derived inbred lines suggested other additional<br />
regions as candidates for grain yield QTLs. These regions are on chromosome 7(5HL) (between<br />
markers ABC717 and ABC718) and on the telomeric region of the short arm of chromosome<br />
2(2HS) (ABG058).<br />
Since RFLP methodology needs large amounts of DNA and entails a complex procedure with<br />
radioactively labeled probes and Southern blotting, which requires several days to produce<br />
results, we converted RFLP targeting those putative QTL regions to PCR-markers: cleaved<br />
amplified polymorphic sites (CAPS) and SNPs. The conversion of RFLP clones to PCR-based<br />
markers rendered a much simpler technique that facilitated the screening of large numbers of<br />
genotypes at the seedling stage, since it requires a small amount of DNA.<br />
Materials and Methods<br />
Genomic DNA extraction was modified from Edwards et al. (1991); the modification added an<br />
extra-step of chloroform-isoamyl alcohol (24:1) extraction. CAPS were developed from RFLP<br />
clones. Primers were designed from the cloned sequences and used to amplify genomic DNA<br />
from the parental cultivars Harrington and Baronesse. If no fragment length polymorphism was<br />
observed, the fragments from both parents were sequenced to discover SNPs. These SNPs were<br />
analyzed to identify restriction enzymes that could be used to develop CAPS, or to design<br />
cultivar-specific primers. CAPS markers were also developed for gene Dhn1 (Choi et al., 2000)<br />
and a candidate gene for seed dormancy and/or pre-harvest sprouting, GA20-oxidase (Li et al.,<br />
2004). Reactions for SNPs were set under stringent conditions (annealing temperatures ~ Tm)<br />
and short cycles (annealing ≤ 15 s). Extensions were 1min or 2min depending on product size.<br />
21
Primer3 software (Rozen and Skaletsky, 2000) assisted the process of primer design. PCR<br />
products were visualized on 1% or 2%, depending on the fragment size, agarose gel under UV<br />
light. PCR products were purified with Exonuclease I (Exo-SAP-IT, UBS, Cleveland, OH).<br />
Sequencing reactions were performed on an Applied Biosystems 3100 Genetic Analyzer (Perkin<br />
Elmer Applied Biosystem Division, Foster City, CA) with the ABI PRISM Big Dye Terminator<br />
v3.1 cycle sequencing kit. Products were confirmed by sequencing in both directions. Analysis<br />
of sequences to find restriction sites and/or SNPs was done with the tools provided by the San<br />
Diego Super Computer Center (SDSC, http://workbench.sdsc.edu).<br />
Results and Discussion<br />
Details of developed PCR markers are listed in Tables 1 (CAPS) and 2 (SNPs). CAPS marker<br />
MWG699 yielded small fragments difficult to visualize after digestion with enzyme Taq1, even<br />
when 3% UltraPure Agarose-1000 was used to run PCR products. To avoid this problem,<br />
cultivar-specific markers were developed taking advantage of SNPs between the parental<br />
genotypes. The two SNP sites are different than the TaqI restriction site, for this reason they are<br />
considered two different markers for locus MWG699.<br />
Table 1. Summary of developed CAPS markers, their location, primers and restriction enzymes<br />
Chr. Locus Bin Forward primer<br />
Reverse primer<br />
3(3H) MWG571A 009 5’-GTATCGTCAACACGGCAGCGT-3’<br />
5’-TACCTGTCAGAAGTGCAGTACC-3’<br />
3(3H) MWG961 012 5’-TCAACTCCAGCCTTCACACACAAC-3’<br />
5’-AAGACGAAGGAGACGTTGTTCATG-3’<br />
2(2H) MWG699 010 5’-ACCCACTGGGTTTGATACTACAAAG-3’<br />
5’-GTGATGTTATTGGTGACTTGAACTC-3’<br />
1(7H)<br />
7(5H)<br />
MWG851A<br />
Dhn1**<br />
001<br />
011<br />
5’-CAAGAACTCCATTCCAATGTACCTG-3’<br />
5’-TACTTCCAGATCCATGACAAGCTAC-3’<br />
5’-TCACTGTTCGTACTTCGTAGCACC-3’<br />
5’-TCCGCAGTTGCTCCTCCAAT-3’<br />
7(5H) ABC309 015 5’-CAGAGATACCACTGGGATTCTAAAC-3’<br />
5’-CGAAAACCCTAGGAGAGCTAATC-3’<br />
7(5H) MWG2249 015 5’-AGCCATGCCGGTCTTGTCAAGAAAG-3’<br />
5’-ATGCATCTGATCCCTGGAGAAGAAC-3’<br />
7(5H) GA20-oxidase 015 5’-GTCCATCATGCGCCTCAACTACTAC-3’<br />
5’-TAGCAAATCTTGCCATCCATCCATG-3’<br />
Enzyme<br />
BamHI<br />
BsgI<br />
TaqI*<br />
HaeIII, MspI<br />
TaqI, HpyCH4 IV<br />
HinfI<br />
* Restriction enzyme (TaqI) previously reported by Tanno et al. (2002) in a different population<br />
** Primers from Choi et al. (TAG 101:350-354)<br />
22<br />
AluI<br />
AvaI
Table 2. Cultivar-specific PCR primers developed exploiting nucleotide polymorphisms<br />
between Harrington (H) and Baronesse (B) cultivars<br />
Chr. Locus Bin Forward primer<br />
Reverse primer<br />
2(2H) MWG699-H* 010 5’-ATGGCTATCGCTTGACCAA-3’<br />
5’-GTGATGTTATTGGTGACTTGAACTC-3’<br />
2(2H) MWG699-B* 010 5’-ATGGCTATCGTTTGACCAG-3’<br />
5’-GTGATGTTATTGGTGACTTGAACTC-3’<br />
2(2H) ABG058-H 001 5’-TCTAGGCTTGCATTTGTCTACAAAG-3’<br />
5’-ATGCTGCTTCGCTGTCTACAATAAC-3’<br />
2(2H) ABG058-B 001 5’-CAATAATCTCTCTTGCCATCATGCC-3’<br />
5’-ATGCTGCTTCGCTGTCTACAATAAC-3’<br />
7(5H) ABC717-H* 009 5’-AACCAAGGCTACCAAGGTAATCCTG-3’<br />
5’-CTCGTACTAACTTCCTACATGGCAA-3’<br />
7(5H) ABC717-B* 009 5’-AACCAAGGCTACCAAGGTAATCCTG-3’<br />
5’-CTCGTACTAACTTCCTACATGGCAC-3’<br />
*Note: These primers require stringent conditions when running PCR<br />
References<br />
Choi, D.W., M.C. Koag and T.J. Close. 2000. Map locations of barley Dhn genes determined by<br />
gene-specific PCR. Theor. Appl. Genet. 101:350-354.<br />
Edwards K., C. Johnstone and C. Thompson. 1991. A simple and rapid method for the<br />
preparation of plant genomic DNA for PCR analysis. Nucleic Acids Res. 19(6):1349.<br />
Li, C., P. Ni, M. Francki, A. Hunter, Y. Zhang, D. Schibeci, H. Li, A. Tarr, J. Wang, M. Cakir, J.<br />
Yu, M. Bellgard, R. Lance and R. Appels. 2004. Genes controlling seed dormancy and<br />
pre-harvest sprouting in a rice-wheat-barley comparison. Funct. Integr. Genomics 4:84-<br />
93.<br />
Schmierer, D.A., N. Kandemir, D.A. Kudrna, B.L. Jones, S.E. Ullrich and A. Kleinhofs. 2004.<br />
Molecular marker-assisted selection for enhanced yield in malting barley. Mol. Breed.<br />
14:463-473.<br />
Rozen, S. and H.J. Skaletsky. 2000. Primer3 on the SSS for general users and for biologist<br />
programmers. In S. Krawetz and S. Misener (ed.) Bioinformatics methods and protocols:<br />
Methods in molecular biology. Humana Press, Totowa, NJ, p. 365-386. Available at<br />
http://www.bioinformatics.nl/cgi-bin/primer3plus/primer3plusAbout.cgi (verified April<br />
2007).<br />
Tanno, K., S. Taketa, K. Takeda and T. Komatsuda. 2002. A DNA marker closely linked to the<br />
vrs1 locus (row-type gene) indicates multiple origins of six-rowed cultivated barley<br />
(Hordeum vulgare L.). Theor. Appl. Genet. 104:54-60.<br />
23
Barley Genetics Newsletter 37: 24-28 (2007)<br />
Genetic Architecture for Yield and Quality Component Traits Over Two<br />
Environments in Barley (Hordeum vulgare L.)<br />
Ambresh Kumar Verma, S.R. Vishwakarma and P.K. Singh<br />
<strong>Department</strong> of Genetics and Plant Breeding<br />
Narendra Deva University of Agriculture and Technology,<br />
Kumarganj, Faizabad, (UP) 224 229, INDIA.<br />
e-mail: ambreshv@gmail.com<br />
Abstract<br />
Triple test-cross analysis involving three testers K-560, Narendra Jau-3 and their hybrid K-560 x<br />
Narendra Jau-3, were cross to 15 strains/varieties of barley to estimate the epistatic, additive and<br />
dominance components of genetic variance. Modified triple test-cross was done for eleven<br />
metric traits in two [normal fertile soil (E1) and saline sodic soil (E2)] environments. Epistasis<br />
was evident for all the characters under study in both environments except plant height in E2,<br />
protein content in E1 and lysine content in both conditions. The i type epistasis was significant<br />
for seed yield per plant in E1 only, while j and l type epistasis were significant for most of the<br />
traits. Additive (D) component was important in all cases in both conditions except number of<br />
effective tillers per plant in E1 and lysine content in E1 & E2 where as non fixable dominance (H)<br />
component were significant for all the characters in both conditions except number of effective<br />
tillers per plant in E1, length of main spike in E2 and lysine content in both environments. The<br />
most of the traits showed partial dominance and directional element, F was non- significant for<br />
all the traits, suggesting ambidirectional nature of dominance.<br />
Key words: barley, modified triple test-cross, genetic variation, protein content, seed yield<br />
Introduction<br />
To enable present barley varieties acceptable in the international market, there is need to develop<br />
better quality genotypes suitable for making product with consumer acceptability. Thus there is<br />
an urgent need to improve grain quality as well as develop better quality genotypes for suitable<br />
processing industry. The consumer acceptability of genotype is affected by chemical constituent<br />
of the grains. It is therefore, desirable to access basic physiochemical characteristics of the grains<br />
so that these can be combined with high yield. The information about nature and magnitude of<br />
genetic components of variance for yield and quality characters is essential for planning an<br />
efficient breeding programme in any crop. The modified triple test-cross analysis of (Ketata et<br />
al., 1976) provide efficient detection and estimation of epistatic variance along with unbiased<br />
estimates of additive and dominance component of genetic variance in determining the<br />
inheritance of eleven traits in barley using modified triple test-cross (TTC) analysis.<br />
Materials and Methods<br />
Two barley pure lines, viz. testers K-560, Narendra Jau-3 and their hybrid (K-560 x Narendra<br />
Jau-3) were cross to 15 lines (Kedar, RD-2552, Narendra Jau-1, Narendra Jau-2, Narendra Jau-4,<br />
RD-2035, BL-2, BH-512, Ratna, Jagrati, DL-88, Azad, K-603, NDB-1173 and RD-2624) of<br />
barley to develop a set of 45 crosses. The experimental materials consisting of 3 testers, 15 lines,<br />
24
30 single crosses and 15 three-way crosses were evaluated in randomized block design with three<br />
replications during rabi 2004-05 in two viz. normal fertile soil (E1) and saline sodic (E2)<br />
condition at Genetics and Plant Breeding Research Farm of Narendra Deva University of<br />
Agriculture & Technology, Kumarganj , Faizabad, U.P. India. Each entry was shown in a 3 m<br />
long single row plant with 10 cm spacing within and 25 cm between rows. Observations were<br />
recorded on five randomly selected competitive plants for eleven quantitative traits (Table 1).<br />
The pelshenke vale, protein content and lysine content were estimated by the method of<br />
(Pelshenke, 1933; Lowery’s, 1951; and Felker et al.; 1978) respectively. Character means were<br />
used for modified triple test- cross analysis (Ketata et al., 1976).<br />
Results and Discussion<br />
The triple test-cross (TTC) analysis revealed that significant epistasis was present for all the<br />
characters in both environments except plant height in E2 , protein content in E1 and lysine<br />
content in both conditions (Table 1). The partitioning of epistasis in to i and j and l types showed<br />
that additive x additive (i) interaction was significant for seed yield per plant in normal fertile<br />
soil condition only. The j and l type epistasis was significant for all the characters in both the<br />
environments except days to maturity and protein content in E1 while lysine content in both<br />
environments. Existence of significant epistasis in inheritance of seed yield and some other yield<br />
components in barley was reported by others also (Gorshkova and Gorodov, 1981). Greater<br />
importance of j and l type epistasis than i component was reported earlier by (Singh, et al., 1984;<br />
Tripati and Singh, 1983; Verma and Yunus, 1986.). On the contrary, (Nanda, et al., 1982)<br />
reported i type epistasis to be more important than j and l type epistasis, and (Singh, 1980)<br />
reported equal importance of these two sub components. The estimates of the components of<br />
genetic variance, additive (D), dominance (H) and F components and the degree of dominance<br />
(H/D 0.5 ) are given in Table 2. The additive (D) component was important in all cases in both<br />
conditions except number of effective tillers per plant in E1 and lysine content in E1 & E2 where<br />
as non fixable dominance (H) component was significant for all the characters in both condition<br />
except number of effective tillers per plant in E1, length of main spike in E2 and lysine content in<br />
both environments. The estimates of D were higher than H for most of the traits, which<br />
suggested partial dominance. The directional element of dominance, F was non-significant for all<br />
the characters in both environments indicated presence of ambidirectional dominance, and alleles<br />
with increasing and decreasing effects appear to be dominant and recessive to the same extent.<br />
The significant of additive component for seed yield and quality component traits, except<br />
number of effective tillers per plant and lysine content indicates that substantial improvement in<br />
yield status can still be achieved by following conventional breeding procedure in barley. The<br />
significant contribution of additive x additive type epistasis for seed yield suggested that this<br />
component should not be ignored while predicting the recombinants extractable from segregating<br />
generations. Further results provided evidence of j and l type epistasis for most of the traits<br />
studied. However, in autogamus crops like barley, where commercial exploitation of hybrid has<br />
started, this type of epistasis is of more use.<br />
References<br />
Felker, C.; Libanauskas, C.K.; Wainer, G. 1978. Estimation of lysine in foods. Crops Sci.<br />
18:480-490.<br />
25
Gorshkova, V.A.; Gorodova, V.T. 1981. Use of top crosses in studying breeding material of<br />
spring barley. Biological Lelektsiyai Semeno Vod Zorm Culture 47:54.<br />
Ketata, H.; Smith, E.L.; Edwards, L.H.; McNew, R.W. 1976. Detection of epistatic, additive and<br />
dominance variance in winter wheat (Triticum aestivum L.). Crop Sci. 16:1- 4.<br />
Lowry’s, O.H.; Roserbrough, N.J.; Farr, A.L.; Randall, R.J. 1951. Protein measurement with the<br />
folin phenol reagent. J. Biol. Chem. 193:265-278.<br />
Nanda, G.S.; Singh, P.; Gill, K.S. 1982. Epistasis, additive and dominance variation in triple test<br />
cross of bread wheat. Theor. Appl. Genet. 62:49-52.<br />
Pelshenke, P.A. 1933. A short method for determination of gluten quality of wheat. Cereal<br />
Chem. 10-90.<br />
Singh, G.; Nanda, G.S.; Gill, K.S. 1984. Inheritance of yield and its components in five crosses<br />
of spring wheat. Indian J. agric. Sci. 54:943-949.<br />
Singh, S. 1980. Detection of component of genetic variance and genotype x environment<br />
interaction in spring wheat. J. agric. Sci. Camb. 95:67-72.<br />
Tripathi, I.D.; Singh, M. 1983. Triple test-cross analysis in three barley populations under saline<br />
alkaline soil conditions. J. agric. Sci. Camb. 101:117-121.<br />
Verma, S.S.; Yunus,M. 1986. Role of epistasis in the analysis of genetic component of variance<br />
in bread wheat. Indian J. agric. Sci. 56:687-689.<br />
26
Table 1. ANOVA (mean square) of triple test-cross to test epistasis for eleven characters in barley under<br />
normal fertile soil (E1) and saline sodic soil condition (E2)<br />
Sources of<br />
variation<br />
Environ-<br />
ments<br />
d.f. Days to<br />
maturity<br />
Plant<br />
height<br />
(cm)<br />
No. of<br />
effective<br />
tillers<br />
/plant<br />
Length<br />
of main<br />
spike<br />
(cm)<br />
Grains<br />
per<br />
spike<br />
Seed<br />
yield/<br />
plant<br />
(g)<br />
1000<br />
seed<br />
weight<br />
(g)<br />
Pelshenke<br />
value<br />
(minutes)<br />
‘i’ Type epistasis<br />
‘j+I’ type pistasis<br />
E1<br />
E2<br />
E1<br />
1<br />
14<br />
7.20<br />
369.80<br />
37.53<br />
3.99<br />
220.81<br />
34.78*<br />
0.17<br />
8.28<br />
8.13**<br />
10.86<br />
3.64<br />
1.64**<br />
428.76<br />
366.35<br />
206.09**<br />
22.36*<br />
300.13<br />
81.08**<br />
9.16<br />
0.793<br />
25.28**<br />
2121.80<br />
1355.77<br />
315.23**<br />
0.61<br />
2.11<br />
4.40<br />
0.75<br />
1.49<br />
0.56<br />
5.83<br />
1.12<br />
205.13**<br />
E2<br />
*<br />
74.66** 43.17*<br />
*<br />
11.57** 1.21** 307.31** 169.98** 23.90* 330.28** 7.46** 0.97 131.66**<br />
Total epistasis E1<br />
E2<br />
15 35.51*<br />
94.33**<br />
32.73*<br />
*<br />
55.01<br />
7.60**<br />
11.35**<br />
2.25**<br />
1.37**<br />
220.93**<br />
311.24**<br />
77.16**<br />
178.65**<br />
24.20**<br />
22.36*<br />
435.66**<br />
398.64**<br />
4.15<br />
7.10**<br />
0.57<br />
0.90<br />
191.84**<br />
122.96**<br />
‘i’ type epistasis<br />
x blocks<br />
E1<br />
E2<br />
2 1.80<br />
92.45**<br />
1.00<br />
55.20*<br />
4.34**<br />
2.07**<br />
2.71**<br />
0.91**<br />
107.19**<br />
91.59*<br />
5.59<br />
75.03**<br />
2.29**<br />
0.198<br />
530.45**<br />
338.94**<br />
0.15<br />
0.527<br />
0.19<br />
3.73<br />
1.46<br />
0.28<br />
‘j+I’ type<br />
E1 28 15.64 7.92 0.14 0.43 24.12 3.34 0.18 32.68 9.62 1.28 2.03<br />
epistasis x blocks E2<br />
2.53 14.17 0.32 0.15 23.78 0.723 12.15 11.18 0.71 1.80 0.31<br />
Total epistasis x E1 30 14.72 7.46 0.14 0.55 29.66 3.49 0.32 36.86 10.00 2.42 1.99<br />
blocks<br />
E2<br />
8.53 16.91 0.43 0.20 28.30 5.68 11.36 33.03 0.69 2.68 0.49<br />
*, ** Significant at 5% and 1% probability levels, respectively.<br />
27<br />
Protein<br />
content<br />
(%)<br />
Lysine<br />
content<br />
(%)<br />
Husk<br />
content<br />
(%)
Table 2. Estimates of additive (D) and dominance (H) components of variance, parameter F, and degree of dominance (H/D) 0.5<br />
in barley under normal fertile soil (E1) and saline sodic soil (E2 ) condition<br />
Sources of<br />
variation<br />
Environ-<br />
ments<br />
D E1<br />
E2<br />
H E1<br />
E2<br />
(H/D) 0.5<br />
E1<br />
E2<br />
F E1<br />
E2<br />
Days to<br />
maturity<br />
Plant<br />
height<br />
(cm)<br />
Number<br />
o f<br />
effective<br />
tillers /<br />
plant<br />
Length<br />
of main<br />
spike<br />
(cm)<br />
Grains per<br />
spike<br />
Seed<br />
yield/<br />
plant<br />
(g)<br />
1000<br />
seed<br />
weight<br />
(g)<br />
Pelshenke<br />
value<br />
(minutes)<br />
Protein<br />
content<br />
(%)<br />
Lysine<br />
content<br />
(%)<br />
Husk<br />
content<br />
(%)<br />
25.51** 429.12** 4.67 4.70** 520.15** 67.72** 17.21** 250.46** 16.77** 1.63 125.83**<br />
40.72** 150.58** 2.64** 2.60** 239.90** 50.43** 22.74** 190.35** 14.75** 1.90 78.26**<br />
20.09** 20.06** 3.11 1.46** 167.30** 33.78** 12.57** 374.09** 4.89** 0.86 77.24**<br />
37.45** 45.28** 4.41** 0.76 224.90** 69.57** 10.56** 237.02** 8.30** 0.65 57.05**<br />
0.60 0.22 0.82 0.56 0.57 0.71 0.85 1.22 0.54 0.74 0.83<br />
0.96 0.55 1.29 0.54 0.81 1.17 0.68 1.12 0.75 0.58 0.85<br />
0.28 0.18 0.31 0.03 0.02 0.04 0.26 -0.08 -0.06 -0.25 0.20<br />
0.10 -0.02 0.15 0.12 0.11 -0.10 0.20 -0.27 0.01 -0.05 0.19<br />
*, ** Significant at 5% and 1% probability levels, respectively.<br />
28
Barley Genetics Newsletter 37: 29–33 (2007)<br />
Line x Tester Analysis in Barley (Hordeum vulgare L.) Across Environments<br />
Ambresh Kumar Verma, S.R. Vishwakarma and P.K. Singh<br />
<strong>Department</strong> of Genetics and Plant Breeding<br />
Narendra Deva University of Agriculture and Technology,<br />
Kumarganj, Faizabad, (UP) 224 229 India.<br />
e-mail: ambreshv@gmail.com<br />
Abstract<br />
A combining ability effects study was conducted through line x tester analysis under normal<br />
fertile and saline sodic soil environments. The results indicated the predominance of nonadditive<br />
gene action for all the traits. The line Kedar and tester K-560 in normal fertile soil and<br />
tester Lakhan in saline sodic soil while RD-2552, Narendra Jau-4 and NDB-1173 under both<br />
environments proved good general combiners for seed yield and quality components characters.<br />
The crosses Kedar x K-560, K-603 x K-560, DL-88 x Lakhan and RD-2035 x K-560 in normal<br />
and RD-2552 x Narendra Jau-3, Narendra Jau-1 x K-560, Narendra Jau-4 x Lakhan, NDB-1173<br />
x K-560, and RD-2624 x K-560, in saline sodic soil while RD-2552 x Lakhan, RD-2035 x<br />
Narendra Jau-3, BL-2 x Lakhan and Jagrati x K-560 in both environment exhibited highest sca<br />
effects for seed yield and other quality traits, showing their desirability to offer transgressive<br />
segregants in succeeding generations.<br />
Key words: barley, combining ability, gene action, protein content, lysine content.<br />
Introduction<br />
Research on barley (Hordeum vulgare L.) bears special significance due to its great elasticity of<br />
adaptation under various stresses and lot of potential both for domestic and industrial uses.<br />
Barley also has been very important winter cereal crop in India , because of its versatile nature,<br />
lower cost of cultivation, superior nutritional qualities and many other uses. The major uses of<br />
barley grains, however are in the production of malt, which is used to make beer, beverage<br />
industrial alcohol, whisky, malt syrups, malted milk and vinegar. The spent malt after brewing is<br />
used as feed. Combining ability analysis helps in identification of desirable parents and crosses<br />
for their further exploitation in breeding programme. Therefore, the present study was<br />
undertaken to estimate combining ability effects for yield and quality components characters and<br />
also to identify suitable parents and crosses in barley under normal fertile and saline soil<br />
environments.<br />
Materials and Methods<br />
The material consisted of 15 lines, namely RD-2552, Narendra Jau-1, Narendra Jau-2, Narendra<br />
Jau-4, RD-2035, BL-2, BH-512, Ratna, Kedar, Jagrati, DL-88, Azad, K-603, NDB-1173 and<br />
RD-2624 with 3 testers viz., Narendra Jau-3, K-560 and Lakhan crosses were attempted in line x<br />
tester fashion. The resulting 45 F1s along with lines and testers were planted in a randomized<br />
block design with three replicates during rabi 2004-2005 under normal fertile soil and saline<br />
sodic environments at Research Farm of Narendra Deva University of Agriculture and<br />
Technology, Kumarganj, Faizabad. Each treatment (genotype) was shown in 3 m length having<br />
row to row and plant to plant distance of 25 cm and 10 cm, respectively. The observations were<br />
recorded on days to maturity, plant height (cm), number of effective tillers/plant, length of main<br />
29
spike (cm), grains per spike, seed yield/plant (g), 1000 seed weight (g), pelshenke value<br />
(minutes), protein content (%), lysine content (%) and husk content (%) on five randomly<br />
selected plants from each replication and environments. The combining ability analysis was<br />
carried out following the method proposed by (Kempthorne,1957).<br />
Results and Discussion<br />
The analysis of variance for combining ability for eleven characters showed that variances due to<br />
gca and sca were significant for the characters like days to maturity, plant height, length of main<br />
spike, grains per spike, seed yield per plant, 1000 seed weight, pelshenke value, protein content<br />
and husk content under both normal fertile and saline sodic soil conditions, suggesting thereby<br />
importance of both additive and non additive gene actions for the inheritance of these characters.<br />
The role of both additive and non-additive effects to grain yield and its component characters in<br />
barley have been reported previously (Choo et al.,1988; Bhatnagar and Sharma1995;1998).<br />
However, the component of variation due to sca was higher than gca for all the characters in all<br />
the environments indicating the predominance of non-additive gene action. Such results infers<br />
that the chosen material had high selection history. Similar results of predominance of sca<br />
variance over gca variance have also been reported by (Guo and Xu,1994; Phogat et al.1995;<br />
Madic,1996; El-Seidy,1997a & 1997b; Bouzerzour and Djakoune,1998).<br />
A perusal of the gca estimates (Table1) showed that the parents RD-2552, Narendra Jau-4, NDB-<br />
1173 in both environments while, Kedar & K-560 in E 1, and Lakhan in E2<br />
were the best<br />
combiners for seed yield and good/ medium combiner for most of the important yield and quality<br />
component characters. Further the parents Ratna, Narendra Jau-1, RD-2552, Narendra Jau-3,<br />
Narendra Jau-1 for early maturity, Lakhan, Narendra Jau-1 for dwarf plant height, Azad, RD-<br />
2624, NDB-1173 and Ratna for high protein content and Narendra Jau-1, Narendra Jau-2 and<br />
Lakhan for high lysine content were found to be good general combiners in both the<br />
environments.<br />
Significant gca values indicated the importance of additive or additive x additive gene effect as<br />
earlier reported by (Griffing,1956). In view of this, these parents offered the best possibilities for<br />
the development of improved lines of barley through hybridization programme. It is, therefore,<br />
recommended that to improve yield one should breed for superior combining ability for the<br />
component traits with an ultimate objective to improve the pace of its genetic improvement.<br />
The estimates of specific combining ability effects of top five ranking crosses for all the<br />
characters are present in Table 2. The perusal of sca effects revealed that crosses Kedar x K-560,<br />
K-603 x K-560, DL-88 x Lakhan RD-2552 x K-560 in normal and RD-2552 x Narendra Jau-3,<br />
Narendra Jau-1 x K-560, Narendra Jau-4 x Lakhan, NDB-1173 x K-560 & RD-2624 x K-560 in<br />
saline sodic and RD-2552 x Lakhan, RD-2035 x Narendra Jau-3, BL-2 x Lakhan and Jagrati x K-<br />
560 under both environments were for seed yield per plant and with other characters. The crosses<br />
RD-2035 x Narendra Jau-3 and BL-2 x Lakhan are excellent crosses for seed yield in both<br />
environments. Therefore, these crosses should be particularly exploited vigorously in future<br />
breeding programmes to obtain good segregants which would lead to buildup a population with<br />
high genetic yield potential with develop salt tolerant genotypes.<br />
30
References<br />
Bhatnagar, V. K., Sharma, S. N. 1995. Diallel analysis for combining ability for grain yield and<br />
its components in barley. Indian J. Genet. 55:228-232.<br />
Bhatnagar, V. K., Sharma, S. N. 1998. Diallel analysis for grain yield and harvest index in barley<br />
under diverse environments. Rachis. 16:22-27.<br />
Bouzerzour, H., Djakoune, A. 1998. Inheritance of grain yield and grain yield components in<br />
barley. Rachis. 16:9-16.<br />
Choo, T. M., Reinbergs, E., Jui, P. Y.1988. Comparison of F2 and F1<br />
diallel analyses in barley.<br />
Genome 30:865-869.<br />
El-Seidy, E. S. H.1997a. Inheritance of earliness and yield in some barley crosses. Ann. Agric.<br />
Sci. Moshtohor. 35:715-30.<br />
El-Seidy, E. S. H. 1997b. Inheritance of plant height, grain yield and its components in three<br />
barley crosses (Hordeum vulgare L.). Ann. Agric. Sci. Moshtohor. 35:63-76.<br />
Guo, Y. Y., Xu, S. Y. 1994. Genetic analysis of yield traits in two-rowed barley. Acta Agric.<br />
Zhejiangensis 6:156-60.<br />
Griffing, B.1956. Concept of general and specific combining ability in relation to diallel crossing<br />
system. Aust. J. Biol. Sci. 9:463-493<br />
Kempthorne, O. 1957. An Introduction to Genetical Statistics. John Wiley & Sons. Inc. New<br />
York.<br />
Madic, M. 1996. Inheritance of spike traits and grain yield in barley (H. vulgare L.) hybrids.<br />
Rev. Res. Work, Fac. Agric. Belgrade 41:53-65.<br />
Phogat, D. S., Singh,D., Dahiya, G. S., Singh, D. 1995. Genetics of yield and yield components<br />
in barley (Hordeum vulgare L.). Crop Res. Hisar. 9:363-369.<br />
31
Table 1. Estimates of general combining ability (gca) effects of parents (lines & testers) for 11 characters in<br />
barley under normal fertile soil condition (E1) & saline sodic soil condition (E2)<br />
Parents<br />
Kedar<br />
K-603<br />
DL-88<br />
RD-2552<br />
Azad<br />
Narendra Jau -1<br />
Narendra Jau -2<br />
Narendra Jau -4<br />
NDB-1173<br />
RD-2035<br />
RD-2624<br />
BL-2<br />
Jagrati<br />
BH-512<br />
Ratna<br />
SE (gi) lines<br />
SE(gi-gj) lines<br />
K-560<br />
Narendra Jau-3<br />
Lakhan<br />
SE (gi) testers<br />
Environments<br />
Days to<br />
maturity<br />
Plant<br />
height<br />
(cm)<br />
No. of<br />
effective<br />
tillers<br />
/plant<br />
Length<br />
of main<br />
spike<br />
(cm)<br />
* Significant at 5% probability level, ** Significant at 1% probability level<br />
32<br />
Grains<br />
/spike<br />
Seed<br />
yield/<br />
plant<br />
(g)<br />
1000<br />
seed<br />
weight<br />
(g)<br />
Pelshenke<br />
value<br />
(minute)<br />
Protein<br />
content<br />
(%)<br />
Lysine<br />
content<br />
(%)<br />
Husk<br />
content<br />
(%)<br />
E1 0.63** 3.45** 0.40* -0.18* -0.71 1.02* 1.10* 6.58** 0.39** -0.12** 2.30**<br />
E2 1.23** 5.76** 0.26 -0.13 -0.72 0.73 0.99** 5.25** 0.27** -0.26** 1.35**<br />
E1 -0.12 -0.62 0.20 -0.26** -3.08** -0.45 -1.04* -1.92* 0.26** -0.18** 0.13<br />
E2 -1.27** 3.62** -0.27 -0.60** -5.51** -1.58** -1.67** -1.67 0.26** -0.13** 0.44*<br />
E1 0.63** 3.85** -1.02** -0.63** -4.14** -3.04** -0.74 0.33 -0.21** 0.02 -1.67**<br />
E2 -0.19 3.40** -0.34 -0.46** -1.82* -0.77 -0.86** 0.58 -0.32** 0.07** -1.23**<br />
E1 -0.62** -4.96** 0.14 0.04 6.76** 1.19* -1.11* -9.92** -0.68** 0.25** -1.88**<br />
E2 -0.69** -2.65** 0.37* 0.11 4.37** 1.83** -0.83** -7.58** -0.52** 0.33** -1.15**<br />
E1 2.21** -6.84** -0.11 0.17* 4.86** 0.53 0.11 7.16** 2.06** -0.59** 1.24**<br />
E2 2.56** 0.42 0.08 0.18 5.20** 0.57 0.08 6.17** 1.94** -0.53** 0.93**<br />
E1 -0.62** -2.76** 0.34* 0.32** -0.54 0.54 0.55 -0.34 -1.38** 0.56** -0.23<br />
E2 -0.52* -4.57** 0.29 0.19 1.50 0.71 0.67** 0.42 -0.49** 0.59** -0.60**<br />
E1 -1.54** 3.59** 0.08 0.80** 1.23 0.48 0.28 -3.42** -1.68** 0.51** 1.50**<br />
E2 0.06 -4.55** 0.33 0.49** 0.34 0.46 0.14 -3.25** -1.77** 0.52** 1.44**<br />
E1 0.29 -2.66** 1.31** 0.65** 4.73** 3.54** -0.32 -6.76** -1.01** 0.34** 0.35<br />
E2 1.14** 2.68** 0.50** 0.50** 1.78 0.90* -0.29 -6.83** -1.05** 0.10** 0.69**<br />
E1 -2.71** -1.98** -0.11 0.71** 6.99** 2.14** 1.75** -1.76* 0.93** -0.28** -1.99**<br />
E2 -2.11** -4.08** 0.00 0.52** 6.08** 1.24** 2.09** -1.67 0.99** -0.32** -1.98**<br />
E1 0.13 0.22 -0.21 -0.23* 4.15** 0.66 -0.99* -2.34** -0.35** 0.11** 1.05**<br />
E2 -0.36 -1.69** 0.09 -0.15 3.43** 0.02 -0.94** -2.08* -0.46** 0.12** 0.98**<br />
E1 0.71** -3.44** -0.45** -0.38** -9.32** -3.52** -1.60** 0.08 1.63** -0.40** -0.63**<br />
E2 -0.19 -0.77 -0.60** -0.05 -3.70** -2.36** -1.47** 0.33 1.54** -0.35** -0.40<br />
E1 1.54** 1.89** 0.43* -0.13 -2.49* 0.63 0.29 3.24** -0.09* -0.03 -1.33**<br />
E2 0.23 -2.29** 0.36* -0.08 -3.35** 0.66 0.03 3.33** -0.18** -0.04** -1.35**<br />
E1 1.13** 4.50** -0.26 -0.32** -4.47** -1.22** 1.20** 1.24 -0.30** 0.15** 0.93**<br />
E2 1.39** -2.44** -0.45* -0.21 -3.45** -0.98* 1.12** 1.58 -0.47** 0.18** 0.69**<br />
E1 -0.12 7.09** -0.31 -0.36** 1.19 -0.74 0.03 0.99 0.02 -0.05 2.49**<br />
E2 0.31 4.29** -0.18 -0.35* 0.12 -0.41 0.12 -0.33 -0.07** -0.03* 2.14**<br />
E1 -1.54** -1.34* -0.44** -0.21* -5.17** -1.75** 0.49 6.83** 0.41** -0.29** -2.26**<br />
E2 -1.61** 2.87** -0.45* 0.04 -4.27** -1.04* 0.83** 5.75** 0.33** -0.25** -1.94**<br />
E1 0.22 0.64 0.17 0.09 0.97 0.45 0.45 0.75 0.04 0.03 0.21<br />
E2 0.22 0.52 0.17 0.13 0.93 0.45 0.21 0.88 0.03 0.01 0.20<br />
E1 0.31 0.90 0.23 0.13 1.37 0.64 0.64 1.05 0.06 0.04 0.30<br />
E2 0.31 0.74 0.25 0.19 1.31 0.64 0.30 1.24 0.04 0.02 0.29<br />
E1 0.88** 1.83** 0.27** 0.18** 0.68 0.76** 0.19 -5.77** 0.18** -0.04** -1.20**<br />
E2 1.31** -0.87** 0.07 0.05 0.41 0.41 -0.22* -5.17** 0.14** -0.06** -0.87**<br />
E1 -0.81** 2.69** 0.31** 0.09 -0.18 0.21 0.24 1.94** 0.07** -0.06** 0.17<br />
E2 -0.36** 0.59* -0.04 0.08 0.48 -0.02 0.17 1.97** 0.34** -0.11** -0.12<br />
E1 0.10 -6.63** -0.90** -0.16** 0.79 -1.11** -0.42 2.30** -0.44** 0.22** 1.37**<br />
E2 0.02 -0.70* 0.17 -0.05 0.09 0.78** 0.15 2.06** -0.60** 0.26** 1.40**<br />
E1 0.11 0.33 0.09 0.05 0.5 0.23 0.23 0.39 0.02 0.02 0.11<br />
E2 0.11 0.27 0.09 0.07 0.48 0.23 0.11 0.45 0.01 0.01 0.10<br />
E1 0.16 0.46 0.12 0.06 0.71 0.33 0.33 0.54 0.03 0.02 0.15<br />
SE(gi-gj)<br />
testers E2 0.16 0.38 0.13 0.10 0.68 0.33 0.15 0.64 0.02 0.01 0.15
Table 2. Promising crosses for seed yield and quality component in barley under normal<br />
fertile soil and saline sodic soil condition.<br />
Characters Normal fertile soil environment Saline sodic soil environment<br />
Days to maturity Kedar x Lakhan, RD-2552 x K-560, RD-<br />
2552 x Narendra Jau-3, Narendra Jau-1 x<br />
K-560, RD-2624 x Lakhan<br />
Plant height (cm) Jagrati x Lakhan, BL-2 x Lakhan,<br />
Narendra Jau-2 x Lakhan, RD-2035 x<br />
Lakhan, Kedar x Narendra Jau-3<br />
K-603 x K-560, RD-2624 x Lakhan,<br />
Narendra Jau-1 x K-560, Kedar x Lakhan,<br />
Jagrati x Narendra Jau-3<br />
Ratna x Narendra Jau-4, BL-2 x Lakhan,<br />
DL-88 x K-560, RD-2035 x Lakhan, RD-<br />
2624 x Lakhan<br />
Number of effective<br />
tillers/plant<br />
RD-2035 x K-560, BL-2 x Narendra Jau-3,<br />
Jagrati x K-560, BL-2 x Lakhan<br />
Narendra Jau-1 x K-560, Narendra Jau-4 x<br />
Lakhan, NDB-1173 x K-560, RD-2035 x<br />
Narendra Jau-3<br />
Length of main spike<br />
(cm)<br />
K-603 x K560, K-603 x Narendra Jau-3,<br />
DL-88 x Lakhan, RD-2035 x K-560,Jagrati<br />
x K-560<br />
Grains/spike BL-2 x Lakhan, Kedar x K-560, K-603 x<br />
K-560, DL-88 x Lakhan, RD-2035 x<br />
Lakhan<br />
Seed yield per plant<br />
(g)<br />
BL-2 x Lakhan, RD-2035 x Narendra Jau-<br />
3, Jagrati x K-560, RD-2552 x Lakhan,<br />
Kedar x K-560<br />
DL-88 x Lakhan, NDB-1173 x Lakhan,<br />
RD-2035 x Lakhan<br />
BL-2 x Lakhan, RD-2624 x K-560, DL-88<br />
x Lakhan, Narendra Jau-4 x Narendra Jau-<br />
3, RD-2035 x Narendra Jau-3<br />
Narendra Jau-1 x K-560, Narendra Jau-4 x<br />
Lakhan, NDB-1173 x K560, RD-2053 x<br />
Narendra Jau-3, BL-2 x Lakhan<br />
1000-seed BL-2 x Lakhan, Jagrati x K-560, BL-2 x Lakhan, Narendra Jau-4 x Lakhan<br />
weight (g) RD-2035 x Narendra Jau-3, Narendra Jau-<br />
4 x Lakhan,<br />
Pelshenke Narendra Jau- 1 x K-560, RD-2552 x K-<br />
value (minute) 560, RD-2624 x Lakhan, Narendra Jau- 4 x<br />
K-560, BL-2 x Narendra Jau-3<br />
Protein RD-2552 x K-560, Narendra Jau-4 x K-<br />
content (%)<br />
560, Kedar x Narendra Jau-3, BH-512 x<br />
Lakhan, Ratna x K-560<br />
Lysine DL-88 x K-560, Kedar x K-560, K-603 x<br />
content (%)<br />
K-560,<br />
RD-2035 x Narendra Jau-3, RD-2552 x<br />
Lakhan<br />
Husk Narendra Jau-2 x Narendra Jau-3, Kedar x<br />
content (%)<br />
Narendra Jau-3, K-603 x K-560, Azad x<br />
K-560, DL-88 x K-560<br />
33<br />
RD-2552 x Narendra Jau-3, RD-2035 x<br />
Narendra Jau-3, Jagrati x Narendra Jau-3<br />
RD-2552 x K-560, Narendra Jau- 4 x<br />
Lakhan, RD-2552 x Narendra Jau-3, RD-<br />
2035 x Narendra Jau-3, Jagrati x Narendra<br />
Jau-3<br />
Narendra Jau-4 x K-560, RD-2552 x K-<br />
560, K-603 x Narendra Jau-3, NDB-1173<br />
x K-560, Kedar x Narendra Jau-3<br />
K-603 x K-560, DL-88 x K-560, Narendra<br />
Jau-4 x Lakhan, Kedar x Lakhan, Narendra<br />
Jau-4 x Narendra Jau-3<br />
Narendra Jau-2 x Narendra Jau-3, Kedar x<br />
Narendra Jau-3, Narendra Jau-4 x<br />
Narendra Jau-3, K-603 x K-560,<br />
DL-88 x K-560
Barley Genetics Newsletter 37: 34–36 (2007)<br />
Four new barley mutants<br />
Sandra AI Wright 1 , Manochehr Azarang 2 , Anders B Falk 3<br />
1 Dipartimento di Scienze Animali, Vegetali e dell’Ambiente, Facoltà di Agraria, Università del<br />
Molise, Campobasso, Italy<br />
2 Maselaboratorierna, Uppsala, Sweden 3 <strong>Department</strong> of Plant Biology and Forest Genetics,<br />
University of Agricultural Sciences, Uppsala, Sweden<br />
Introduction<br />
We screened a fast-neutron mutated barley population to isolate and characterize barley lesion<br />
mimic mutants. The albino mutant frequency in this population was found to be around 2%,<br />
suggesting that a sufficiently high level of mutants could potentially be found. From a screen of<br />
about 5000 M2 spikes, we found four mutants that were characterized by the presence of necrotic<br />
or chlorotic leaf areas in the form of spots or stripes. The four mutants were designated 1661,<br />
2721, 3091 and 3550. The mutants were characterized with respect to their visual appearance. In<br />
order to relate these mutants properly to previously described barley lesion mimic mutants, we<br />
cultivated all previously described barley mutants with aberrant leaf phenotypes (Davis et al..<br />
1997) and compared the phenotypes to those of the newly obtained mutants.<br />
Materials and Methods<br />
The near-isogenic barley line Bowman(Rph3) was obtained from Dr Jerry D. Franckowiak,<br />
North Dakota, <strong>US</strong>A. It was constructed by introgression of the Rph3 resistance gene from the<br />
cultivar Estate into the cultivar Bowman, and represents the 7th backcross. Mutation was<br />
performed with fast neutrons at the International Atomic Energy Agency (IAEA), Vienna,<br />
Austria, with a dose of 5 Gy. Five thousand M2 spikes were sown and the mutant plants were<br />
screened for aberrant leaf phenotype. Selected mutants were backcrossed twice to wildtype<br />
Bowman (Rph3), and their phenotypes were analyzed using the backcrossed material. Plants<br />
were grown either in a greenhouse or in caged outdoor compartments during the summer.<br />
Experiments were done in controlled growth chambers at 22°C with 16/8 hours of light/darkness<br />
(long day conditions), or 8/16 hours of light/darkness (short day conditions). Allelism tests were<br />
done as inspections of leaf phenotypes of F1 plants resulting from crosses between relevant<br />
mutants. An AFLP-based procedure was used to screen for molecular markers linked to the<br />
mutants (Castiglioni et al. 1998).<br />
Results and Discussion<br />
Mutant 1661 displays chlorotic stripes (Figure 1). These stripes are most pronounced on the first<br />
leaf. Under long day conditions, the stripes do not appear on the later emerging leaves and the<br />
plants eventually seem to recover from the phenotype conferred by the mutation. Under short<br />
day conditions the mutation is semi-lethal since all leaves develop the characteristic chlorotic<br />
stripes and the plants fail to reach maturity and produce seeds. However, the phenotype initially<br />
proves to be more severe under long day conditions. Mutant 1661 is phenotypically similar to the<br />
previously described mutants mottled leaf 1 and mottled leaf 5, which have clearly marked white<br />
34
ands across the leaves (Davis et al. 1997). Allelism tests indicated that 1661 is not allelic to<br />
these. None of the pre-mapped Proctor-Nudinka AFLP markers were linked to mutant 1661.<br />
Mutant 2721 is characterized by chlorotic leaf spots and streaks that coalesce and eventually<br />
form large white patches on the leaves (Figure 1). The phenotype is displayed on all leaves and<br />
as the leaves mature, the white regions gradually become darker, seeming to undergo necrosis<br />
and death. Often the necrosis affects the leaf edges, resulting in wrinkled leaf edges. Under short<br />
day conditions, the leaf phenotype is delayed by several days and is reduced in severity.<br />
Eventually all leaves display the phenotype even under short day conditions. The mutant initially<br />
appears to be similar to the mottled leaf 2 (Davis et al. 1997) and mottled leaf 6 (Franckowiak<br />
2002) mutants, which display yellow bands on the leaves. However, these mutants do not display<br />
the necrosis of 2721. Allelism tests suggest that 2721 is not allelic to the mottled leaf mutants.<br />
None of the pre-mapped Proctor-Nudinka AFLP markers were linked to mutant 2721.<br />
Mutant 3091 has brown spots towards the leaf tips and leaf edges, particularly on the first leaf<br />
(Figure 1). Short days do not significantly alter the phenotype. The leaf phenotype resembles<br />
those of mutants nec4 and nec5 (Davis et al.. 1997). However, allelism tests indicate that mutant<br />
3091 is not allelic to either of these two. For the mutation in 3091, linkage was detected to the<br />
AFLP marker E37M33-6 on barley chromosome 3 (3H).<br />
Mutant 3550 has conspicuous black or brown spots on the leaves (Figure 1). The spots usually<br />
do not coalesce. They appear on all above-ground parts of the plant including the bristles. Mutant<br />
3550 is delayed in maturation and ripening, with seeds being ready for harvest about four weeks<br />
later than in the wildtype. Short day conditions lead to a slightly less pronounced phenotype.<br />
Mutant 3550 is similar to the nec1 mutant (Davis et al. 1997), but allelism could be ruled out due<br />
to different mapping positions. The mutation in 3550 was localized on chromosome 7 (5H).<br />
Linkage was detected to the AFLP markers E40M38-7, E36M36-5, E42M36-14, E42M40-2 and<br />
E41M32-5.<br />
References:<br />
Castiglioni, P., Pozzi, C., Heun, M., Terzi, V., Müller, K.J., Rodhe, W. and Salamini, F. 1998.<br />
An AFLP-based procedure for the efficient mapping of mutations and DNA probes in<br />
barley. Genetics 149:2039–2056.<br />
Davis, M.P., J.D. Franckowiak, T. Konishi, and U. Lundqvist, 1997. New and revised<br />
descriptions of barley genes. Barley Genetics Newsletter 26: 22-516.<br />
Franckowiak, J.D. 2002., BGS 629 Mottled leaf 6. Barley Genetics Newsletter 32:170.<br />
35
Bowman(Rph3)<br />
1661 2721 3091 3550<br />
Figure 1. Leaf phenotypes of the parent cultivar Bowman(Rph3) and four lesion mimic mutants; 1661, 2721, 3091,<br />
3550.<br />
36
Barley Genetics Newsletter 37: 37 – 43 (2007)<br />
The Scandinavian Barley Chlorophyll Mutation Collection<br />
Mats Hansson<br />
<strong>Department</strong> of Biochemistry, Lund University<br />
P.O. Box 124, SE-221 00 Sweden<br />
Barley (Hordeum vulgare L.) albina, striata, chlorina, tigrina, viridis and xantha mutants, which<br />
can be obtained from the <strong>Department</strong> of Biochemistry, Lund University, Sweden.<br />
The collection was previously held at the Carlsberg Laboratory, Copenhagen, Denmark, by<br />
Professor Diter von Wettstein.<br />
Please contact the cordinator for Nuclear genes affecting the chloroplast:<br />
Mats Hansson<br />
<strong>Department</strong> of Biochemistry<br />
Lund University<br />
Box 124<br />
SE-22100 Lund, Sweden<br />
Tel. +46 46 2220105<br />
Fax +46 46 2224116<br />
Email mats.hansson@biochemistry.lu.se<br />
Available mutants are marked by “×”. If missing, the mutant might be present in the collection of<br />
the Nordic Gene Bank (www.nordgen.org).<br />
Albina mutants<br />
alb 7 ×<br />
alb 10<br />
alb 11 ×<br />
alb 12<br />
alb 13 ×<br />
alb 14<br />
alb 15<br />
alb 16 ×<br />
alb 17 ×<br />
alb 18 ×<br />
alb 19 ×<br />
alb 20<br />
alb 21<br />
37<br />
alb 22 ×<br />
alb 24 ×<br />
alb 25 ×<br />
alb 26 ×<br />
alb 27 ×<br />
alb 28 ×<br />
alb 29<br />
alb 30<br />
alb 32 ×<br />
alb 33<br />
alb 34<br />
alb 35<br />
alb 36
alb 37<br />
alb 38<br />
alb 40<br />
alb 41<br />
alb 42 ×<br />
alb 43<br />
alb 44<br />
alb 45 ×<br />
alb 46 ×<br />
alb 47<br />
alb 48<br />
alb 49<br />
alb 50 ×<br />
alb 51<br />
alb 52 ×<br />
alb 53<br />
alb 54<br />
alb 55 ×<br />
alb 56<br />
alb 57<br />
alb 58<br />
alb 59 ×<br />
alb 60 ×<br />
alb 61<br />
alb 62<br />
alb 63<br />
alb 64<br />
alb 65<br />
alb 66 ×<br />
alb 67<br />
alb 68<br />
alb 69<br />
alb 70<br />
alb 71<br />
alb 72<br />
alb 73<br />
alb 74<br />
alb 75<br />
alb 76 ×<br />
alb 77<br />
alb 78 ×<br />
alb 80<br />
alb 81 ×<br />
alb 82<br />
38<br />
alb 83<br />
alb 84<br />
alb 85<br />
alb 86<br />
alb 87<br />
alb 88<br />
alb 89 ×<br />
alb 90 ×<br />
alb 91<br />
alb 92<br />
alb 94<br />
alb 95 ×<br />
alb 96<br />
alb 97<br />
alb 98<br />
alb 99<br />
alb 100<br />
alb 101<br />
alb 102<br />
alb 103<br />
alb 104<br />
alb 105<br />
alb 106<br />
alb 107<br />
alb 108<br />
alb 109<br />
alb 110<br />
alb 111 ×<br />
alb 112<br />
alb 113 ×<br />
alb 114<br />
alb 115<br />
alb 116<br />
alb 117<br />
alb 118<br />
alb 119<br />
alb 120<br />
alb 122 ×<br />
alb 123<br />
alb 124<br />
alb 125<br />
alb 126<br />
alb 127<br />
alb 128
alb 129<br />
alb 130<br />
alb 131<br />
alb 132<br />
alb 133 ×<br />
Striata mutants<br />
Arnason ×<br />
Arnason grøn<br />
striata 4 ×<br />
striata 6 ×<br />
striata 7 ×<br />
striata 8<br />
striata 11<br />
striata 12<br />
striata 13<br />
striata 15<br />
striata 16<br />
striata 17<br />
striata 19<br />
striata 21<br />
striata 22 ×<br />
striata 23<br />
striata 26<br />
striata 33 ×<br />
striata 34 ×<br />
striata 35 ×<br />
striata 36 ×<br />
striata 37<br />
striata 38<br />
striata 39<br />
striata 104 ×<br />
striata 105 ×<br />
39<br />
alb 134 ×<br />
alb 135
Chlorina mutants<br />
clo 101 ×<br />
clo 102 ×<br />
clo 103 ×<br />
clo 104 ×<br />
clo 105 ×<br />
clo 106 L ×<br />
clo 106 L (A+B) ×<br />
clo 106 line LA<br />
clo 106 line LB<br />
clo 106 line M ×<br />
clo 106 line ML ×<br />
clo 107 ×<br />
clo 108 ×<br />
clo 109 ×<br />
clo 110 ×<br />
clo 111 ×<br />
clo 112 ×<br />
clo 113 ×<br />
clo 114 ×<br />
clo 115 ×<br />
clo 116 ×<br />
clo 117 ×<br />
clo 118a ×<br />
clo 118b ×<br />
clo 119 ×<br />
clo 121 ×<br />
clo 122 ×<br />
clo 123 ×<br />
clo 124 ×<br />
clo 125 = Xan-h.Clo125 ×<br />
clo 126 ×<br />
clo 127 ×<br />
clo 130 ×<br />
clo 131a<br />
clo 131b<br />
clo133a ×<br />
clo133b ×<br />
clo 134 ×<br />
clo135 ×<br />
40<br />
clo 136 ×<br />
clo 137 ×<br />
clo 138 ×<br />
clo 140 ×<br />
clo 141 ×<br />
clo 142 ×<br />
clo 143 ×<br />
clo 144 ×<br />
clo 145 ×<br />
clo 146 ×<br />
clo 147 ×<br />
clo 148 ×<br />
clo149 ×<br />
clo 150 ×<br />
clo 151 ×<br />
clo 152 ×<br />
clo 153 ×<br />
clo 154 ×<br />
clo 155 ×<br />
clo 157 = Xan-h.Clo157 ×<br />
clo 158 ×<br />
clo 159 ×<br />
clo 160 ×<br />
clo 161 = Xan-h.Clo161 ×<br />
clo 164 ×<br />
clo 165 ×<br />
clo 166 ×<br />
clo 167 het<br />
clo170 ×<br />
clo 171 ×<br />
clo 173 ×<br />
clo 174 ×<br />
clo 175 ×<br />
clo 176 ×<br />
clo 177<br />
clo 179 ×<br />
clo 180 ×
Tigrina mutants<br />
tig 1 ×<br />
tig 3<br />
tig 6 ×<br />
tig 7 ×<br />
tig 11 ×<br />
tig 12 ×<br />
tig 13<br />
tig 14<br />
tig 15 ×<br />
tig 17<br />
tig 18<br />
tig 19 ×<br />
tig 20 ×<br />
tig 21 ×<br />
tig 22<br />
Viridis mutants<br />
vir 10 ×<br />
vir 11 ×<br />
vir 12 ×<br />
vir 13 ×<br />
vir 14 ×<br />
vir 15 ×<br />
vir 17 ×<br />
vir 18 ×<br />
vir 19 ×<br />
vir 21 ×<br />
vir 23 ×<br />
vir 24 ×<br />
vir 25 ×<br />
vir 27 ×<br />
vir 29 ×<br />
xan 75 = vir 30 ×<br />
vir 33 ×<br />
vir 34 ×<br />
vir 35 ×<br />
vir 38 ×<br />
vir 39 ×<br />
41<br />
tig 23 ×<br />
tig 24 ×<br />
tig 24 hom<br />
tig 25 ×<br />
tig 26 ×<br />
tig 27 ×<br />
tig 28 ×<br />
tig 29 hom ×<br />
tig 30 ×<br />
tig 31 ×<br />
tig 32 ×<br />
tig 33 ×<br />
tig 34 ×<br />
vir 41 ×<br />
vir 42 ×<br />
vir 43 ×<br />
vir 44 ×<br />
vir 45 ×<br />
vir 46 ×<br />
vir 47 ×<br />
vir 49 ×<br />
vir 50 ×<br />
vir 51<br />
vir 52 ×<br />
vir 55 ×<br />
xan 76 = vir 56 ×<br />
vir 59<br />
vir 60 ×<br />
vir 61 ×<br />
vir 63 ×<br />
vir 64 ×<br />
vir 65 ×<br />
vir 68 ×<br />
vir 69 ×
vir 101 ×<br />
vir 102 ×<br />
vir 103<br />
vir 104 ×<br />
vir 105<br />
vir 106<br />
vir 107<br />
vir 108<br />
vir 109<br />
vir 109<br />
vir 110<br />
vir 111<br />
vir 112<br />
vir 113 light<br />
vir 113 dark<br />
vir 114<br />
vir 115 ×<br />
vir 119<br />
vir 120 ×<br />
vir 121 ×<br />
vir 122 ×<br />
vir 123 ×<br />
vir 129<br />
vir 130 ×<br />
vir 131<br />
vir 132 ×<br />
vir 133 ×<br />
vir 134<br />
Xantha mutants<br />
xan 3 ×<br />
xan 10 ×<br />
xan 11 ×<br />
xan 12 ×<br />
xan 13 ×<br />
xan 14<br />
xan 15 ×<br />
xan 16 ×<br />
xan 17 ×<br />
xan 18 ×<br />
xan 19 ×<br />
42<br />
vir 135<br />
vir 137 ×<br />
vir 138<br />
vir 139<br />
vir 141<br />
vir 142 hom<br />
vir 142<br />
vir 143 ×<br />
vir 144<br />
vir 145 hom<br />
vir 145<br />
vir 149<br />
vir 152 ×<br />
vir 156 ×<br />
vir 157 ×<br />
vir 158<br />
vir 159 ×<br />
vir 160 ×<br />
vir 165<br />
vir 166 ×<br />
vir 167<br />
vir 168 ×<br />
vir 169<br />
vir 170 = xan 83 ×<br />
vir 519 ×<br />
xan 20 ×<br />
xan 21 ×<br />
xan 22<br />
xan 23 ×<br />
xan 24<br />
xan 25 ×<br />
xan 26 ×<br />
xan 27 ×<br />
xan 28 ×<br />
xan 29<br />
xan 30 ×
xan 31 ×<br />
xan 32<br />
xan 33<br />
xan 35 ×<br />
xan 37 ×<br />
xan 38 ×<br />
xan 39 ×<br />
xan 40 ×<br />
xan 41 ×<br />
xan 42 ×<br />
xan 43<br />
xan 44 ×<br />
xan 45 ×<br />
xan 46 ×<br />
xan 47 ×<br />
xan 48 ×<br />
xan 49 ×<br />
xan 50 ×<br />
xan 51 ×<br />
xan 52 ×<br />
xan 53 ×<br />
xan 54<br />
xan 55 ×<br />
xan 56 ×<br />
xan 57 ×<br />
xan 58 ×<br />
xan 59 ×<br />
xan 60 ×<br />
xan 62 ×<br />
xan 63 ×<br />
xan 64 ×<br />
xan 65<br />
xan 66<br />
xan 68 ×<br />
xan 69<br />
xan 70<br />
xan 71 ×<br />
xan 72 ×<br />
xan 73 ×<br />
xan 74 ×<br />
xan-q.75 = vir30<br />
xan-q.76 = vir56<br />
xan-q.77 = vir-xa1<br />
43<br />
xan-q.78 = vir-alb1<br />
xan-q.79 = xa-alb3<br />
xan-q.80 = alb-l.26<br />
xan 81 = Proto 1 ×<br />
xan 82 = Proto 2 ×<br />
xan 83 = vir 170 (Proto 3) ×<br />
xan 84 = Proto 4 ×<br />
xan 101<br />
xan 102<br />
xan 103<br />
xan 104<br />
xan 105<br />
xan 106<br />
xan 107
Barley Genetics Newsletter (2007) 37: 44-46<br />
CAPS markers targeting barley Rpr1 region<br />
Ling Zhang* and Andris Kleinhofs<br />
<strong>Department</strong> of Crop and Soil Sciences<br />
Washington State University, Pullman, WA, 99164-6420<br />
*E-mail: lzhang1@wsu.edu<br />
Abstract<br />
Eight cleaved amplified polymorphic sequences (CAPS) markers were developed around Rpr1<br />
genetic region. Eight markers were co-dominant between barley cultivars Morex and Steptoe<br />
after digestion with restriction enzymes.<br />
Introduction<br />
In barley, resistance to Puccinia graminis f. sp. tritici pathotype MCC requires the presence of at<br />
least of two host genes, Rpg1 (Brueggeman et al., 2002) and Rpr1 (Zhang et al., 2006).<br />
Mutational analysis and transcript-based cloning were used to isolate 3 candidate Rpr1 genes.<br />
These 3 candidate Rpr1 genes, HU03D17U_s_at, Contig4901_s_at and Contig7061_s_at were<br />
mapped to chromosome 4 bin 5. Screening recombinants between the three candidate genes will<br />
identify the real Rpr1 gene. Therefore, molecular markers in this region are needed.<br />
Cleaved amplified polymorphic sequences (CAPS) markers are PCR-based markers, requires a<br />
small amount of genomic DNA, which will facilitate the screening of large numbers of<br />
genotypes at the seedling stage. 141 probesets representing a major Rpr1 eQTL served as a<br />
starting point to develop CAPS markers. Here we report the development and mapping of CAPS<br />
markers in the Rpr1 region.<br />
Materials and Methods<br />
CAPS markers development<br />
Plant genomic DNA extraction was modified from Edwards et al. (1991); the modification added<br />
an extra-step of chloroform-isoamyl alcohol (24:1) extraction. Barley EST unigene sequences<br />
(HarvEST assembly#21; http://harvest.ucr.edu/) were used as templates for primer design. RFLP<br />
clone MWG058 was sequenced using primers T3 and T7 with the BigDye terminator system on<br />
ABI Prizm 377 DNA sequencer (Applied Biosystems) at the Bioanalytical Center, Washington<br />
State University, Pullman. A pair of primers was designed from the MWG058 sequence. All the<br />
primer pairs listed in Table 1 were used to amplify genomic DNA from the parent cultivars<br />
Morex and Steptoe. All PCRs of 20μl contained 20-50ng of genomic DNA, 0.1mMdNTP mix,<br />
12.5 pmol of each primer, 1μl of RedTaq DNA polymerase (Sigma), and 2 μl of 10xRedTaq<br />
reaction buffer. Amplification was performed in a PTC-100 programmable thermal controller<br />
(MJ Research, Cambridge, MA) at 95°C for 4 min, followed by 35 cycles of 95°C for 1 min,<br />
60°C for 1 min, and 72°C for 1 min; this was followed by 7 min at 72°C. PCR products were<br />
purified using the Gel Extraction Kit (Qiagen, Valencia, CA) and sequenced. Steptoe sequence<br />
was compared to the Morex sequence in order to identify single nucleotide polymorphisms<br />
(SNPs) that could be utilized for CAPS marker development. Sequence analysis was done by<br />
VectorNTI software (Invitrogen). SNPs were identified and restriction enzymes (New England<br />
44
BioLabs) were selected (Table 1). All the PCR products were digested directly using restriction<br />
enzymes correspondingly. Cleaved PCR products were then separated on 1% agarose gel.<br />
Genetic mapping<br />
The Steptoe x Morex "minimapper" population consisting of 35 selected doubled-haploid lines<br />
(DHL), was used to map the molecular markers to the barley Bin map (Kleinhofs and Graner<br />
2002). CAPS marker genetic order and the distance between snp_3139 and LZ2502 was<br />
estimated based on segregation data from Steptoe x Rpr1 F2 population.<br />
Results and Discussion<br />
Details of developed CAPS markers are listed in Tables 1 and Fig. 1. Molecular mapping in<br />
Steptoe x Morex population with CAPS markers showed that LZ6641, LZ13393 and LZ10152<br />
co-segregated with LZMWG058, ABG484 and BCD453B, respectively. Markers snp_3139<br />
(Druka, personal communication) and LZ2502 are the closest to Rpr1 delimiting the 3 markers<br />
LZ17u, LZ4901 and LZ7061 that co-segregate with Rpr1. LZ2502 and sn3139 are about 1cM<br />
apart and can be used to screen recombinants in Steptoe x rpr1 F2 population.<br />
Figure 1. Chromosome locations of eight CAPS marker in barley chr. 4 (4H).<br />
45
Table 1. Summary of developed CAPS markers, primers, restriction enzymes and annotation. All<br />
the CAPS markers were mapped to Chr 4 (4H), except LZ15227 was mapped to Chr 7 (5H).<br />
Markers<br />
Affymetrix<br />
Probesets<br />
LZ2502 Contig<br />
2502_at<br />
LZ6641 Contig<br />
6641_at<br />
LZ6435 Contig<br />
6435_at<br />
LZ13393 Contig<br />
13393_at<br />
LZ15227 Contig<br />
15227_s_at<br />
LZ1321 Contig<br />
1321_at<br />
LZ10152 Contig<br />
10152_at<br />
LZMWG058 a<br />
snp_3139 b<br />
Forward Primer<br />
Reverse Primer<br />
2502F: AGCTTCAGCTTCAGGTCGAT<br />
2502R: GAAACTTAGAACCTGAACC<br />
6641F: TGATTGATCCTTTGCTGTCT<br />
6641R: CTGGAAAGCGTTCAAATGCT<br />
6435F:ACACCAGGAAGATCATCGAC<br />
6435R:ACAATGGAGAACACATGGTT<br />
13393F:AAGTGGACCGCGAAGCACGT<br />
13393R:GCAGCATGTCAGGTTATACA<br />
15227F:ATGGACTAATGACCCCAACA<br />
15227R:TGCAACACACAAAGCCAGTC<br />
1321F: CACTATCGACTTCCCGGAAT<br />
1321R: ACTGCAATCAGGGTTCATCA<br />
10152F: AGATCTCCGGCTACGTGCTG<br />
10152R: CGTACATCAGCTCGAAGAAA<br />
MWG058F: ATTCATGCATCTACCCATCTCA<br />
MWG058R: TTGGATTGGCTAGAATCCTGGA<br />
3139F: AACCACGCAGCAAGCCTAT<br />
3139R: CTCGCTTCCTCCGTCATCAT<br />
Enzyme Annotation<br />
HpyCH4V putative IAA1 protein<br />
AvaII putative expressed SLT1 protein<br />
DdeI phosphoenolpyruvate carboxykinase<br />
(ATP) -like protein<br />
AvaII hypothetical protein<br />
AseI microtubule associated protein<br />
Sau3A or<br />
MboI<br />
Sau3A or<br />
MboI<br />
calmodulin<br />
BtsI unknown<br />
DdeI unknown<br />
putative membrane protein<br />
a<br />
LZMWG058 was developed from RFLP clone MWG058.<br />
b<br />
snp_3139 sequence provided by Druka A, Scottish Crop Research Institute (SCRI), Invergowrie,<br />
Dundee DD2 5DA, UK<br />
References<br />
Brueggeman R., Rostoks N., Kudrna D., Kilian A., Han F., Chen J., Druka A., Steffenson B.,<br />
Kleinhofs A. 2002. The barley stem rust-resistance gene Rpg1 is a novel diseaseresistance<br />
gene with homology to receptor kinases. Proc. Natl. Acad. Sci. <strong>US</strong>A 99: 9328-<br />
9333.<br />
Edwards K., Johnstone C., Thompson C. 1991. A simple and rapid method for the preparation of<br />
plant genomic DNA for PCR analysis. Nucleic Acids Res. 19 (6):1349.<br />
Kleinhofs A., Graner A. 2002. An integrated map of the barley genome. In: R. L. Phillips and I.<br />
Vasil, eds., DNA-Based Markers in Plants, 2nd Edition. Kluwer Academic Publishers,<br />
Boston. pp. 187-199.<br />
Zhang L., Fetch T., Nirmala J., Schmierer D., Brueggeman R., Steffenson B., Kleinhofs A. 2006.<br />
Rpr1, a gene required for Rpg1-dependent resistance to stem rust in barley. Theor Appl<br />
Genet 113: 847-855<br />
46
Barley Genetics Newsletter (2007) 37: 47-49<br />
Tolerance to high copper ions concentration in the nutrient medium of some<br />
Bulgarian barley cultivars<br />
J. Stoinova 1 , S. Phileva 1 , M. Merakchyiska 2 and S. Paunova 2<br />
1 D. Kostov Institute of Genetics, Bulgarian Academy of Sciences, 1113 Bulgaria<br />
2 M. Popov Institute of Plant Physiology, Bulgarian Academy of<br />
Sciences, 1113 Bulgaria<br />
Abstract. The winter two-rowed barley cultivars Obzor, Krasii 2, Vihren and Karan were<br />
examined for tolerance to environmental copper ions pollution. The plants were treated with 10 -6<br />
M and 10 -5 M CuSO4.5H2O for 7 days. It was found that under 10 -6 M the roots of cultivars<br />
Krassi 2 and Vihren were longer by 9% and 16% respectively, while plant green part was<br />
identical to that of the control. The root of cultivar Karan was severely inhibited its length<br />
reaching 77% that of the control. The green part was less affected. The data suggest that cultivars<br />
Krasii 2 and Vihren are tolerant while cultivar Karan is susceptible to increased copper ions<br />
concentration in the environment.<br />
Key words: barley, tolerance, copper ions<br />
Barley is an economically important crop being efficiently used in animal husbandry and<br />
brewing. Resistant to abiotic and biotic stress barley cultivars are a reliable tool to produce high<br />
quality stuff avoiding diverse chemical substances for pest control and other agricultural<br />
practices. Acevedo and Fereres (1993 ) concluded that breeding for abiotic stress resistance is<br />
becoming more promising with the recognition that selection showed be carried out in the target<br />
environment and that it is related to narrow adaption.<br />
During the last decades barley resistance to certain heavy metals has been examined in<br />
different aspects. It was shown that increased boron concentration in the environment reduce<br />
plant growth rate, while relative susceptibility of the genotypes to boron toxicity was not<br />
affected ( Nable et al.,1999). The effect of high boron concentration on barley was studied on<br />
cell level ( Jenkin et al.,1993). Data from studies with leaf protoplasts revealed that lack of cell<br />
walls prevents manifestation of differences between tolerant and susceptible barley genotypes.<br />
Treatment of H. vulgare seeds with nickel sulphate resulted into morphological alterations and<br />
increased chlorophyll and protein percentage, as well as that of the aberrant cells (Mishra and<br />
Singh, 1999). Addition of copper (2-4 kg/ha) led to higher grain yield from wheat, rye and oats<br />
(Piening et al., 1989). Tang et al. (2000) studied the location of genes for tolerance to<br />
aluminium and found that the latter is disposed to the long arm of 4H chromosome, while<br />
according to Rigin and Yakovleva (2001) the tolerance is coded by two polygenes.<br />
The purpose of this investigation was to examine the tolerance to increased copper ions<br />
concentration in the nutrient medium of four Bulgarian barley cultivars.<br />
Materials and Methods<br />
The winter two-rowed cultivars Obzor, Krassi 2, Vihren and Karan were studied. The seeds<br />
were germinated in moist filter paper rols in a semi-dark chamber at 18° C. After germination<br />
(2.5 – 3.5 cm root length) the shoots were transffered on a solution containing 10 -6 M or 10 -5 M<br />
CuSO4.5H2O at a temperature of while the control plants were grown in distilled water. All<br />
47
plants were grown at 12 h illumination/12 h dark and 20°- 24° C. After 7 days treatment<br />
biometric data were collected. To determine copper tolerance of the plants usually the tolerance<br />
index (Simon, 1978) was used, i.e. the plant growth in a heavy metal polluted medium was<br />
compared to that under control conditions:<br />
growth in a polluted medium<br />
IT =<br />
100<br />
growth under control conditions<br />
The data of copper influence on length of root and shoot have been processed by the variation<br />
statistical method.<br />
Results and Discussion<br />
A three-fold treatment with 10 -6 M copper ions concentration of the root (cultivar Obzor)<br />
caused inhibition (93-94% as compared to the control ) or stimulation up to 103%. In the case of<br />
the cultivar Karan the roots were strongly inhibited their length varying from 62% to 94% of the<br />
control. The plants of the cultivar Krassi 2 and cultivar Vihren responded to two of the<br />
treatments through stimulated root growth from 122% to 135% and from 116% to 129%,<br />
respectively. The shoot length was less affected, some inhibition varying from 1% to 6% or<br />
stimulation up to 2 – 4% being manifested by the cultivars Obzor, Krassi 2 and Vihren. More<br />
significant inhibition was manifested by cultivar Karan.<br />
The higher (10 -5 M) copper ions concentration severely inhibited both root and shoot of the<br />
plants from all cultivars after the three treatment accomplished.<br />
The mean data (Table 1) show that 10 -6 M ions stimulate (9-16%) root growth of the plants<br />
from cultivars Krassi 2 and Vihren while the roots of the plants from Karan reaced only 77% of<br />
the control. The green part of the plants was equal to that of the control ones. The variation<br />
coefficients revealed slight heterogenic variation, while the plant green part length variation was<br />
homogenic, exept that for cultivar Karan. Higher (10 -5 M) copper ions concentration highly<br />
reduced root growth, root length reaching 32-41% that of the control. The aboveground part of<br />
the plants from cultivar Krassi 2 and cultivar Vihren was identical to that of the control while<br />
those of cultivar Karan and Obzor were 69-75% that of the control. The variation coefficients<br />
revealed slight heterogenic variation.<br />
Under 10 -6 M copper ions concentration root mass of cultivar Obzor and cultivar Karan<br />
reaching 90-91% that of the control, the negative effect being stronger than that exerted on<br />
cultivars Krassi 2 and Vihren. The green part mass increased (1-3%) in the case of cultivar<br />
Obzor, cultivar Krassi 2 and cultivar Vihren, while that of cultivar Karan was slightly inhibited.<br />
10 -5 M copper ions concentration redused plant green part and root mass to 75-79% and 52-<br />
61%, respectively.<br />
To conclude, the cultivars Krassi 2 and Vihren manifested tolerance to increased copper ions<br />
concentration in the environment as under 10 -6 M the root was 9-16% longer than that of the<br />
control combined with less reduction of the mass (94%) of the control. The grain yield was 672<br />
kg/da and 677 kg/da, respectively. Therefore, they are suitable for cultivation in highly polluted<br />
regions.<br />
Cultivar Karan proved to be susceptible to copper pollution its root being severely reduced.<br />
Reference<br />
48
Barley Genetics Newsletter (2007) 37: 47-49<br />
Avecedo,E. and Fereres, E. 1993. Resistance to abiotic stress. In: Plant Breeding: principles and prospects<br />
(ed. by Hayward, M .D; Bosemark, N. O., Romagoza, I.). London, UK, Chapman and Hall Ltd,<br />
406-421.<br />
Jenkin, M., Hu, H. N.,Brown, P., Graham, R., Lance, R. and Sparrow, D. 1993. Investigation on boron<br />
uptake at the cellular level. Plant and Soil, 155/56, 143-146.<br />
Mishra, K. and Singh, R. J. 1999. Nickel genotoxicity assessment in Hordeum vulgare. J of<br />
Environmental Biology, 20, 71-72.<br />
Nable, R. O., Lance, R. C. H. and Cartwright, B. 1990. Uptake of boron and silicon by barley genotypes<br />
with differing susceptibilities to boron toxicity. Annals of Botany, 66, 83-90.<br />
Piening, L. J., Mac Pherson, D. J. and Malhi, S. S. 1989. Stem melanosis of some wheat, barley and oat<br />
cultivars on a copper deficient soil. Can. J of Plant Pathology, 11, 65-67.<br />
Rigin, B.V. and Yakovleva, O. V. 2001. Genetic aspects of barley tolerance to toxic aluminium ions.<br />
International research-practic conference Sankt-Peterburg, 13-16 november, 2001, p. 397.<br />
Simon, E. 1978. Heavy metals in soils, vegetation development and heavy metal tolerance in plant<br />
populations from metalliferous areas. New. Phytol., 81, 175-188.<br />
Tang, Y., Sorre, M. E., Kohain, L.V. and Garvin, D. F. 2000. Identification of RFLP markers linked to<br />
the barley aluminium tolerance gene Alp. Crop science, 40, 778-782.<br />
Table 1. Influence of copper ions on the length of root and shoot<br />
Variant Cultivar<br />
Length root Length shoot<br />
%control M ± m Vc% %control M ± m Vc%<br />
Control 100 12.0 ± 0.13 10.57 100 14.30 ± 0.13 8.93<br />
10 -6 M 97 11.7 ± 0.18 12.84 99 14.20 ±.0.13 7.88<br />
10 -5 Obzor<br />
M<br />
38 4.6 ±.0.36 14.82 75 10.67 ±.0.13 11.91<br />
Control 100 12.8 ± 0.21 15.28 100 14.40 ±.0.13 9.62<br />
10 -6 M 116 14.8 ± 0.23 15.51 100 14.40 ± 0.12 8.49<br />
10 -5 Krassi 2<br />
M<br />
41 5.3 ± 0.25 12.40 73 10.50 ± 0.11 11.11<br />
Control 100 13.7 ± 0.29 19.65 100 16.40 ± 0.15 8.40<br />
10 -6 M 109 15.0 ± 0.23 15.29 99 16.20 ±.0.16 9.47<br />
10 -5 Vihren<br />
M<br />
40 5.5 ± 0.08 15.75 72 11.80 ± 0.16 12.85<br />
Control 100 14.5 ± 0.23 14.65 100 15.40 ± 0.18 10.26<br />
10 -6 M 77 11.2 ± 0.22 16.50 98 15.10 ± 0.21 12.16<br />
10 -5 Karan<br />
M<br />
32 4.6 ± 0.08 16.49 69 10.60 ± 0.19 16.15<br />
49
Barley Genetics Newsletter (2007) 37: 100 - 104<br />
Rules for Nomenclature and Gene Symbolization in Barley<br />
Jerome D. Franckowiak 1 and Udda Lundqvist 2<br />
1 HermitageResearch Station<br />
Queensland <strong>Department</strong> of Primary Industries and Fisheries<br />
Warwick, Queensland 4370, Australia<br />
2 Nordic Gene Bank<br />
P.O. Box 41, SE-230 53 Alnarp, Sweden<br />
In this volume of the Barley Genetics Newsletter the recommended rules for nomenclature and<br />
gene symbolization in barley as reported in BGN 2:11-14, BGN 11:1-16, BGN 21:11-14, BGN<br />
26:4-8, and BGN 31:76-79 are again reprinted. Also, the current lists of new and revised BGS<br />
descriptions are presented by BGS number order (Table 1) and by locus symbol in alphabetic<br />
order (Table 2) in this issue.<br />
1. In naming hereditary factors, the use of languages of higher internationality should be<br />
given preference.<br />
2. Symbols of hereditary factors, derived from their original names, should be written in<br />
Roman letters of distinctive type, preferably in italics, and be as short as possible.<br />
AMENDMENT: The original name should be as descriptive as possible of the<br />
phenotype. All gene symbols should consist of three letters.<br />
COMMENTS: All new gene symbols should consist of three letters. Existing<br />
gene symbols of less than three letters should be converted to the three-letter<br />
system whenever symbols are revised. When appropriate, one or two letters<br />
should be added to existing symbols.<br />
For example, add the letters "ap" to "K" to produce the symbol "Kap" to replace<br />
"K" as the symbol for Kapuze (hooded). As another example, add the letters "ud"<br />
to "n" to produce the symbol "nud" to replace "n" as the symbol for naked seed.<br />
Similarly the letter "g" can be added to "ms" to produce the symbol "msg" for<br />
genetic male sterility and the letter "e" can be added to "ds" to produce the<br />
symbol "des" for desynapsis. When inappropriate or when conflicts arise,<br />
questions should be referred to the Committee on Genetic Marker Stocks,<br />
Nomenclature, and Symbolization of the International Barley Genetics<br />
Symposium for resolution.<br />
3. Whenever unambiguous, the name and symbol of a dominant begin with a capital letter<br />
and those of a recessive with a small letter.<br />
100
AMENDMENT: When ambiguous (co-dominance, incomplete dominance, etc.) all<br />
symbols should consist of a capital letter followed by two small letters that designate the<br />
character, a number that represents a particular locus, and a letter or letters that represents<br />
a particular allele or mutational event at that particular locus.<br />
COMMENTS: As an example, the letters "Mdh" can be used to designate the<br />
character malate dehydrogenase, "Mdh1" would represent a particular locus for<br />
malate dehydrogenase and "Mdh1a", "Mdh1b", "Mdh1c", etc. would represent<br />
particular alleles or mutational events at the "Mdh1" locus. Row number can be<br />
used as an example of symbolizing factors showing incomplete dominance. At the<br />
present time, the symbol "v" is used to represent the row number in Hordeum<br />
vulgare, "V" is used to represent the row number in Hordeum distichum, and "V t "<br />
is used to represent the row number in Hordeum deficiens. According to the<br />
amendment to Rule 3, if row number were to be designated by the letters "Vul",<br />
the designation of the locus on chromosome 2 would then become "Vul1" and the<br />
alleles "v", "V", and "V t " would be designated "Vul1a", "Vul1b", and "Vul1c".<br />
SUPPLEMENTARY AMENDMENT: A period should be placed before the allele<br />
symbol in the complete gene symbol.<br />
COMMENTS: Since DNA sequences similar to those of the original locus may<br />
occur at several positions in the Hordeum vulgare genome, a three-letter symbol<br />
plus a number is inadequate to represent all potential loci. Also, both numbers and<br />
letters have been assigned to specific mutants and isozymes in Hordeum vulgare.<br />
The six-rowed spike locus is used as an example although the symbol Vul1 for<br />
row number in Hordeum vulgare is not recommended because the botanical<br />
classification of Hordeum spp has changed. The locus symbol vrs1 and the name<br />
six-rowed spike 1 are recommended for the v locus. Gene symbols recommended<br />
for common alleles at the vrs1 locus are vrs1.a, vrs1.b, vrs1.c, and vrs1.t for the<br />
"v", "V", "v lr ", and "V t " genes, respectively.<br />
4. Literal or numeral superscripts are used to represent the different members of an allelic<br />
series.<br />
AMENDMENT: All letters and numbers used in symbolization should be written on one<br />
line; no superscripts or subscripts should be used.<br />
5. Standard or wild type alleles are designated by the gene symbols with a + as a superscript<br />
or by a + with the gene symbol as a superscript. In formulae, the + alone may be used.<br />
AMENDMENT: This rule will not be used in barley symbolization.<br />
6. Two or more genes having phenotypically similar effects are designated by a common<br />
basic symbol. Non-allelic loci (mimics, polymeric genes, etc.) are distinguished by an<br />
additional letter or Arabic numeral either on the same line after a hyphen or as a<br />
subscript. Alleles of independent mutational origin may be indicated by a superscript.<br />
101
AMENDMENT: Barley gene symbols should consist of three letters that designate the<br />
character, a number that represents a particular locus, and a letter or letters that represents<br />
a particular allele or mutational event at that particular locus. All letters and numbers<br />
should be written on the same line without hyphens or spaces. Alleles or mutational<br />
events that have not been assigned to a locus should be symbolized by three letters that<br />
designate the character followed by two commas used to reserve space for the locus<br />
number when determined, followed by a letter or letters representing the particular allele<br />
or mutational event. After appropriate allele testing, the correct locus number will be<br />
substituted for the commas. Where appropriate (when assigning new symbols or when<br />
revising existing symbols) letters representing alleles or mutational events should be<br />
assigned consecutively without regard to locus number or priority in discovery or<br />
publication.<br />
COMMENTS: The use of the proposed system of symbolization can be<br />
illustrated by the desynaptic mutants. Two loci are known: lc on chromosome 1<br />
(7H) and ds on chromosome 3 (3H). These will be resymbolized as des1a and<br />
des2b. A large number of desynaptic mutants have been collected. They will be<br />
designated des,,c, des,,d, des,,e, etc. If allele tests show that des,,c is at a different<br />
locus than des1 and des2, des,,c will become des3c. If allele tests show that des,,d<br />
is at the same locus as des2, des,,d will become des2d. In practical use, the<br />
symbol des will be used when speaking of desynapsis in general or if only one<br />
locus was known for the character. The symbol des2 will be used when speaking<br />
of that particular locus, and the symbol des2b will be used only when speaking of<br />
that particular allele or mutational event. If additional designation is needed in<br />
particular symbolization, it can be obtained by adding numbers behind the allele<br />
letters, and, if still further designation is needed, letters can be added to the<br />
symbol behind the last number. Symbolization consisting of alternation of letters<br />
and numbers written on the same line without hyphens or spaces will allow for<br />
the expansion of the symbol as future needs arise. In any work with large numbers<br />
of polymeric gene mutants, every mutant has to be given a designation not shared<br />
by any other mutant of this polymeric group and this designation should become a<br />
part of the permanent symbol representing that particular allele or mutational<br />
event. This requirement can be met by assigning allele designations in<br />
consecutive order without regard to locus number.<br />
SUPPLEMENTARY AMENDMENT: A period should be used instead of two commas<br />
in gene symbols for mutants within a polymeric group that can not be assigned to a<br />
specific locus.<br />
COMMENTS: The des symbol should be used when referring to desynapsis in<br />
general; des1 and des2, for specific loci; des1.a and des2.b for specific genes or<br />
alleles at their respective loci; and des.c, des.d, des.e etc., for desynaptic mutants<br />
not assigned to a specific locus.<br />
SUPPLEMENTARY AMENDMENT:<br />
Even if the locus in question is the only one known that affects a given phenotype, the<br />
three-letter basic symbol is followed by a serial number.<br />
102
7. Inhibitors, suppressors, and enhancers are designated by the symbols I, Su, and En, or by<br />
i, su, and en if they are recessive, followed by a hyphen and the symbol of the allele<br />
affected.<br />
AMENDMENT<br />
This rule is no longer appicable and will not be used in barley symbolization.<br />
8. Whenever convenient, lethals should be designated by the letter l or L and sterility and<br />
incompatibility genes by s or S.<br />
AMENDMENT: This rule will not be used in barley symbolization.<br />
COMMENTS: J.G. Moseman (BGN 2:145-147) proposed that the first of the<br />
three letters for designating genes for reaction to pests should be R. The second<br />
and third letters will be the genus and species names of the pest.<br />
SUPPLEMENTARY COMMENT: A motion was passed during the workshop on<br />
"Linkage Groups and Genetic Stock Collections" at the Fifth International Barley<br />
Genetics Symposium in 1986 (Barley Genetics V:1056-1058, BGN 17:1-4), that<br />
the International Committee for Nomenclature and Symbolization of Barley<br />
Genes should "recommend use of Ml as the designation of genes for resistance to<br />
powdery mildew.”<br />
9. Linkage groups and corresponding chromosomes are preferably designated by Arabic<br />
numerals.<br />
SUPPLEMENTARY AMENDMENT: The current wheat homoeologous group<br />
numbering scheme (the Triticeae system) is recommended for Hordeum vulgare<br />
chromosomes. Arabic numerals followed by an H will indicate specific barley<br />
chromosomes. The H. vulgare chromosomes should be 7H, 2H, 3H, 4H, 1H, 6H, and 5H<br />
instead of 1, 2, 3, 4, 5, 6, and 7, respectively.<br />
10. The letter X and Y are recommended to designate sex chromosomes.<br />
AMENDMENT: This rule will not be used in barley symbolization.<br />
11. Genic formulae are written as fractions with the maternal alleles given first or above.<br />
Each fraction corresponds to a single linkage group. Different linkage groups written in<br />
numerical sequence are separated by semicolons. Symbols of unlocated genes are placed<br />
within parenthesis at the end of the formula. In euploids and aneuploids, the gene<br />
symbols are repeated as many times as there are homologous loci.<br />
12. Chromosomal aberrations should be indicated by abbreviations: Df for deficiency, Dp for<br />
duplication, In for inversion, T for translocation, Tp for transposition.<br />
13. The zygotic number of chromosomes is indicated by 2n, the gametic number by n, and<br />
basic number by x.<br />
103
14. Symbols of extra-chromosomal factors should be enclosed within brackets and precede<br />
the genic formula.<br />
The following recommendations made by the International Committee for Nomenclature and<br />
Symbolization of Barley Genes at the Fourth International Barley Genetics Symposium in 1981<br />
(Barley Genetics IV:959-961) on gene and mutation designations were as follows.<br />
AMENDMENT:<br />
A. Present designations for genes and mutations. - Most of the present designations should<br />
be maintained. However, new designations may be given, when additional information<br />
indicates that new designations would aid in the identification of genes and mutations.<br />
B. New designations for genes and mutations. - New genes or mutations will be designated<br />
by characteristic, locus, allele, and then the order of identification or mutational event.<br />
Three letters will be used to identify new characteristics. Consecutive numbers will be<br />
used to identify the order of identification or mutational event. Loci will be designated by<br />
numbers and alleles by letters when they are identified. For example, des-6 indicates that<br />
this is the sixth gene or mutation identified for the characteristic des (desynaptic). des 1-6<br />
and des 2-7 indicate that gene or mutational events 6 and 7 for the desynaptic<br />
characteristic have been shown to be at different loci and those loci are then designated 1<br />
and 2, respectively. des 1a6 and des 1b8, indicate that the gene or mutational events 6<br />
and 8 for the characteristic desynaptic have been shown to be at different alleles at locus<br />
1 and those alleles are then designated a and b.<br />
SUPPLEMENTARY COMMENT:<br />
A motion was passed during the workshop of the "Nomenclature and Gene<br />
Symbolization Committee" at the Fifth International Barley Genetics Symposium in 1986<br />
(Barley Genetics V:1056-1058) that "the recommended systems for Nomenclature and<br />
Gene Symbolization of the International Committee be published annually in the Barley<br />
Genetics Newsletter."<br />
SUPPLEMENTARY COMMENT 2:<br />
At the workshop for ”Recommendations of Barley Nomenclature” held at Saskatoon,<br />
July 31, 1996 and adopted at the General Meeting of the Seventh International Barley<br />
Genetics Symposium, it was recommended that a period instead of a dash be used to<br />
designate the allele portion of the gene symbol. Consequently, the first gene symbol for<br />
the characteristic des (desynapsis) should be expressed as des1.a. The code des1<br />
identifies a specific locus. The period indicates that the symbol a identifies a specific<br />
allele or mutational event that produces a desynaptic phenotype. (The allele symbol a<br />
will be always associated with this specific desynaptic mutant even if the locus symbol is<br />
changed based on subsequent research results.)<br />
104
Barley Genetics Newsletter (2007) 37: 105-153<br />
REPORTS OF THE COORDINATORS<br />
Overall coordinator’s report<br />
Udda Lundqvist<br />
Nordic Gene Bank<br />
P.O. Box 41. SE-230 53 Alnarp<br />
e-mail: udda@nordgen.org<br />
Since the latest overall coordinator’s report in Barley Genetics Newsletter Volume 36, some<br />
changes of the coordinators have taken place. I do hope that most of you are willing to continue<br />
with this work and provide us with new important information and literature search in the future.<br />
A replacement was found for Chromosome 4H, namely Arnis Druka, Genetics Programme at the<br />
Scottish Crop Research Institute, Invergowrie, Dundee, United Kingdom. Please observe some<br />
address changes have taken place since the last volume of BGN. Jerry Franckowiak, the<br />
Coordinator for chromosome 2H, the semi-dwarf collection and all his immense efforts creating<br />
isogenic lines in the Bowman genetic background of many different barley genetic stocks has<br />
moved from North Dakota State University to Warwick, Queensland, Australia. The Curator, An<br />
Hang, for the Barley Genetics Stock Center at the <strong>US</strong>DA-ARS station at Aberdeen, Idaho, <strong>US</strong>A,<br />
has retired during the year 2007. Dr. Harold Bockelman from the same station is nominated as<br />
successor. An Hang has been involved and engaged in Barley Genetics since many decades, first<br />
together with Tak Tsuchiya at Fort Collins, Colorado and since 1990 at Aberdeen, Idaho. He<br />
took care of the move of all genetic barley stocks from Fort Collins to Aberdeen, has been<br />
evaluating and increasing most of them. He has been a considerable collaborator and colleague<br />
to the barley community, handled with big carefulness all the different barley types and<br />
transferred a large knowledge to all of us. I take this opportunity to thank him for all his<br />
kindness, helpfulness, enthusiasm and inspiration during all these years. All the best wishes to<br />
him in the future and his retirement. But I want to thank those who have resigned for their good<br />
corporation and the reliability of sending informative reports during all the years.<br />
In this connection I also want to call upon the barley community to pay attention on the AceDB<br />
database for ’Barley Genes and Barley Genetic Stocks’. It contains much information connected<br />
with images and is useful for barley research groups inducing barley mutants and looking for<br />
new characters. It gets updated continuously and some more images are added to the original<br />
version. The searchable address is: www.untamo.net/bgs<br />
In some months the 10th International Barley Genetics Symposium will be organized in<br />
Alexandria, Egypt. I hope that many of you will be to participate in the meetings. It is of big<br />
importance to discuss the future of different items, especially the coordination system and the<br />
future of Barley Genetics Newsletter. I would like to encourage the coordinators and their<br />
colleagues already to-day to provide me with suggestions, ideas, items or topics to be brought up<br />
during the meetings.<br />
105
Barley Genetics Newsletter (2007) 37: 105-153<br />
List of Barley Coordinators<br />
Chromoosome 1H (5): Gunter Backes, The University of Copenhagen, Faculty of Life Science,<br />
<strong>Department</strong> of Agricultural Sciences, Thorvaldsensvej 40, DK-1871 Fredriksberg C, Denmark.<br />
FAX: +45 3528 3468; e-mail: <br />
Chromosome 2H (2): Jerry. D. Franckowiak, Hermitage Research Station, Queensland<br />
<strong>Department</strong> of Primary Industries and Fisheries, Warwick, Queensland 4370, Australia, FAX:<br />
+61 7 4660 3600; e-mail: <br />
Chromosome 3H (3): Luke Ramsey, Genetics Programme, Scottish Crop Research Institute,<br />
Invergowrie, Dundee, DD2 5DA, United Kingdom. FAX: +44 1382 562426. E-mail:<br />
<br />
Chromosome 4H (4): Arnis Druka, Genetics Programme, Scottish Crop Research Institute,<br />
Invergowrie, Dundee, DD2 5DA, United Kingdom. FAX: +44 1382 562426. e-mail:<br />
<br />
Chromosome 5H (7): George Fedak, Eastern Cereal and Oilseed Research Centre, Agriculture<br />
and Agri-Food Canada, ECORC, Ottawa, ON, Canada K1A 0C6, FAX: +1 613 759 6559; email:<br />
<br />
Chromosome 6H (6): Duane Falk, <strong>Department</strong> of Crop Science, University of Guelph, Guelph,<br />
ON, Canada, N1G 2W1. FAX: +1 519 763 8933; e-mail: <br />
Chromosome 7H (1): Lynn Dahleen, <strong>US</strong>DA-ARS, State University Station, P.O. Box 5677,<br />
Fargo, ND 58105, <strong>US</strong>A. FAX: + 1 701 239 1369; e-mail: <br />
Integration of molecular and morphological marker maps: Andy Kleinhofs, <strong>Department</strong> of<br />
Crop and Soil Sciences, Washington State University, Pullman, WA 99164-6420, <strong>US</strong>A. FAX:<br />
+1 509 335 8674; e-mail: <br />
Barley Genetics Stock Center: An Hang, <strong>US</strong>DA-ARS, National Small Grains Germplasm<br />
Research Facility, 1691 S. 2700 W., Aberdeen, ID 83210, <strong>US</strong>A. FAX: +1 208 397 4165; e-mail:<br />
<br />
and<br />
Harold Bockelmann, National Small Grains Collection, U.S. <strong>Department</strong> of Agriculture –<br />
Agricultural Research Service, 1691 S. 2700 W., Aberdeen, ID 83210, <strong>US</strong>A. FAX: +1 208 397<br />
4165. e-mail: <br />
Trisomic and aneuploid stocks: An Hang, <strong>US</strong>DA-ARS, National Small Grains Germplasm<br />
Research Facility, 1691 S. 2700 W., Aberdeen, ID 83210, <strong>US</strong>A. FAX: +1 208 397 4165; e-mail:<br />
<br />
Translocations and balanced tertiary trisomics: Andreas Houben, Institute of Plant Genetics<br />
and Crop Plant Research, Corrensstrasse 3, DE-06466 Gatersleben, Germany. FAX: +49 39482<br />
5137; e-mail: <br />
106
Barley Genetics Newsletter (2007) 37: 105-153<br />
List of Barley Coordinators (continued)<br />
Desynaptic genes: Andreas Houben, Institute of Plant Genetics and Crop Plant Research,<br />
Corrensstrasse 3, DE-06466 Gatersleben, Germany. FAX: +49 39482 5137; e-mail:<br />
<br />
Autotetraploids: Wolfgang Friedt, Institute of Crop Science and Plant Breeding, Justus-Liebig-<br />
University, Heinrich-Buff-Ring 26-32, DE-35392 Giessen, Germany. FAX: +49 641 9937429; email:<br />
<br />
Disease and pest resistance genes: Brian Steffenson, <strong>Department</strong> of Plant Pathology,<br />
University of Minnesota, 495 Borlaug Hall, 1991 Upper Buford Circle, St. Paul, MN 55108-<br />
6030, <strong>US</strong>A. FAX: +1 612 625 9728; e-mail: <br />
Eceriferum genes: Udda Lundqvist, Nordic Gene Bank, P.O. Box 41, SE-230 53 Alnarp,<br />
Sweden. FAX:.+46 40 536650; e-mail: < udda@nordgen.org><br />
Chloroplast genes: Mats Hansson, <strong>Department</strong> of Biochemistry, University of Lund, Box 124,<br />
SE-221 00 Lund, Sweden. FAX: +46 46 222 4534 e-mail: <br />
Genetic male sterile genes: Mario C. Therrien, Agriculture and Agri-Food Canada, P.O. Box<br />
1000A, R.R. #3, Brandon, MB, Canada R7A 5Y3, FAX: +1 204 728 3858; e-mail:<br />
<br />
Ear morphology genes: Udda Lundqvist, Nordic Gene Bank, P.O. Box 41, SE-230 53 Alnarp,<br />
Sweden. FAX: +46 40 536650; e-mail: < udda@nordgen.org><br />
or<br />
Antonio Michele Stanca: Istituto Sperimentale per la Cerealicoltura, Sezione di Fiorenzuola<br />
d’Arda, Via Protaso 302, IT-29017 Fiorenzuola d’Arda (PC), Italy. FAX +39 0523 983750, email:<br />
<br />
Semi-dwarf genes: Jerry D. Franckowiak, Hermitage Research Station, Queensland <strong>Department</strong><br />
of Primary Industries and Fisheries, Warwick, Queensland 4370, Australia, FAX: +61 7 4660<br />
3600; e-mail: < Jerome.franckowiak@dpi.qld.gpv.au ><br />
Early maturity genes: Udda Lundqvist, Nordic Gene Bank, P.O. Box 41 SE-230 53 Alnarp,<br />
Sweden. FAX: +46 40 536650; e-mail: <br />
Biochemical mutants - Including lysine, hordein and nitrate reductase: Andy Kleinhofs,<br />
<strong>Department</strong> of Crop and Soil Sciences, Washington State University, Pullman, WA 99164-6420,<br />
<strong>US</strong>A. FAX: +1 509 335 8674; e-mail: <br />
Barley-wheat genetic stocks: A.K.M.R. Islam, <strong>Department</strong> of Plant Science, Waite Agricultural<br />
Research Institute, The University of Adelaide, Glen Osmond, S.A. 5064, Australia. FAX: +61 8<br />
8303 7109; e-mail: <br />
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Coordinator’s Report: Barley Chromosome 1H (5)<br />
Gunter Backes<br />
The University of Copenhagen<br />
Faculty of Life Sciences<br />
<strong>Department</strong> of Agricultural Sciences<br />
Thorvaldsensvej 40<br />
DK-1871 Frederiksberg C, Denmark<br />
guba@life.ku.dk<br />
In Arabidopsis, HLM1 encodes the cyclic nucleotide-gated ion channel 4. A mutant plant for this<br />
gene shows necrotic lesions and thereby similarities to the hypersensitive response (HR) to<br />
pathogens. Rostoks et al. (2006) isolated the homolog of this gene from the previously<br />
characterized barley mutant nec1 and localized the gene in the ‘Steptoe’ x ‘Morex’ “minimapper<br />
population” to chromosome 1H, Bin9.<br />
In an attempt to localize transcription factors (TFs ) belonging to the gene family of C-repeat<br />
binding factors (CBF), their regulators and MYB-TFs, altogether known to regulate plant<br />
response to cold and drought stress, Tondelli et al. (2006) localized several homologs to the<br />
respective Arabidopsis genes in a joined map of the three populations ‘Nure’ × ‘Tremois’,<br />
‘Proctor’ × ‘Nudinka’ and ‘Steptoe’ × ‘Morex’. Further, they compared the loci of these putative<br />
TFs with the position of published QTLs. They localized 9 homologs and assigned two further<br />
homologs by wheat-barley addition lines to the respective chromosomes. On chromosome 1H<br />
they localized HvMYB4 to Bin6, a homolog to AtMyb2 from Arabidopsis and OsMYB4 from rice,<br />
both known to be part of the regulation processes during abiotic stresses.<br />
In a similar effort, Skinner et al. (2006) localized the barley homologs of 14 Arabidopsis CBF-<br />
TFs and 2 further TFs in the barley populations ‘Steptoe’ × ‘Morex’ and 88Ab536 ×<br />
‘Strider’.The authors also tested the ‘Steptoe’ × ‘Morex’ population in climatic chambers for<br />
cold tolerance and localized QTLs based on these data. On chromosome 1H, they localized<br />
HvZFP16-1, a homolog to AtZAT12, to Bin4. A further homolog of the same Arabidopsis gene,<br />
HvZFPR16-2 was assigned to 1H by wheat-barley addition lines. A QTL for cold tolerance was<br />
localized on 1H, Bin11, by a LOD score of 5.7. No co-localization between the QTLs and the<br />
TFs localized in this study was found. Nevertheless, comparison with literature indicated QTLs<br />
at the position of two candidate gene loci on 5H.<br />
The same group (Szücs et al. 2006) published results describing the localization of photoreceptor<br />
genes and vernalization-related genes together with QTLs for photoperiod response. Mapping of<br />
both, the candidate genes and QTLs, was carried out in DH populations of the crosses ‘Dicktoe’<br />
× ‘Morex’ and ‘Dicktoe’ × ‘Kompolti korai’. While none of the candidate genes was localized on<br />
chromosome 1H, two QTL were detected with the ‘Dicktoo’ × ‘Morex’ population: a major QTL<br />
in Bin11 and a further QTL in Bin12.<br />
In order to localize qualitative and quantitative resistance against rice blast in barley, Inukai et al.<br />
(2006) analyzed a segregating DH population from the cross ‘Baroness’ × BCD47 with two<br />
different rice blast isolates in a greenhouse experiment. For one of the isolates, a qualitative<br />
segregation was found and consequently a new resistance gene, RMo1, was localized on<br />
chromosome 1H, Bin2 at or near the position of the Mla-locus. For the other isolate, a<br />
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quantitative segregation was found and a major QTL was detected at the position of RMo1, while<br />
further 3 QTLs were localized on the chromosomes 3H, 4H and 7H.<br />
Jafary et al. (2006) investigated the inheritance and specificity of plant factors that determine the<br />
degree of basal defence by host- and nonhost pathogens. For this purpose, they analyzed 152<br />
RILs from the cross ‘Vada’ x ‘SusPtrit’ with 2 rust isolates from barley rusts and 8 isolates from<br />
rusts with no barley-specificity, isolated from cultivated and wild Poaceae. ‘SusPtrit’ is an<br />
experimental barley accession selected for susceptibility to the wheat leaf rust fungus Puccinia<br />
triticina. On chromosome 1H, an R-gene against the fungus Puccinia hordei-secalini was<br />
localized. P. hordei-secalini has no host-specificity for H. vulgare. Furthermore, three different<br />
QTLs were detected. One of them conferred resistance against P. hordei-murini, one against P.<br />
graminis f.sp. lolii and one against P. graminis f.sp. tritici. Only the latter has barley-specificity.<br />
As the linkage map for 1H in this analysis purely consisted of AFLP marker, it was not possible<br />
to assign the R-gene or QTLs to the Bin-map.<br />
A new qualitative resistance gene against spot blotch, Rcs6, caused by Cochliobolus sativus, was<br />
localized on chromosome 1H either proximal on Bin1 or distal on Bin2 by Bilgic et al. (2006).<br />
They tested the DH population ‘Calcuchima-sib’ × ‘Bowman-BC’ with two different isolates<br />
both on seedlings in the greenhouse and on adult plants in the field. While one isolate identified<br />
the above mentioned resistance gene both in the seedlings and the adult plants, the other isolate<br />
detected different quantitative resistance loci for the greenhouse compared with the field, none of<br />
them on the position of Rcs6.<br />
Rsp2 and Rsp3, originally designated Sep2 and Sep3, are barley resistance genes against speckled<br />
leaf blotch in barley, caused by Septoria passerinii. These genes were mapped by Zhong et al.<br />
(2006) in an F2:3 population of the cross 'Foster' x 'Clho 4780' based on seedling tests with a<br />
specific isolate. These two genes are either closely linked or allelic and are localized on<br />
chromosome 1H and, as estimated by the flanking markers, more exactly in Bin3.<br />
Sameri et al. (2006) localized QTLs for different agronomic traits in an RIL population derived<br />
from a cross between two Japanese barley varieties ‘Azumamugi’ and ‘Kanto Nakate Gold’.<br />
‘Azumamugi’ is an oriental type barley, while ‘Kanto Nakate Gold’ belongs to the occidental<br />
type of barley varieties in Japan. The agronomic traits were evaluated in a field experiment on<br />
one location over two years. On chromosome 1H, one QTL for days to heading was localized<br />
near the position of Ppd-H2 (photoperiod sensitivity, Bin9/10) and one QTL for days to heading<br />
and days to maturity was detected near the position of eam8 (‘early maturity’, Bin14).<br />
In an F2:4 population from a cross between two wild barleys (H. v. ssp. spontaneum) from Israel,<br />
Vanhala and Stam (2006) localized QTL for seed dormancy. One of the lines (‘Mehola’)<br />
originates from the Jordan valley with low humidity and shows high seed dormancy, while the<br />
other line (‘Ashkelon’) originates from the Mediterranean coast with relatively high humidity<br />
and shows low dormancy. The germination rate was tested after 0 days, 14 days, 28 days and 42<br />
days of after-ripening at + 40˚ C. On chromosome 1H, the only QTL where the ‘Ashkelon’-allele<br />
prolongated the dormancy was found, while for the four other QTLs, on chromosomes 2H, 5H,<br />
6H and 7H, ‘Mehola’ contributed the allele with the higher dormancy. As the map of 1H was<br />
solely based on AFLPs, it was not possible to assign the position to a Bin.<br />
QTLs for grain yield (Bin11/12), heading date (Bin7), plant height (Bin14), ear length (Bin9,<br />
Bin13), spikelets/spike (Bin8/9), grain/spike (Bin8/9), spikes/plant (Bin12) and 1000-grain mass<br />
(Bin9, Bin11/12) were detected on chromosome 1H in an advanced-backcross experiment (Li et<br />
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al. 2006). The wild barley parent was the line ‘HS584’, and the recurrent cultivated parent was<br />
the variety ‘Brenda’. The field trials were carried out on 2 locations during four years. The map<br />
positions of the marker were based on the ‘Igri’ × ‘Franka’ and ‘Steptoe’ × ‘Morex’ SSR maps<br />
((Li et al., 2003).<br />
In another advanced-backcross with the wild barley line ‘ISR42-8’ and the recurrent cultivated<br />
parent ‘Scarlett’, von Korff et al. (2005) analyzed agronomic traits in a field experiments (four<br />
locations during two years). On 1H, QTLs were found for ears per m² (Bin14), heading date<br />
(Bin13), plant height (Bin13), harvest index (Bin13, Bin14) and yield (Bin6-8).<br />
References:<br />
Bilgic, H., B. J. Steffenson, and P. M. Hayes. 2006. Molecular mapping of loci conferring<br />
resistance to different pathotypes of the spot blotch pathogen in barley. Phytopathology<br />
96(7): 699-708.<br />
Inukai, T., M. I. Vales, K. Hori, K. Sato, and P. M. Hayes. 2006. RMo1 confers blast<br />
resistance in barley and is located within the complex of resistance genes containing Mla,<br />
a powdery mildew resistance gene. Mol. Plant Microbe Interact. 19(9): 1034-1041.<br />
Jafary, H., L. J. Szabo, and R. E. Niks. 2006. Innate nonhost immunity in barley to different<br />
heterologous rust fungi is controlled by sets of resistance genes with different and<br />
overlapping specificities. Mol. Plant Microbe Interact. 19(11): 1270-1279.<br />
Li, J. Z., X. Q. Huang, F. Heinrichs, M. W. Ganal, and M. S. Roder. 2006. Analysis of QTLs<br />
for yield components, agronomic traits, and disease resistance in an advanced backcross<br />
population of spring barley. Genome 49(5): 454-466.<br />
Li, J. Z., T. G. Sjakste, M. S. Röder, and M. W. Ganal. 2003. Development and genetic<br />
mapping of 127 new microsatellite markers in barley. Theor. Appl. Genet. 107(6): 1021-<br />
1027.<br />
Rostoks, N., D. Schmierer, S. Mudie, T. Drader, R. Brueggeman, D. G. Caldwell, R.<br />
Waugh, and A. Kleinhofs. 2006. Barley necrotic locus nec1 encodes the cyclic<br />
nucleotide-gated ion channel 4 homologous to the Arabidopsis HLM1. Mol. Genet.<br />
Genom 275(2): 159-168.<br />
Sameri, M., K. Takeda, and T. Komatsuda. 2006. Quantitative trait loci controlling agronomic<br />
traits in recombinant inbred lines from a cross of oriental- and occidental-type barley<br />
cultivars. Breed. Sci. 56(3): 243-252.<br />
Skinner, J. S., P. Szücs, J. von Zitzewitz, L. Marquez-Cedillo, T. Filichkin, E. J. Stockinger,<br />
M. F. Thomashow, T. H. H. Chen, and P. M. Hayes. 2006. Mapping of barley<br />
homologs to genes that regulate low temperature tolerance in Arabidopsis. Theor. Appl.<br />
Genet. 112(5): 832-842.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Szücs, P., I. Karsai, J. von Zitzewitz, K. Meszaros, L. L. D. Cooper, Y. Q. Gu, T. H. H.<br />
Chen, P. M. Hayes, and J. S. Skinner. 2006. Positional relationships between<br />
photoperiod response QTL and photoreceptor and vernalization genes in barley.<br />
Theor. Appl. Genet. 112(7): 1277-1285.<br />
Tondelli, A., E. Francia, D. Barabaschi, A. Aprile, J. S. Skinner, E. J. Stockinger, A. M.<br />
Stanca, and N. Pecchioni. 2006. Mapping regulatory genes as candidates for cold<br />
and drought stress tolerance in barley. Theor. Appl. Genet. 112(3): 445-454.<br />
Vanhala, T. K. and P. Stam. 2006. Quantitative trait loci for seed dormancy in wild barley<br />
(Hordeum spontaneum c. Koch). Genet. Resour. Crop. Evol. 53(5): 1013-1019.<br />
von Korff, M., H. Wang, and J. Léon. 2005. AB-QTL analysis in spring barley. I. Detection of<br />
resistance genes against powdery mildew, leaf rust and scald introgressed from wild<br />
barley. Theor. Appl. Genet. 111(3): 583-590.<br />
Zhong, S. B., H. Toubia-Rahme, B. J. Steffenson, and K. P. Smith. 2006. Molecular mapping<br />
and marker-assisted selection of genes for septoria speckled leaf blotch resistance in<br />
barley. Phytopathology 96(9): 993-999.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Chromosome 2H (2)<br />
J.D. Franckowiak<br />
Hermitage Research Station<br />
Queensland <strong>Department</strong> of Primary Industries and Fisheries<br />
Warwick, Queensland 4370, Australia<br />
e-mail: jerome.franckowiak@dpi.qld.gpv.au<br />
Komatsuda et al. (2007) cloned the six-rowed spike 1 (vrs1) gene located on chromosome 2HL<br />
of barley. Expression of the Vrs1 was strictly localized in the lateral-spikelet primordia of<br />
immature spikes and suggests that the VRS1 protein suppresses development of lateral spikelets.<br />
Phylogenetic analysis of the six-rowed cultivars and mutants demonstrated that six-rowed spike<br />
trait originated repeatedly from two-rowed barley, at least three different origins among<br />
domesticated accessions. Also, the DNA sequence defects in many of vrs1 mutants held in the<br />
Nordic Gene Bank were identified.<br />
When the DNA sequence of vrs1 was determined, Pourkheirandish et al. (2007) found that the<br />
region around the vrs1 locus was collinear with rice chromosome 4. However, the rice<br />
orthologue for the vrs1 sequence was found on rice chromosome 7. The authors speculated that a<br />
transposition of the chromosomal segment Vrs1 to chromosome 2H occurred during the<br />
evolution of barley. Pourkheirandish et al. (2007) also reported that the vrs1 locus is a region of<br />
suppressed recombination based on the study of more than 13,000 gametes.<br />
Řepková et al. (2006) reported on the mapping of four new sources of resistance to powdery<br />
mildew, caused by Blumeria graminis f. sp. hordei, that were identified in accessions of wild<br />
barley, Hordeum vulgare ssp. spontaneum. Accession PI 466197 was found to have two<br />
dominant resistance genes. One is an allelic at the mla locus and the other was located on<br />
chromosome 2HS based on a highly significant linkage with molecular marker Bmac0134.<br />
Dahleen and Franckowiak (2006) found that cer-zt locus is located on chromosome 2HS based<br />
on linkage to molecular marker Bmac0134 in bin 2H-1. The cert-zt.389 mutant has very little<br />
surface wax on the spike (Lundqvist and Franckowiak, 1997), but little effect on other<br />
agronomic traits except a slightly increased number of kernels per spike (Dahleen and<br />
Franckowiak, 2006).<br />
Based on the analysis of 134 recombinant chromosome substitution lines (RCLs) from the BC3<br />
generation of the backcross of wild barley line (OUH602) into ‘Haurna Nijo’, Hori et al. (2005)<br />
found that QTLs for short spike and lax spike are on chromosome 2HL near the closed flowering<br />
(cleistogamy, cly1/Cly2) locus of Haurna Nijo. In a previous paper, Hori et al. (2003) reported<br />
that these QTLs plus one for short culm were observed in a population of doubled-haploid lines<br />
from a Haurna Nijo/OUH602 cross.<br />
Using recombinant inbred lines, Yun et al. (2005) found a QTL for resistance to Septoria<br />
speckled leaf blotch (SSLB, caused by Septoria passerinii Sacc.) from H. vulgare subsp.<br />
spontaneum, located in bins 7 to 11 of chromosome 2H. They examined a recombinant inbred<br />
line (RIL) population developed from a cross between wild barley accession OUH602 and the<br />
two-rowed malting cultivar ‘Harrington’ for reaction to SSLB. About 40% of the variation in<br />
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response to SSLB was explained by the QTL on 2H, named QTL Rsp-2H-7-ll. The mapped<br />
disease resistances were validated using an advanced backcross population (BC2F6:8) from the<br />
same donor parent, but having two more backcrosses to Harrington (Yun et al., 2006).<br />
A QTL regulating synthesis of cell wall (1,3;1,4)-beta-D-glucans was located between the<br />
markers Adh8 bin 6 and ABG019 bin 7 with the peak closer to ABG019 on 2H (Burton et al.,<br />
2006). The cellulose synthase-like (CslF) gene cluster in cereals was identified as candidates<br />
responsible for mediating cell wall (1,3;1,4)- ß -D-glucan synthesis using of rice synteny and by<br />
transforming Arabidopsis (Burton et al., 2006). The research was based on the map location of a<br />
major QTL for (1,3;1,4)-ß-D-glucan content of un-germinated barley grains on 2H. This report is<br />
believed the first example of a map-based cloning of a QTL in barley.<br />
Korff et al. (2006) reported on a large number of QTLs for agronomic traits detected in doubledhaploid<br />
lines from the second backcross of ‘Scarlett’ backcrossed to Hordeum vulgare ssp.<br />
spontaneum accession ISR42-8. Using a population 301 BC2DH in eight environments, they<br />
reported detection of 86 QTLs for nine agronomic traits. The QTLs having large effects that<br />
were associated with chromosome 2H included: ears/m2, days to head (Eam1 or Ppd-H1), plant<br />
height (sdw1 from Scarlett), and yield.<br />
Yin et al. (2005) confirmed that a QTL having an important effect on preflowering duration in<br />
the ‘Apex’/‘Prisma’ population of 94 recombinant inbred lines (RILs) was located on the long<br />
arm of chromosome 2H. The other QTL having a large effect was located on chromosome 3H at<br />
the same position as the sdw1 gene from Prisma.<br />
Dragan et al. (2007) located two members of the nicotianamine synthase (NAS) family of genes<br />
on the short arm of chromosome 2H (2HS). Nicotianamine is involved chelation of iron and<br />
other heavy metals and their transport in the plant.<br />
The number of molecular markers located on chromosome 2H has been increased by several<br />
studies. Beaubien and Smith (2006) placed 7 of the 60 new mapped SSR markers on 2H at bin<br />
positions that previously had been identified as being poorly covered by SSR markers currently<br />
available. Stein et al. (2007) published an expressed sequence tag (EST)-based map for barley<br />
based 200 anchor markers from three previously published maps. The map contained 1,055 loci<br />
and a map size of 1,118.3 cM. The map for 2H contained 179 EST loci and a map length of<br />
165.1 cM. Using barley-wheat addition lines and the Barley1 Affymetrix GeneChip probe array,<br />
Cho et al. (2006) associated 1,787 of 4,104 transcript accumulation patterns detected in Betzes,<br />
but not Chinese Spring, with specific barley chromosomes. Of these 271 were associated with<br />
the 2H addition line of Chinese Spring.<br />
Takahashi et al. (2006) mapped in barley miniature inverted-repeat transposable elements<br />
(MITEs), which represent a large superfamily of transposons that is moderately to highly<br />
repetitive and frequently found near or within plant genes. To elucidate the organization of<br />
MITEs in the barley genome, MITEs were integrated into the genetic map of barley using 93<br />
doubled haploid lines from a Haruna Nijo by H. vulgare ssp. spontaneum accession OUH602<br />
cross. They described the use of MITEs in amplified fragment length polymorphism (AFLP)<br />
mapping and demonstrate their superiority over conventional AFLP mapping. A total of 214 loci<br />
covered a total map distance of 1,165 cM, and 39 were placed on 2H.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
References:<br />
Beaubien, K.A., and K. P. Smith. 2006. New SSR markers for barley derived from the EST<br />
database. Barley Genet. Newsl. 36:30-36.<br />
Burton, R.A., S.M. Wilson, M. Hrmova, A.J. Harvey, N.J. Shirley, A. Medhurst, B.A.<br />
Stone, E.J. Newbigin, A. Bacic, and G.B. Fincher. 2006. Cellulose synthase-like CslF<br />
genes mediate the synthesis of cell wall (1,3;1,4)-beta-D-glucans. Science 311:1940-<br />
1943.<br />
Cho, S., D.F. Garvin, and G.J. Muehlbauer. 2006. Transcriptome analysis and physical<br />
mapping of barley genes in wheat–barley chromosome addition lines. Genetics 172:1277-<br />
1285.<br />
Dahleen, L.S. and J.D. Franckowiak. 2006. SSR linkages to eight additional morphological<br />
marker traits. Barley Genet. Newsl. 36:12-16.<br />
Dragan Perovic, D., P. Tiffin, D. Douchkov, H. Bäumlein, and A. Graner. 2007. An<br />
integrated approach for the comparative analysis of a multigene family: The<br />
nicotianamine synthase genes of barley. Funct. Integr. Genomics 7:169-179.<br />
Hori, K., T. Kobayashi, A. Shimizu, K. Sato, K. Takeda, and S. Kawasaki. 2003. Efficient<br />
construction of high-density linkage map and its application to QTL analysis in barley.<br />
Theor. Appl. Genet. 107:806-813.<br />
Hori, K., K. Sato, N. Nankaku, and K. Takeda. 2005. QTL analysis in recombinant<br />
chromosome substitution lines and doubled haploid lines derived from a cross between<br />
Hordeum vulgare ssp. vulgare and Hordeum vulgare ssp. spontaneum. Mol. Breeding<br />
16(4):295-311.<br />
Komatsuda, T., M. Pourkheirandish, C. He, P. Azhaguvel, H. Kanamori, D. Perovic, N.<br />
Stein, A. Graner, T. Wicker, A. Tagiri, U. Lundqvist, T. Fujimura, M. Matsuoka, T.<br />
Matsumoto, and M. Yano. 2007. Six-rowed barley originated from a mutation in a<br />
homeodomain-leucine zipper I-class homeobox gene. PNAS 104:1424-1429.<br />
von Korff, M., H. Wang, J. Léon, and K. Pillen. 2006. AB-QTL analysis in spring barley: II.<br />
Detection of favourable exotic alleles for agronomic traits introgressed from wild barley<br />
(H. vulgare ssp. spontaneum). Theor. Appl. Genet. 112:1221-1231.<br />
Lundqvist, U., and J.D. Franckowiak. 1997. BGS 437, Eceriferum-zt, cer-zt. Barley Genet<br />
Newsl. 26:389<br />
Pourkheirandish, M., T. Wicker, N. Stein, T. Fujimura, and T. Komatsuda. 2007. Analysis<br />
of the barley chromosome 2 region containing the six-rowed spike gene vrs1 reveals a<br />
breakdown of the rice-barley micro collinearity by a transposition. Theor. Appl. Genet.<br />
114:1357-1365.<br />
Řepková, J., A. Dreiseitl, P. Lízal, Z. Kyjovská, K. Teturová, R. Psotková, and A. Jahoor.<br />
2006. Identification of resistance genes against powdery mildew in four accessions of<br />
Hordeum vulgare ssp. spontaneum. Euphytica 151:23-30.<br />
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Stein, N., M. Prasad, U. Scholz, T. Thiel, H. Zhang, M. Wolf, R. Kota, R.K. Varshney, D.<br />
Perovic, I. Grosse, and A. Graner. 2007. A 1,000-loci transcript map of the barley<br />
genome: new anchoring points for integrative grass genomic. Theor. Appl. Genet.<br />
114:823-839.<br />
Takahashi, H., H. Akagi, K. Mori, K. Sato, and K. Takeda. 2006. Genomic distribution of<br />
MITEs in barley determined by MITE-AFLP mapping. Genome 49:1616- 1618.<br />
Yin, X., P.C. Struik, F.A. van Eeuwijk, P. Stam, and J. Tang. 2005. QTL analysis and QTLbased<br />
prediction of flowering phenology in recombinant inbred lines of barley J. Exp.<br />
Bot. 56 (413):967-976.<br />
Yun, S.J., L. Gyenis, E. Bossolini, P.M. Hayes, I. Matus, K.P. Smith, B.J. Steffenson, R.<br />
Tuberosa, and G.J. Muehlbauer. 2006. Validation of quantitative trait loci for multiple<br />
disease resistance in barley using advanced backcross lines developed with a wild barley.<br />
Crop Sci. 46:1179-1186.<br />
Yun, S.J., L. Gyenis, P.M. Hayes, I. Matus, K.P. Smith, B.J. Steffenson, and G.J.<br />
Muehlbauer. 2005. Quantitative trait loci for multiple disease resistance in wild barley.<br />
Crop Sci. 45:2563-2572.<br />
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Coordinator’s Report: Barley Chromosome 3H<br />
L. Ramsay<br />
Genetics Programme<br />
Scottish Crop Research Institute<br />
Invergowrie, Dundee, DD2 5DA, Scotland, UK.<br />
e-mail: Luke.Ramsay@scri.ac.uk<br />
Over the last year there have been a number of publications reporting the mapping of genes and<br />
QTL on barley chromosome 3H. One of the highlights of this reporting period was the genetic<br />
mapping of over 1000 genes, including 179 on 3H, by Stein et al. (2007). This worked<br />
confirmed the syntenic relationship of 3H to rice chromosome 1 and importantly put the detailed<br />
EST information underlying this and previous reports in the public domain. This includes the<br />
details of which genes are represented by the EST derived microsatellites reported by Varshney<br />
et al. (2006) that included 35 that map to 3H. Another report of EST derived microsatellites with<br />
the associated EST information, including 11 on 3H, was that of Beaubien and Smith (2006).<br />
Another important general mapping paper was the development of a consensus map derived<br />
from DArT marker loci (Wenzl et al, 2006) that opens up the possibility of using loci derived<br />
from this technology as proxies for more expensive genic markers. Also of importance is the<br />
detailed consensus map presented by Marcel et al. (2007) which brings together standard AFLP,<br />
microsatellite and RFLP loci and that will allow additional alignment of past work with the<br />
positions of genic loci.<br />
A range of QTL on chromosome 3H were again reported this year. In a RIL population derived<br />
from a cross between Azumamugi and Kanto Nakate Gold studied by Sameri et al. (2006) QTL<br />
were found on 3H for a range of agronomic characters including plant height, spike length and<br />
awn length. The position of the QTL found indicates that they are due to the segregation of uzu<br />
in the population. Li et al. (2006) reported the positions of QTL for a range of agronomic traits<br />
using recombinant chromosome substation lines derived from a Hordeum vulgare subsp. vulgare<br />
(cltv. Brenda) by Hordeum vulgare subsp. spontaneum (accession HS584) cross to delineate<br />
association with genomic regions. The QTL found on 3H included those for yield and<br />
components such as spikelet no. per spike, grain no. per spike, thousand-grain mass as well as<br />
other traits such as heading date, plant height, ear length, leaf length and leaf area. A QTL for<br />
resistance on 3H was also found to leaf rust in two trials which may relate to the two QTL for<br />
leaf rust resistance found on chromosome 3H in a consensus map by Marcel et al. (2007) in a<br />
summary of work on six mapping populations. One of the populations used in the construction<br />
of this consensus map, L94 x Vada, was also tested for mildew and scald resistance and a novel<br />
powdery mildew resistance QTL designated Rbgq2 was detected on 3H which did not map to a<br />
region where a major gene for powdery mildew has previously been reported (Shtaya et al.<br />
2006). Another of the populations included in the report of Marcel et al. 2007 was that derived<br />
from a cross between an experimental line SusPrit and Vada to study the inheritance of non-host<br />
immunity to rusts (Jafary et al. 2006). This work found three QTL on 3H associated with host<br />
and non-host resistance to Puccinia spp. Other disease QTL reported on 3H included the<br />
improved resolution of spot blotch resistance QTL on the Calicuchima-sib / Bowman BC<br />
population by Bilgic et al. (2006) and a scald resistance QTL on the long arm of 3H identified<br />
using a partial map of a doubled haploid population derived from a Mundah/Keel cross (Cheong<br />
et al., 2006).<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
References:<br />
Beaubien, K.A. and K.P. Smith. 2006 New SSR markers for barley derived from the EST<br />
database. Barley Genetics Newsletter 36: 30-43.<br />
Bilgic, H., B.J. Steffenson, and P.M. Hayes. 2006. Molecular mapping of loci conferring<br />
resistance to different pathotypes of the spot blotch pathogen in barley. Phytopathology<br />
96: 699-708.<br />
Cheong, J., K. Williams, and H. Wallwork, 2006. The identification of QTLs for adult plant<br />
resistance to leaf scald in barley. Australian Journal of Agricultural Research 57: 961-<br />
965.<br />
Jafary, H., L.J. Szabo, and R.E. Niks, 2006. Innate nonhost immunity in barley to different<br />
heterologous rust fungi is controlled by sets of resistance genes with different and<br />
overlapping specificities. Molecular Plant-Microbe Interactions 19: 1270-1279.<br />
Li, J.Z., X.Q. Huang, F. Heinrichs, M.W. Ganal, and M.S. Roder, 2006. Analysis of QTLs<br />
for yield components, agronomic traits, and disease resistance in an advanced backcross<br />
population of spring barley. Genome 49: 454-466.<br />
Marcel, T.C., R.K. Varshney, M. Barbieri, H. Jafary, M.J.D. de Kock, A. Graner and R.E.<br />
Niks, 2007. A high-density consensus map of barley to compare the distribution of<br />
QTLs for partial resistance to Puccinina hordei and of defence gene homologues. Theor<br />
Appl. Genet 114: 487-500.<br />
Samuri, M., K. Takeda and Komatsuda, T. 2006. Quantitative trait loci controlling<br />
agronomic traits in recombinant inbred lines from a cross of oriental- and occidental-type<br />
barley cultivars. Breeding Science 56: 243-252.<br />
Shtaya, M.J.Y., T.C. Marcel, J.C. Sillero, R.E. Niks, and D. Rubiales, 2006. Identification of<br />
QTLs for powdery mildew and scald resistance in barley. Euphytica 151: 421-429.<br />
Stein, N., M. Prassa, U. Scholz, T. Thiel, H. Zhang, M. Wolf, R. Kota, R.K. Varshney, D.<br />
Perovic, I. Grosse and A. Graner, 2007. A 1,000-loci transcript map of the barley<br />
genome: new anchoring points for integrative grass genomics. Theor Appl Genet on-line<br />
preprint DOI 10.1007/s00122-006-0480-2<br />
Varshney, R.K., I. Grosse, U. Hahnel, R. Siefken, M. Prasad, N. Stein, P. Langridge, L.<br />
Altschmied, and A. Graner, 2006. Genetic mapping and BAC assignment of ESTderived<br />
SSR markers shows non-uniform distribution of genes in the barley genome.<br />
Theor Appl Genet 113: 239-250.<br />
Wenzl, P., H.B. Li, J. Carling, M.X. Zhou, H. Raman, E. Paul, P. Hearnden, C. Maier, L.<br />
Xia, V. Caig, J. Ovesna, M. Cakir, D. Poulsen, J.P. Wang, R. Raman, K.P. Smith,<br />
G.J. Muehlbauer, K.J. Chalmers, A. Kleinhofs, E. Huttner, and A. Kilian, 2006. A<br />
high-density consensus map of barley linking DArT markers to SSR, RFLP and STS loci<br />
and agricultural traits. BMC Genomics 7:206.<br />
117
Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s Report: Chromosome 4H<br />
Arnis Druka<br />
Genetics Programme<br />
Scottish Crop Research Institute<br />
Invergowrie, Dundee, DD2 5DA, Scotland, UK.<br />
e-mail: adruka@scri.sari.ac.uk<br />
Several papers that relate to the genes on chromosome 4H have been published in 2006 - 2007.<br />
At least four of them combine mRNA abundance analyses with phenotypic trait genetic analyses<br />
clearly showing added value of such approach (Malatrasi et al., 2006; Zhang et al., 2006; Wang<br />
et al., 2007 and Walia et al., 2007).<br />
The HvMATE gene, encoding a multidrug and toxic compound extrusion protein has been<br />
identified as a candidate controlling aluminium (Al) tolerance in barley. The gene itself was<br />
found not to be polymorphic between Al-tolerant and sensitive cultivars, but it accumulates<br />
mRNA 30 times more in the Al tolerant cultivar. HvMATE mRNA accumulation was measured<br />
in the F(2:3) families and was found significantly correlated with the Al tolerance and Alactivated<br />
citrate efflux phenotypes that have been mapped on the long arm of chromosome 4H<br />
(Wang et al., 2007).<br />
A different study addressed the salt tolerance in barley by analysing single feature<br />
polymorphisms (SFPs) and an oligonucleotide pool assay for single nucleotide polymorphisms<br />
(SNPs) in the salt tolerant cultivar Golden Promise and intolerant cultivar Maythorpe. Golden<br />
Promise has been generated by inducing mutation in the cultivar Maythorpe. The transcriptome<br />
analysis indicates that the response of the two genotypes to the salinity stress is quite different..<br />
This study identified 3 haplotype blocks spanning 6.4 cM on chromosome 1H, 23.7 cM on<br />
chromosome 4H and 3.0 cM on 5H suggesting that Golden Promise is not isogenic (Walia et al.,<br />
2007).<br />
A gene encoding the branched-chain amino acid aminotransferase (HvBCAT-1) that mapped on<br />
chromosome 4H, was identified by using differential mRNA display applied to ABA, drought<br />
and cold treated barley seedling shoots. Transcript levels of Hvbcat-1 increased in response to<br />
drought stress. The complementation of a yeast double knockout strain revealed that HvBCAT-1<br />
can function as the mitochondrial (catabolic) BCATs in vivo. This allowed to put forward the<br />
hypothesis, that under drought stress conditions, one of the detoxification mechanisms could be<br />
associated with degradation of the branched-chain amino acids (Malatrasi et al., 2006).<br />
Zhang et al. (2006) have reported a novel locus that is required for Rpg1 gene mediated<br />
resistance to the stem rust (Puccinia graminis f. sp. tritici) fungus. It was identified by inducing<br />
the irradiation mutations in the resistant barley cultivar and selecting for susceptible individuals<br />
in the M2 progeny. Rpg1 gene in one such susceptible mutant plants was found to be intact and<br />
the following mutation mapping identified a locus on chromosome 4H, that was named Rpr1<br />
(Required for P. graminis resistance). Several candidate genes or novel markers for this locus<br />
were identified by using large scale parallel transcript profiling approach.<br />
Other papers that related to chromosome 4H were describing either characterization and<br />
mapping gene families and the candidate genes for certain QTLs (Brueggeman et al., 2006;<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Skinner et al., 2006) or mapping novel QTLs (Friesen et al., 2006; Richardson et al., 2006; Yan<br />
and Chen 2006; von Korff et al., 2006).<br />
Thus, Brueggeman et al. (2006) reported mapping of members of the serine/threonine kinaselike<br />
protein family that encode at least one predicted catalytically active kinase domain. One of<br />
them was localized to chromosome 4H. In a different study, allelic nature and map locations of<br />
barley homologs to three classes of Arabidopsis low temperature regulatory genes-CBFs, ICE1,<br />
and ZAT12 were investigated for associations with the LT tolerance QTLs. In the same study,<br />
phenotyping of the Dicktoo x Morex (DxM) mapping population under controlled freezing<br />
conditions identified three new low temperature tolerance (LT) QTLs on 1H-L, 4H-S, and 4H-L<br />
in addition to the previously reported 5H-L Fr-H1 QTL. (Skinner et al., 2006).<br />
Barley interaction with the net blotch fungi, Pyrenophora teres f. teres (net-type net blotch<br />
(NTNB)) and Pyrenophora teres f. maculata (spot-type net blotch (STNB)) was studied using a<br />
doubled-haploid population derived from the lines SM89010 and Q21861. Major QTLs for<br />
NTNB and STNB resistance were located on chromosomes 6H and 4H, respectively (Friesen et<br />
al., 2006).<br />
Barley and the stripe rust fungus (Puccinia striiformis f. sp. hordei) interaction phenotypes, such<br />
as latency period, infection efficiency, lesion size and pustule density were mapped using i-<br />
BISON lines (intermediate barley near-isogenic lines). The (i-BISON) lines represented disease<br />
resistance QTL combined in one-, two-, and three-way combinations in a susceptible<br />
background. The 4H QTL allele had the largest effect followed by the alleles on<br />
chromosomes1H and 5H (Richardson et al., 2006).<br />
In a different study Yan and Chen (2006) reported population of 182 recombinant inbred lines<br />
(RILs) (F8) derived from cultivars Steptoe and GZ that was generated to map the resistance to<br />
two barley stripe rust fungus strains on the long arm of barley chromosome 4H.<br />
The BC2DH population derived from a cross between the spring barley cultivar Scarlett and the<br />
wild barley accession ISR42-8 (Hordeum vulgare ssp. spontaneum) was developed to evaluate<br />
nine agronomic traits. Favourable ISR42-8 alleles were detected for the yield-related traits that<br />
have QTLs on the long arm of chromosome 4H (von Korff et al., 2006).<br />
References:<br />
Brueggeman R. T. Drader, and A. Kleinhofs 2006. The barley serine/threonine kinase gene<br />
Rpg1 providing resistance to stem rust belongs to a gene family with five other members<br />
encoding kinase domains. Theor. Appl. Genet. 113(6):1147-1158.<br />
Friesen T.L., J.D. Faris, Z. Lai, and B.J. Steffenson. 2006. Identification and chromosomal<br />
location of major genes for resistance to Pyrenophora teres in a doubled-haploid barley<br />
population. Genome 49(7):855-859.<br />
Malatrasi M., M. Corradi, J.T. Svensson, T.J. Close, M. Gulli, and N. Marmiroli 2006. A<br />
branched-chain amino acid aminotransferase gene isolated from Hordeum vulgare is<br />
differentially regulated by drought stress. Theor. Appl. Genet. 113(6):965-976.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Richardson K.L., M.I. Vales, J.G. Kling, C.C. Mundt, and P.M. Hayes. 2006. Pyramiding<br />
and dissecting disease resistance QTL to barley stripe rust. Theor. Appl. Genet.<br />
113(3):485-495.<br />
Skinner J.S., P. Szucs, J. von Zitzewitz, L. Marquez-Cedillo, T. Filichkin, E.J. Stockinger,<br />
M.F. Thomashow, T.H. Chen, and P.M. Hayes. 2006. Mapping of barley homologs to<br />
genes that regulate low temperature tolerance in Arabidopsis. Theor. Appl. Genet.<br />
112(5):832-842.<br />
von Korff M., H. Wang, J. Léon, and K. Pillen. 2006. AB-QTL analysis in spring barley: II.<br />
Detection of favourable exotic alleles for agronomic traits introgressed from wild barley<br />
(H. vulgare ssp. spontaneum). Theor. Appl. Genet. 112(7):1221-1231.<br />
Walia H., C. Wilson, P. Condamine, A.M. Ismail, J. Xu, X. Cui, and T.J. Close. 2007.<br />
Array-based genotyping and expression analysis of barley cv. Maythorpe and Golden<br />
Promise. BMC Genomics. Mar 30; 8:87.<br />
Wang J., H. Raman, M. Zhou, P.R. Ryan, E. Delhaize, D.M. Hebb, N. Coombes, and N.<br />
Mendham. 2007. High-resolution mapping of the Alp locus and identification of a<br />
candidate gene HvMATE controlling aluminium tolerance in barley (Hordeum vulgare<br />
L.). Theor. Appl. Genet. Jun 6 [Epub ahead of print].<br />
Yan G.P. and X.M. Chen. Molecular mapping of a recessive gene for resistance to stripe rust in<br />
barley. Theor. Appl. Genet. 2006 113(3):529-537.<br />
Zhang L., T. Fetch, J. Nirmala, D. Schmierer, R. Brueggeman, B. Steffenson, and A.<br />
Kleinhofs. 2006. Rpr1, a gene required for Rpg1-dependent resistance to stem rust in<br />
barley. Theor. Appl. Genet. 113(5):847-855.<br />
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Coordinator’s Report: Chromosome 5H<br />
George Fedak<br />
Eastern Cereals and Oilseeds Research Centre<br />
Agriculture and Agri-Food Canada<br />
Ottawa ON K1A 0C6<br />
e-mail: fedakga@agr.gc.ca<br />
The grain hardness locus (Ha) of barley consists of a cluster of genes located on the short arm of<br />
5H designated as Hina, Hinb-1, Hinb-2 and GSP. Eighty diverse barley genotypes were screened<br />
for kernel hardness, ruminant digestibility and haplotypes of the four alleles. The highest level of<br />
genetic variation was obtained with GSP followed by Hina, Hinb-2. Hina was significantly<br />
related to grain hardness while Hinb-1 and Hinb-2 were significantly associated with dry water<br />
digestibility. (Turuspekov et al., 2007).<br />
Using the Nure (winter) x Tremois (spring) mapping population, two low temperature QTL were<br />
located on the long arm of chromosome 5H. FrHi was located in a distal position and Fr-H2 in a<br />
proximal location. The location of the latter coincided with the location of a QTL regulating the<br />
accumulation of two COR proteins; COR14b and TMC-Ap3. Six barley genes for the CBF<br />
transcription factor have been mapped in a single cluster in this region and they represent<br />
candidate genes for Fr-H2. (Francia et.al., 2007)<br />
In a related study, Lombda phage libraries were constructed from 2 spring (Morex and Tremois)<br />
and two winter (Dicktos, Nure) cultivars. Clones containing CBF genes were sequenced. It was<br />
found that the winter varieties have a large duplication at the Fr-H2 gene resulting in an increased<br />
number of CBF genes at this locus. The spring barley Tremois, however, has a significant<br />
deletion at this locus. This suggests that the relative numbers of CBF in the cluster contributes to<br />
different levels of winterhardiness (Knox et.al.,2007).<br />
References:<br />
Francia, E., D. Baraboschi, A. Tondelli, G. Laido, A.M. Stanca, E. Stockinger and N.<br />
Pecchioni, 2007. Fine mapping of a Hv CBF gene cluster, at Fr-H2, a QTL controlling<br />
frost resistance in barley. PAG - XV Poster 331. Plant and Animal Genomes XV<br />
Conference, Jan 13-17, 2007. San Diego CA.<br />
Knox, A.K., H. Chang . and E.I. Stockinger, 2007. Comparative Sequence analysis of CBF<br />
genes at the Fr-H2 locus in four barley cultivars. PAG – XV Poster 321. Plant and Animal<br />
Genomes XV Conference, Jan 13-17, 2007. San Diego CA.<br />
Turuspekov Y., B. Beecher, Y. Darlington, J. Bowman, T. Blake and M. Gioux.<br />
2007.Genetic variation of hardness locus in barley. PAG-XV Poster 329. Plant and<br />
Animal Genomes XV Conference, Jan 13-17, 2007, San Diego CA.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s Report: Chromosome 7H<br />
Lynn S. Dahleen<br />
<strong>US</strong>DA-Agricultural Research Service<br />
Fargo, ND 58105, <strong>US</strong>A<br />
e-mail: DAHLEEN@fargo.ars.usda.gov<br />
Barley gene mapping in 2006 showed a greater emphasis on using candidate gene approaches in<br />
addition to standard qualitative and quantitative trait mapping. Increased use of public EST and<br />
BAC libraries was evident, providing tools to better understand the barley genome.<br />
Efforts to map morphological genes have continued. Roder et al. (2006) mapped the shrunken<br />
endosperm gene seg8 to a 4.6 cM interval near the centromere of chromosome 7H, while Taketa<br />
et al. (2006) developed a fine map of the naked caryopsis nud locus, placing it in a 0.66 cM<br />
region. Rostoks et al. (2006) found that the barley homolog of the Arabidopsis HLM1 gene<br />
corresponded to the nec1 locus on chromosome 7H. Allelic variation was uncovered at the locus<br />
that causes necrotic spotting of nec1 plants. Rossini et al. (2006) examined candidate rice genes<br />
in regions syntenous with markers linked to various barley morphological mutants. On<br />
chromosome 7H they found brh1 candidate genes on rice chromosome 6 and candidates for<br />
suKF-76, suKE-74 (suppressors of Hooded) and sld4 on rice chromosomes 6 and 8. The low<br />
resolution of the barley maps in this region resulted in selection of rather large rice regions and<br />
numerous candidate genes. Yan et al. (2006) identified the AtFT flowering locus as an ortholog<br />
of the barley and wheat vernalization gene VRN3. The barley gene VRN-H3 was located on the<br />
short arm of chromosome 7H, not chromosome 1H as previously thought based on loose linkage<br />
with BLP. Szucs et al. (2006) mapped genes for photoreceptor gene families and vernalization<br />
regulation, and compared their locations to QTL for photoperiod response. The barley ortholog<br />
to a wheat flowering repressor, HvVRT-2 mapped to the short arm of chromosome 7H. This<br />
locus coincided with a photoperiod QTL with small effects mapped in the Dicktoo x Morex<br />
population. Tondelli et al. (2006), using a similar approach, mapped candidate genes for cold or<br />
drought response based on sequences identified in other plants. Two orthologs of Arabidopsis<br />
genes (AtFRY1 and AtICE1) that have a prominent role in cold acclimation were identified on<br />
chromosome 7H.<br />
QTL analyses for a variety of traits were reported this year. Chloupek et al. (2006) mapped root<br />
system size traits in a population segregating for two semidwarf genes, sdw1 and ari-e.GP. On<br />
chromosome 7H, they identified a region associated with height, and another region associated<br />
with harvest index, plant weight, root system size at grain filling and total root system size.<br />
Advanced backcross QTL analysis continued, with von Korff et al. (2006) detecting favorable<br />
alleles from wild barley in crosses with Scarlett. Out of the 86 QTL identified for 9 traits, the H.<br />
spontaneum alleles improved performance for 31. QTL for height, heading date, harvest index,<br />
lodging at flowering, vegetative dry biomass, thousand grain weight, brittleness and yield were<br />
located on chromosome 7H. Li et al. (2006), in a similar study of a wild barley x Brenda<br />
advanced backcross, found 100 QTL. Chromosome 7H QTL included yield, heading date,<br />
height, ear length, spikelets per spike, seed per spike, spikes per plant, thousand grain weight,<br />
leaf length, and leaf area loci. Yun et al. (2006) also used advanced backcross lines from a cross<br />
of H. spontaneum with Harrington to validate QTLs for disease resistance loci. A QTL for spot<br />
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blotch resistance previously identified in a RIL population was confirmed to be located on<br />
chromosome 7H.<br />
Additional disease resistance genes were located in several studies. Cheong et al. (2006) located<br />
a QTL for adult plant resistance to leaf scald using two populations. One of these QTL was<br />
located on the short arm of chromosome 7H. Rossi et al. (2006) located QTL for barley strip rust<br />
and leaf rust resistance plus a powdery mildew resistance QTL on chromosome 7H. Jafary et al.<br />
(2006) located genes involved in nonhost immunity to rust pathogens, including four QTL on<br />
chromosome 7H controlling reactions to seven rust species. Brueggeman et al. (2006) identified<br />
five additional members of the Rpg1 gene family, including one that is closely linked to Rpg1 on<br />
chromosome 7H.<br />
Kilian et al. (2006) examine haplotype structure at seven loci, including the Adh3 and Waxy loci<br />
on chromosome 7H, to compare sequence diversity between 20 domesticated and 25 wild<br />
barleys. As expected, more haplotypes were identified in the wild barley than the domesticated<br />
barley. At Adh3, wild barley showed 15 haplotypes while domesticated barley had three and at<br />
Waxy, the wilds had 17 haplotypes compared to 4 in the domesticated barley. This diversity was<br />
also evident in nucleotide sequence, with more polymorphic sites in the wild barley than in the<br />
domesticated barley. Pickering et al. (2006) examined associations between chromosomes in two<br />
H. vulgare x H. bulbosum hybrids. Chromosome 7HS-7H b S associations were higher than the<br />
average for other chromosome arms in both hybrids examined.<br />
References.<br />
Brueggeman R, T Drader and A Kleinhofs. 2006. The barley serine/threonine kinase gene<br />
Rpg1 providing resistance to stem rust belongs to a gene family with five other members<br />
encoding kinase domains. Theor Appl Genet 113:1147-1158.<br />
Cheong J, K Williams and H Wallwork. 2006. The identification of QTLs for adult plant<br />
resistance to leaf scald in barley. Austral J Agric Res 57:961-965.<br />
Chloupek O, BP Forster and WTB Thomas. 2006. The effect of semi-dwarf genes on root<br />
system size in field-grown barley. Theor Appl Genet 112:779-786.<br />
Jafary H, LJ Szabo and RE Niks. 2006. Innate nonhost immunity in barley to different<br />
heterologous rust fungi is controlled by sets of resistance genes with different and<br />
overlapping specificities. MPMI 19:1270-1279.<br />
Kilian B, H Ozkan, J Kohl, A von Haeseler, F Borale, O Deusch, A Brandolini, C Yucel, W<br />
Martin and F Salamini. 2006. Haplotype structure at seven barley genes: relevance to<br />
gene pool bottlenecks, phylogeny of ear type and site of barley domestication. Mol Gen<br />
Genomics 276:230-241.<br />
Korff M von, H Wang, J Leon and K Pillen. 2006. AB-QTL analysis in spring barley: II.<br />
Detection of favourable exotic alleles for agronomic traits introgressed from wild barley<br />
(H. vulgare ssp. spontaneum). Theor Appl Genet 112:1221-1231.<br />
Li JZ, XQ Huang, F Heinrichs, MW Ganal and MS Roder. 2006. Analysis of QTLs for yield<br />
components, agronomic traits, and disease resistance in an advanced backcross<br />
population of spring barley. Genome 49:454-466.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Pickering R, S Klatte and RC Butler. 2006. Identification of all chromosome arms and their<br />
involvement in meiotic homoeologous associations at metaphase 1 in 2 Hordeum vulgare<br />
L. x Hordeum bulbosum L. hybrids. Genome 49:73-78.<br />
Roder MS, C Kaiser and W Weschke. 2006. Molecular mapping of the shrunken endosperm<br />
genes seg8 and sex1 in barley (Hordeum vulgare L.). Genome 49:1209-1214.<br />
Rossi C, A Cuesta-Marcos, I Vales, L Gomez-Pando, G Orjeda, R Wise, K Sato, K Hori, F<br />
Capettini, H Vivar, X Chen and P Hayes. 2006. Mapping multiple disease resistance<br />
genes using a barley mapping population evaluated in Peru, Mexico, and the <strong>US</strong>A. Mol<br />
Breed 18:355-366.<br />
Rossini L, A Vecchietti, L Nicoloso, N Stein, S Franzago, F Salamini and C Pozzi. 2006.<br />
Candidate genes for barley mutants involved in plant architecture: an in silico approach.<br />
Theor Appl Genet 112:1073-1085.<br />
Rostoks N, D Schmierer, S Mudie, T Drader, R Brueggeman, DG Caldwell, R Waugh and<br />
A Kleinhofs. 2006. Barley necrotic locus nec1 encodes the cyclic nucleotide-gated ion<br />
channel 4 homologous to the Arabidopsis HLM1. Mol Gen Genomics 275:159-168.<br />
Szucs P, I Karsai, J von Zitzewitz, K Meszarus, LLD Cooper, YQ Gu, THH Chen, PM<br />
Hayes and JS Skinner. 2006. Positional relationships between photoperiod response<br />
QTL and photoreceptor and vernalization genes in barley. Theor Appl Genet 112:1277-<br />
1285.<br />
Taketa S, T Awayama, S Amano, Y Sakurai and M Ichii. 2006. High-resolution mapping of<br />
the nud locus controlling the naked caryopsis in barley. Plant Breed 125:337-342.<br />
Tondelli A, E Francia, D Barbaschi, A Aprile, JS Skinner, EJ Stockinger, AM Stanca and<br />
N Pecchioni. 2006. Mapping regulatory genes as candidates for cold and drought stress<br />
tolerance in barley. Theor Appl Genet 112:445-454.<br />
Yan L, D Fu, C Li, A Blechl, G Tranquilli, M Bonafede, A Sanchez, M Valarik, S Yasuda<br />
and J Dubcovsky. 2006. The wheat and barley vernalization gene VRN3 is an orthologue<br />
of FT. PNAS 103:19581-19586.<br />
Yun SJ, L Gyenis, E Bossolini, PM Hayes, I Matus, KP Smith, BJ Steffenson, R Tuberosa<br />
and GJ Muehlbauer. 2006. Validation of quantitative trait loci for multiple disease<br />
resistance in barley using advanced backcross lines developed with a wild barley. Crop<br />
Sci 46:1179-1186.<br />
124
Barley Genetics Newsletter (2007) 37: 105-153<br />
Integrating Molecular and Morphological/Physiological Marker Maps<br />
A. Kleinhofs<br />
Dept. Crop and Soil Sciences and<br />
School of Molecular Biosciences<br />
Washington State University<br />
Pullman, WA 99164-6420, <strong>US</strong>A<br />
e-mail: andyk@wsu.edu<br />
There has been only limited progress in mapping morphological markers during the past year.<br />
However, there is a major effort under way in Europe to map the morphological mutant isolines<br />
developed by Jerry Franckowiak (for details see<br />
http://www.smallgraincereals.org/SGCNewsletterSummer2007.pdf). This effort, when<br />
completed, should provide a map of 1000 morphological markers with accurate reference to<br />
molecular markers.<br />
A recessive barley stripe rust resistance gene rpsGZ (from Grannenlose Zweizeilige) was<br />
mapped to chromosome 4H bin9 (Yan and Chen, 2006). This gene cosegregated with several<br />
RGAP markers identified in the publication complete with primer information. It is also closely<br />
linked to SSR markers EBmac0679 and EBmac0701.<br />
Septoria speckled leaf blotch resistance genes Rsp1, Rsp2, and Rsp3 were mapped (Lee and<br />
Neate, 2007a). Rsp2 cosegregated with MWG938 placing it on chromosome 5(1H) bin2. Rsp3<br />
was closely linked to Rsp2 on chromosome 5S(1HS). Rsp2 was flanked by RAPD markers<br />
OPBA12314C and OPB17451R at 2.4 and 3.5 cM, respectively. I was not able to locate these to<br />
a specific bin. Rsp1 was mapped to chromosome 3H short arm flanked by RAPD markers<br />
OPC2441R (3.0 cM) and UBC285158R (4.3 cM). In addition closely linked DArT markers were<br />
also identified. Specific bin location was not possible. The RAPD markers were converted into<br />
sequence tagged markers and primer sequences published (Lee and Neate, 2007b).<br />
Aluminum tolerance gene Alp was mapped to chromosome 4H bin7 closely linked to ABG715<br />
and cosegregating with several markers including HvMATE (AV942930) which was proposed<br />
as a candidate gene for the Alp locus (Wang et al., 2007)<br />
Five barley flowering locus T-like (FT-like) genes were mapped (Faure et al., 2007). HvFT1<br />
maps on chromosome 1(7H) between markers AF022725 and Bmac31 closely linked to VRN-H3<br />
and ABC158 placing it in bin4. The VRN-H3 gene was previously believed to be located on<br />
chromosome 5(1H), but more recently shown to be on chromosome 1(7H) (Yan et al., 2006).<br />
HvFT2 was mapped on chromosome 3H between markers Bmac067 and MWG985 placing it in<br />
bin6. HvHT3 was mapped to chromosome 7(5H) cosegregating with PSR162 placing it in bin11.<br />
HvFT4 was mapped to the short arm of chromosome 2H proximal to cMWG663 placing it in<br />
bin6. HvHT5 was mapped to chromosome 4H long arm cosegregating with scsnp20989. It was<br />
not possible for me to determine the bin placement.<br />
125
Barley Genetics Newsletter (2007) 37: 105-153<br />
The naked caryopsis gene (nud) has been previously mapped to chromosome 1(7H) bin 7. It has<br />
now been mapped at a high resolution (Taketa et al., 2006). The closest SCAR marker SKT9,<br />
mapped 0.06 cM from the nud locus based on 4,760 gametes from 6 mapping populations.<br />
Barley lipoxygenase (Lox-1) thermostability factor was shown to cosegregate with the structural<br />
gene LoxA (Hirota et al., 2006). The LoxA and LoxC loci were previously mapped to<br />
chromosome 4H bin3 and chromosome 7(5H) probably bin 10 (van Mechelen et al., 1999).<br />
Barley homologs of large number of Arabidopsis low temperature regulatory genes were mapped<br />
assigning either linkage map or chromosome locations to 1 ICE1, 2 ZAT12 and 17 CBF<br />
homologs (Skinner et al., 2006). Eleven of the CBF genes with assigned linkage map positions<br />
formed two tandem clusters on 5HL(7L). These were coincident with reported Triticeae low<br />
temperature tolerance and CO R gene accumulation QTL and suggest that one or more of the<br />
CBF genes may be candidates for Fr-H2 QTL.<br />
Bin Assignments for Morphological Map Markers and closest molecular marker<br />
* - indicates that the gene has been cloned<br />
red - indicates that the gene is very accurately mapped with molecular markers<br />
yellow - indicates that it is fairly accurately mapped with molecular markers<br />
blue - indicates that the gene has been approximately mapped mainly using Bulked<br />
Segregant Analysis<br />
Chr.1(7H)<br />
BIN1 ABG704<br />
*Rpg1 RSB228 Brueggeman et al., PNAS 99:9328, ‘02<br />
Run1<br />
Rdg2a MWG851A Bulgarelli et al., TAG 108:1401, ‘04<br />
Rrs2 MWG555A Schweizer et al., TAG 90:920, ‘95<br />
mlt<br />
brh1 MWG2074B Li et al., 8 th IBGS 3:72, ‘00<br />
BIN2 ABG320<br />
Est5 iEst5 Kleinhofs et al., TAG 86:705, ‘93<br />
fch12 BCD130 Schmierer et al., BGN 31:12, ‘01<br />
*wax Wax Kleinhofs BGN 32:152, ‘02<br />
gsh3 His3A Kleinhofs BGN 32:152, ‘02<br />
BIN3 ABC151A<br />
fch5 ABC167A Kleinhofs BGN 32:152, ‘02<br />
Rcs5 KAJ185 Johnson & Kleinhofs, unpublished<br />
yvs2<br />
cer-ze ABG380 Kleinhofs BGN 27:105, ‘96<br />
BIN4 ABG380<br />
wnd<br />
*HvFT1 ABC158 Faure et al., Genetics 176:599, '07<br />
VrnH3 ABC158 Yan et al., PNAS 103:19581, '06<br />
Lga BE193581 Johnson & Kleinhofs, unpublished<br />
abo7<br />
126
Barley Genetics Newsletter (2007) 37: 105-153<br />
BIN5 ksuA1A<br />
ant1<br />
nar3 MWG836 Kleinhofs BGN 32:152, ‘02<br />
ert-m<br />
ert-a<br />
BIN6 ABC255<br />
ert-d<br />
fch8<br />
fst3<br />
cer-f<br />
msg14<br />
BIN7 ABG701<br />
dsp1 cMWG704 Sameri & Komatsuda JARQ 41:195, '07<br />
msg10<br />
rsm1 ABC455 Edwards & Steffenson, Phytopath. 86:184,’96<br />
sex6<br />
seg5<br />
seg2<br />
pmr ABC308 Kleinhofs BGN 27:105, ‘96<br />
mo6b Hsp17 Soule et al., J Her. 91:483, ‘00<br />
nud sKT9 Taketa et al., Plant Breeding 125:337, '06<br />
fch4 MWG003 Kleinhofs BGN 27:105, ‘96<br />
BIN8 *Amy2 Amy2 Kleinhofs et al., TAG 86:705, ‘93<br />
lks2 WG380B Costa et al., TAG 103:415, ‘01<br />
ubs4<br />
blx2<br />
BIN9 RZ242<br />
lbi3<br />
Rpt4 Psr117D Williams et al., TAG 99:323, ‘99<br />
xnt4<br />
lpa2 ? Larson et al., TAG 97:141, ‘98<br />
msg50<br />
Rym2<br />
seg4<br />
BIN10 ABC310B<br />
Xnt1 BF626025 Hansson et al., PNAS 96:1744, ‘99<br />
xnt-h BF626025 Hansson et al., PNAS 96:1744, ‘99<br />
BIN11 ABC305<br />
Rph3<br />
Tha2 Toojinda et al., TAG 101:580, ‘00<br />
BIN12 ABG461A<br />
Mlf<br />
xnt9<br />
seg1<br />
msg23<br />
BIN13 Tha<br />
Rph19 Rlch4(Nc) Park & Karakousis Plt. Breed. 121:232. ‘02<br />
127
Barley Genetics Newsletter (2007) 37: 105-153<br />
Chr.2(2H)<br />
BIN1 MWG844A<br />
sbk<br />
brh3 Bmac0134 Dahleen et al., J. Heredity 96:654, ‘05<br />
BIN2 ABG703B<br />
BIN3 MWG878A gsh6 Kleinhofs BGN 32:152, ‘02<br />
gsh1<br />
gsh8<br />
BIN4 ABG318<br />
Eam1<br />
Ppd-H1 MWG858 Laurie et al., Heredity 72:619, ‘94<br />
sld2<br />
rtt<br />
flo-c<br />
sld4<br />
BIN5 ABG358<br />
fch15<br />
brc1<br />
com2<br />
BIN6 Pox<br />
msg9<br />
abo2<br />
Rph15 P13M40 Weerasena et al., TAG 108:712 ‘04<br />
rph16 MWG874 Drescher et al., 8thIBGS II:95, ‘00<br />
BIN7 Bgq60<br />
yst4 CDO537 Kleinhofs BGN 32:152, ‘02<br />
Az94 CDO537 Kleinhofs BGN 32:152, ‘02<br />
gai MWG2058 Börner et al., TAG 99:670, ‘99<br />
msg33<br />
*HvCslF (barley Cellulose synthase-like) Burton et al., Science 311:1940 ‘06<br />
*Bmy2<br />
msg3<br />
fch1<br />
BIN8 ABC468<br />
Eam6 ABC167b Tohno-oka et al., 8thIBGS III:239, ‘00<br />
gsh5<br />
msg2<br />
eog ABC451 Kleinhofs BGN 27:105, ‘96<br />
abr<br />
cer-n<br />
BIN9 ABC451<br />
Gth<br />
hcm1<br />
wst4<br />
*vrs1 MWG699 Komatsuda et al., Genome 42:248, ‘00<br />
BIN10 MWG865<br />
cer-g<br />
Lks1<br />
mtt4<br />
Pre2<br />
128
Barley Genetics Newsletter (2007) 37: 105-153<br />
Chr. 3(3H)<br />
msg27<br />
BIN11 MWG503<br />
Rha2 AWBMA21 Kretschmer et al., TAG 94:1060, ‘97<br />
Ant2 MWG087 Freialdenhoven et al., Plt. Cell 6:983, ’94<br />
*Rar1 AW983293B Freialdenhoven et al., Plt. Cell 6:983, ’94<br />
fol-a<br />
gal MWG581A Börner et al., TAG 99:670, ‘99<br />
fch14<br />
Pau<br />
BIN12 ksuD22<br />
Pvc<br />
BIN13 ABC252<br />
lig BCD266 Pratchett & Laurie Hereditas 120:35, ‘94<br />
nar4 Gln2 Kleinhofs BGN 27:105, ‘96<br />
Zeo1 cnx1 Costa et al., TAG 103:415, ‘01<br />
lpa1 ABC157 Larson et al., TAG 97:141, ‘98<br />
BIN14 ABC165<br />
BIN15 MWG844B<br />
gpa CDO036 Kleinhofs BGN 27:105, ‘96<br />
wst7 MWG949A Costa et al., TAG 103:415, ‘01<br />
MlLa Ris16 Giese et al., TAG 85:897, ‘93<br />
trp<br />
BIN1 Rph5 ABG070 Mammadov et al., TAG 111:1651, ‘05<br />
Rph6 BCD907 Zhong et al., Phytopath. 93:604, ‘03<br />
Rph7 MWG848 Brunner et al., TAG 101:783, ‘00<br />
BIN2 JS195F BI958652; BF631357; BG369659<br />
ant17<br />
sld5<br />
mo7a ABC171A Soule et al., J. Hered. 91:483, ‘00<br />
brh8<br />
BIN3 ABG321<br />
xnt6<br />
BIN4 MWG798B<br />
btr1 Senthil & Komatsuda Euphytica 145:215, ‘05<br />
btr2 Senthil & Komatsuda Euphytica 145:215, ‘05<br />
lzd<br />
alm ABG471 Kleinhofs BGNL 27:105, ‘96<br />
BIN5 BCD1532<br />
abo9<br />
sca<br />
yst2<br />
dsp10<br />
BIN6 ABG396<br />
Rrs1 Graner et al., TAG 93: 421 ´96<br />
Rh/Pt ABG396 Smilde et al., 8th IBGS 2:178, ‘00<br />
Rrs.B87 BCD828 Williams et al., Plant Breed. 120:301, ‘01<br />
129
Barley Genetics Newsletter (2007) 37: 105-153<br />
AtpbB<br />
abo6<br />
xnt3<br />
HvHT2 Bmac067 Faure et al., Genetics 176:599, '07<br />
msg5<br />
ari-a<br />
yst1<br />
zeb1<br />
ert-c<br />
ert-ii<br />
cer-zd<br />
Ryd2 WG889B Collins et al., TAG 92:858, ‘96<br />
*uzu AB088206 Saisho et al., Breeding Sci. 54:409, ‘04<br />
BIN7 MWG571B<br />
cer-r<br />
BIN8 ABG377<br />
wst6<br />
cer-zn<br />
sld1<br />
BIN9 ABG453<br />
wst1<br />
BIN10 CDO345<br />
vrs4<br />
Int1<br />
gsh2<br />
BIN11 CDO113B<br />
als<br />
sdw1 PSR170 Laurie et al., Plant Breed. 111:198, ‘93<br />
BIN12 His4B<br />
sdw2<br />
BIN13 ABG004<br />
Pub ABG389 Kleinhofs et al., TAG 86:705, ‘93<br />
BIN14 ABC161<br />
cur2<br />
BIN15 ABC174<br />
Rph10<br />
fch2<br />
BIN16 ABC166<br />
eam10<br />
Est1/2/3<br />
*rym4 eIF4E Stein et al.,Plt. J. 42:912, ‘05<br />
*rym5 eIF4E and Kanyuka et al., Mol. Plant Path. 6:449, ’05<br />
Est4<br />
ant28<br />
Chr.4(4H)<br />
BIN1 MWG634<br />
BIN2 JS103.3<br />
fch9<br />
130
Barley Genetics Newsletter (2007) 37: 105-153<br />
sln<br />
BIN3 Ole1<br />
Dwf2 Ivandic et al., TAG 98:728, ‘99<br />
*LoxA MWG011b van Mechelen et al., Plt. Mol. Biol. 39:1283, '99<br />
LoxB van Mechelen et al., Plt. Mol. Biol. 39:1283, '99<br />
Lox-1 thermo Hirota et al., Plant breeding 125:231, '06<br />
Ynd<br />
int-c MWG2033 Komatsuda, TAG 105:85, ‘02<br />
Zeo3<br />
glo-a<br />
rym1 MWG2134 Okada et al., Breeding Sci. 54:319, ‘04<br />
BIN4 BCD402B<br />
*Kap X83518 Müller et al., Nature 374:727, ‘95<br />
lbi2<br />
zeb2<br />
lgn3<br />
BIN5 BCD808B<br />
lgn4<br />
lks5<br />
eam9<br />
msg24<br />
BIN6 ABG484<br />
glf1<br />
rym11 MWG2134 Bauer et al., TAG 95:1263, ‘97<br />
Mlg MWG032 Kurth et al., TAG 102:53, ‘01<br />
cer-zg<br />
brh2<br />
BIN7 bBE54A<br />
glf3<br />
Alp HvMATE Wang et al., TAG 115:265 '07<br />
frp<br />
min1<br />
blx4<br />
sid<br />
blx3<br />
BIN8 BCD453B<br />
blx1<br />
BIN9 ABG319A<br />
ert1<br />
rpsGZ EBmac0679 Yan & Chen, TAG 113:529, '06<br />
BIN10 KFP221<br />
*mlo P93766 Bueschges et al., Cell 88:695, ‘97<br />
BIN11 ABG397<br />
BIN12 ABG319C<br />
Hsh HVM067 Costa et al., TAG 103:415, ‘01<br />
Hln<br />
*sgh1(ZCCT-H; HvSnf2) Zitzewitz et al., PMB 59:449, ‘05<br />
yhd1<br />
BIN13 *Bmy1 pcbC51 Kleinhofs et al., TAG 86:705, ‘93<br />
rym8 MWG2307 Bauer et al., TAG 95:1263, ‘97<br />
131
Barley Genetics Newsletter (2007) 37: 105-153<br />
rym9 MWG517 Bauer et al., TAG 95:1263, ‘97<br />
Wsp3<br />
Chr. 5(1H)<br />
BIN1 Tel5P<br />
Rph4<br />
Mlra<br />
Cer-yy<br />
Sex76 Hor2 Netsvetaev BGN 27:51, ‘97<br />
*Hor5 Hor5 Kleinhofs et al., TAG 86:705, ‘93<br />
BIN2 MWG938<br />
Rsp2 MWG938 Lee & Neate, Phytopath. 97:155, '07<br />
*Hor2 Hor2 Kleinhofs et al., TAG 86:705, ‘93<br />
Rrs14 Hor2 Garvin et al., Plant Breed. 119:193-196, ‘00<br />
*Mla6 AJ302292 Halterman et al., Plt J. 25:335, ‘01<br />
BIN3 MWG837<br />
*Hor1 Hor1 Kleinhofs et al., TAG 86:705, ‘93<br />
Rps4<br />
Mlk<br />
BIN4 ABA004<br />
Lys4<br />
BIN5 BCD098<br />
Mlnn;<br />
msg31;<br />
sls;<br />
msg4;<br />
fch3;<br />
BIN6 Ica1<br />
amo1<br />
BIN7 JS074<br />
clh<br />
vrs3<br />
Ror1 ABG452 Collins et al., Plt. Phys. 125:1236, ‘01<br />
BIN8 Pcr2<br />
fst2<br />
cer-zi<br />
cer-e<br />
ert-b<br />
MlGa<br />
msg1<br />
xnt7<br />
BIN9 Glb1<br />
*nec1 BF630384 Rostoks et al., MGG 275:159, '06<br />
BIN10 DAK123B<br />
abo1<br />
Glb1<br />
BIN11 PSR330<br />
*HvFT3 PSR162 Faure et al., Genetics 176:599, '07<br />
PpdH2 PSR162 Laurie et al., Genome 38:575, '95<br />
wst5<br />
132
Barley Genetics Newsletter (2007) 37: 105-153<br />
cud2<br />
BIN12 MWG706A<br />
rlv<br />
lel1<br />
BIN13 BCD1930<br />
Blp ABC261 Costa et al., TAG 103:415, ‘01<br />
BIN14 ABC261<br />
fch7<br />
trd<br />
eam8<br />
Chr. 6(6H)<br />
BIN1 ABG062<br />
*Nar1 X57845 Kleinhofs et al., TAG 86:705, ‘93<br />
abo15<br />
BIN2 ABG378B<br />
nar8 ABG378B Kleinhofs BGN 27:105, ‘96<br />
nec3<br />
Rrs13<br />
BIN3 MWG652A<br />
BIN4 DD1.1C<br />
msg36<br />
BIN5 ABG387B<br />
nec2<br />
ant21<br />
msg6<br />
eam7<br />
BIN6 Ldh1<br />
rob HVM031 Costa et al., TAG 103:415, ‘01<br />
sex1<br />
gsh4<br />
ant13<br />
cul2 Crg4(KFP128) Babb & Muehlbauer BGN 31:28, ‘01<br />
fch11<br />
mtt5<br />
abo14<br />
BIN7 ABG474<br />
BIN8 ABC170B<br />
BIN9 *Nar7 X60173 Warner et al., Genome 38:743, ‘95<br />
*Amy1 JR115 Kleinhofs et al., TAG 86:705, ‘93<br />
*Nir pCIB808 Kleinhofs et al., TAG 86:705, ‘93<br />
mul2<br />
cur3<br />
BIN10 MWG934<br />
lax-b<br />
raw5<br />
cur1<br />
BIN11 Tef1<br />
BIN12 xnt5<br />
133
Barley Genetics Newsletter (2007) 37: 105-153<br />
Aat2<br />
BIN13 Rph11 Acp3 Feuerstein et al., Plant breed. 104:318, ‘90<br />
lax-c<br />
BIN14 DAK213C<br />
dsp9<br />
Chr. 7(5H)<br />
BIN1 DAK133<br />
abo12<br />
msg16<br />
ddt<br />
BIN2 MWG920.1A<br />
dex1<br />
msg19<br />
nld<br />
fch6<br />
glo-b<br />
BIN3 cud1 ABG705A<br />
lys3<br />
fst1<br />
blf1<br />
vrs2<br />
BIN4 ABG395<br />
cer-zj<br />
cer-zp<br />
msg18<br />
wst2<br />
Rph2 ITS1 Borovkova et al., Genome 40:236, ‘97<br />
lax-a PSR118 Laurie et al., TAG 93:81, ‘96<br />
com1<br />
ari-e<br />
ert-g<br />
ert-n<br />
BIN5 Ltp1<br />
rym3 MWG028 Saeki et al., TAG 99:727, ‘99<br />
BIN6 WG530<br />
BIN7 ABC324<br />
BIN8 ABC302A<br />
BIN9 BCD926<br />
srh ksuA1B Kleinhofs et al., TAG 86:705, ‘93<br />
cer-i<br />
mtt2<br />
lys1<br />
cer-t<br />
dsk<br />
var1<br />
cer-w<br />
Eam5<br />
134
Barley Genetics Newsletter (2007) 37: 105-153<br />
BIN10 ABG473<br />
raw1<br />
msg7<br />
BIN11 MWG514B<br />
Rph9/12ABG712 Borokova et al., Phytopath. 88:76, ‘98<br />
*Sgh2 (HvBM5A) Zitzewitz et al., PMB 59:449, ‘05<br />
*Ror2 AY246906 Collins et al., Nature 425:973, ‘03<br />
lbi1<br />
Rha4<br />
raw2<br />
BIN12 WG908<br />
none<br />
BIN13 ABG496<br />
rpg4 ARD5303 Druka et al., unpublished<br />
RpgQ ARD5304 Druka et al., unpublished<br />
BIN14 ABG390<br />
var3<br />
BIN15 ABG463<br />
135
Barley Genetics Newsletter (2007) 37: 105-153<br />
References:<br />
Babb, S.L. and G.J. Muehlbauer. 2001. Map location of the Barley Tillering Mutant uniculm2<br />
(cul2) on Chromosome 6H. BGN31:28.<br />
Bauer, E., J. Weyen, A. Schiemann, A. Graner, and F. Ordon. 1997. Molecular mapping of<br />
novel resistance genes against Barley Mild Mosaic Virus (BaMMV). Theor. Appl. Genet.<br />
95:1263-1269.<br />
Borovkova, I.G., Y. Jin, B.J. Steffenson, A. Kilian, T.K. Blake, and A. Kleinhofs. 1997.<br />
Identification and mapping of a leaf rust resistance gene in barley line Q21861. Genome<br />
40:236-241.<br />
Borokova, I.G., Y. Jin, and B.J. Steffenson. 1998. Chromosomal Location and Genetic<br />
Relationship of Leaf Rust Resistance Genes Rph9 and Rph12 in Barley. Phytopathology<br />
88:76-80.<br />
Börner, A., V. Korzun, S. Malyshev, V. Ivandic, and A. Graner. 1999. Molecular mapping of<br />
two dwarfing genes differing in their GA response on chromosome 2H of barley. Theor.<br />
Appl. Genet. 99:670-675.<br />
Brueggeman, R., N. Rostoks, D. Kudrna, A. Kilian, F. Han, J. Chen, A. Druka, B.<br />
Steffenson, and A. Kleinhofs. 2002. The barley stem-rust resistance gene Rpg1 is a<br />
novel disease-resistance gene with homology to receptor kinases. Proc. Natl. Acad. Sci.<br />
<strong>US</strong>A 99:9328-9333.<br />
Brunner, S., B. Keller, and C. Feuillet. 2000. Molecular mapping of the Rph7.g leaf rust<br />
resistance gene in barley (Hordeum vulgare L.). Theor. Appl. Genet. 101:763-788.<br />
Büschges, R., K. Hollricher, R. Panstruga, G. Simons, M. Wolter, A. Frijters, R. van<br />
Daelen, T. van der Lee, P. Diergaarde, J. Groenendijk, S. Töpsch, P. Vos, F.<br />
Salamini, and P. Schulze-Lefert. 1997. The barley mlo gene: A novel control element<br />
of plant pathogen resistance. Cell 88:695-705.<br />
Bulgarelli, D., N.C. Collins, G. Tacconi, E. Dellaglio, R. Brueggeman, A. Kleinhofs, A.M.<br />
Stanca, and G. Vale. 2004. High resolution genetic mapping of the leaf stripe resistance<br />
gene Rdg2a in barley. Theor. Appl. Genet. 108:1401-1408.<br />
Burton, R.A., S.M. Wilson, M. Hrmova, A.J. Harvey, N.J. Shirley, A. Medhurst, B.A.<br />
Stone, E.J. Newbigin, A. Bacic, and G.B. Fincher. 2006. Cellulose synthase-like CslF<br />
genes mediate the synthesis of cell wall (1,3; 1,4)-b-D-glucans. Science 311:1940-1943.<br />
Collins, N.C., N.G. Paltridge, C.M. Ford, and R.H. Symons.1996. The Yd2 gene for barley<br />
yellow dwarf virus resistance maps close to the centromere on the long arm of barley<br />
chromosome 3. Theor. Appl. Genet. 92:858-864.<br />
Collins, N.C., T. Lahaye, C. Peterhänsel, A. Freialdenhoven, M. Corbitt, and P. Schulze-<br />
Lefert. 2001. Sequence haplotypes revealed by sequence-tagged site fine mapping of the<br />
Ror1 gene in the centromeric region of barley chromosome 1H. Plant Physiology<br />
125:1236-1247.<br />
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Collins, N.C., H. Thordal-Christensen, V. Lipka, S. Bau, E. Kombrink, J-L. Qiu, R.<br />
Hückelhoven, N. Stein, A. Freialdenhoven, S.C. Somerville, and P. Schulze-Lefert.<br />
2003. Snare-protein-mediated disease resistance at the plant cell wall. Nature 425:973-<br />
976.<br />
Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch, S.F. Kramer,<br />
D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda, M.I. Vales,<br />
and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe Barleys: a<br />
phenotypically polymorphic doubled-haploid population. Theor. Appl. Genet. 103:415-<br />
424.<br />
Drescher, A., V. Ivandic, U. Walther, and A. Graner. 2000. High-resolution mapping of the<br />
Rph16 locus in barley. p. 95-97. In: S. Logue (ed.) Barley Genetics VIII. Volume II.<br />
Proc. Eigth Int. Barley Genet. Symp. Adelaide. Dept. Plant Science, Waite Campus,<br />
Adelaide University, Glen Osmond, South Australia.<br />
Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization and molecular<br />
mapping of genes determining semidwarfism in barley. J. Heredity 96:654-662.<br />
Edwards, M.C. and B.J. Steffenson. 1996. Genetics and mapping of barley stripe mosaic virus<br />
resistance in barley. Phytopath. 86;184-187.<br />
Faure S, J. Higgins, A. Turner, and D.A. Laurie. 2007. The FLOWERING LOC<strong>US</strong> T-like<br />
gene family in barley (Hordeum vulgare). Genetics 176:599-609.<br />
Feuerstein, U., A.H.D. Brown, and J.J. Burdon. 1990. Linkage of rust resistance genes from<br />
wild barley (Hordeum spontaneum) with isoenzyme markers. Plant. Breed. 104:318-324.<br />
Freialdenhoven, A., B. Scherag, K. Hollrichter, D.B. Collinge, H. Thordal-Christensen, and<br />
P. Schulze-Lefert. 1994. Nar-1 and Nar-2, two loci required for Mla12-specified racespecific<br />
resistance to powdery mildew in barley. Plant Cell 6:983-994.<br />
Garvin, D.F., A.H.D. Brown, H. Raman, and B.J. Read. 2000. Genetic mapping of the barley<br />
Rrs14 scald resistance gene with RLFP, isozyme and seed storage protein marker. Plant<br />
Breeding 119:193-196.<br />
Giese, H., A.G. Holm-Jensen, H.P. Jensen, and J. Jensen. 1993. Localisation of the<br />
Laevigatum powdery mildew resistance gene to barley chromosome 2 by the use of<br />
RLFP markers. Theor. Appl. Genet. 85:897-900.<br />
Graner, A. and A. Tekauz. 1996. RFLP mapping in barley of a dominant gene conferring to<br />
scald (Rynchosporium secalis). Theor. Appl. Genet. 93:421-425.<br />
Haltermann, D., F. Zhou, F. Wei, R.P. Wise, and P. Schulze-Lefert. 2001. The MLA6 coiled<br />
coil, NBS-LRR protein confers AvrMla6-dependent resistance specificity to Blumeria<br />
graminis f. sp. hordei in barley and wheat. Plt. J. 25:335-348.<br />
137
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Hansson, A., C.G. Kannangara, D. von Wettstein, and M. Hansson. 1999. Molecular basis<br />
for semidomiance of missense mutations in the XANTHA-H (42-kDa) subunit of<br />
magnesium chelatase. Proc. Natl. Acad. Sci. <strong>US</strong>A 96:1744-1749.<br />
Hirota, N., T. Kaneko, K. Ito, and K. Takeda. 2006. Mapping a factor controlling the<br />
thermostability of seed lipoxygenase-1 in barley. Plant Breeding 125:231-235.<br />
Ivandic, V., S. Malyshev, V. Korzun, A. Graner, and A. Börner. 1999. Comparative mapping<br />
of a gibberellic acid-insensitive dwarfing gene (Dwf2) on chromosome 4HS in barley.<br />
Theor. Appl. Genet. 98:728-731.<br />
Kanyuka, K., A. Druka, D..G. Caldwell, A. Tymon, N. McCallum, R. Waugh, and M. J.<br />
Adams. 2005. Evidence that the recessive bymovirus resistance locus rym4 in barley<br />
corresponds to the eukaryotic translation initiation factor 4E gene. Molecular Plant<br />
pathology 6:449-458.<br />
Kleinhofs, A., A. Kilian, M.A. Saghai Marrof, R.M. Biyashev, P. Hayes, F.Q. Chen, N.<br />
Lapitan, A. Fenwick, T.K. Blake, V. Kanazin, E. Ananiev, L. Dahleen, D. Kudrna, J.<br />
Bollinger, S.J. Knapp, B. Liu, M. Sorells, M. Heun, J.D. Franckowiak, D. Hoffman,<br />
R. Skadsen, and B.J. Steffenson. 1993. A molecular, isozyme and morphological map<br />
of the barley (Hordeum vulgare) genome. Theor. Appl. Genet. 86:705-712.<br />
Kleinhofs, A. 1996. Integrating Barley RFLP and Classical Marker Maps. Coordinator’s report.<br />
BGN27:105-112.<br />
Kleinhofs, A. 2002. Integrating Molecular and Morphological/Physiological Marker Maps.<br />
Coordinator’s Report. BGN32:152-159.<br />
Komatsuda, T., W. Li, F. Takaiwa, and S. Oka. 1999. High resolution map around the vrs1<br />
locus controlling two- and six-rowed spike in barley. (Hordeum vulgare). Genome<br />
42:248-253.<br />
Komatsuda, T., and Y. Mano. 2002. Molecular mapping of the intermedium spike-c (int-c) and<br />
non-brittle rachis 1 (btr1) loci in barley (Hordeum vulgare L.). Theor. Appl Genet.<br />
105:85-90.<br />
Kretschmer, J.M., K.J. Chalmers, S. Manning, A. Karakousis, A.R. Barr, A.K.M.R. Islam,<br />
S.J. Logue, Y.W. Choe, S.J. Barker, R.C.M. Lance, and P. Langridge. 1997. RFLP<br />
mapping of the Ha2 cereal cyst nematode resistance in barley.Theor. Appl. Genet.<br />
94:1060-1064.<br />
Kurth, J., R. Kolsch, V. Simons, and P. Schulze-Lefert. 2001. A high-resolution genetic map<br />
and a diagnostic RFLP marker for the Mlg resistance locus to powdery mildew in barley.<br />
Theor. Appl. Genet. 102:53-60.<br />
Larson, S.R., K.A. Young, A. Cook, T.K. Blake, and V. Raboy. 1998. Linkage mapping of<br />
two mutations that reduce phytic acid content of barley grain. Theor. Appl. Genet.<br />
97:141-146.<br />
138
Barley Genetics Newsletter (2007) 37: 105-153<br />
Laurie, D.A., N. Pratchett, C. Romero, E. Simpson, and J.W. Snape. 1993. Assignment of<br />
the denso dwarfing gene to the long arm of chromosome 3 (3H) of barley by use of RFLP<br />
markers. Plant. Breed. 111:198-203.<br />
Laurie, D.A., N. Pratchett, J.H. Bezan, and J.W. Snape. 1994. Genetic analysis of a<br />
photoperiod response gene on the short arm of chromosome 2 (2H) of Hordeum vulgare<br />
(barley). Heredity 72:619-627.<br />
Laurie, D.A., N. Pratchett, J.H. Bezan, and J.W. Snape. 1995. RFLP mapping of five major<br />
genes and eight quantitative trait loci controlling flowering time in a winter x spring<br />
barley (Hordeum vulgare L.) cross. Genome 38:575-585.<br />
Laurie, D.A., N. Pratchett, R.A. Allen, and S.S. Hantke. 1996. RFLP mapping of the barley<br />
homeotic mutant lax-a. Theor. Appl. Genet. 93:81-85.<br />
Lee S.H. and S.M. Neate. 2007a. Molecular mapping of Rsp1, Rsp2, and Rsp3 genes conferring<br />
resistance to Septoria speckled leaf blotch in barley. Phytopathology 97:155-161.<br />
Lee S.H. and S.M. Neate. 2007b. Sequence tagged site markers to Rsp1, Rsp2, and Rsp3 genes<br />
for resistance to Septoria speckled leaf blotch in barley. Phytopathology 97:161-169.<br />
Li. M., D. Kudrna, and A. Kleinhofs. 2000. Fine mapping of a Semi-dwarf gene Brachytic1 in<br />
barley. p. 72-74. In: S. Logue (ed.) Barley Genetics VIII. Volume III. Proc. Eigth Int.<br />
Barley Genet. Symp. Adelaide. Dept. Plant Science, Waite Campus, Adelaide University,<br />
Glen Osmond, South Australia.<br />
Mammadov, J.A., B. J. Steffenson, and M.A. Saghai Maroof. 2005. High resolution mapping<br />
of the barley leaf rust resistance gene Rph5 using barley expressed sequence tags (ESTs)<br />
and synteny with rice. Theor. Appl. Genet. 111:1651-1660.<br />
Mechelen J.R. van, R.C. Schuurink, M. Smits, A. Graner, A.C. Douma, N.J.A. Sedee, N.F.<br />
Schmitt, and B.E Valk. 1999. Molecular characterization of two lipoxygenases from<br />
barley. Plant Molecular Biology 39:1283-1298.<br />
Müller, K.J., N. Romano, O. Gerstner, F. Gracia-Maroto, C. Pozzi, F. Salamini, and W.<br />
Rhode. 1995. The barley Hooded mutation caused by a duplication in a homeobox gene<br />
intron. Nature 374:727-730.<br />
Netsvetaev, V.P. 1997. High lysine mutant of winter barley - L76. BGN27:51-54.<br />
Okada, Y., R. Kanatani, S. Arai, and I. Kazutoshi. 2004. Interaction between barley mosaic<br />
disease-resistance genes rym1 and rym5, in the response to BaYMV strains. Breeding<br />
Science 54 (4):319-324.<br />
Park, R.F. and A. Karakousis. 2002. Characterization and mapping Rph19 conferring<br />
resistance to Puccinia hordei in the cultivar ’Rea1’ and several Australian barley. Plant<br />
Breeding 121:232-236.<br />
Pratchett, N. and D.A. Laurie. 1994. Genetic map location of the barley developmental mutant<br />
liguleless in relation to RFLP markers. Hereditas 120:35-39.<br />
139
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Rostoks, N., D. Schmierer, S. Mudie, T. Drader, R. Brueggeman, D. Caldwell, R. Waugh,<br />
and A. Kleinhofs. 2006. Barley necrotic locus nec1 encodes the cyclic nucleotide-gated<br />
ion channel 4 homologous to the Arabidopsis Hlm1. Mol.Gen. Genomics 275:159-168.<br />
Saeki, K., C. Miyazaki, N. Hirota, A. Saito, K. Ito, and T. Konishi. 1999. RFLP mapping of<br />
BaYMV resistance gene rym3 in barley (Hordeum vulgare). Theor. Appl. Genet. 99:727-<br />
732.<br />
Saisho, D., K.-I. Tanno, M. Chono, I. Honda, H. Kitano, and K. Takeda. 2004. Spontaneous<br />
Brassinolide-insensitive barley mutants ’uzu’ adapted to East Asia. Breeding Science<br />
54(4): 409-416.<br />
Sameri, M. and T. Komatsuda. 2007. Location of quantitative trait loci for yield components in<br />
a cross oriental x occidental barley cultivar (Hordeum vulgare L.). Japan Agricultural<br />
Research Quaterly 41(3):195-199.<br />
Schmierer, D., A. Druka, D. Kudrna, and A. Kleinhofs. 2001. Fine Mapping of the fch12<br />
chlorina seedling mutant. BGN31:12-13.<br />
Schweizer, G.F., M. Baumer, G. Daniel, H. Rugel, and M.S. Röder. 1995. RFLP markers<br />
linked to scald (Rhynchosporium secalis) resistance gene Rh2 in barley. Theor. Appl.<br />
Genet. 90:920-922.<br />
Skinner J.S., P. Szucs, J. von Zitzewitz, L. Marquez-Cedillo, T. Filichkin, E.J. Stockinger,<br />
M.F. Thomashow, T.H.H. Chen, and P.M. Hayes. 2006. Mapping of barley homologs<br />
to genes that regulate low temperature tolerance in Arabidopsis. Theor. Appl. Genet.<br />
112:832-842.<br />
Smilde, W.D., A. Tekauz, and A. Graner. 2000. Development of a high resolution map for the<br />
Rh and Pt resistance on barley Chromosome 3H. p. 178-180. In: S. Logue (ed.) Barley<br />
Genetics VIII. Volume II. Proc. Eigth Int. Barley Genet. Symp. Adelaide. Dept. Plant<br />
Science, Waite Campus, Adelaide University, Glen Osmond, South Australia.<br />
Soule, J.D., D.A. Kudrna, and A. Kleinhofs. 2000. Isolation, mapping, and characterization of<br />
two barley multiovary mutants. J. Heredity 91:483-487.<br />
Senthil, N. and T. Komatsuda. 2005. Inter-subspecific maps of non-brittle rachis gene btr1/btr2<br />
using occidental, oriental and wild barley lines. Euphytica 145:215-220.<br />
Stein, N., D. Perovic, J. Kumlehn, B. Pellio, S. Stracke, S. Streng, F. Ordon, and A. Graner.<br />
2005. The eukaryotic translation initian factor 4E confers multiallelic recessive<br />
Bymovirus resistance in Hordeum vulgare (L.). The Plant Journal 42:912-922.<br />
Taketa, S., T. Awayama, S. Amano, Y. Sakurai, and M. Ichii. 2006. High-resolution mapping<br />
of the nud locus controlling the naked caryopsis in barley. Plant Breeding 125:337-342.<br />
Tohno-oka, T., M. Ishii, R. Kanatani, H. Takahashi, and K. Takeda. 2000. Genetic Analysis<br />
of photoperiotic response of barley in different daylength conditions. p.239-241. In: S.<br />
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Logue (ed.) Barley Genetics VIII. Volume III. Proc. Eigth Int. Barley Genet. Symp.<br />
Adelaide. Dept. Plant Science, Waite Campus, Adelaide University, Glen Osmond, South<br />
Australia.<br />
Toojinda, T., L.H. Broers, X.M. Chen, P.M. Hayes, A. Kleinhofs, J. Korte, D. Kudrna, H.<br />
Leung, R.F. Line, W. Powell, L. Ramsey, H. Vivar, and R. Waugh. 2000. Mapping<br />
quantitative and qualitative disease resistance genes in a doubled haploid population of<br />
barley (Hordeum vulgare). Theor. Appl. Genet. 101:580-589.<br />
Wang, J., H. Raman, M. Zhou, P.R. Ryan, E. Delhaize, D.M. Hebb, N. Coombes, and N.<br />
Mendham. 2007. High-resolution mapping of the Alp locus and identification of a<br />
candidate gene HvMATE controlling aluminum tolerance in barley (Hordeum vulgare<br />
L.). Theor. Appl. Genet. 115:265-276.<br />
Warner, R.L., D.A. Kudrna, and A. Kleinhofs. 1995. Association of the NAD(P)H-bispecific<br />
nitrate reductase structural gene with the Nar7 locus in barley. Genome 38:743-746.<br />
Weerasena, J.S., B.J. Steffenson, and A.B. Falk. 2004. Conversion of an amplified fragment<br />
length polymorphism marker into a c-dominant marker in mapping Rph15 gene<br />
conferring resistance to barley leaf rust, Puccinia hordei Otth. Theor. Appl. Genet.<br />
108:712-719.<br />
Williams, K.J., A. Lichon, P. Gianquitto, J.M. Kretschmer, A. Karakousis, S. Manning, P.<br />
Langridge, and H. Wallwork. 1999. Identification and mapping of a gene conferring<br />
resistance to the spot form of net blotch (Pyrenophora teres f. maculata) in barley. Theor.<br />
Appl. Genet. 99: 323-327.<br />
Williams, K., P. Bogacki, L. Scott, A. Karakousis, and H. Wallwork. 2001. Mapping of a<br />
gene for leaf scald resistance in barley line ’B87/14’ and validation of microsatellite and<br />
RFLP markers for marker-assisted selection. Plant Breed. 120:301-304.<br />
Yan G.P., and X.M. Chen. 2006. Molecular mapping of a recessive gene for resistance to stripe<br />
rust in barley. Theor. Appl. Genet. 113:529-537.<br />
Yan L., D. Fu, C. Li, A. Blechl, G. Tranquilli, M. Bonafede, A. Sanchez, M. Valarik, S.<br />
Yasuda, and J. Dubcovsky. 2006. The wheat and barley vernalization gene VRN3 is an<br />
orthologue of FT. Proc. Natl. Acad. Sci. <strong>US</strong>A 103:19581-19586.<br />
Zhong, S.B., R.J. Effertz, Y. Jin, J.D. Franckowiak, and B.J. Steffenson. 2003. Molecular<br />
mapping of the leaf rust resistance gene Rph6 in barley and its linkage relationships with<br />
Rph5 and Rph7. Phytopathology 93 (5):604-609.<br />
Zitzewitz, J. von, P. Scucs, J. Dobcov, L. Yan, E. Francia, N. Pecchioni, A. Casas, T.H.H.<br />
Chen, P.M. Hayes, and J.S. Skinner. 2005. Molecular and structural characterization of<br />
barley vernalization genes. Plant Molecular Biology 59:449-467.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Barley Genetic Stock Collection<br />
A. Hang and K. Satterfield<br />
<strong>US</strong>DA-ARS, National Small Grains Germplasm Research Facility,<br />
Aberdeen, Idaho 83210, <strong>US</strong>A<br />
e-mail: anhang@uidaho.edu<br />
In 2006, 655 barley genetic stocks were planted in the field and in the greenhouse for evaluation<br />
and for seed increase.<br />
Two mapping populations, including SSD F6 seed OSU 11/Harrington and SSD F6 seed<br />
OSU 15/Harrington, derived from single seed descent (SSD) of crosses between Hordeum<br />
vulgare subps. Spontaneum and cultivar “Harrington” obtained from Dr. Pat Hayes, Oregon<br />
State University (OSU), were planted in the field for seed increase.<br />
Four necrotic or lesion mimic mutants obtained from Dr. Anders Falk, Biological Research<br />
Center, Sweden, were also grown in the greenhouse for observation and for seed increase.<br />
Three hundred forty-five samples of barley genetic stocks were shipped to researchers in<br />
2006.<br />
Coordinator’s report: Trisomic and aneuploid stocks<br />
An Hang<br />
<strong>US</strong>DA-ARS, National Small Grains Germplasm Research Facility,<br />
Aberdeen, Idaho 83210, <strong>US</strong>A<br />
e-mail: anhang@uidaho.edu<br />
There is no new information about trisomic and aneuploid stocks. Lists of these stocks are<br />
available in BGN 25:104. Seed request for these stocks should be sent to the coordinator.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Translocations and balanced tertiary trisomics<br />
Andreas Houben<br />
Institute of Plant Genetics and Crop Plant Research<br />
DE-06466 Gatersleben, Germany<br />
email: houben@ipk-gatersleben.de<br />
Restructured barley chromosomes have been used by Nasuda et al. (2005) to elucidate the<br />
function of centromere-localized DNA sequences. The satellite sequences (AGGGAG)(n) and<br />
Ty3/gypsy-like retrotransposons are known to localize at the barley centromeres. Using a<br />
gametocidal system, which induces chromosomal mutations in barley chromosomes added to<br />
common wheat, the authors obtained an isochromosome for the short arm of barley chromosome<br />
7H (7HS) that lacked the barley-specific satellite sequence (AGGGAG)(n). Two telocentric<br />
derivatives of the isochromosome arose in the progeny: 7HS* with and 7HS** without the<br />
pericentromeric C-band. FISH analysis demonstrated that both truncated telosomes lacked not<br />
only the barley-specific centromeric repeats but also any of the known wheat centromeric<br />
tandem repeats. Although they lacked these centromeric repeats, both truncated<br />
telochromosomes showed normal mitotic and meiotic transmission. Indirect immunostaining<br />
revealed that centromere-specific proteins localized at the centromeric region of 7HS*. The<br />
authors conclude that the barley centromeric repeats are neither sufficient nor obligatory to<br />
assemble kinetochores, and discussed the possible formation of a novel centromere in a barley<br />
chromosome.<br />
The collection is being maintained in cold storage. To the best knowledge of the coordinator,<br />
there are no new publications dealing with balanced tertiary trisomics in barley. Limited seed<br />
samples are available any time, and requests can be made to the coordinator.<br />
Reference:<br />
Nasuda, S., Hudakova, S., Schubert, I., Houben, A., and Endo, T.R. 2005. Stable barley<br />
chromosomes without centromeric repeats. Proc Natl Acad Sci U S A 102: 9842-9847.<br />
Coordinator’s report: Autotetraploids<br />
Wolfgang Friedt<br />
Institute of Crop Science and Plant Breeding I.<br />
Justus-Liebig-University, Heinrich-Buff-Ring 26-32<br />
DE-35392 Giessen, Germany<br />
e-mail: wolfgang.friedt@agrar.uni-giessen.de<br />
Fax: +49(0)641-9937429<br />
The collection of barley autotetraploids (exclusively spring types) described in former issues of<br />
BGN is maintained at the Giessen Field Experiment Station of our institute. The set of stocks,<br />
i.e. autotetraploids (4n) and corresponding diploid (2n) progenitors (if available) have last been<br />
grown in the field for seed multiplication in summer 2000. Limited seed samples of the stocks<br />
are available for distribution.<br />
143
Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Eceriferum Genes<br />
Udda Lundqvist<br />
Nordic Gene Bank<br />
P.O. Box 41, SE-230 53 Alnarp, Sweden<br />
e-mail: udda@nordgen.org<br />
Dahleen and Franckowiak (2006) could localize the eceriferum-zt locus on chromosome 2HS<br />
based on their molecar linkage studies and found linkage in bin 2H-01 d 16.8 distal from the<br />
SSR molecular marker Bmac 0134. Surface wax coating on the spike appears slightly reduced<br />
with eceriferum-zt. The wax code for this mutant gene is + ++ ++ .<br />
No further research work on gene localization has been reported on these collections of<br />
Eceriferum and Glossy genes. All descriptions in Barley Genetics Newsletter (BGN) Volume 26<br />
are valid and still up-to-date. All Swedish Eceriferum alleles can be seen in the SESTO database<br />
of the Nordic Gene Bank. Descriptions, images and graphic chromosome map displays of the<br />
Eceriferum and Glossy genes are available in the AceDB database for Barley Genes and Barley<br />
Genetic Stocks, and they get currently updated. Its address is found by: www.untamo.net/bgs<br />
As my possibilities in searching literature are very limited, I apologize if I am missing any<br />
important papers. Please send me notes of publications and reports to include in next year’s<br />
reports.<br />
Every research of interest in the field of Eceriferum genes, ‘Glossy sheath’ and ‘Glossy leaf’<br />
genes can be reported to the coordinator as well. Seed requests regarding the Swedish mutants<br />
can be forwarded to the coordinator udda@nordgen.org or to the Nordic Gene Bank,<br />
www.nordgen.org/ngb, all others to the Small Grain Germplasm Research Facility (<strong>US</strong>DA-<br />
ARS), Aberdeen, ID 83210, <strong>US</strong>A, nsgchb@ars-grin.gov or to the coordinator at any time.<br />
Reference:<br />
Dahleen, L.S. and J.D. Franckowiak. 2006. SSR linkages to eight additional morphological<br />
marker traits. Barley Genetic Newsletter 36:12+16.<br />
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Coordinator’s report: Nuclear genes affecting the chloroplast<br />
Mats Hansson<br />
<strong>Department</strong> of Biochemistry,<br />
Lund University, Box 124,<br />
SE-22100 Lund, Sweden<br />
E-mail: mats.hansson@biochemistry.lu.se<br />
Barley nuclear mutants deficient in chlorophyll biosynthesis and chloroplast development are<br />
named albina, xantha, viridis, chlorina, tigrina and striata depending on their colour and colour<br />
pattern. In the albina mutants the leaves are completely white due to lack of both chlorophyll and<br />
carotene pigments. The xantha mutants are yellow and produce carotene, but no chlorophyll. The<br />
chlorina and viridis mutants are both pale green, but differ in chlorina being viable. The tigrina<br />
and striata mutants are stripped transverse and along the leaves, respectively.<br />
Although the mutations are generally lethal, the large endosperm of barley seeds supports plant<br />
growth for several weeks, allowing analysis of the mutants at a seedling stage. This has been<br />
utilized in three studies concerning cold acclimation (Svensson et al. 2006), photosystem II<br />
(Morosinotto et al. 2006) and dominance/recessivity in chlorophyll biosynthesis (Axelsson et al.<br />
2006), respectively.<br />
Svensson and collaborators (2006) used the Affymetrix Barley1 GeneChip in combination with<br />
albina-e.16, albina-f.17, xantha-s.46 and xantha-b.12 to assess the effect of the chloroplast on<br />
the expression of cold-regulated genes. About 67% of wild-type cold-regulated genes were not<br />
regulated by cold in any mutant (chloroplast-dependent cold-regulated genes).They found that<br />
the lack of cold regulation in the mutants is due to the presence of signalling pathway(s)<br />
normally cold activated in wild type but constitutively active in the mutants, as well as to the<br />
disruption of low-temperature signalling pathway(s) due to the absence of active chloroplasts.<br />
They also found that photooxidative stress signalling pathway is constitutively active in the<br />
mutants. These results demonstrate the major role of the chloroplast in the control of the<br />
molecular adaptation to cold.<br />
The barley mutant viridis-zb.63 lacks photosystem I and was employed by Morosinotto et al.<br />
(2006) to mimic extreme and chronic overexcitation of photosystem II. The mutation was shown<br />
to reduce the photosystem II antenna to a minimal size of about 100 chlorophylls per<br />
photosystem II reaction centre, which was not further reducible. The minimal photosystem II<br />
unit was found to consist of a dimeric photosystem II reaction centre core surrounded by<br />
monomeric Lhcb4 (chlorophyll protein 29), Lhcb5 (chlorophyll protein 26) and trimeric lightharvesting<br />
complex II antenna proteins. This minimal photosystem II unit forms arrays in vivo,<br />
possibly to increase the efficiency of energy distribution and provide photoprotection. In wildtype<br />
plants, an additional antenna protein, chlorophyll protein 24 (Lhcb6), which is not<br />
expressed in viridis-zb.63, is proposed to associate to this minimal unit and stabilize larger<br />
antenna systems when needed. The analysis of the mutant also revealed the presence of two<br />
distinct signalling pathways activated by excess light absorbed by photosystem II: one,<br />
dependent on the redox state of the electron transport chain, is involved in the regulation of<br />
antenna size, and the second, more directly linked to the level of photoinhibitory stress perceived<br />
by the cell, participates in regulating carotenoid biosynthesis.<br />
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Axelsson et al. (2006) studied the enzyme Mg-chelatase, which catalyzes the insertion of Mg 2+ into<br />
protoporphyrin IX at the first committed step of the chlorophyll biosynthetic pathway. It consists of<br />
three subunits; I, D and H. The I-subunit belongs to the AAA-protein superfamily (ATPases<br />
associated with various cellular activities) that is known to form hexameric ring structures in an<br />
ATP-dependant fashion. Dominant mutations in the Xantha-h gene, encoding the I-subunit, revealed<br />
that it functions in a cooperative manner. Axelsson et al. demonstrated that the D-subunit, encoded<br />
by Xantha-g, forms ATP-independent oligomeric structures and should also be classified as an<br />
AAA-protein. Furthermore, the question of cooperativity of the D-subunit was addressed by<br />
characterizing xantha-g.28, -g.37, -g.44, -g.45 and -g.65 at the molecular level. The recessive<br />
behavior in vivo was explained by the absence of mutant proteins in the barley cell. The identified<br />
mutations were constructed in the corresponding gene of Rhodobacter capsulatus and the resulting<br />
D-proteins were studied in vitro. Mixtures of wild-type and mutant R. capsulatus D-subunits showed<br />
a lower activity as compared to wild-type subunits assayed alone. Thus, the mutant D-subunits<br />
displayed a dominant behavior in vitro thus revealing cooperativity between the D-subunits in the<br />
oligomeric state. Based on these results, they proposed a model where the D-oligomer forms a<br />
platform for the stepwise assembly of the I-subunits. The cooperative behavior suggests that the Doligomer<br />
takes an active part in the conformational dynamics between the subunits of the enzyme.<br />
The stock list of barley mutants defective in chlorophyll biosynthesis and chloroplast development is<br />
found elsewhere in this issue of BGN and at<br />
http://www.mps.lu.se/fileadmin/mps/People/Hansson/Barley_mutants_web.pdf<br />
Dr. Mats Hansson<br />
<strong>Department</strong> of Biochemistry<br />
Lund University<br />
Box 124<br />
SE-22100 Lund, SWEDEN<br />
Phone: +46-46-222 0105<br />
Fax: +46-46-222 4534<br />
E-mail: Mats.Hansson@biochemistry.lu.se<br />
New references:<br />
Axelsson, E., A. Sawicki, S. Nilsson, I. Schröder, S. Al-Karadaghi, R. D. Willows and M.<br />
Hansson. 2006. Recessiveness and dominance in barley mutants deficient in Mgchelatase<br />
subunit D, an AAA protein involved in chlorophyll biosynthesis. Plant Cell 18:<br />
3606-3616.<br />
Morosinotto, T., R. Bassi, S. Frigerio, G. Finazzi, E. Morris and J. Barber. 2006.<br />
Biochemical and structural analyses of a higher plant photosystem II supercomplex of a<br />
photosystem I-less mutant of barley. Consequences of a chronic over-reduction of the<br />
plastoquinone pool. FEBS J. 273: 4616-4630.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Svensson, J.T., C. Crosatti, C. Campoli, R. Bassi, A. Michele Stanca, T.J. Close, and L.<br />
Cativelli. 2006. Transcriptions analysis of cold acclimation in barley albina and<br />
xantha mutants. Plant Physiol. 141: 257-270.<br />
Coordinator’s report: The Genetic Male<br />
Sterile Barley Collection<br />
M.C. Therrien<br />
Agriculture and Agri-Food Canada<br />
Brandon Research Centre<br />
Box 1000A, RR#3, Brandon, MB<br />
Canada R7A 5Y3<br />
E-mail: MTherrien@agr.gc.ca<br />
The GMSBC has been at Brandon since 1992. If there are any new sources of male-sterile genes<br />
that you are aware of, please advice me, as this would be a good time to add any new source to<br />
the collection. For a list of the entries in the collection, simply E-mail me at the above adress. I<br />
can send the file (14Mb) in Excel format. We continue to store the collection at -20 o C and will<br />
have small (5 g) samples available for the asking. Since I have not received any reports or<br />
requests the last years, there is absolutely no summary in my report.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Ear morphology genes.<br />
Udda Lundqvist<br />
Nordic Gene Bank<br />
P.O. Box 41 SE-230 53 Alnarp, Sweden<br />
e-mail: udda@nordgen.org<br />
A. Michele Stanca and Valeria Terzi<br />
C.R.A., Genomic Research Centre<br />
Via San Protaso 302<br />
29017-Fiorenzuola d’Arda, Italy<br />
e-mail: michele@stanca.it; v.terzi@iol.it<br />
The studies on barley development in these last years have taken advantages both from genetics<br />
and genomics approaches. Some barley genes involved in the ear morphology development<br />
have been mapped on high density molecular linkage maps and this strategy has been<br />
accompanied by candidate gene approaches (Pozzi et al., 2002).<br />
In the work of Pozzi et al. (2003) 29 genetic loci for which mutant alleles exist were placed on a<br />
restriction fragment length polymorphism- amplified fragment length polymorphism (RFLP-<br />
AFLP) map. Among the 29 loci considered in the work, some specifically affect ear morphology<br />
and assume characteristics proper to phytomers of other regions (like third outer glume and<br />
awned palea) or are characterized by the presence of modified organs (like liguleless,<br />
bracteatum, triple awned lemma and awned lemma). In Table 1 the map positions individuated<br />
by Pozzi et al. (2003) for some loci involved in ear morphology are reported.<br />
Table 1. Position of 12 developmental mutant loci in a Proctor X Nudinka AFLP map (from<br />
Pozzi et al, 2003), but revised regarding symbols and nomenclature.<br />
Mutant symbol and name Map position Closest marker<br />
adp1, awned palea1 Chr. 3H/27 E3634-8<br />
als1, absent lower laterals1 Chr 3H/28 E4234-11<br />
bra-d.7, bracteatum-d.7 Chr. 1H E3634-7<br />
dub.1, double seed.1 Chr 5H/66 and 67 E4038-4<br />
hex-v.3, hexastichon-v.3 Chr. 2H/19-21 E4343-7<br />
hex-v.4, hexastichon-v.4 Chr. 2H/19 and 20 E3438-3<br />
int- c.5, intermedium-c.5 Chr. 4H/8 E4143-5<br />
Kap1, Hooded lemma1 Chr. 4H/36 and 37 E4140-1<br />
lks2, short awn2 Chr. 7H/6 E4138-3<br />
lks5, short awn5 Chr. 4H/38 E4143-5<br />
trp1, triple awned lemma1 Chr. 2H/22 and 23 E3644-13<br />
trd1, third outher glume1 Chr.1H/52 E3634-7<br />
The genetics of barley Hooded suppression has been studied by Roig et al. (2004). The genetic<br />
basis of this phenotype is a mutation in the homeobox Bkn3. After chemical mutagenesis and<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
complementation tests, five suK (suppressor of K) loci were identified and mapped on<br />
chromosomes 5H and 7H.<br />
Comparative genetic studies across species has revealed syntenous conservation in the order of<br />
genes and markers along grass chromosomes. Starting from this observation, Rossini et al.<br />
(2006) used a synteny approach comparing barley and rice genomes to individuate candidate<br />
genes for a set of barley developmental mutants.<br />
The gene vrs1 (six-rowed spike 1) responsible for the six-rowed spike in barley has been recently<br />
isolated by means of positional cloning by Komatsuda et al. (2007). The wild type Vrs1 gene,<br />
present in two-rowed barley, encodes a transcription factor that includes a homeodomain with a<br />
closely linked leucine zipper motif. VRS1 protein suppresses lateral rows and give two-rowed<br />
spike, whereas a mutation in the homeodomain-leucine zipper of Vrs1 resulted into loss of<br />
function and development of six-rowed phenotype.<br />
The conservation and implementation of the barley morphological mutant collections is essential<br />
for future studies, ranging from the use of computer graphic L-system- models to simulate the<br />
final morphology of a plant (Buck-Sorlin et al., 2004) to the use of genomic tools for the<br />
elucidation of the gene functions.<br />
Regarding the Swedish mutation collection two new additional mutant loci could be mapped<br />
based on molecular mapping studies using simple sequences repeat (SSR) markers (Dahleen et<br />
al. 2005, Dahleen and Franckowiak. 2006).<br />
(1). The intermedium spike-k (int-k) gene could be localized in the centromeric region of<br />
chromosome 7H, closely linked to Bmag0217 and Bmac0162 in bins 6 to 7. This spike mutant<br />
has a short and dense spike and the lateral spikelets are enlarged with a pointed apex.<br />
Occasionally they have a short awn. The central spikelets are semi-sterile and there is no seed set<br />
in the lateral spikelets. Plants have a dense coating of wax surface. They also have significantly<br />
reduced height, peduncle length, awn length, kernels per spike, leaf length, kernel weight and<br />
yield.<br />
(2). The erectoides-t (ert-t) gene, one of the dense spike mutant loci, could be localized near the<br />
tip of chromosome 2HS, approximately 11.4 cM distal from SSR marker Bmac0134, near the<br />
boundary between bins 2H-01 and 2H-02. The spikes of this mutant gene are semicompact,<br />
rachis internode length is about 2.7 mm and culm length is about 2/3 of normal. These<br />
phenotypic traits plus short awns are inheritated together. Based on general appearance of the<br />
plants, ert-t can be placed in the brachytic class and by diallelic crosses three earlier identified<br />
Brachytic 3 (brh3) phenotypes were found to be allelic at the ert-t locus (Franckowiak, 2006).<br />
References<br />
Buck-Sorlin, G., O. Kniemeyer, and W. Kurth. 2004. Integrated grammar representation of<br />
genes, metabiolites and morphology: the example of hordeomorphs. 2004. Fourth<br />
International Workshop on Functional Structural Plant Models, Montpellier, France pp.<br />
386-389.<br />
Dahleen, L.S., L.J.Vander Val, and J.D. Franckowiak. 2005. Characterixation and molecular<br />
mapping of genes determining semidwarfism in barley. J. Hered. 96:654-662.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Dahleen, L.S., and J.D. Franckowiak. 2006. SSR Linkages to Eight Additional Morphological<br />
Marker Traits. Barley Genetics Newsletter (BGN) 36:12-16.<br />
Franckowik, J,D. 2006. Coordinator’s report: Chromosome 2H (2). BGN 36:54-56<br />
Komatsuda, T., M. Pourkheirandish, C. He, P. Azhaguvel, H. Kanamori, D. Perovic, N.<br />
Stein, A. Graner, T. Wicker , A. Tagiri, U. Lundqvist, T. Fujimura, M. Matsuoka,<br />
T. Matsumoto, and M. Yano M. 2007. Six-rowed barley originated from a mutation in a<br />
homeodomain-leucine zipper I-class homeobox gene. PNAS 104: 1424-1429.<br />
Pozzi, C., L. Rossini, L. Santi, M.R. Stile, L. Nicoloso, F. Barale, D. Vandoni, I. Decimo, C.<br />
Roig, P. Faccioli, V. Terzi, Y. Wang, and F. Salamini, F. 2002. Molecular genetics of<br />
barley development: from genetics to genomics. In: From biodiversity to genomics:<br />
breeding strategies for small grain cereals in the third millenium. Eucarpia Cereal Section<br />
Meeting, 2002. pp 463- 466.<br />
Pozzi, C., D. Di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of a barley<br />
(Hordeum vulgare) molecular linkage map with the position of genetic loci hosting 29<br />
developmental mutants. Heredity, 90: 390-396.<br />
Roig, C., C. Pozzi, L. Santi, J. Muller, Y. Wang, M.R. Stile, L. Rossini, A.M. Stanca, and F.<br />
Salamini. 2004. Genetics of barley Hooded suppression. Genetics, 167: 439-448.<br />
Rossini L., A. Vecchietti, L. Nicoloso, N. Stein, S. Franzago, F. Salamini, C. and Pozzi.<br />
2006. Candidate genes for barley mutants involved in plant architecture: an in silico<br />
approach. Theoretical and Applied Genetics, 112: 1073-1085.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Semidwarf genes<br />
J.D. Franckowiak<br />
Hermitage Research Station<br />
Queensland <strong>Department</strong> of Primary Industries and Fisheries<br />
Warwick, Queensland 4370, Australia<br />
e-mail: jerome.franckowiak@dpi.qld.gpv.au<br />
The sdw1 (denso) gene for the semidwarfism in barley was shown to be orthologous to the sd1 of<br />
rice based on similar map positions and linkage of both semidwarf genes to RFLP maker R1545<br />
(Zhang et al., 2005). Both genes are sensitive to gibberellic acid (GA) treatments and encode a<br />
GA20-oxidase mutant (HV20ox2), which produces lower levels of GA and causes the dwarf<br />
phenotype. Zhang et al. (2005) reported that the barley and rice genes shared 88% sequence<br />
similarity and 89% amino acid identity.<br />
Yin et al. (2005) confirmed that QTLs that lengthening the preflowering duration in the<br />
‘Apex’/‘Prisma’ population of 94 recombinant inbred lines (RILs) were located in the long arms<br />
of chromosome 2H and 3H and originated from Prisma. The QTL on 3HL was associated with<br />
presence of the sdw1 gene from Prisma and likely is a pleiotropic effect of sdw1 gene.<br />
Korff et al. (2006) detected the presence of the sdw1 gene from ‘Scarlett’ in doubled-haploid<br />
lines from the second backcross of Scarlett to Hordeum vulgare ssp. spontaneum accession<br />
ISR42-8. QTLs for plant height were detected also on other chromosomes by Korff et al. (2006).<br />
Gruszka et al. (2006) reported that a semidwarf mutant 093AR, which was produced by MNU<br />
(N-methyl-N-nitrosourea) treatment of variety Aramir, is allelic to the uzu (uzu1 on chromosome<br />
3HL) dwarfing gene. Their analysis of the DNA sequence of the HvBRI1 gene of 093AR showed<br />
a single-nucleotide substitution of the C to A substitutions at the positions 1760 and 1761.<br />
Gruszka et al. (2006) also confirmed that the uzu1 mutation was an A to G change at position<br />
2612 of the HvBRI1 gene. Chono et al. (2003) previously reported that the mutation resulted in<br />
an amino acid change at the highly conserved residue (His-857 to Arg-857) of the kinase domain<br />
of BRI1 (brassinosteroids) receptor protein. This change caused reduced sensitivity to BRs and<br />
reduced plant height (Chono et al., 2003).<br />
References:<br />
Gruszka, D., J. Zbieszczyk, M. Kwasniewski, I. Szarejko and M. Maluszynski. 2006. A new<br />
allele in a uzu gene encoding brassinosteroid receptor. Barley Genet. Newsl 36:1-2.<br />
author: wintergrun@op.pl<br />
Chono, M., I. Honda, H. Zeniya, K. Yoneyama, D. Saisho, K. Takeda, S. Takatsuto, T.<br />
Hoshino and Y. Watanabe. 2003. A semidwarf phenotype of barley uzu results from a<br />
nucleotide substitution in the gene encoding a putative brassinosteroid receptor. Plant<br />
Physiol. 133:1209-1219.<br />
von Korff, M., H. Wang, J. Léon, and K. Pillen. 2006. AB-QTL analysis in spring barley: II.<br />
Detection of favourable exotic alleles for agronomic traits introgressed from wild barley<br />
(H. vulgare ssp. spontaneum). Theor. Appl. Genet. 112:1221-1231.<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Yin, X., P.C. Struik, F.A. van Eeuwijk, P. Stam, and J. Tang. 2005. QTL analysis and QTLbased<br />
prediction of flowering phenology in recombinant inbred lines of barley J. Exp.<br />
Bot. 56(413):967-976.<br />
Zhang, J., X. Yang, P. Moolhuijzen, C. Li, M. Bellgard, R. Lance, and R. Appels. 2005.<br />
Towards isolation of the barley green revolution gene. Australian Barley Technical<br />
Symposium 2005.<br />
http://www.cdesign.com.au/proceedings_abts2005/posters%20(pdf)/poster_li.pdf.<br />
Verified 4 July 2007.<br />
Coordinator’s report : Wheat-barley genetic stocks<br />
A.K.M.R. Islam<br />
Faculty of Agriculture, Food & Wine, The University of Adelaide, Waite Campus,<br />
Glen Osmond, SA 5064, Australia<br />
e-mail: rislam@waite.adelaide.edu.com<br />
The production of five different disomic addition lines (1Hm, 2Hm, 4Hm, 5Hm and 7Hm) of<br />
Hordeum marinum chromosomes to Chinese Spring wheat has been reported earlier. It has now<br />
been possible to isolate a monosomic addition for chromosome 6Hm. Amphiploids have also<br />
been produced between H. marinum and more cultivars of commercial wheat (Islam and Colmer,<br />
unpublished).<br />
References:<br />
Islam,S; Malik, AI; Islam, AKMR; Colmer TD. 2007. Salt tolerance in a Hordeum marinum-<br />
Triticum aestivum amphiploid, and its parents. J Experimental Botany (in Press).<br />
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Barley Genetics Newsletter (2007) 37: 105-153<br />
Coordinator’s report: Early maturity genes<br />
Udda Lundqvist<br />
Nordic Gene Bank<br />
P-O. Box 41<br />
SE-23 053 Alnarp, Sweden<br />
e-mail: udda@nordgen.org<br />
Not much new research on gene localization has been reported on the Early maturity or<br />
Praematurum genes since the latest reports in Barley Genetic Newsletter (BGN) or in the AceDB<br />
database for Barley Genes and Barley Genetic Stocks.<br />
During the last years many reports are published on various QTLs detected in populations<br />
derived from wild x cultivated barley crosses with the goal of transferring desirable genes into<br />
cultivated barley lines.<br />
Korff et al. (2006) transferred favourable genes from wild barley to cultivated barleys and made<br />
evaluations in backcrosses of a doubled haploid population. A QTL for Early heading was<br />
associated with the Early maturity 1 (Eam1 or Ppd-H1) gene in the bin 3 region of 2HS.<br />
Several QTLs were found in crosses between two- and six-rowed cultivars. One QTL for early<br />
heading is reported and found in bin 8 of 2HL and is probably the Eam6 gene from a six-rowed<br />
parent (Franckowiak 2006).<br />
All information and descriptions made in the Barley Genetics Newsletter are valid and up-todate.<br />
As my possibilities in searching literature are very limited, I apologize if I am missing any<br />
important papers and reports. I would like to call on the barley community to assist me by<br />
sending notes of publications and reports to include in next year’s report. Descriptions, images<br />
and graphic chromosome map displays of the Early maturity or Praematurum genes are available<br />
in the AceDB database for Barley Genes and Barley Genetic Stocks. They get currently updated<br />
and are searchable under the address: www.untamo.net/bgs<br />
Every research of interest in the field of Early maturity genes can be reported to the coordinator<br />
as well. Seed requests regarding the Swedish mutants can be forwarded to the coordinator or<br />
directly to the Nordic Gene Bank, www.nordgen.org/ngb, all others to the Small Grain<br />
Germplasm Research Facility (<strong>US</strong>DA-ARS), Aberdeen, ID 83210, <strong>US</strong>A, nsgchb@ars-grin.gov<br />
or to the coordinator at any time.<br />
References:<br />
Franckowiak, J.D. 2006. Coordinator’s report: Chromosome 2H. Barley Genetics Newsletter<br />
36:54-56.<br />
Korff, M. von, H. Wang, J. Léon, and K. Pilen. 2006. AB-QTL analysis in spring barley: II.<br />
Detection of favourable exotic alleles for agronomic traits introgressed from wild barley<br />
(Hordeum vulgare ssp. spontaneum). Theor. Appl. Genet. 112:1221-1231.<br />
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Barley Genetics Newsletter (2007) 37: 154-187<br />
Descriptions of barley genetic stocks for 2007.<br />
Lynn Dahleen, 1 , Jerome D. Franckowiak 2 , Udda Lundqvist 3<br />
1 <strong>US</strong>DA-ARS, State University Station, P.O. Box 5677,<br />
Fargo, ND 58105-5051, <strong>US</strong>A<br />
2 Hermitage Research Station,<br />
Queensland <strong>Department</strong> of Primary Industries and Fisheries<br />
Warwick, Queensland 4370, Australia<br />
3 Nordic Gene Bank<br />
P.O. Box 41, SE-230 53 Alnarp, Sweden<br />
In this volume of the Barley Genetics Newsletter, seventy six new and revised Barley Genetic<br />
Stock descriptions are published (Table 1). The current list of new and revised BGS descriptions,<br />
including those in Table 1, are again presented by BGS number order (Table 2) and by locus<br />
symbol in alphabetic order (Table 3) in this issue. Information on the description location,<br />
recommended locus name, chromosomal location, previous gene symbols, and the primary<br />
genetic stock (GSHO number) are included in these lists. The GSHO stocks are held in the<br />
<strong>US</strong>DA-ARS Barley Genetic Stocks collection at the National Small Grains collection, (U.S.<br />
<strong>Department</strong> of Agriculture – Agricultural Research Service), Aberdeen, Idaho 83210, <strong>US</strong>A. The<br />
NGB stocks are held in the Nordic Gene Bank, P.O. Box 41, SE-230 53 Alnap, Sweden. This<br />
information is available through the Internet at the following addresses:<br />
(1) www.ars.usda.gov.PacWest/Aberdeen<br />
(2) www.ars-grin.gov:7000/npgs/descriptors/barley-genetics (GRIN)<br />
(3) http://wheat.pw.usda.gov/ggpages/bgn/<br />
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Table 1. A listing of Barley Genetic Stock (BGS) descriptions published in this issue of the<br />
Barley Genetics Newsletter, giving recommended locus symbols and names, and stock<br />
location information.<br />
BGS Locus symbol* Chr. Locus name or phenotype Descr. GSHO<br />
no. Rec Prev. loc. † vol.p. no. ‡<br />
1 brh1 br, ari-i 7HS Brachytic 1 37:188 25<br />
2 fch12 fc, clo-fc 7HS Chlorina seedling 12 37:190 36<br />
6 vrs1 v, Int-d 2HL Six-rowed spike 1 37:192 196<br />
7 nud1 n, h 7HL Naked caryopsis 1 37:195 115<br />
10 lks2 lk2, lk4 7HL Short awn 2 37:197 1232<br />
22 Rsg1 Grb 7H Reaction to Schizaphis gramineum 1 37:199 1317<br />
32 Rph9 Pa9 5HL Reaction to Puccinia hordei 9 37:201 1601<br />
41 brh7 brh.w 5HS Brachytic 7 37:203 1687<br />
44 brh16 brh.v 7HL Brachytic 16 37:204 1686<br />
60 lig1 li, aur-a 2HL Liguleless 1 37:205 6<br />
79 wst7 rb 2HL White streak 7 37:207 247<br />
82 Zeo1 Knd 2HL Zeocriton 1 37:209 1613<br />
85 yst4 yst4 2HL Yellow streak 4 37:210 2502<br />
87 fch14 f14 2HL Chlorina seedling 14 37:211 1739<br />
88 Rph2 Pa2, A 5HS Reaction to Puccinia hordei 2 37:212 1593<br />
96 Rph15 Rph16 2HS Reaction to Puccinia hordei 15 37:214 1586<br />
98 Eam6 Ea6 2HS Early maturity 6 37:216<br />
100 sld4 sld.d 7HS Slender dwarf 4 37:218 2479<br />
101 als1 als 3HL Absent lower laterals 1 37:219 1065<br />
102 uzu1 uz, u 3HL Uzu 1 or semi-brachytic 1 37:220 1300<br />
108 alm1 al, ebu-a 3HS Albino lemma 1 37:222 270<br />
122 Rph5 Pa5, Rph6 3HS Reaction to Puccinia hordei 5 37:224 1597<br />
130 eam10 easp 3HL Early maturity 10 37:226 2504<br />
136 Rph7 Pa7, Pa5 3HS Reaction to Puccinia hordei 7 37:228 1318<br />
142 brh8 brh.ad 3HS Brachytic 8 37:230 1671<br />
148 brh14 brh.q 3HL Brachytic 14 37:231 1682<br />
149 Rpc1 3H Reaction to Puccinia coronata var hordei 1 37:232 1601<br />
155 glf1 gl, cer-zh 4HL Glossy leaf 1 37:233 98<br />
157 brh2 br2, ari-l 4HL Brachytic 2 37:235 573<br />
178 int-c i, v5 4HS Intermedium spike-c 37:237 776<br />
179 Hsh1 Hs 4HL Hairy leaf sheath 1 37:240 986<br />
185 brh5 brh.m 4HS Brachytic 5 37:242 1678<br />
186 sld3 sld.e 4HS Slender dwarf 3 37:243 2480<br />
187 brh9 brh.k 4HS Brachytic 9 37:244 1676<br />
203 Blp1 B 1HL Black lemma and pericarp 1 37:245 988<br />
214 eam8 eak, mat-a 1HL Early maturity 8 37:247 765<br />
222 nec1 sp,,b 1HL Necrotic leaf spot 1 37:251 989<br />
253 cul2 uc2 6HL Uniculm 2 37:253 531<br />
155
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 1 (continued)<br />
BGS Locus symbol* Chr. Locus name or phenotype Descr. GSHO<br />
no. Rec Prev. loc. † vol.p. no. ‡<br />
254 rob1 o, rob-o 6HS Orange lemma 1 37:255 707<br />
266 ert-e, dsp9 ert-e 6HL Erectoides-e 37:257 477<br />
306 var1 va 5HL Variegated 1 37:259 1278<br />
348 Eam5 Ea5 5HL Early maturity 5 37:260<br />
349 brh4 brh.j 2HL Brachytic 4 37:262 1675<br />
350 brh6 brh.s 5HS Brachytic 6 37:263 1683<br />
377 seg1 se1 7HL Shrunken endosperm genetic 1 37:264 750<br />
379 seg3 se3 3HS Shrunken endosperm genetic 3 37:265 752<br />
380 seg4 se4 7HL Shrunken endosperm genetic 4 37:267 753<br />
396 seg6 se6 3HL Shrunken endosperm genetic 6 37:268 2467<br />
397 seg7 se7 Shrunken endosperm genetic 7 37:269 2468<br />
437 cer-zt cer-zt 2HS Eceriferum-zt 37:270 1527<br />
449 cer-yf cer-yf Eceriferum-yf 37:271 1539<br />
455 seg8 seg8 7H Shrunken endosperm genetic 8 37:272 2469<br />
474 lax-a lax-a 5HL Laxatum-a 37:273 1775<br />
516 Rsp2 Sep2 1HS Reaction to septoria passerinii 2 37:275 2511<br />
517 Rsp3 Sep3 1HS Reaction to septoria passerinii 3 37:276 2512<br />
518 sdw1 denso 3HL Semidwarf 1 37:277 2513<br />
546 int-k int-k 7H Intermedium spike-k 37:279 1770<br />
547 int-m int-m Intermedium spike-m 37:280 1772<br />
566 ert-t ert-t, brh3 2HS Erectoides-t 37:281 494<br />
577 Rsg2 Rsg2 Reaction to Schizaphis gramineum 2 37:283 2513<br />
586 bra-d 1HL Bracteatum-d 37:284 1696<br />
593 adp1 adp 3HL Awned palea 1 37:285 1950<br />
599 ant17 ant17 3HS Proanthocyanin-free 17 37:286<br />
617 cul4 uc-5 3HL Uniculme 4 37:289 2493<br />
623 eli-a lig-a Eligulum-a 37:290<br />
633 mnd6 den-6 5HL Many noded dwarf 6 37:291 1713<br />
636 tst2 lin2 Tip sterile 2 37:292 1781<br />
653 brh10 brh.l 2HS Brachytic 10 37:293 1677<br />
654 brh11 brh.n 5HS Brachytic 11 37:294 1679<br />
655 brh12 brh.o 5HS Brachytic 12 37:295 1680<br />
656 brh13 brh.p 5HS Brachytic 13 37:296 1681<br />
657 brh15 brh.u Brachytic 15 37:297 1685<br />
658 brh17 brh.ab 5HS Brachytic 17 37:298 1669<br />
659 brh18 brh.ac 5HS Brachytic 18 37:299 1670<br />
660 nld2 Narrow leafed dwarf 2 37:300<br />
661 dub1 5HL Double seed 1 37:301<br />
156
Barley Genetics Newsletter (2007) 37: 154-187<br />
* Recommended locus symbols are based on utilization of a three letter code for barley genes as<br />
approved at the business meeting of the Seventh International Barley Genetics Symposium at<br />
Saskatoon, Saskatchewan, Canada, on 05 August 1996.<br />
† Chromosome numbers and arm designations are based on a resolution passed at the business<br />
meeting of the Seventh International Barley Genetics Symposium at Saskatoon, Saskatchewan,<br />
Canada, on 05 August 1996. The Burnham and Hagberg (1956) designations of barley<br />
chromosomes were 1 2 3 4 5 6 and 7 while new designations based on the Triticeae system are<br />
7H 2H 3H 4H 1H 6H and 5H, respectively.<br />
‡ The seed stock associated with each BGS number is held as a GSHO stock number in the<br />
Barley Genetics Stock Collection at the <strong>US</strong>DA-ARS National Grains Germplasm Research<br />
Facility, Aberdeen, Idaho, <strong>US</strong>A.<br />
157
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2. A listing of Barley Genetic Stock (BGS) descriptions in recent issues of the Barley<br />
Genetics Newsletter recommended locus symbols and names, and stock location<br />
information.<br />
BGS Locus symbol * Chr. Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. † vol. p. no. ‡<br />
1 brh1 br, ari-i 7HS Brachytic 1 37:188 25<br />
2 fch12 fc, clo-fc 7HS Chlorina seedling 12 37:190 36<br />
3 yvs2 yc 7HS Virescent seedling 2 26: 46 41<br />
4 abo8 ac2, alb-m 7HS Albino seedling 8 26: 47 61<br />
5 fch8 f8 7HS Chlorina seedling 8 26: 48 40<br />
6 vrs1 v, Int-d 2HL Six-rowed spike 1 37:192 196<br />
7 nud1 n, h 7HL Naked caryopsis 1 37:195 115<br />
9 dsp1 l 7HS Dense spike 1 26: 53 1232<br />
10 lks2 lk2, lk4 7HL Short awn 2 37:197 566<br />
11 ubs4 u4, ari-d 7HL Unbranched style 4 26: 56 567<br />
12 des1 lc 7H Desynapsis 1 26: 57 592<br />
13 des4 des4 7H Desynapsis 4 26: 58 595<br />
14 des5 des5 7H Desynapsis 5 26: 59 596<br />
15 blx1 bl 4HL Non-blue aleurone xenia 1 26: 60 185<br />
16 wax1 wx, glx 7HS Waxy endosperm 1 26: 61 908<br />
17 fch4 f4, yv 7HL Chlorina seedling 4 26: 63 1214<br />
18 fch5 f5, yv2 7HS Chlorina seedling 5 26: 64 1215<br />
19 blx2 bl2 7HS Non-blue aleurone xenia 2 26: 65 209<br />
20 Rym2 Ym2 7HL Reaction to BaYMV 2 26: 66 984<br />
21 Run1 Un 7HS Reaction to Ustilago nuda 1 26: 67 1324<br />
22 Rsg1 Grb 7H Reaction to Schizaphis graminum 1 37:199 1317<br />
23 wnd1 wnd 7HS Winding dwarf 1 26: 69 2499<br />
24 fst3 fs3 7HS Fragile stem 3 26: 70 1746<br />
25 Xnt1 Xa 7HL Xantha seedling 1 26: 71 1606<br />
26 snb1 sb 7HS Subnodal bract 1 26: 72 1217<br />
27 lbi3 lb3 7HL Long basal rachis internode 3 26: 73 536<br />
28 ert-a ert-a 7HS Erectoides-a 26: 74 468<br />
29 ert-d ert-d 7HS Erectoides-d 26: 76 475<br />
30 ert-m ert-m 7HS Erectoides-m 26: 78 487<br />
31 sex6 sex6 7HS Shrunken endosperm xenia 6 26: 80 2476<br />
32 Rph9 Pa9 5HL Reaction to Puccinia hordei 9 37:201 1601<br />
33 ant1 rs, rub-a 7HS Anthocyanin-less 1 26: 82 1620<br />
34 msg50 msg,,hm 7HL Male sterile genetic 50 26: 83 2404<br />
35 rsm1 sm 7HS Reaction to BSMV 1 26: 84 2492<br />
36 xnt4 xc2 7HL Xantha seedling 4 26: 85 42<br />
37 xnt9 xan,,i 7HL Xantha seedling 9 26: 86 584<br />
38 smn1 smn 7HS Seminudoides 1 32: 78 1602<br />
39 mss2 mss2 7HS Midseason stripe 2 32: 79 2409<br />
158
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
40 prm1 prm 7HS Premature ripe 1 32: 80 2429<br />
41 brh7 brh.w 5HS Brachytic 7 37:203 1687<br />
42 Pyr1 Pyr1 7HS Pyramidatum 1 32: 82 1581<br />
43 mov1 mo5 7HL Multiovary 1 35:185<br />
44 brh16 brh.v 7HL Brachytic 16 37:204 1686<br />
51 rtt1 rt 2HS Rattail spike 1 26: 87 216<br />
52 fch15 or 2HS Chlorina seedling 15 26: 88 49<br />
53 abo2 a2 2HS Albino seedling 2 26: 89 70<br />
55 fch1 f, lg 2HS Chlorina seedling 1 26: 90 112<br />
56 wst4 wst4 2HL White streak 4 26: 91 568<br />
57 eog1 e, lep-e 2HL Elongated outer glume 1 26: 92 29<br />
58 vrs1 lr, v lr 2HL Six-rowed spike 1 26: 94 153<br />
59 gpa1 gp, gp2 2HL Grandpa 1 26: 95 1379<br />
60 lig1 li, aur-a 2HL Liguleless 1 37:205 6<br />
61 trp1 tr 2HL Triple awned lemma 1 26: 97 210<br />
62 sbk1 sk, cal-a 2HS Subjacent hood 1 32: 83 267<br />
63 yvs1 yx, alb-c2 2HS Virescent seedling 1 26: 99 68<br />
64 des7 des7 2H Desynapsis 7 26:100 598<br />
65 Eam1 Ea, Ppd-H1 2HS Early maturity 1 26:101 1316<br />
66 vrs1 V d 2HL Two-rowed spike 1 26:103 346<br />
67 vrs1 V t 2HL Deficiens 1 26:104 684<br />
68 Pvc 1 Pc 2HL Purple veined lemma 1 26:105 132<br />
69 Gth 1 G 2HL Toothed lemma 1 26:106 309<br />
70 Rph1 Pa 2H Reaction to Puccinia hordei 1 26:107 1313<br />
71 com2 bir2 2HS Compositum 2 26:108 1703<br />
72 glo-c glo-c 2H Globosum-c 26:109 1329<br />
73 fol-a fol-a 2HL Angustifolium-a 26:110 1744<br />
74 flo-c flo-c 2HS Extra floret-c 26:111 1743<br />
75 Lks1 Lk 2HL Awnless 1 26:112 44<br />
76 Pre2 Re2, P 2HL Red lemma and pericarp 2 26:113 234<br />
77 hcm1 h 2HL Short culm 1 26:115 2492<br />
78 mtt4 mt,,e, mt 2HL Mottled leaf 4 26:116 1231<br />
79 wst7 rb 2HL White streak 7 37:207 247<br />
80 ant2 pr, rub 2HL Anthocyanin-less 2 26:118 1632<br />
81 gsh7 gs7 Glossy sheath 7 26:119 1759<br />
82 Zeo1 Knd 2HL Zeocriton 1 37:209 1613<br />
83 sld2 sld2 2HS Slender dwarf 2 26:121 2491<br />
84 mss1 mss 2H Midseason stripe 1 26:122 1404<br />
85 yst4 yst4 2HL Yellow streak 4 37:210 2502<br />
86 fch13 f13 Chlorina seedling 13 26:124 16<br />
159
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
87 fch14 f14 2HL Chlorina seedling 14 37:211 1739<br />
88 Rph2 Pa2, A 5HS Reaction to Puccinia hordei 2 37:212 1593<br />
89 ari-g ari-g, lk10 Breviaristatum-g 26:128 1655<br />
90 ert-j ert-j 2H Erectoides-j 26:129 484<br />
91 ert-q ert-q 2H Erectoides-q 26:130 1562<br />
92 ert-u ert-u, br5 2H Erectoides-u 26:131 496<br />
93 ert-zd ert-zd, br7 2H Erectoides-zd 26:132 504<br />
94 abo4 a4 2H Albino seedling 4 26:133 167<br />
95 abo13 alb,,p 2HL Albino seedling 13 26:134 585<br />
96 Rph15 Rph16 2HS Reaction to Puccinia hordei 15 37:214 1586<br />
97 acr1 acr 2HL Accordion rachis 1 32: 85 1617<br />
98 Eam6 Ea6, Ea 2HS Early maturity 6 37:216<br />
99 lin1 s, rin 2HL Lesser internode number 1 32: 88 2492<br />
100 sld4 sld.d 7HS Slender dwarf 4 37:218 2479<br />
101 als1 als 3HL Absent lower laterals 1 37:219 1065<br />
102 uzu1 uz, u 3HL Uzu 1 or semi brachytic 1 37:220 1300<br />
104 yst1 yst, ys 3HS Yellow streak 1 26:138 1140<br />
105 xnt3 xc, vir-l 3HS Xantha seedling 3 26:139 66<br />
106 abo6 ac 3HS Albino seedling 6 26:140 30<br />
107 wst1 wst, wst3 3HL White streak 1 26:141 159<br />
108 alm1 al, ebu-a 3HS Albino lemma 1 37:222 270<br />
109 yst2 yst2 3HS Yellow streak 2 26:144 570<br />
111 dsp10 lc 3HS Dense spike 10 26:145 71<br />
112 abo9 an 3HS Albino seedling 9 26:146 348<br />
113 xnt6 xs 3HS Xantha seedling 6 26:147 117<br />
114 cur2 cu2 3HL Curly 2 26:148 274<br />
115 btr1 bt1 3HS Non-brittle rachis 1 26:149 1233<br />
116 btr2 bt2 3HS Non-brittle rachis 2 26:150 842<br />
117 fch2 f2, lg5 3HL Chlorina seedling 2 26:151 107<br />
118 lnt1 rnt, int-l 3HL Low number of tillers 1 26:153 833<br />
119 des2 ds 3H Desynapsis 2 26:154 593<br />
120 zeb1 zb 3HL Zebra stripe 1 26:155 1279<br />
121 Rph3 Pa3 7HL Reaction to Puccinia hordei 3 26:156 1316<br />
122 Rph5 Pa5, Pa6 3HS Reaction to Puccinia hordei 5 37:224 1597<br />
123 Ryd2 Yd2 3HL Reaction to BYDV 2 26:158 1315<br />
124 vrs4 mul, int-e 3HL Six-rowed spike 4 26:159 775<br />
125 lzd1 lzd, dw4 3HS Lazy dwarf 1 26:161 1787<br />
126 sld1 dw1 3HL Slender dwarf 1 26:162 2488<br />
127 Pub1 Pub 3HL Pubescent leaf blade 1 26:163 1576<br />
128 sca1 sca 3HS Short crooked awn 1 26:164 2439<br />
160
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
129 wst6 wst,,j 3HL White streak 6 26:165 2500<br />
130 eam10 easp 3HL Early maturity 10 37:226 2504<br />
131 gra-a gran-a 3HL Granum-a 26:167 1757<br />
132 ari-a ari-a 3HS Breviaristatum-a 26:168 1648<br />
133 sdw2 sdw-b 3HL Semidwarf 2 26:169 2466<br />
134 ert-c ert-c 3HL Erectoides-c 26:170 471<br />
135 ert-ii ert-ii 3HL Erectoides-ii 26:172 483<br />
136 Rph7 Pa7, Pa5 3HS Reaction to Puccinia hordei 7 37:228 1318<br />
137 Rph10 Rph10 3HL Reaction to Puccinia hordei 10 26:174 1588<br />
138 nec4 nec4 3H Necrotic leaf spot 4 26:175<br />
139 nec5 nec5 3H Necrotic leaf spot 5 26:176<br />
140 xnt8 xan,,h 3HS Xantha seedling 8 26:177 582<br />
141 rym5 Ym 3HL Reaction to Barley yellow mosaic virus 5 32: 90<br />
142 brh8 brh.ad 3HS Brachytic 8 37:230 1671<br />
143 sex8 sex.j 3HS Shrunken endosperm 8 32: 93 2471<br />
144 sld5 sld5 3HS Slender dwarf 5 32: 94 2483<br />
146 cal-d cal-d 3H Calcaroides-d 32: 95 1697<br />
147 mov2 mo 3HS Multiovary 2 35:190<br />
148 brh14 brh.q 3HL Brachytic 14 37:231 1682<br />
149 Rpc1 3H Reaction to Puccinia coronata var. hordei 1 37:232 1601<br />
151 fch9 f9 4HS Chlorina seedling 9 26:178 571<br />
152 Kap1 K 4HS Hooded lemma 1 26:179 985<br />
155 glf1 gl, cer-zh 4HL Glossy leaf 1 37:233 98<br />
156 lbi2 lb2, ert-i 4HL Long basal rachis internode 2 26:183 572<br />
157 brh2 br2, ari-l 4HL Brachytic 2 37:235 573<br />
158 yhd1 yh 4HL Yellow head 1 26:185 574<br />
160 min2 en-min Enhancer of minute 1 26:186 266<br />
161 min1 min 4HL Semi-minute dwarf 1 26:187 987<br />
163 sgh1 sh1 4HL Spring growth habit 1 26:188 575<br />
164 Hln1 Hn 4HL Hairs on lemma nerves 1 26:189 576<br />
165 glf3 gl3, cer-j 4HL Glossy leaf 3 26:190 577<br />
166 msg25 msg,,r 4HL Male sterile genetic 25 26:192 744<br />
167 rym1 Ym 4HL Reaction to barley yellow mosaic virus 1 32: 96<br />
168 glo-a glo-a 4HS Globosum-a 26:194 1328<br />
170 lgn3 lg3 4HL Light green 3 26:195 171<br />
171 lgn4 lg4, lg9 4HL Light green 4 26:196 681<br />
172 lks5 lk5, ari-c 4HL Short awn 5 26:197 1297<br />
173 blx3 bl3 4HL Non-blue aleurone xenia 3 26:198 2506<br />
174 blx4 bl4 4HL Non-blue (pink) aleurone xenia 4 26:199 2507<br />
176 ovl1 ovl 4H Ovaryless 1 35:191<br />
161
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
178 int-c i, v5 4HS Intermedium spike-c 37:237 776<br />
179 Hsh1 Hs 4HL Hairy leaf sheath 1 37:240 986<br />
180 sid1 nls 4HL Single internode dwarf 1 26:203 2477<br />
181 eam9 ea,,c 4HL Early maturity 9 26:204 1732<br />
182 flo-a flo-a Extra floret-a 26:205 1741<br />
183 Ynd1 Yn 4HS Yellow node 1 32:98<br />
184 Zeo3 Zeo.h 4HL Zeocriton 3 32:99 1611<br />
185 brh5 brh.m 4HS Brachytic 5 37:242 1678<br />
186 sld3 ant17.567 4HS Slender dwarf 3 37:243 2480<br />
187 brh9 brh.k 4HS Brachytic 9 37:244 1676<br />
201 fch7 f7 1HL Chlorina seedling 7 26:206 4<br />
202 trd1 t, bra-c 1HL Third outer glume 1 26:207 227<br />
203 Blp1 B 1HL Black lemma and pericarp 1 37:245 988<br />
207 abo1 at 1HL Albino seedling 1 26:210 51<br />
208 fst2 fs2 1HL Fragile stem 2 26:211 578<br />
213 Sgh3 Sh3 1HL Spring growth habit 3 26:212 764<br />
214 eam8 eak, mat-a 1HL Early maturity 8 37:247 765<br />
215 des6 des6 1H Desynapsis 6 26:216 597<br />
218 Rph4 Pa4 1HS Reaction to Puccinia hordei 4 26:217 1314<br />
220 fch3 f3 1HS Chlorina seedling 3 26:218 851<br />
221 wst5 wst5 1HL White streak 5 26:219 591<br />
222 nec1 sp,,b 1HL Necrotic leaf spot 1 37:251 989<br />
223 zeb3 zb3, zbc 1HL Zebra stripe 3 26:221 1451<br />
224 ert-b ert-b 1HL Erectoides-b 26:222 470<br />
225 clh1 clh 1HL Curled leaf dwarf 1 26:223 1212<br />
226 rvl1 rvl 1HL Revoluted leaf 1 26:224 608<br />
227 sls1 sls 1HS Small lateral spikelet 1 26:225 2492<br />
228 Sil1 Sil 1HS Subcrown internode length 1 26:226 1604<br />
229 cud2 cud2 1HL Curly dwarf 2 26:227 1712<br />
230 glo-e glo-e 1HL Globosum-e 26:228 1755<br />
231 cur5 cu5 1HS Curly 5 26:229 1710<br />
232 Lys4 sex5 1HS High lysine 4 26:230 2475<br />
233 xnt7 xan,,g 1HL Xantha seedling 7 26:231 581<br />
234 mov3 mo-a 1H Multiovary 3 32:102<br />
235 lel1 lel 1HL Leafy lemma 1 32:103 1780<br />
251 mul2 mul2 6HL Multiflorus 2 26:232 1394<br />
252 eam7 ea7, ec 6HS Early maturity 7 26:233 579<br />
253 cul2 uc2 6HL Uniculm 2 37:253 531<br />
254 rob1 o, rob-o 6HS Orange lemma 1 37:255 707<br />
255 xnt5 xn 6HL Xantha seedling 5 26:237 43<br />
162
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
257 raw5 r,,e 6HL Smooth awn 5 26:238 785<br />
258 dsp9 l9, ert-e 6HL Dense spike 9 26:239 1774<br />
260 fch11 f11 6HL Chlorina seedling 11 26:240 1738<br />
261 nec2 nec2 6HS Necrotic leaf spot 2 26:241 1224<br />
262 cur1 cu1 6HL Curly 1 26:242 1705<br />
263 cur3 cu3 6HL Curly 3 26:243 1707<br />
264 mtt5 mt,,f 6HL Mottled leaf 5 26:244 2410<br />
265 nec3 nec3 6HS Necrotic leaf spot 3 26:245 1330<br />
266 ert-e ert-e, dsp9 6HL Erectoides-e 37:257 477<br />
267 Rph11 Rph11 6HL Reaction to Puccinia hordei 11 26:247 1589<br />
268 lax-b lax-b 6HL Laxatum-b 26:248 1776<br />
269 lys6 lys6 6H High lysine 6 26:249 1786<br />
270 abo14 alb,,q 6HL Albino seedling 14 26:250 586<br />
271 abo15 alb,,t 6HS Albino seedling 15 26:251<br />
301 fst1 fs 5HL Fragile stem 1 26:252 629<br />
302 mtt2 mt2 5HL Mottled leaf 2 26:253 1398<br />
303 var3 va3 5HL Variegated 3 26:254 1277<br />
304 wst2 wst2 5HL White streak 2 26:255 766<br />
305 crm1 cm 5HL Cream seedling 1 26:256 20<br />
306 var1 va 5HL Variegated 1 37:259 1278<br />
308 lbi1 lb, rac-a 5HL Long basal rachis internode 1 26:258 580<br />
309 Sgh2 Sh2 5HL Spring growth habit 2 26:259 770<br />
311 dex1 sex2 5HS Defective endosperm xenia 1 26:260<br />
312 raw1 r 5HL Smooth awn 1 26:261 27<br />
313 fch6 f6, yv 5HL Chlorina seedling 6 26:262 1390<br />
314 vrs2 v2 5HL Six-rowed spike 2 26:263 773<br />
315 vrs3 v3, int-a 1HL Six-rowed spike 3 26:264 774<br />
317 ddt1 ddt 5HS Reaction to DDT 1 26:266 331<br />
319 rpg4 rpg4 5HL Reaction to Puccinia graminis 4 26:267 2438<br />
320 int-b int-b 5HL Intermedium spike-b 26:268 1764<br />
321 srh1 s, l 5HL Short rachilla hair 1 26:269 27<br />
322 dsk1 dsk 5HL Dusky 1 26:270 1714<br />
323 nld1 nld 5HL Narrow leafed dwarf 1 26:271 769<br />
324 cud1 cud 5HL Curly dwarf 1 26:272 1711<br />
325 crl1 crl, cl Curly lateral 1 26:273 1211<br />
326 blf1 bb 5HL Broad leaf 1 26:274 1393<br />
327 flo-b flo-b 5HL Extra floret-b 26:275 1742<br />
328 ari-e ari-e 5HL Breviaristatum-e 26:276 1653<br />
329 ari-h ari-h 5HL Breviaristatum-h 26:277 1656<br />
330 ert-g ert-g, br3 5HL Erectoides-g 26:278 479<br />
163
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
331 ert-n ert-n 5HL Erectoides-n 26:279 488<br />
332 Ert-r Ert-r Erectoides-r 26:280 492<br />
333 Rph12 Rph12 5HL Reaction to Puccinia hordei 12 26:281 1590<br />
334 raw6 r6 5HL Smooth awn 6 26:282 2437<br />
335 msg49 msg,,jw 5HL Male sterile genetic 49 26:283 2402<br />
336 glo-b glo-b 5HL Globosum-b 26:284 1326<br />
337 blf2 bb2, nlh 5HL Broad leaf 2 26:285 1667<br />
338 lys1 lys 5HL High lysine 1 26:286 1784<br />
339 lys3 sex3 5HL High lysine 3 26:287 1785<br />
340 raw2 r2 5HL Smooth awn 2 26:289 27<br />
341 abo12 alb,,o 5HS Albino seedling 12 26:290 583<br />
342 glo-f glo-e 5HL Globosum-f 26:291<br />
343 Lfb1 Lfb 5HL Leafy bract 1 28: 30 1577<br />
344 var2 va2 5HL Variegated 2 32:104 2496<br />
345 rym3 ym3 5HS Reaction to barley yellow mosaic virus 3 32:105<br />
346 yst5 yst5 5HL Yellow streak 5 32:107 2501<br />
347 mnd4 m4 5HL Many noded dwarf 4 32:108 1798<br />
348 Eam5 Ea5 5HL Early maturity 5 37:260<br />
349 brh4 brh.j 2HL Brachytic 4 37:262 1675<br />
350 brh6 brh.s 5HS Brachytic 6 37:263 1683<br />
351 gsh1 gs1, cer-q 2HS Glossy sheath 1 26:292 735<br />
352 gsh2 gs2, cer-b 3HL Glossy sheath 2 26:294 736<br />
353 gsh3 gs3, cer-a 7HS Glossy sheath 3 26:296 737<br />
354 gsh4 gs4, cer-x 6HL Glossy sheath 4 26:298 738<br />
355 gsh5 gs5, cer-s 2HL Glossy sheath 5 26:300 739<br />
356 gsh6 gs6, cer-c 2HS Glossy sheath 6 26:302 740<br />
357 msg1 ms1 1HL Male sterile genetic 1 26:304 1810<br />
358 msg2 ms2 2HL Male sterile genetic 2 26:306 2371<br />
359 msg3 ms3 2HS Male sterile genetic 3 26:307 1130<br />
360 msg4 ms4 1H Male sterile genetic 4 26:308 2392<br />
361 msg5 ms5 3HS Male sterile genetic 5 26:309 2403<br />
362 msg6 ms6 6HS Male sterile genetic 6 26:310 2405<br />
363 msg7 ms7 5HL Male sterile genetic 7 26:311 2406<br />
364 msg8 ms8 5HL Male sterile genetic 8 26:312 2407<br />
365 msg9 ms9 2HS Male sterile genetic 9 26:313 2408<br />
366 msg10 ms10 7HS Male sterile genetic 10 26:314 1811<br />
367 msg11 ms11 Male sterile genetic 11 26:315 1812<br />
368 msg13 ms13 Male sterile genetic 13 26:316 1813<br />
369 msg14 ms14 7HS Male sterile genetic 14 26:317 1814<br />
370 msg15 ms15 Male sterile genetic 15 26:318 1815<br />
164
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
371 msg16 ms16 5HS Male sterile genetic 16 26:319 1816<br />
372 msg17 ms17 Male sterile genetic 17 26:320 1817<br />
373 msg18 ms18 5HL Male sterile genetic 18 26:321 1818<br />
374 msg19 ms19 5HS Male sterile genetic 19 26:322 1819<br />
375 msg20 ms20 1H Male sterile genetic 20 26:323 2372<br />
376 msg21 ms21 Male sterile genetic 21 26:324 2373<br />
377 seg1 se1 7HL Shrunken endosperm genetic 1 37:264 750<br />
378 seg2 se2 7HS Shrunken endosperm genetic 2 26:326 751<br />
379 seg3 se3 3H Shrunken endosperm genetic 3 37:265 752<br />
380 seg4 se4 7HL Shrunken endosperm genetic 4 37:267 753<br />
381 seg5 se5 7HS Shrunken endosperm genetic 5 26:329 754<br />
382 sex1 lys5 6HL Shrunken endosperm xenia 1 26:330 755<br />
383 msg22 ms22 7H Male sterile genetic 22 26:331 741<br />
384 msg23 ms23 7HL Male sterile genetic 23 26:332 2375<br />
385 msg24 ms24 4HL Male sterile genetic 24 26:333 2376<br />
386 des3 des3 Desynapsis 3 26:334 594<br />
387 des8 des8 Desynapsis 8 26:335 599<br />
388 des9 des9 Desynapsis 9 26:336 600<br />
389 des10 des10 Desynapsis 10 26:337 601<br />
390 des11 des11 Desynapsis 11 26:338 602<br />
391 des12 des12 Desynapsis 12 26:339 603<br />
392 des13 des13 Desynapsis 13 26:340 604<br />
393 des14 des14 Desynapsis 14 26:341 605<br />
394 des15 des15 Desynapsis 15 26:342 606<br />
395 msg26 msg,,u 7HS Male sterile genetic 26 26:343 745<br />
396 seg6 se6 3HL Shrunken endosperm genetic 6 37:268 2467<br />
397 seg7 se7 Shrunken endosperm genetic 7 37:269 2468<br />
399 cer-d cer-d Eceriferum-d 26:346 425<br />
400 cer-e cer-e 1HL Eceriferum-e 26:347 1518<br />
401 cer-f cer-f 7HS Eceriferum-f 26:348 427<br />
402 cer-g cer-g 2HL Eceriferum-g 26:349 428<br />
403 cer-h cer-h Eceriferum-h 26:351 429<br />
404 cer-i cer-i 5HL Eceriferum-i 26:352 430<br />
405 cer-k cer-k 7HS Eceriferum-k 26:354 432<br />
406 cer-l cer-l Eceriferum-l 26:355 433<br />
407 cer-m cer-m Eceriferum-m 26:356 434<br />
408 cer-n gs9 2HL Eceriferum-n 26:357 435<br />
409 cer-o cer-o Eceriferum-o 26:359 436<br />
410 cer-p cer-p Eceriferum-p 26:360 437<br />
411 cer-r cer-r 3HL Eceriferum-r 26:361 439<br />
165
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
412 cer-t cer-t 5HL Eceriferum-t 26:362 441<br />
413 gsh8 cer-u, gs8 2HS Glossy sheath 8 26:364 442<br />
414 cer-v cer-v 2HS Eceriferum-v 26:366 443<br />
415 cer-w cer-w 5HL Eceriferum-w 26:367 1519<br />
417 cer-y cer-y Eceriferum-y 26:368 446<br />
418 cer-z cer-z 7HS Eceriferum-z 26:369 447<br />
419 cer-za cer-za 5HL Eceriferum-za 26:370 1521<br />
420 cer-zb cer-zb Eceriferum-zb 26:371 1522<br />
421 cer-zc cer-zc Eceriferum-zc 26:372 450<br />
422 cer-zd cer-zd 3HL Eceriferum-zd 26:373 451<br />
423 cer-ze gl5 7HS Eceriferum-ze 26:374 452<br />
424 cer-zf cer-zf Eceriferum-zf 26:376 453<br />
425 cer-zg cer-zg 4HL Eceriferum-zg 26:377 454<br />
427 cer-zi cer-zi 1HL Eceriferum-zi 26:378 456<br />
428 cer-zj cer-zj 5HL Eceriferum-zj 26:379 457<br />
429 cer-zk cer-zk 2H Eceriferum-zk 26:381 458<br />
430 cer-zl cer-zl Eceriferum-zl 26:382 459<br />
431 cer-zn cer-zn 3HL Eceriferum-zn 26:383 1523<br />
432 cer-zo cer-zo Eceriferum-zo 26:384 462<br />
433 cer-zp cer-zp 5HL Eceriferum-zp 26:385 463<br />
434 cer-zq cer-zq Eceriferum-zq 26:386 1524<br />
435 cer-zr cer-zr Eceriferum-zr 26:387 1525<br />
436 cer-zs cer-zs Eceriferum-zs 26:388 1526<br />
437 cer-zt cer-zt 2HS Eceriferum-zt 37:270 1527<br />
438 cer-zu cer-zu Eceriferum-zu 26:390 1528<br />
439 cer-zv cer-zv Eceriferum-zv 26:391 1529<br />
440 cer-zw cer-zw Eceriferum-zw 26:392 1530<br />
441 cer-zx cer-zx Eceriferum-zx 26:393 1531<br />
442 cer-zy cer-zy Eceriferum-zy 26:394 1532<br />
443 cer-zz cer-zz Eceriferum-zz 26:395 1533<br />
444 cer-ya cer-ya 3HS Eceriferum-ya 26:396 1534<br />
445 cer-yb cer-yb 2HL Eceriferum-yb 26:397 1535<br />
446 cer-yc cer-yc Eceriferum-yc 26:398 1536<br />
447 cer-yd cer-yd 3HS Eceriferum-yd 26:399 1537<br />
448 cer-ye cer-ye 5HL Eceriferum-ye 26:400 1538<br />
449 cer-yf cer-yf Eceriferum-yf 37:271 1539<br />
450 cer-yg cer-yg 7HS Eceriferum-yg 26:402 1540<br />
451 cer-yh cer-yh 3HS Eceriferum-yh 26:403 1541<br />
454 blx5 bl5 7HL Non-blue aleurone xenia 5 26:404 2509<br />
455 seg8 seg8 7H Shrunken endosperm genetic 8 37:272 2469<br />
166
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
460 cur4 cu4, glo-d 2HL Curly 4 26:406 1708<br />
461 zeb2 zb2, f10 4HL Zebra stripe 2 26:407 93<br />
462 yst3 yst,,c 3HS Yellow streak 3 26:409 48<br />
463 gig1 gig, sf 2H? Gigas 1 26:410 1650<br />
464 msg27 msg,,ae 2HL Male sterile genetic 27 26:411 2379<br />
465 msg28 msg,,as 6H Male sterile genetic 28 26:412 2380<br />
466 msg29 msg,,a 5HL Male sterile genetic 29 26:413 2381<br />
467 msg30 msg,,c 7HL Male sterile genetic 30 26:414 2382<br />
468 msg31 msg,,d 1HS Male sterile genetic 31 26:415 2383<br />
469 msg32 msg,,w 7H Male sterile genetic 32 26:416 2384<br />
470 msg33 msg,,x 2HS Male sterile genetic 33 26:417 2385<br />
471 msg34 msg,,av 6H Male sterile genetic 34 26:418 2386<br />
472 abr1 abr 2HL Accordion basal rachis internode 1 26:419 1563<br />
473 com1 bir1 5HL Compositum 1 26:420 1702<br />
474 lax-a lax-a 5HL Laxatum-a 37:273 1775<br />
475 lax-c lax-c 6HL Laxatum-c 26:423 1777<br />
498 msg35 msg,,dr 2HL Male sterile genetic 35 26:424 2387<br />
499 msg36 msg,,bk 6HS Male sterile genetic 36 26:425 2388<br />
500 msg37 msg,,hl Male sterile genetic 37 26:426 2389<br />
501 msg38 msg,,jl Male sterile genetic 38 26:427 2390<br />
502 msg39 msg,,dm 6H Male sterile genetic 39 26:428 2391<br />
503 msg40 msg,,ac 6H Male sterile genetic 40 26:429 2393<br />
504 msg41 msg,,aj Male sterile genetic 41 26:430 2394<br />
505 msg42 msg,,db 3H Male sterile genetic 42 26:431 2395<br />
506 msg43 msg,,br Male sterile genetic 43 26:432 2396<br />
507 msg44 msg,,cx Male sterile genetic 44 26:433 2397<br />
508 msg45 msg,,dp Male sterile genetic 45 26:434 2398<br />
509 msg46 msg,,ec Male sterile genetic 46 26:435 2399<br />
510 msg47 msg,,ep Male sterile genetic 47 26:436 2400<br />
511 Rpg1 T 7HS Reaction to Puccinia graminis 1 26:437 701<br />
512 Rpg2 T2 Reaction to Puccinia graminis 2 26:439 187<br />
513 xnt2 xb Xantha seedling 2 26:440 2<br />
515 Rsp1 Sep Reaction to Septoria passerinii 1 26:441 2510<br />
516 Rsp2 Sep2 Reaction to Septoria passerinii 2 37:275 2511<br />
517 Rsp3 Sep3 Reaction to Septoria passerinii 3 37:276 2512<br />
518 sdw1 denso 3HL Semidwarf 1 37:277 2513<br />
519 mnd1 m Many-noded dwarf 1 26:446 253<br />
520 msg48 msg,,jt 2H Male sterile genetic 48 26:447 2401<br />
521 mtt1 mt 1HS Mottled leaf 1 26:448 622<br />
522 cer-yi cer-yi Eceriferum-yi 26:449 1542<br />
167
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
523 cer-yj cer-yj Eceriferum-yj 26:450 1543<br />
524 cer-yk cer-yk Eceriferum-yk 26:451 1544<br />
525 cer-yl cer-yl Eceriferum-yl 26:452 1545<br />
526 cer-ym cer-ym Eceriferum-ym 26:453 1546<br />
527 cer-yn cer-yn Eceriferum-yn 26:454 1547<br />
528 cer-yo cer-yo Eceriferum-yo 26:455 1548<br />
529 cer-yp cer-yp Eceriferum-yp 26:456 1549<br />
530 cer-yq cer-yq Eceriferum-yq 26:457 1550<br />
531 cer-yr cer-yr Eceriferum-yr 26:458 1551<br />
532 cer-ys cer-ys Eceriferum-ys 26:459 1552<br />
533 cer-yt cer-yt Eceriferum-yt 26:460 1553<br />
534 cer-yu cer-yu Eceriferum-yu 26:461 1554<br />
535 cer-yx cer-yx Eceriferum-yx 26:462 1555<br />
536 Cer-yy Gle1 1HS Eceriferum-yy 26:463 1556<br />
537 cer-yz cer-yz Eceriferum-yz 26:464 1557<br />
538 cer-xa cer-xa Eceriferum-xa 26:465 1558<br />
539 cer-xb cer-xb Eceriferum-xb 26:466 1559<br />
540 cer-xc cer-xc Eceriferum-xc 26:467 1560<br />
541 cer-xd cer-xd Eceriferum-xd 26:468 1561<br />
542 Dwf2 Dwf2 Dominant dwarf 2 24:170<br />
543 int-f int-f Intermedium spike-f 26:469 1767<br />
544 int-h int-h Intermedium spike-h 26:470 1768<br />
545 int-i int-i Intermedium spike-i 26:471 1769<br />
546 int-k int-k 7H Intermedium spike-k 37:279 1770<br />
547 int-m int-m Intermedium spike-m 37:280 1772<br />
548 Fol-b Ang Angustifolium-b 26:474 17<br />
549 Lga1 Log Long glume awn 1 26:475 835<br />
550 ari-b ari-b Breviaristatum-b 26:476 1649<br />
551 ari-f ari-f Breviaristatum-f 26:477 1654<br />
552 ari-j ari-j Breviaristatum-j 26:478 1658<br />
553 ari-k ari-k Breviaristatum-k 26:479 1659<br />
554 ari-m ari-m Breviaristatum-m 26:480 1661<br />
555 ari-n ari-n Breviaristatum-n 26:481 1662<br />
556 ari-o ari-o Breviaristatum-o 26:482<br />
557 ari-p ari-p Breviaristatum-p 26:483 1664<br />
558 ari-q ari-q Breviaristatum-q 26:484 1665<br />
559 ari-r ari-r Breviaristatum-r 26:485 1666<br />
560 ert-f ert-f Erectoides-f 26:486 478<br />
561 ert-h ert-h Erectoides-h 26:487 481<br />
562 ert-k ert-k Erectoides-k 26:488 485<br />
168
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
563 ert-l ert-l Erectoides-l 26:489 486<br />
564 ert-p ert-p Erectoides-p 26:490 490<br />
565 ert-s ert-s Erectoides-s 26:491 493<br />
566 ert-t ert-t, brh3 2HS Erectoides-t 37:281 494<br />
567 ert-v ert-v Erectoides-v 26:493 497<br />
568 ert-x ert-x Erectoides-x 26:494 498<br />
569 ert-y ert-y Erectoides-y 26:495 499<br />
570 ert-z ert-z Erectoides-z 26:496 500<br />
571 ert-za ert-za Erectoides-za 26:497 501<br />
572 ert-zb ert-zb Erectoides-zb 26:498 502<br />
573 ert-zc ert-zc Erectoides-zc 26:499 503<br />
574 ert-ze ert-ze Erectoides-ze 26:500 505<br />
575 Rph6 Pa6 Reaction to Puccinia hordei 6 26:501 1598<br />
576 Rph8 Pa8 Reaction to Puccinia hordei 8 26:502 1600<br />
577 Rsg2 Rsg2 Reaction to Schizaphis graminum 2 37:283 2513<br />
578 mat-b mat-b Praematurum-b 26:504 1788<br />
579 mat-c mat-c Praematurum-c 26:506 1789<br />
580 mat-d mat-d Praematurum-d 26:507 1790<br />
581 mat-e mat-e Praematurum-e 26:508 1791<br />
582 mat-f mat-f Praematurum-f 26:509 1792<br />
583 mat-g mat-g Praematurum-g 26:510 1793<br />
584 mat-h mat-h Praematurum-h 26:511 1794<br />
585 mat-i mat-i Praematurum-i 26:512 1795<br />
586 bra-d bra-d 1HL Bracteatum-d 37:284 1696<br />
587 abo3 a2, alb-za Albino seedling 3 26:514 165<br />
588 abo10 at2 Albino seedling 10 26:515 57<br />
589 abo11 at3, alb t Albino seedling 11 26:516 233<br />
590 Rph13 Rph13 Reaction to Puccinia hordei 13 28: 31 1591<br />
591 Rph14 Rph14 Reaction to Puccinia hordei 14 28: 32 1592<br />
592 yhd2 yh2 Yellow head 2 28: 33 757<br />
593 adp1 adp Awned palea 1 37:285 1618<br />
594 ant3 rub Anthocyanin-deficient 3 29: 82 1641<br />
595 ant4 ant4 Anthocyanin-deficient 4 29: 83 1642<br />
596 ant5 rs2 Anthocyanin-deficient 5 29: 84 1643<br />
597 ant6 ant6 Anthocyanin-deficient 6 29: 85 1644<br />
598 ant13 ant13 6HL Proanthocyanin-free 13 29: 86 1624<br />
599 ant17 ant17 3HS Proanthocyanin-free 17 37:286<br />
600 ant18 ant18 7HL Proanthocyanin-free 18 29: 90 1630<br />
601 ant19 ant19 Proanthocyanin-free 19 29: 92 1631<br />
602 ant20 ant20 Anthocyanin-rich 20 29: 93 1633<br />
169
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
603 ant21 ant21 6H Proanthocyanin-free 21 29: 94 1634<br />
604 ant22 ant22 7HL Proanthocyanin-free 22 29: 95 1635<br />
605 ant25 ant25 Proanthocyanin-free 25 29: 96 1638<br />
606 ant26 ant26 Proanthocyanin-free 26 29: 97 1639<br />
607 ant27 ant27 Proanthocyanin-free 27 29: 98 1640<br />
608 ant28 ant28 3HL Proanthocyanin-free 28 29: 99<br />
609 ant29 ant29 Proanthocyanin-free 29 29:100<br />
610 ant30 ant30 Proanthocyanin-free 30 29:101<br />
611 Nec6 Sp Necrotic leaf spot 6 32:112 2424<br />
612 gig2 gig2 Gigas 2 32:113 1750<br />
613 brc1 brc-5 2HS Branched 1 32:114<br />
614 Zeo2 Zeo2 Zeocriton 2 32:115 637<br />
615 wxs1 wxs1 Waxy spike 1 32:116<br />
616 cul3 cul3 Uniculme 3 32:117 2494<br />
617 cul4 uc-5 3HL Uniculme 4 37:289 2493<br />
618 mnd3 mn3, m3 3H Many noded dwarf 3 32:119 1797<br />
619 bra-a bra-a 7HS Bracteatum-a 32:120 1693<br />
620 cal-b cal-b 5H Calcaroides-b 32:121 1697<br />
621 Cal-c Cal-c 5HL Calcaroides-c 32:122 1567<br />
622 cal-e cal-23 5HS Calcaroides-e 32:123<br />
623 eli-a lig-a Eligulum-a 37:290<br />
624 ops1 op-3 Opposite spikelets 1 32:125 2427<br />
625 sci-a sci-3 Scirpoides 1 32:126<br />
626 scl-a scl-6 Scirpoides leaf-a 32:127<br />
627 viv-a viv-5 Viviparoides-a 32:128 2498<br />
628 sex7 sex.i 5HL Shrunken endosperm 7 32:129 2470<br />
629 mtt6 mtt6 Mottled leaf 6 32:130 2411<br />
630 Ari-s ari-265 Breviaristatum-s 32:131<br />
631 brh3 brh.g, ert-t Brachytic 3 32:132 1672<br />
632 mnd5 mnd5 Many noded dwarf 5 32:133<br />
633 mnd6 den-6 5HL Many noded dwarf 6 37:291 1713<br />
634 pmr2 nec-50 Premature ripe 2 32:135 2421<br />
635 nec7 nec-45 Necroticans 7 32:136 2420<br />
636 tst2 lin2 Tip sterile 2 37:292 1781<br />
637 nar1 nar1 6HS NADH nitrate reductase-deficient 1 35:194<br />
638 nar2 nar2 5HL NADH nitrate reductase-deficient 2 35:195<br />
639 nar3 nar3 7HS NADH nitrate reductase-deficient 3 35:196<br />
640 nar4 nar4 2HL NADH nitrate reductase-deficient 4 35:197<br />
641 nar5 nar5 5HL NADH nitrate reductase-deficient 5 35:198<br />
642 nar6 nar6 2HL NADH nitrate reductase-deficient 6 35:199<br />
170
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 2 (continued)<br />
BGS Locus symbol * Chr. † Locus name or phenotype Descr. GSHO<br />
no. Rec. Prev. loc. vol. p. no. ‡<br />
643 nar7 nar7 6HL NADH nitrate reductase-deficient 7 35:200<br />
644 nar8 nar8 6HS NADH nitrate reductase-deficient 8 35:201<br />
645 bsp1 bsp1 Bushy spike 1 35:202<br />
646 ovl2 ovl2 Ovaryless 2 35:204<br />
647 tst1 tst1 Tip sterile 1 35:205<br />
648 mov4 mo8 Multiovary 4 35:206<br />
649 asp1 asp1 Aborted spike 1 35:207<br />
650 sun1 sun1 Sensitivity to Ustilago nuda 1 35:208<br />
651 lam1 lam1 Late maturity 1 35:209<br />
652 ylf1 ylf1 Yellow leaf 1 35:210<br />
653 brh10 brh.l 2HS Brachytic 10 37:293 1677<br />
654 brh11 brh.n 5HS Brachytic 11 37:294 1679<br />
655 brh12 brh.o 5HS Brachytic 12 37:295 1680<br />
656 brh13 brh.p 5HS Brachytic 13 37:296 1681<br />
657 brh15 brh.u Brachytic 15 37:297 1685<br />
658 brh17 brh.ab 5HS Brachytic 17 37:298 1669<br />
659 brh18 brh.ac 5HS Brachytic 18 37:299 1670<br />
660 nld2 Narrow leafed dwarf 2 37:300<br />
661 dub1 5HL Double seed 1 37:301<br />
* Recommended locus symbols are based on utilization of a three-letter code for barley genes as<br />
approved at the business meeting of the Seventh International Barley Genetics Symposium at<br />
Saskatoon, Saskatchewan, Canada, on 05 August 5 1996.<br />
† Chromosome numbers and arm designations are based on a resolution passed at the business<br />
meeting of the Seventh International Barley Genetics Symposium at Saskatoon, Saskatchewan,<br />
Canada, on August 05 1996. The Burnham and Hagberg (1956) designations of barley<br />
chromosomes were 1 2 3 4 5 6 and 7 while new designations based on the Triticeae system are<br />
7H 2H 3H 4H 1H 6H and 5H, respectively.<br />
‡ The seed stock associated with each BGS number is held as a GSHO stock number in the<br />
Barley Genetics Stock Collection at the <strong>US</strong>DA-ARS National Small Grains Germplasm<br />
Research Facility, Aberdeen, Idaho, <strong>US</strong>A.<br />
171
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3. An alphabetic listing of recently published Barley Genetic Stock (BGS) descriptions for<br />
loci in barley (Hordeum vulgare), including information on chromosomal locations,<br />
recommended locus names, and original cultivars.<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
abo1 at 207 1HL Albino seedling 1 26:210 Trebi<br />
abo2 a2 53 2HS Albino seedling 2 26:89 Nilsson-Ehle No 2<br />
abo3 alb-za 587 Albino seedling 3 26:514 Unknown cultivar<br />
abo4 a4 94 2H Albino seedling 4 26:133 Unknown cultivar<br />
abo6 ac 106 3HS Albino seedling 6 26:140 Colsess<br />
abo8 ac2 4 7HS Albino seedling 8 26:47 Coast<br />
abo9 an 112 3HS Albino seedling 9 26:146 Nigrinudum<br />
abo10 at2 588 Albino seedling 10 26:515 Canadian Thorpe<br />
abo11 at3 589 Albino seedling 11 26:516 Trebi<br />
abo12 alb,,o 341 5HS Albino seedling 12 26:290 Titan<br />
abo13 alb,,p 95 2HL Albino seedling 13 26:134 Titan<br />
abo14 alb,,q 270 6HL Albino seedling 14 26:250 Shabet<br />
abo15 alb,,t 271 6HS Albino seedling 15 26:251 Betzes<br />
abr1 abr 472 2HL Accordion basal rachis 26:419 Bonus<br />
internode 1<br />
acr1 acr 97 2HL Accordion rachis 1 32:85 Burma Girl<br />
adp1 adp 593 3HL Awned palea 1 37:285 Unknown cultivar<br />
alm1 al 108 3HS Albino lemma 1 37:222 Russia 82<br />
als1 als 101 3HL Absent lower laterals 1 37:219 Montcalm<br />
ant1 rs 33 7HS Anthocyanin-less 1 26:82 Bonus<br />
ant2 pr 80 2HL Anthocyanin-less 2 26:118 Foma<br />
ant3 594 Anthocyanin-deficient 3 29:82 Bonus<br />
ant4 595 Anthocyanin-deficient 4 29:83 Foma<br />
ant5 596 Anthocyanin-deficient 5 29:84 Bonus<br />
ant6 597 Anthocyanin-deficient 6 29:85 Foma<br />
ant13 598 6HL Proanthocyanidin-free 13 29:86 Foma<br />
ant17 599 3HS Proanthocyanidin-free 17 37:286 Nordal<br />
ant18 600 7HL Proanthocyanidin-free 18 29:90 Nordal<br />
ant19 601 Proanthocyanidin-free 19 29:92 Alf<br />
ant20 602 Anthocyanidin-rich 20 29:93 Foma<br />
ant21 603 6H Proanthocyanidin-free 21 29:94 Georgie<br />
ant22 604 7HS Proanthocyanidin-free 22 29:95 Hege 802<br />
ant25 605 Proanthocyanidin-free 25 29:96 Secobra 18193<br />
ant26 606 Proanthocyanidin-free 26 29.97 Grit<br />
ant27 607 Proanthocyanidin-free 27 29:98 Zebit<br />
ant28 608 3HL Proanthocyanidin-free 28 29:99 Grit<br />
ant29 609 Proanthocyanidin-free 29 29:100 Ca 708912<br />
ant30 610 Proanthocyanidin-free 30 29:101 Gunhild<br />
172
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
ari-a 132 3HS Breviaristatum-a 26:168 Bonus<br />
ari-b 550 Breviaristatum-b 26:476 Bonus<br />
ari-e 328 5HL Breviaristatum-e 26:276 Bonus<br />
ari-f 551 Breviaristatum-f 26:477 Bonus<br />
ari-g 89 Breviaristatum-g 26:128 Bonus<br />
ari-h 329 5HL Breviaristatum-h 26:277 Foma<br />
ari-j 552 Breviaristatum-j 26:478 Bonus<br />
ari-k 553 Breviaristatum-k 26:479 Bonus<br />
ari-m 554 Breviaristatum-m 26:480 Bonus<br />
ari-n 555 Breviaristatum-n 26:481 Bonus<br />
ari-o 556 Breviaristatum-o 26:482 Bonus<br />
ari-p 557 Breviaristatum-p 26:483 Foma<br />
ari-q 558 Breviaristatum-q 26:484 Kristina<br />
ari-r 559 Breviaristatum-r 26:485 Bonus<br />
Ari-s ari-265 630 Breviaristatum-s 32:131 Kristina<br />
asp1 649 Aborted spike 1 35:207 Steptoe<br />
blf1 bb 326 5HL Broad leaf 1 26:274 Bonus<br />
blf2 bb2 337 5HL Broad leaf 2 26:285 Hannchen<br />
Blp1 B 203 1HL Black lemma and pericarp 1 37:245 Nigrinudum<br />
blx1 bl 15 4HL Non-blue aleurone xenia 1 26:60 Goldfoil<br />
blx2 bl2 19 7HS Non-blue aleurone xenia 2 26:65 Nepal<br />
blx3 bl3 173 4HL Non-blue aleurone xenia 3 26:198 Blx<br />
blx4 bl4 174 4HL Non-blue (pink) aleurone 26:199 Ab 6<br />
xenia 4<br />
blx5 bl5 454 7HL Non-blue aleurone xenia 5 26:404 BGM 122<br />
bra-a 619 7HS Bracteatum-a 32:120 Bonus<br />
bra-d 586 1HS Bracteatum-d 37:284 Foma<br />
brc1 brc-5 613 2HS Branched 1 32:114<br />
brh1 br 1 7HS Brachytic 1 37:188 Himalaya<br />
brh2 br2 157 4HL Brachytic 2 37:235 Svanhals<br />
brh3 brh.g, ert-t 631 Brachytic 3 32:132 Birgitta<br />
brh4 brh.j 349 5HS Brachytic 4 37:262 Birgitta<br />
brh5 brh.m 185 4HS Brachytic 5 37:242 Birgitta<br />
brh6 brh.s 350 5HS Brachytic 6 37:263 Akashinriki<br />
brh7 brh.w 41 5HS Brachytic 7 37:203 Volla<br />
brh8 brh.ad 142 3HS Brachytic 8 37:230 Birgitta<br />
brh9 brh.k 187 4HS Brachytic 9 37:244 Birgitta<br />
brh10 brh.l 653 2HS Brachytic 10 37:293 Birgitta<br />
brh11 brh.n 654 5HS Brachytic 11 37:294 Birgitta<br />
brh12 brh.o 655 5HS Brachytic 12 37:295 Birgitta<br />
173
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
brh13 brh.p 656 5HS Brachytic 13 37:296 Birgitta<br />
brh14 brh.q 148 3HL Brachytic 14 37:231 Akashinriki<br />
brh15 brh.u 657 Brachytic 15 37:297 Julia<br />
brh16 brh.v 44 7HL Brachytic 16 37:204 Korál<br />
brh17 brh.ab 658 5HS Brachytic 17 37:298 Morex<br />
brh18 Brh.ac 659 5HS Brachytic 18 37:299 Triumph<br />
bsp1 645 Bushy spike 1 35:203 Morex<br />
btr1 bt 115 3HS Non-brittle rachis 1 26:149 A 222<br />
btr2 bt2 116 3HS Non-brittle rachis 2 26:150 Sakigoke<br />
cal-b 620 5H Calcaroides-b 32:121 Bonus<br />
Cal-c 621 5HL Calcaroides-c 32:122 Bonus<br />
cal-d 146 3H Calcaroides-d 32:95 Foma<br />
cal-e 622 5HS Calcaroides-e 32:123 Semira<br />
cer-d 399 Eceriferum-d + ++ ++ 26:346 Bonus<br />
cer-e 400 1HL Eceriferum-e -/+ ++ ++ 26:347 Bonus<br />
cer-f 401 7HS Eceriferum-f + + ++ 26:348 Bonus<br />
cer-g 402 2HL Eceriferum-g + + ++ 26:349 Bonus<br />
cer-h 403 Eceriferum-h - ++ ++ 26:351 Bonus<br />
cer-i 404 5HL Eceriferum-i - ++ ++ 26:352 Bonus<br />
cer-k 405 7HS Eceriferum-k + ++ ++ 26:354 Bonus<br />
cer-l 406 Eceriferum-l + ++ ++ 26:355 Bonus<br />
cer-m 407 Eceriferum-m +/++ + ++ 26:356 Bonus<br />
cer-n gs9 408 2HL Eceriferum-n - - ++ & 26:357 Bonus<br />
- +/- ++<br />
cer-o 409 Eceriferum-o -/+ ++ ++ 26:359 Bonus<br />
cer-p 410 Eceriferum-p ++ ++ + 26:360 Bonus<br />
cer-r 411 3HL Eceriferum-r +/- + ++ 26:361 Bonus<br />
cer-t 412 5HL Eceriferum-t +/- ++ ++ 26:362 Bonus<br />
cer-v 414 2HS Eceriferum-v +/- ++ ++ 26:366 Bonus<br />
cer-w 415 5HL Eceriferum-w +/- ++ ++ 26:367 Bonus<br />
cer-y 417 Eceriferum-y + +/++ ++ 26:368 Bonus<br />
cer-z 418 7HS Eceriferum-z - - ++ 26:369 Bonus<br />
cer-za 419 5HL Eceriferum-za ++ ++ - 26:370 Foma<br />
cer-zb 420 Eceriferum-zb - ++ ++ 26:371 Bonus<br />
cer-zc 421 Eceriferum-zc +/- ++ ++ 26:372 Bonus<br />
cer-zd 422 3HL Eceriferum-zd ++ ++ - 26:373 Bonus<br />
cer-ze gl5 423 7HS Eceriferum-ze ++ ++ - 26:374 Bonus<br />
cer-zf 424 Eceriferum-zf ++ ++ + 26:376 Bonus<br />
cer-zg 425 4HL Eceriferum-zg ++ ++ + 26:377 Foma<br />
cer-zi 427 1HL Eceriferum-zi + + ++ 26:378 Bonus<br />
174
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
cer-zj 428 5HL Eceriferum-zj ++ ++ - 26:379 Bonus<br />
cer-zk 429 2H Eceriferum-zk + + +/- 26:381 Bonus<br />
cer-zl 430 Eceriferum-zl - - ++ 26:382 Bonus<br />
cer-zn 431 3HL Eceriferum-zn +/- ++ ++ 26:383 Foma<br />
cer-zo 432 Eceriferum-zo - ++ ++ 26:384 Foma<br />
cer-zp 433 5HL Eceriferum-zp ++ ++ - 26:385 Bonus<br />
cer-zq 434 Eceriferum-zq ++ ++ - 26:386 Foma<br />
cer-zr 435 Eceriferum-zr +/- ++ ++ 26:387 Foma<br />
cer-zs 436 Eceriferum-zs + ++ ++ 26:388 Foma<br />
cer-zt 437 2HS Eceriferum-zt + ++ ++ 37:270 Foma<br />
cer-zu 438 Eceriferum-zu - + ++ 26:390 Bonus<br />
cer-zv 439 Eceriferum-zv - - - 26:391 Foma<br />
cer-zw 440 Eceriferum-zw + + ++ 26:392 Foma<br />
cer-zx 441 Eceriferum-zx + + ++ 26:393 Bonus<br />
cer-zy 442 Eceriferum-zy ++ ++ + 26:394 Bonus<br />
cer-zz 443 Eceriferum-zz ++ ++ - 26:395 Bonus<br />
cer-ya 444 3HS Eceriferum-ya ++ ++ - 26:396 Bonus<br />
cer-yb 445 2HL Eceriferum-yb ++ ++ - 26:397 Bonus<br />
cer-yc 446 Eceriferum-yc - ++ ++ 26:398 Bonus<br />
cer-yd 447 3HS Eceriferum-yd - ++ ++ 26:399 Bonus<br />
cer-ye 448 5HL Eceriferum-ye ++ ++ - 26:400 Foma<br />
cer-yf 449 Eceriferum-yf ++ ++ + 37:271 Bonus<br />
cer-yg 450 7HS Eceriferum-yg - - - 26:402 Carlsberg II<br />
cer-yh 451 3HS Eceriferum-yh - ++ ++ 26:403 Bonus<br />
cer-yi 522 Eceriferum-yi ++ ++ - 26:449 Foma<br />
cer-yj 523 Eceriferum-yj ++ ++ - 26:450 Bonus<br />
cer-yk 524 Eceriferum-yk + + ++ 26:451 Bonus<br />
cer-yl 525 Eceriferum-yl - - ++ 26:452 Bonus<br />
cer-ym 526 Eceriferum-ym - - - 26:453 Bonus<br />
cer-yn 527 Eceriferum-yn + + ++ 26:454 Kristina<br />
cer-yo 528 Eceriferum-yo ++ ++ + 26:455 Bonus<br />
cer-yp 529 Eceriferum-yp ++ ++ + 26:456 Bonus<br />
cer-yq 530 Eceriferum-yq ++ ++ - 26:457 Kristina<br />
cer-yr 531 Eceriferum-yr -/+ + ++ 26:458 Foma<br />
cer-ys 532 Eceriferum-ys ++ ++ - 26:459 Bonus<br />
cer-yt 533 Eceriferum-yt - ++ ++ 26:460 Bonus<br />
cer-yu 534 Eceriferum-yu ++ ++ - 26:461 Bonus<br />
cer-yx 535 Eceriferum-yx + + ++ 26:462 Foma<br />
Cer-yy Gle1 536 1HS Eceriferum-yy - ++ ++ 26:463 Bonus<br />
cer-yz 537 Eceriferum-yz + + ++ 26:464 Bonus<br />
175
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
cer-xa 538 Eceriferum-xa ++ ++ - 26:465 Foma<br />
cer-xb 539 Eceriferum-xb - ++ ++ 26:466 Bonus<br />
cer-xc 540 Eceriferum-xc + + ++ 26:467 Bonus<br />
cer-xd 541 Eceriferum-xd + + ++ 26:468 Bonus<br />
clh1 clh 225 1HL Curled leaf dwarf 1 26:223 Hannchen<br />
com1 bir1 473 5HL Compositum 1 26:420 Foma<br />
com2 bir2 71 2HS Compositum 2 26:108 CIMMYT freak<br />
crl1 cl 325 Curly lateral 1 26:273 Montcalm<br />
crm1 cm 305 5HL Cream seedling 1 26:256 Black Hulless<br />
cud1 cud 324 5HL Curly dwarf 1 26:272 Akashinriki<br />
cud2 229 1HL Curly dwarf 2 26:227 Akashinriki<br />
cul2 uc2 253 6HL Uniculm 2 37:253 Kindred<br />
cul3 616 Uniculme 3 32:117 Donaria<br />
cul4 uc-5 617 3HL Uniculme 4 37:289 Bonus<br />
cur1 cu1 262 6HL Curly 1 26:242 48-cr cr-17<br />
cur2 cu2 114 3HL Curly 2 26:148 Choshiro<br />
cur3 cu3 263 6HL Curly 3 26:243 Akashinriki<br />
cur4 cu4 460 2HL Curly 4 26:406 Asahi 5<br />
cur5 cu5 231 1HS Curly 5 26:229 Glenn<br />
ddt1 ddt 317 5HS Reaction to DDT 1 26:266 Spartan<br />
des1 lc 12 7H Desynapsis 1 26:57 Mars<br />
des2 ds 119 3H Desynapsis 2 26:154 Husky<br />
des3 386 Desynapsis 3 26:334 Betzes<br />
des4 13 7H Desynapsis 4 26:58 Betzes<br />
des5 14 7H Desynapsis 5 26:59 Betzes<br />
des6 215 1H Desynapsis 6 26:216 Betzes<br />
des7 64 2H Desynapsis 7 26:100 Betzes<br />
des8 387 Desynapsis 8 26:335 Betzes<br />
des9 388 Desynapsis 9 26:336 Betzes<br />
des10 389 Desynapsis 10 26:337 Betzes<br />
des11 390 Desynapsis 11 26:338 Betzes<br />
des12 391 Desynapsis 12 26:339 Betzes<br />
des13 392 Desynapsis 13 26:340 Betzes<br />
des14 393 Desynapsis 14 26:341 Betzes<br />
des15 394 Desynapsis 15 26:342 Ingrid<br />
dex1 sex2 311 5HS Defective endosperm xenia 1 26:260 BTT 63-j-18-17<br />
dsk1 dsk 322 5HL Dusky 1 26:270 Chikurin-Ibaraki 1<br />
dsp1 l 9 7HS Dense spike 1 26:53 Honen 6<br />
dsp9 l9, ert-e 258 6HL Dense spike 9 26:239 Akashinriki<br />
176
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
dsp10 lc 111 3HS Dense spike 10 26:145 Club Mariout<br />
dub1 661 6HL Double seed 1 37:301 Bonus<br />
Dwf2 542 Dominant dwarf 2 24:170 Klages/Mata<br />
Eam1 Ea 65 2HS Early maturity 1 26:101 Estate<br />
Eam5 Ea5 348 5HL Early maturity 5 37:260 Higuerilla*2/<br />
Gobernadora<br />
eam6 Ea6, Ea 98 2HS Early maturity 6 37:216 Morex<br />
eam7 ea7 252 6HS Early maturity 7 26:233 California Mariout<br />
eam8 eak, ert-o 214 1HL Early maturity 8 37:247 Kinai 5<br />
eam9 ea,,c 181 4HL Early maturity 9 26:204 Tayeh 8<br />
eam10 easp 130 3HL Early maturity 10 37:226 Super Precoz<br />
eli-a lig-a 623 Eligulum-a 37:290 Foma<br />
eog 1 e 57 2HL Elongated outer glume 1 26:92 Triple Bearded<br />
Club Mariout<br />
ert-a ert-6 28 7HS Erectoides-a 26:74 Gull<br />
ert-b ert-2 224 1HS Erectoides-b 26:222 Gull<br />
ert-c ert-1 134 3HL Erectoides-c 26:170 Gull<br />
ert-d ert-7 29 7HS Erectoides-d 26:76 Gull<br />
ert-e dsp9 266 6HL Erectoides-e 37:257 Bonus<br />
ert-f ert-18 560 Erectoides-f 26:486 Bonus<br />
ert-g ert-24 330 5HL Erectoides-g 26:278 Bonus<br />
ert-h ert-25 561 Erectoides-h 26:487 Bonus<br />
ert-ii ert-79 135 3HL Erectoides-ii 26:172 Bonus<br />
ert-j ert.31 90 2H Erectoides-j 26:129 Bonus<br />
ert-k ert-32 562 Erectoides-k 26:488 Bonus<br />
ert-l ert-12 563 Erectoides-l 26:489 Maja<br />
ert-m ert-34 30 7HS Erectoides-m 26:78 Bonus<br />
ert-n ert-51 331 5HL Erectoides-n 26:279 Bonus<br />
ert-p ert-44 564 Erectoides-p 26:490 Bonus<br />
ert-q ert-101 91 2H Erectoides-q 26:130 Bonus<br />
Ert-r Ert-52 332 Erectoides-r 26:280 Bonus<br />
ert-s ert-50 565 Erectoides-s 26:491 Bonus<br />
ert-t brh3 566 2HS Erectoides-t 37:281 Bonus<br />
ert-u ert-56 92 2H Erectoides-u 26:131 Bonus<br />
ert-v ert-57 567 Erectoides-v 26:493 Bonus<br />
ert-x ert-58 568 Erectoides-x 26:494 Bonus<br />
ert-y ert-69 569 Erectoides-y 26:495 Bonus<br />
ert-z ert-71 570 Erectoides-z 26:496 Bonus<br />
ert-za ert-102 571 Erectoides-za 26:497 Bonus<br />
ert-zb ert-132 572 Erectoides-zb 26:498 Bonus<br />
177
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
ert-zc ert-149 573 Erectoides-zc 26:499 Bonus<br />
ert-zd ert-159 93 2H Erectoides-zd 26:132 Bonus<br />
ert-ze ert-105 574 Erectoides-ze 26:500 Bonus<br />
fch1 f 55 2HS Chlorina seedling 1 26:90 Minn 84-7<br />
fch2 f2 117 3HL Chlorina seedling 2 26:151 28-3398<br />
fch3 f3 220 1HS Chlorina seedling 3 26:218 Minn 89-4<br />
fch4 f4 17 7HL Chlorina seedling 4 26:63 Montcalm<br />
fch5 f5 18 7HS Chlorina seedling 5 26:64 Gateway<br />
fch6 f6 313 5HL Chlorina seedling 6 26:262 Himalaya<br />
fch7 f7 201 1HL Chlorina seedling 7 26:206 Smyrna<br />
fch8 f8 5 7HS Chlorina seedling 8 26:48 Comfort<br />
fch9 f9 151 4HS Chlorina seedling 9 26:178 Ko A<br />
fch11 f11 260 6HL Chlorina seedling 11 26:240 Himalaya<br />
fch12 fc 2 7HS Chlorina seedling 12 37:190 Colsess<br />
fch13 f13 86 Chlorina seedling 13 26:124 Niggrinudum<br />
fch14 f14 87 2HL Chlorina seedling 14 37:211 Shyri<br />
fch15 or 52 2HS Chlorina seedling 15 26:88 Trebi IV<br />
flo-a 182 Extra floret-a 26:205 Foma<br />
flo-b 327 5HL Extra floret-b 26:275 Foma<br />
flo-c 74 2HS Extra floret-c 26:111 Foma<br />
fol-a 73 2HL Angustifolium-a 26:110 Proctor<br />
Fol-b Ang 548 Angustifolium-b 26:474 Unknown<br />
fst1 fs 301 5HL Fragile stem 1 26:252 Kamairazu<br />
fst2 fs2 208 1HL Fragile stem 2 26:211 Oshichi<br />
fst3 fs3 24 7HS Fragile stem 3 26:70 Kobinkatagi 4<br />
gig1 gig 463 2H? Gigas 1 26:410 Tochigi Golden<br />
Melon<br />
gig2 612 Gigas 2 32:113 ND12463<br />
glf1 gl 155 4HL Glossy leaf 1 ++ ++ - 37:233 Himalaya<br />
glf3 gl3 165 4HL Glossy leaf 3 ++ ++ - 26:190 Goseshikoku<br />
glo-a 168 4HS Globosum-a 26:194 Proctor<br />
glo-b 336 5HL Globosum-b 26:284 Villa<br />
glo-c 72 2H Globosum-c 26:109 Villa<br />
glo-e 230 1HL Globosum-e 26:228 Foma<br />
glo-f 342 5HL Globosum-f 26:291 Damazy<br />
gpa1 gp 59 2HL Grandpa 1 26:95 Lyallpur<br />
gra-a gran-a 131 3HL Granum-a 26:167 Donaria<br />
gsh1 gs1 351 2HS Glossy sheath 1 - - ++ 26:292 CIho 5818<br />
gsh2 gs2 352 3HL Glossy sheath 2 - - ++ 26:294 Atlas<br />
gsh3 gs3 353 7HS Glossy sheath 3 - - ++ 26:296 Mars<br />
178
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
gsh4 gs4 354 6HL Glossy sheath 4 - - ++ 26:298 Gateway<br />
gsh5 gs5 355 2HL Glossy sheath 5 + - ++ 26:300 Jotun<br />
gsh6 gs6 356 2HS Glossy sheath 6 - - ++ 26:302 Betzes<br />
gsh7 gs7 81 Glossy sheath 7 - - ++ 26:119 Akashinriki<br />
gsh8 cer-u 413 2HS Glossy sheath 8 + + ++ 26:364 Bonus<br />
Gth1 G 69 2HL Toothed lemma 1 26:106 Machine<br />
(Wexelsen)<br />
hcm1 h 77 2HL Short culm 1 26:115 Morex<br />
Hln1 Hn 164 4HL Hairs on lemma nerves 1 26:189 Kogane-mugi<br />
Hsh1 Hs 179 4HL Hairy leaf sheath 1 37:240 Kimugi<br />
int-b 320 5HL Intermedium spike-b 26:268 Bonus<br />
int-c i 178 4HS Intermedium spike-c 37:237 Gamma 4<br />
int-f 543 Intermedium spike-f 26:469 Foma<br />
int-h 544 Intermedium spike-h 26:470 Kristina<br />
int-i 545 Intermedium spike-i 26:471 Kristina<br />
int-k 546 7H Intermedium spike-k 37:279 Kristina<br />
int-m 547 Intermedium spike-m 37:280 Bonus<br />
Kap1 K 152 4HS Hooded lemma 1 26:179 Colsess<br />
lam1 651 Late maturity 1 35:209 Steptoe<br />
lax-a 474 5HL Laxatum-a 37:273 Bonus<br />
lax-b 268 6HL Laxatum-b 26:248 Bonus<br />
lax-c 475 6HL Laxatum-c 26:423 Bonus<br />
lbi1 lb 308 5HL Long basal rachis internode<br />
1<br />
26:258 Wisconsin 38<br />
lbi2 lb2 156 4HL Long basal rachis internode<br />
2<br />
26:183 Montcalm<br />
lbi3 lb3 27 7HL Long basal rachis internode 26:73 Montcalm<br />
3<br />
lel1 lel 235 1HL Leafy lemma 1 32:103 G7118<br />
Lfb1 Lfb 343 5HL Leafy bract 1 28:30 Montcalm<br />
Lga1 Log 549 Long glume awn 1 26:475 Guy Mayle<br />
lgn3 lg3 170 4HL Light green 3 26:195 No 154<br />
lgn4 lg4 171 4HL Light green 4 26:196 Himalaya /<br />
Ingrescens<br />
lig1 li 60 2HL Liguleless 1 37:205 Muyoji<br />
lin1 s, rin 99 2HL Lesser internode number 1 32:88 Natural occurence<br />
Lks1 Lk 75 2HL Awnless 1 26:112 Hordeum inerme<br />
lks2 lk2 10 7HL Short awn 2 37:197 Honen 6<br />
lks5 lk5 172 4HL Short awn 5 26:197 CIho 5641<br />
lnt1 lnt 118 3HL Low number of tillers 1 26:153 Mitake<br />
179
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
lys1 lys 338 5HL High lysine 1 26:286 Hiproly<br />
lys3 sex3 339 5HL High lysine 3 26:287 Bomi<br />
Lys4 sex5 232 1HS High lysine 4 26:230 Bomi<br />
lys6 269 6H High lysine 6 26:249 Bomi<br />
lzd1 lzd 125 3HS Lazy dwarf 1 26:161 Akashinriki<br />
mat-b 578 Praematurum-b 26:504 Bonus<br />
mat-c 579 Praematurum-c 26:506 Bonus<br />
mat-d 580 Praematurum-d 26:507 Bonus<br />
mat-e 581 Praematurum-e 26:508 Bonus<br />
mat-f 582 Praematurum-f 26:509 Bonus<br />
mat-g 583 Praematurum-g 26:510 Bonus<br />
mat-h 584 Praematurum-h 26:511 Bonus<br />
mat-i 585 Praematurum-i 26:512 Bonus<br />
min1 min 161 4HL Semi-minute dwarf 1 26:187 Taisho-mugi<br />
min2 en-min 160 Enhancer of minute 1 26:186 Kaiyo Bozu<br />
mnd1 m 519 Many-noded dwarf 1 26:446 Mesa<br />
mnd3 m3 618 3H Many noded dwarf 3 32:119 Montcalm<br />
mnd4 m4 347 5HL Many noded dwarf 4 32:108 Akashinriki<br />
mnd5 632 Many noded dwarf 5 32:133 C2-95-199<br />
mnd6 den-6 633 5HL Many noded dwarf 6 37:291 Bonus<br />
mov1 mo6b 43 7HL Multiovary 1 35:185 Steptoe<br />
mov2 mo7a 147 3HS Multiovary 2 35:190 Steptoe<br />
mov3 mo-a 234 1H Multiovary 3 32:102 Akashinriki<br />
mov4 648 Multiovary 4 35:206 Steptoe<br />
msg1 357 1HL Male sterile genetic 1 26:304 CIho 5368<br />
msg2 358 2HL Male sterile genetic 2 26:306 Manchuria<br />
msg3 359 2HS Male sterile genetic 3 26:307 Gateway<br />
msg4 360 1H Male sterile genetic 4 26:308 Freja<br />
msg5 361 3HS Male sterile genetic 5 26:309 Carlsberg II<br />
msg6 362 6HS Male sterile genetic 6 26:310 Hanna<br />
msg7 363 5HL Male sterile genetic 7 26:311 Dekap<br />
msg8 364 5HL Male sterile genetic 8 26:312 Betzes<br />
msg9 365 2HS Male sterile genetic 9 26:313 Vantage<br />
msg10 366 7HS Male sterile genetic 10 26:314 Compana<br />
msg11 367 Male sterile genetic 11 26:315 Gateway<br />
msg13 368 Male sterile genetic 13 26:316 Haisa II<br />
msg14 369 7HS Male sterile genetic 14 26:317 Unitan<br />
msg15 370 Male sterile genetic 15 26:318 Atlas/2*Kindred<br />
msg16 371 5HS Male sterile genetic 16 26:319 Betzes<br />
msg17 372 Male sterile genetic 17 26:320 Compana<br />
180
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
msg18 373 5HL Male sterile genetic 18 26:321 Compana<br />
msg19 374 5HS Male sterile genetic 19 26:322 CIho 14393<br />
msg20 375 1H Male sterile genetic 20 26:323 Hannchen<br />
msg21 376 Male sterile genetic 21 26:324 Midwest Bulk<br />
msg22 383 7H Male sterile genetic 22 26:331 Glacier/Compana<br />
msg23 384 7HL Male sterile genetic 23 26:332 Betzes<br />
msg24 385 4HL Male sterile genetic 24 26:333 Betzes<br />
msg25 166 4HL Male sterile genetic 25 26:192 Betzes<br />
msg26 395 7HS Male sterile genetic 26 26:343 Unitan<br />
msg27 464 2HL Male sterile genetic 27 26:411 Firlbecks III<br />
msg28 465 6H Male sterile genetic 28 26:412 York<br />
msg29 466 5HL Male sterile genetic 29 26:413 Ackermans MGZ<br />
msg30 467 7HL Male sterile genetic 30 26:414 Compana<br />
msg31 468 1HS Male sterile genetic 31 26:415 51Ab4834<br />
msg32 469 7H Male sterile genetic 32 26:416 Betzes<br />
msg33 470 2HS Male sterile genetic 33 26:417 Betzes<br />
msg34 471 6H Male sterile genetic 34 26:418 Paragon<br />
msg35 498 2HL Male sterile genetic 35 26:424 Karl<br />
msg36 499 6HS Male sterile genetic 36 26:425 Betzes<br />
msg37 500 Male sterile genetic 37 26:426 Clermont<br />
msg38 501 Male sterile genetic 38 26:427 Ingrid<br />
msg39 502 6H Male sterile genetic 39 26:428 CIho 15836<br />
msg40 503 6H Male sterile genetic 40 26:429 Conquest<br />
msg41 504 Male sterile genetic 41 26:430 Betzes<br />
msg42 505 3H Male sterile genetic 42 26:431 Betzes<br />
msg43 506 Male sterile genetic 43 26:432 Betzes<br />
msg44 507 Male sterile genetic 44 26:433 HA6-33-02<br />
msg45 508 Male sterile genetic 45 26:434 RPB439-71<br />
msg46 509 Male sterile genetic 46 26:435 Hector<br />
msg47 510 Male sterile genetic 47 26:436 Sel 12384CO<br />
msg48 520 2H Male sterile genetic 48 26:447 Simba<br />
msg49 335 5HL Male sterile genetic 49 26:283 ND7369<br />
msg50 34 7HL Male sterile genetic 50 26:83 Berac<br />
mss1 mss 84 2H Midseason stripe 1 26:122 Montcalm<br />
mss2 39 7HS Midseason stripe 2 32:79 ND11258<br />
mtt1 mt 521 1HS Mottled leaf 1 26:448 Montcalm<br />
mtt2 mt2 302 5HL Mottled leaf 2 26:253 Montcalm<br />
mtt4 mt,,e 78 2HL Mottled leaf 4 26:116 Victorie<br />
mtt5 mt,,f 264 6HL Mottled leaf 5 26:244 Akashinriki<br />
mtt6 629 Mottled leaf 6 32:130 ND6809<br />
181
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
mul2 251 6HL Multiflorus 2 26:232 Montcalm<br />
nar1 637 6HS NADH ntrate reductasedeficient<br />
1<br />
35:194 Steptoe<br />
nar2 638 5HL NADH ntrate reductasedeficient<br />
2<br />
35:196 Steptoe<br />
nar3 639 7HS NADH ntrate reductasedeficient<br />
3<br />
35:197 Winer<br />
nar4 640 2HL NADH ntrate reductasedeficient<br />
4<br />
35:198 Steptoe<br />
nar5 641 5HL NADH ntrate reductasedeficient<br />
5<br />
35:199 Steptoe<br />
nar6 642 2HL NADH ntrate reductasedeficient<br />
6<br />
35:200 Steptoe<br />
nar7 643 6HL NADH ntrate reductasedeficient<br />
7<br />
35:201 Steptoe<br />
nar8 644 6HS NADH ntrate reductasedeficient<br />
8<br />
35:202 Steptoe<br />
nec1 222 1HL Necrotic leaf spot 1 37:251 Carlsberg II<br />
nec2 261 6HS Necrotic leaf spot 2 26:241 Carlsberg II<br />
nec3 265 6HS Necrotic leaf spot 3 26:245 Proctor<br />
nec4 138 3H Necrotic leaf spot 4 26:175 Proctor<br />
nec5 139 3H Necrotic leaf spot 5 26:176 Diamant<br />
Nec6 Sp 611 Necrotic leaf spot 6 32:112 Awnless Atlas<br />
nec7 nec-45 635 Necroticans 7 32:136 Kristina<br />
nld1 nld 323 5HL Narrow leafed dwarf 1 26:271 Nagaoka<br />
nld2 660 Narrow leafed dwarf 2 37:300 Steptoe<br />
nud1 n, nud 7 7HL Naked caryopsis 1 37:195 Himalaya<br />
ops1 op-3 624 Opposite spikelets 1 32:125 Bonus<br />
ovl1 176 4H Ovaryless 1 35:191 Kanto Bansei Gold<br />
ovl2 646 Ovaryless 2 35:204 Harrington<br />
pmr1 pmr 40 7HS Premature ripe 1 32:80 Glenn<br />
pmr2 nec-50 634 Premature ripe 2 32:135 Bonus<br />
Pre2 Re2 76 2HL Red lemma and pericarp 2 26:113 Buckley 3277<br />
Pub1 Pub 127 3HL Pubescent leaf blade 1 26:163 Multiple Dominant<br />
Pvc1 Pc 68 2HL Purple veined lemma 1 26:105 Buckley 2223-6<br />
Pyr1 42 7HS Pyramidatum 1 32:82 Pokko/Hja80001<br />
raw1 r 312 5HL Smooth awn 1 26:261 Lion<br />
raw2 r2 340 5HL Smooth awn 2 26:289 Lion<br />
raw5 r,,e 257 6HL Smooth awn 5 26:238 Akashinriki<br />
raw6 r6 334 5HL Smooth awn 6 26:282 Glenn<br />
182
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
rob1 o 254 6HS Orange lemma 1 37:255 CIho 5649<br />
Rpc1 149 3H Reaction to Puccinia<br />
coronata var. hordei 1<br />
37:232 Hor 2596<br />
Rpg1 T 511 7HS Reaction to Puccinia<br />
graminis 1<br />
26:437 Chevron<br />
Rpg2 T2 512 Reaction to Puccinia<br />
graminis 2<br />
26:439 Hietpas 5<br />
rpg4 319 5HL Reaction to Puccinia<br />
graminis 4<br />
26:267 Q21861<br />
Rph1 Pa 70 2H Reaction to Puccinia hordei<br />
1<br />
26:107 Oderbrucker<br />
Rph2 Pa2 88 5HS Reaction to Puccinia hordei<br />
2<br />
37:212 Peruvian<br />
Rph3 Pa3 121 7HL Reaction to Puccinia hordei<br />
3<br />
26:156 Estate<br />
Rph4 Pa4 218 1HS Reaction to Puccinia hordei<br />
4<br />
26:217 Gull<br />
Rph5 Pa5 122 3HS Reaction to Puccinia hordei<br />
5<br />
37:224 Magnif 102<br />
Rph6 Pa6 575 3HS Reaction to Puccinia hordei<br />
6<br />
26:501 Bolivia<br />
Rph7 Pa7 136 3HS Reaction to Puccinia hordei<br />
7<br />
37:228 Cebada Capa<br />
Rph8 Pa8 576 Reaction to Puccinia hordei<br />
8<br />
26:502 Egypt 4<br />
Rph9 Pa9 32 5HL Reaction to Puccinia hordei<br />
9<br />
37:201 HOR 2596<br />
Rph10 137 3HL Reaction to Puccinia hordei<br />
10<br />
26:174 Clipper C8<br />
Rph11 267 6HL Reaction to Puccinia hordei<br />
11<br />
26:247 Clipper C67<br />
Rph12 333 5HL Reaction to Puccinia hordei<br />
12<br />
26:281 Triumph<br />
Rph13 590 Reaction to Puccinia hordei<br />
13<br />
28:31 PI 531849<br />
Rph14 591 Reaction to Puccinia hordei<br />
14<br />
28:32 PI 584760<br />
Rph15 Rph16 96 2HL Reaction to Puccinia hordei<br />
15<br />
37:214 PI 355447<br />
183
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
Rsg1 Grb 22 7H Reaction to Schizaphis<br />
graminum 1<br />
37:199 Omugi<br />
Rsg2 577 Reaction to Schizaphis<br />
graminum 2<br />
37:283 PI 426756<br />
rsm1 sm 35 7HS Reaction to BSMV 1 26:84 Modjo 1<br />
Rsp1 Sep 515 Reaction to Septoria<br />
passerinii 1<br />
26:441 CIho 14300<br />
Rsp2 Sep2 516 Reaction to Septoria<br />
passerinii 2<br />
37:275 PI 70837<br />
Rsp3 Sep3 517 Reaction to Septoria<br />
passerinii 3<br />
37:276 CIho 10644<br />
rtt1 rt 51 2HS Rattail spike 1 26:87 Goldfoil<br />
Run1 Un 21 7HS Reaction to Ustilago nuda 1 26:67 Trebi<br />
rvl1 rvl 226 1HL Revoluted leaf 1 26:224 Hakata 2<br />
Ryd2 Yd2 123 3HL Reaction to BYDV 2 26:158 CIho 2376<br />
Rym1 Ym 167 4HL Reaction to BaYMV 1 32:96 Mokusekko 3<br />
Rym2 Ym2 20 7HL Reaction to BaYMV 2 26:66 Mihori Hadaka 3<br />
rym3 ym3 345 5HS Reaction to BaYMV 3 32:105 Chikurin Ibaraki<br />
rym5 Ym 141 3HL Reaction to BaYMV 5 32:90 Mokusekko 3<br />
sbk1 sk, cal-a 62 2HS Subjacent hood 1 32:83 Tayeh 13<br />
sca1 sca 128 3HS Short crooked awn 1 26:164 Akashinriki<br />
sci-a sci-3 625 Scirpoides-a 32:126 Bonus<br />
scl-a scl-6 626 Scirpoides leaf-a 32:127 Foma<br />
sdw1 sdw 518 3HL Semidwarf 1 37:277 M21<br />
sdw2 sdw-b 133 3HL Semidwarf 2 26:169 Mg2170<br />
seg1 se1 377 7HL Shrunken endosperm genetic<br />
1<br />
37:264 Betzes<br />
seg2 se2 378 7HS Shrunken endosperm genetic<br />
2<br />
26:326 Betzes<br />
seg3 se3 379 3H Shrunken endosperm genetic<br />
3<br />
37:265 Compana<br />
seg4 se4 380 7HL Shrunken endosperm genetic<br />
4<br />
37:267 Compana<br />
seg5 se5 381 7HS Shrunken endosperm genetic<br />
5<br />
26:329 Sermo/7*Glacier<br />
seg6 se6 396 3HL Shrunken endosperm genetic<br />
6<br />
37:268 Ingrid<br />
seg7 se7 397 Shrunken endosperm genetic<br />
7<br />
37:269 Ingrid<br />
184
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
seg8 455 7H Shrunken endosperm genetic<br />
8<br />
37:272 60Ab1810-53<br />
sex1 lys5 382 6HL Shrunken endosperm xenia 1 26:330 Compana<br />
sex6 31 7HS Shrunken endosperm xenia 6 26:80 K6827<br />
sex7 sex.i 628 5HL Shrunken endosperm xenia 7 32:129 I90-374<br />
sex8 sex.j 143 6HS Shrunken endosperm xenia 8 32:93 I89-633<br />
sgh1 sh1 163 4HL Spring growth habit 1 26:188 Iwate Mensury C<br />
Sgh2 Sh2 309 5HL Spring growth habit 2 26:259 Indian Barley<br />
Sgh3 Sh3 213 1HL Spring growth habit 3 26:212 Tammi/Hayakiso 2<br />
sid1 nls 180 4HL Single internode dwarf 1 26:203 Akashinriki<br />
Sil1 Sil 228 1HS Subcrown internode length 1 26:226 NE 62203<br />
sld1 dw-1 126 3HL Slender dwarf 1 26:162 Akashinriki<br />
sld2 83 2HS Slender dwarf 2 26:121 Akashinriki<br />
sld3 ant-567 186 4HS Slender dwarf 3 37:243 Manker<br />
sld4 100 7HS Slender dwarf 4 37:218 Glacier<br />
sld5 144 3HS Slender dwarf 5 32:94 Indian Dwarf<br />
sls1 sls 227 1HS Small lateral spikelet 1 26:225 Morex<br />
smn1 smn 38 7HS Seminudoides 1 32:225 Haisa<br />
snb1 sb 26 7HS Subnodal bract 1 26:72 L50-200<br />
srh1 s 321 5HL Short rachilla hair 1 26:269 Lion<br />
sun1 650 Sensitivity to Ustilago nuda 1 35:208 Steptoe<br />
trd1 trd 202 1HL Third outer glume 1 26:207 Valki<br />
trp1 tr 61 2HL Triple awned lemma 1 26:97 CIho 6630<br />
tst1 647 Tip sterile 1 35:205 Steptoe<br />
tst2 lin2 636 Tip sterile 2 37:292 Donaria<br />
ubs4 u4 11 7HL Unbranched style 4 26:56 Ao-Hadaka<br />
uzu1 uz 102 3HL Uzu 1 or semi brachytic 1 37:220 Baitori<br />
var1 va 306 5HL Variegated 1 37:259 Montcalm<br />
var2 va2 344 5HL Variegated 2 32:104 Montcalm<br />
var3 va3 303 5HL Variegated 3 26:254 Montcalm<br />
viv-a viv-5 627 Viviparoides-a 32:128 Foma<br />
vrs1 v 6 2HL Six-rowed spike 1 37:192 Trebi<br />
vrs1 lr 58 2HL Six-rowed spike 1 26:94 Nudihaxtoni<br />
vrs1 V d 66 2HL Two-rowed spike 1 26:103 Svanhals<br />
185
Barley Genetics Newsletter (2007) 37: 154-187<br />
Table 3 (continued)<br />
Locus symbol* BGS Chr. Locus name or phenotype Descr. Parental cultivar<br />
Rec. Prev. no. loc. † vol. p.<br />
vrs1 V t 67 2HL Deficiens 1 26:104 White Deficiens<br />
vrs2 v2 314 5HL Six-rowed spike 2 26:263 Svanhals<br />
vrs3 v3 315 1HL Six-rowed spike 3 26:264 Hadata 2<br />
vrs4 v4 124 3HL Six-rowed spike 4 26:159 MFB 104<br />
wax1 wx 16 7HS Waxy endosperm 1 26:61 Oderbrucker<br />
wnd1 wnd 23 7HS Winding dwarf 1 26:69 Kogen-mugi<br />
wst1 wst 107 3HL White streak 1 26:141 CIho 11767<br />
wst2 304 5HL White streak 2 26:255 Manabe<br />
wst4 56 2HL White streak 4 26:91 Kanyo 7<br />
wst5 221 1HL White streak 5 26:219 Carlsberg II<br />
wst6 wst,,j 129 3HL White streak 6 26:165 Akashinriki<br />
wst7 rb 79 2HL White streak 7 37:207 GS397<br />
Xnt1 Xa 25 7HL Xantha seedling 1 26:71 Akanshinriki<br />
xnt2 xb 513 Xantha seedling 2 26:440 Black Hulless<br />
xnt3 xc 105 3HS Xantha seedling 3 26:139 Colsess<br />
xnt4 xc2 36 7HL Xantha seedling 4 26:85 Coast<br />
xnt5 xn 255 6HL Xantha seedling 5 26:237 Nepal<br />
xnt6 xs 113 3HS Xantha seedling 6 26:147 Smyrna<br />
xnt7 xan,,g 233 1HL Xantha seedling 7 26:231 Erbet<br />
xnt8 xan,,h 140 3HS Xantha seedling 8 26:177 Carlsberg II<br />
xnt9 xan,,i 37 7HL Xantha seedling 9 26:86 Erbet<br />
yhd1 yh 158 4HL Yellow head 1 26:185 Kimugi<br />
yhd2 yh2 592 Yellow head 2 28:34 Compana<br />
ylf1 652 Yellow leaf 1 35:210 Villa<br />
Ynd1 Yn 183 4HS Yellow node 1 32:98 Morex<br />
yst1 yst 104 3HS Yellow streak 1 26:138 Gateway<br />
yst2 109 3HS Yellow streak 2 26:144 Kuromugi 148/<br />
Mensury C<br />
yst3 yst,,c 462 3HS Yellow streak 3 26:409 Lion<br />
yst4 85 2HL Yellow streak 4 37:210 Glenn<br />
yst5 346 5HL Yellow streak 5 32:107 Bowman / ant10.30<br />
yvs1 yx 63 2HS Virescent seedling 1 26:99 Minn 71-8<br />
yvs2 yc 3 7HS Virescent seedling 2 26:46 Coast<br />
zeb1 zb 120 3HL Zebra stripe 1 26:155 Mars<br />
zeb2 zb2 461 4HL Zebra stripe 2 26:407 Unknown<br />
zeb3 zb3 223 1HL Zebra stripe 3 26:221 Utah 41<br />
Zeo1 Knd 82 2HL Zeocriton 1 37:209 Donaria<br />
Zeo2 614 Zeocriton 2 32:115 36Ab51<br />
Zeo3 Mo1 184 4HL Zeocriton 3 32:99 Morex<br />
186
Barley Genetics Newsletter (2007) 37: 154-187<br />
* Recommended locus symbols are based on utilization of a three-letter code for barley genes as<br />
approved at the business meeting of the Seventh International Barley Genetics Symposium at<br />
Saskatoon, Saskatchewan, Canada, on 05 August 1996.<br />
† Chromosome numbers and arm designations are based on the Triticeae system. Utilization of<br />
this system for naming of barley chromosomes was at the business meeting of the Seventh<br />
International Barley Genetics Symposium at Saskatoon, Saskatchewan, Canada, on 05 August<br />
1996. The Burnham and Hagberg (1956) designations of barley chromosomes were 1 2 3 4 5 6<br />
and 7 while new designations based on the Triticeae system are 7H 2H 3H 4H 1H 6H and 5H,<br />
respectively.<br />
187
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 1<br />
Locus name: Brachytic 1<br />
Locus symbol: brh1<br />
BGS 1, Brachytic 1, brh1<br />
Previous nomenclature and gene symbolization:<br />
Brachytic = br (10, 12).<br />
Breviaristatum-i = ari-i (5, 8).<br />
Dwarf x = dx1 (6).<br />
Inheritance:<br />
Monofactorial recessive (10, 12).<br />
Located in chromosome 7HS [1S] (3), about 9.3 cM distal from the fch12<br />
(chlorina seedling 12) locus (12), 0.8 cM distal from RFLP marker BCD129 (9),<br />
about 5.0 cM from AFLP marker E4134-8 in subgroup 1 of the Proctor/Nudinka<br />
map (11), and about 13.6 cM proximal from SSR marker HVM04 in bin 1H-02 (2).<br />
Description:<br />
Plants have short leaves, culms, spikes, awns, and kernels. The seedling leaf is<br />
about 2/3 normal length. A similar reduction in the size of other organs is<br />
observed, but the awns are less than 1/2 normal length (6). The mutant<br />
phenotype is easy to classify at all stages of growth. The approximately 20%<br />
reduction in kernels size is caused primarily by a reduction in kernel length. The<br />
yields of the brh1 mutants are about 2/3 normal and lodging is greatly reduced in<br />
the Bowman brh1 lines (2). Börner (1) reported that ari-i.38 seedlings are<br />
sensitive to gibberellic acid. Powers (10) states that the assigned gene symbol<br />
for this mutant is br and that L.J. Stadler selected this symbol.<br />
Origin of mutant:<br />
A spontaneous mutant in Himalaya (CIho 1312) (10, 12).<br />
Mutational events:<br />
brh1.a in Himalaya (12); brh1.c (GSHO 229) in Moravian (PI 539135) (13); arii.38<br />
(NGB 115888, GSHO 1657) in Bonus (PI 189763) (8, 14); brh1.e (GSHO<br />
1690) in Aramir (PI 467786) (14); brh1.f (dx1, GSHO 1422) in Domen (CIho<br />
9562) (6); brh1.t (OUM136, GSHO 1691) in Akashinriki (PI 467400, OUJ659);<br />
brh1.x (7125, DWS1224, GSHO 1692) in Volla (PI 280423); brh1.z (Hja80001) in<br />
Aapo; brh1.aa (Hja80051) in a Hja80001 cross (4, 7); and brh1.ae (FN53) in<br />
Steptoe (CIho 15229) (4).<br />
Mutant used for description and seed stocks:<br />
brh1.a in Himalaya (GSHO 25); brh1.a in Bowman (PI 483237)*7 (GSHO 1820);<br />
ari-i.38 in Bowman*6 (GSHO 1821); brh1.e in Bowman*7 (GSHO 1822); brh1.t in<br />
Bowman*7 (GSHO 1823); brh1.x in Bowman*7 (GSHO 1824); brh1.z in<br />
Bowman*7 (GSHO 2179).<br />
References:<br />
1. Börner, A. 1996. GA response in semidwarf barley. Barley Genet. Newsl.<br />
25:24-26.<br />
2. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
3. Fedak, G., T. Tsuchiya, and S.B. Helgason. 1972. Use of monotelotrisomics<br />
for linkage mapping in barley. Can. J. Genet. Cytol. 14:949-957.<br />
4. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
188
Barley Genetics Newsletter (2007) 37: 188-301<br />
5. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
6. Holm, E., and K. Aastveit. 1966. Induction and effects of the brachytic allele in<br />
barley. Adv. Front Plant Sci. 17:81-94.<br />
7. Kivi, E. 1986. (personal communications).<br />
8. Kucera, J., U. Lundqvist, and Å. Gustafsson. 1975. Inheritance of<br />
breviaristatum mutants in barley. Hereditas 80:263-278.<br />
9. Li , M., D. Kudrna, and A. Kleinhofs. 2000. Fine mapping of a semi-dwarf gene<br />
brachytic 1 in barley. p. 72-74. In S. Logue (ed.) Barley Genetics VIII. Volume III,<br />
Proc. Eighth Int. Barley Genet. Symp., Adelaide, Dept. Plant Science, Waite<br />
Campus, Adelaide University, Glen Osmond, South Australia.<br />
10. Powers, L. 1936. The nature of the interactions of genes affecting four<br />
quantitative characters in a cross between Hordeum deficiens and vulgare.<br />
Genetics 21:398-420.<br />
11. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration<br />
of a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
12. Swenson, S.P. 1940. Genetic and cytological studies on a brachytic mutant in<br />
barley. J. Agric. Res. 60:687-713.<br />
13. Szarejko, I., and M. Maluszynski. 1984. New brachytic mutant of spring<br />
barley variety Aramir. Barley Genet. Newsl. 14:33-35.<br />
14. Tsuchiya, T. 1974. Allelic relationships of genes for short-awned mutants in<br />
barley. Barley Genet. Newsl. 4:80-81.<br />
Prepared:<br />
T. Tsuchiya and T.E. Haus. 1971. BGN 1:104.<br />
Revised:<br />
T. Tsuchiya. 1980. BGN 10:100.<br />
J.D. Franckowiak. 1997. BGN 26:44.<br />
J.D. Franckowiak and L. S. Dahleen. 2007. BGN 37:188-189.<br />
189
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 2<br />
Locus name: Chlorina seedling 12<br />
Locus symbol: fch12<br />
BGS 2, Chlorina seedling 12, fch12<br />
Previous nomenclature and gene symbolization:<br />
Chlorina seedling-c = fc (3).<br />
Chlorina seedling-fc = clo-fc (7).<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Located in chromosome 7HS [1S] (1, 4), about 3.6 cM distal from the gsh3<br />
(glossy sheath 3) locus (6), and about 9.3 cM proximal from the brh1 (brachytic<br />
1) locus (8), in bin 7H-02 about 2.3 cM from RFLP marker KFP027 and cosegregating<br />
with markers BCD130 and ABC327 (5).<br />
Description:<br />
Seedling leaves are yellow with green tips and new leaves show a yellow base<br />
and a green tip. As the plant develops, leaf color changes to pale green (3).<br />
Plants are vigorous, but anthesis is delayed and seed yield may be low.<br />
Origin of mutant:<br />
A spontaneous mutant in Colsess (CIho 2792) (3).<br />
Mutational events:<br />
fch12.b (fc) in Colsess (Colsess V) (3); fch12.l (Trebi chlorina 453, GSHO 155),<br />
fch12.m (Trebi V, GSHO 158), fch12.n (Trebi IX, GSHO 18), fch12.o (Trebi XI,<br />
GSHO 163) in Trebi (PI 537442) (2); clo-fc.110 in Bonus (PI 189763) (7); fch12.b<br />
may be present in the brachytic chlorina stocks (GSHO 124 and GSHO 174) (9).<br />
Mutant used for description and seed stocks:<br />
fch12.b in Colsess (GSHO 36); fch12.b in Bowman (PI 483237)*7 (GSHO 1826).<br />
References:<br />
1. Fedak, G., T. Tsuchiya, and S.B. Helgason. 1972. Use of monotelotrisomics<br />
for linkage mapping in barley. Can. J. Genet. Cytol. 14:949-957.<br />
2. McMullen, M. 1972. Allelism testing of seven chlorina mutants in Trebi barley.<br />
Barley Genet. Newsl. 2:76-79.<br />
3. Robertson, D.W., and G.W. Deming. 1930. Genetic studies in barley. J. Hered.<br />
21:283-288.<br />
4. Robertson, D.W., G.W. Deming, and D. Koonce. 1932. Inheritance in barley. J.<br />
Agric. Res. 44:445-466.<br />
5. Schmierer, D., A. Druka, D. Kudrna, and A. Kleinhofs. 2001. Fine mapping of<br />
the fch12 chlorina seedling mutant. Barley Genet. Newsl. 31:12-13.<br />
6. Shahla, A., and T. Tsuchiya. 1987. Cytogenetic studies in barley chromosome<br />
1 by means of telotrisomic, acrotrisomic and conventional analysis. Theor. Appl.<br />
Genet. 75:5-12.<br />
7. Simpson, D.J., O. Machold, G. Høyer-Hansen, and D. von Wettstein. 1985.<br />
Chlorina mutants of barley (Hordeum vulgare L.). Carlsberg Res. Commun.<br />
50:223-238.<br />
8. Swenson, S.P. 1940. Genetic and cytological studies on a brachytic mutation<br />
in barley. J. Hered. 31:213-214.<br />
9. Wang, S., and T. Tsuchiya. 1991. Genetic analysis of the relationship between<br />
new chlorina mutants in genetic stocks and established f series stocks in barley.<br />
Barley Genet. Newsl. 20:63-65.<br />
Prepared:<br />
190
Barley Genetics Newsletter (2007) 37: 188-301<br />
T. Tsuchiya and T. E. Haus. 1971. BGN 1:105.<br />
Revised:<br />
T. Tsuchiya. 1980. BGN 10:101.<br />
J.D. Franckowiak and A. Hang. 1997. BGN 26:45.<br />
J.D. Franckowiak. 2007. BGN 37:190-191.<br />
191
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 6<br />
Locus name: Six-rowed spike 1<br />
Locus symbol: vrs1<br />
BGS 6, Six-rowed spike 1, vrs1<br />
Previous nomenclature and gene symbolization:<br />
Two-row vs six-row = Zz (21).<br />
Six-row vs two-row = Aa (6).<br />
Two-rowed = D (17).<br />
Six-row vs two-row = Vv (3).<br />
Six-row vs two-row (distichon) vs two-row (deficiens) = A, a s , a f (8).<br />
Reduced lateral spikelet appendage on the lemma = lr (9).<br />
Allelic series v, V d , V, and V t (22).<br />
Hexastichon mutants = hex-v (5, 6).<br />
Intermedium spike-d = Int-d (4).<br />
Reduced lateral spikelet appendage on the lemma = v lr (19).<br />
The vrs1 DNA sequence identified as HvHox1 (10).<br />
Inheritance:<br />
A multiple allelic series, incomplete dominant allele interactions based on the<br />
size and shape of lateral spikelets (1, 19, 22).<br />
Located in chromosome 2HL (3, 6, 12, 14), about 30.5 cM distal from the eog1<br />
(elongated outer glume 1) locus (18), in bin 2H-09 and in a 0.90-cM interval<br />
between markers cMWG699 and MWG865 (11).<br />
Description:<br />
Alleles at this complex locus modify development of the lateral spikelets and the<br />
associated lemma awn. The vrs1.a allele (v gene) is present in most six-rowed<br />
cultivars and produces well-developed lateral spikelets (6). Based on<br />
phylogenetic analysis of the six-rowed cultivars, the six-rowed gene originated<br />
independently at least three times (vrs1.a1, vrs1.a2, and vrs1.a3) from different<br />
wild type (Vrs1.b) alleles (10). The lemma awn of lateral spikelets will vary from<br />
3/4 to nearly as long as those of central spikelets, depending upon alleles<br />
present at other loci. The Vrs1.b allele (V gene, distichon) is present in many<br />
two-rowed cultivars and reduces lateral spikelets to sterile bracts with a rounded<br />
tip. The Vrs1.t allele (V t gene, deficiens) causes an extreme reduction in the size<br />
of lateral spikelets. The lr or v lr (vrs1.c) gene in Nudihaxtoni and Bozu types will<br />
not recombine with the vrs1.a allele (12, 19) and produces phenotypes similar to<br />
the Vrs1.d allele (V d gene) of Svanhals (22). The series of induced mutants in<br />
two-rowed barley called hex-v and Int-d mutants differ in the size of lateral<br />
spikelets, but they interact with the vrs1.a allele as incomplete dominants (5).<br />
Many heterozygous combinations with vrs1.a have a pointed tip on the lemma of<br />
sterile lateral spikelets. Alleles at the int-c (intermedium spike-c) locus modify<br />
lateral size in the presence of vrs1.a, Vrs1.b, and Vrs1.d, but not when Vrs1.t is<br />
present (22). Multiple origins of vrs1 alleles in six-rowed barley have been<br />
confirmed by molecular analysis (20). Komatsuda et al. (10) found that<br />
expression of the Vrs1 gene was strictly localized in the lateral-spikelet primordia<br />
of immature spikes and suggested that the VRS1 protein suppresses<br />
development of lateral spikelets.<br />
Origin of mutant:<br />
Natural occurrence in six-rowed barley and induced frequently by mutagenic<br />
agents (10, 14).<br />
192
Barley Genetics Newsletter (2007) 37: 188-301<br />
Mutational events:<br />
vrs1.a1 in most six-rowed cultivars (1, 10, 22); vrs1.a2 in Dissa and Valenci (10),<br />
vrs1.a3 in Natsudaikon Mugi (OUK735) (10), Vrs1.b in wild barley (10), Vrs1.b2<br />
in Pamella Blue (OUH630) (10), Vrs1.b3 in Bonus (PI 189763) (10), Vrs1.t in a<br />
few two-rowed cultivars (10, 22); vrs1.c or lr in Nudihaxtoni (PI 32368) (12, 19);<br />
Vrs1.d in Svanhals (PI 5474) (22); 23 induced mutants from programs in<br />
Belgium, Germany, and Hungary (2); hex-v.3 (NGB 115545), -v.4 (NGB 115546),<br />
-v.6 (NGB 115547), -v.7 (NGB115548), -v.8 (NGB 115549), -v.9 (NGB 115550), -<br />
v.10 (NGB 115551), -v.11 (NGB 115552), -v.12 (NGB 115553), -v.18 (NGB<br />
115559), -v.44 (NGB 115581), -v.45 (NGB 115582 ), -v.46 (NGB 115583 ), -v.47<br />
(NGB 115584), -v.48 (NGB 115585), in Bonus, -v.13 (NGB 115554), -v.14 (NGB<br />
115555), -v.15 (NGB 115556), -v.16 (NGB 115557), -v.17 (NGB 115558), -v.19<br />
(NGB 115560), -v.21 (NGB 115562), -v.22 (NGB 115563), -v.23 (NGB 115564),<br />
-v.24 (NGB 115565), -v.25 (NGB 115566), -v.26 (NGB 115567), -v.27 (NGB<br />
115568), -v.28 (NGB 115569), -v.29 (NGB 115570), -v.30 (NGB 115571), -v.31<br />
(NGB 115572), -v.35 (NGB 115574) in Foma (CIho 11333), -v.20 (NGB 115561)<br />
in Ingrid (CIho 10083), -v.33 (NGB 115573), -v.36 (NGB 115575), -v.38 (NGB<br />
115576), -v.39 (NGB 115577), -v.41 (NGB 115578), -v.42 (NGB 115579), -v.43<br />
(NGB 115580) in Kristina (NGB 1500) (5, 14); hex-v.49 (NGB 115586) in Bonus,<br />
-v.50 (NGB 115587), -v.51 (NGB 115588) in Sv 79353, -v.52 (NGB 119353) in<br />
Golf (PI 488529) (13); Int-d.11 (NGB 115429), -d.12 (NGB 115430), -d.22 (NGB<br />
115440), -d.24 (NGB 115442), -d.28 (NGB 115446), -d.36 (NGB 115454) in<br />
Foma, -d.40 (NGB 115458), -d.41 (NGB 115459), -d.50 (NGB 115468), -d.57<br />
(NGB 115475), -d.67 (NGB 115485), -d.68 (NGB 115486), -d.69 (NGB 115487)<br />
in Kristina (5, 15); Int-d.73 (NGB 115491), -d.80 (NGB 115498), -d.82 (NGB<br />
115500) in Bonus, -d.93 (NGB 115511), -d.94 (NGB 115512), -d.96 (NGB<br />
115514), -d.97 (NGB 115515), -d.100 (NGB 115518) in Hege (NGB 13692) (13);<br />
vrs1.o (v1b) in New Golden (16).<br />
Mutant used for description and seed stock:<br />
vrs1.a in Trebi (PI 537442, GSHO 196); vrs1.a in Bonneville (CIho 7248) (7);<br />
vrs1.a from Glenn (CIho 15769) in Bowman (PI 483237)*8 (GSHO 1907); Intd.12<br />
in Bowman*7 (GSHO 1910).<br />
References:<br />
1. Biffen, R.H. 1906. Experiments on the hybridization of barleys. Proc. Camb.<br />
Phil. Soc. 13:304-308.<br />
2. Fukuyama, T., J. Hayashi, I. Moriya, and R. Takahashi. 1972. A test for<br />
allelism of 32 induced six-rowed mutants. Barley Genet. Newsl. 2:25-27.<br />
3. Griffee, F. 1925. Correlated inheritance of botanical characters in barley, and<br />
manner of reaction to Helminthosporium sativum. J. Agric. Res. 30:915-935.<br />
4. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
5. Gustafsson, Å., and U. Lundqvist. 1980. Hexastichon and intermedium<br />
mutants in barley. Hereditas 92:229-236.<br />
6. Harlan, H.V., and H.K. Hayes. 1920. Occurrence of the fixed intermediate,<br />
Hordeum intermedium haxtoni, in crosses between H. vulgare pallidium and H.<br />
distichum palmella. J. Agric. Res.19:575-591.<br />
7. Hockett, E.A. 1985. Registration of two- and six-rowed isogenic Bonneville<br />
barley germplasm. Crop Sci. 25:201.<br />
8. Hor, K.S. 1924. Interrelations of genetic factors in barley. Genetics 9:151-180.<br />
9. Immer, F.R., and M.T. Henderson. 1943. Linkage studies in barley. Genetics<br />
193
Barley Genetics Newsletter (2007) 37: 188-301<br />
28:419-440.<br />
10. Komatsuda, T., M. Pourkheirandish, C. He, P. Azhaguvel, H. Kanamori, D.<br />
Perovic, N. Stein, A. Graner, T. Wicker, A. Tagiri, U. Lundqvist, T. Fujimura, M.<br />
Matsuoka, T. Matsumoto, and M. Yano. 2007. Six-rowed barley originated from a<br />
mutation in a homeodomain-leucine zipper I-class homeobox gene. PNAS<br />
104:1424-1429.<br />
11. Komatsuda, T., and K. Tanno. 2004. Comparative high resolution map of the<br />
six-rowed locus 1 (vrs1) in several populations of barley, Hordeum vulgare L.<br />
Hereditas 141:68-73.<br />
12. Leonard, W.H. 1942. Inheritance of reduced lateral spikelet appendages in<br />
the Nudihaxtoni variety of barley. J. Am. Soc. Agron. 34:211-221.<br />
13. Lundqvist, U. (unpublished).<br />
14. Lundqvist, U., and A. Lundqvist. 1987. Barley mutants - diversity and<br />
genetics. p. 251-257. In S. Yasuda and T. Konishi (eds.) Barley Genetics V.<br />
Proc. Fifth Int. Barley Genet. Symp., Okayama, 1986. Sanyo Press Co.,<br />
Okayama.<br />
15. Lundqvist, U., and A. Lundqvist. 1988. Induced intermedium mutants in<br />
barley: origin, morphology and inheritance. Hereditas 108:13-26.<br />
16. Makino, T., M. Furusho, and T. Fukuoka. 1995. A mutant having six-rowed<br />
gene allelic to v locus. Barley Genet. Newsl. 24:122.<br />
17. Miyake, K., and Y. Imai. 1922. [Genetic studies in barley. 1.] Bot. Mag.,<br />
Tokyo 36:25-38. [In Japanese.]<br />
18. Swenson, S.P., and D.G. Wells. 1944. The linkage relation of four genes in<br />
chromosome 1 of barley. J. Am. Soc. Agron. 36:429-435.<br />
19. Takahashi, R., J. Hayashi, I. Moriya, and S. Yasuda. 1982. Studies on<br />
classification and inheritance of barley varieties having awnless or short-awned<br />
lateral spikelets (Bozu barley). I. Variation of awn types and classification.<br />
Nogaku Kenyu 60:13-24. [In Japanese with English summary.]<br />
20. Tanno, K., S. Taketa, K. Takeda, and T. Komatsuda. 2002. A DNA marker<br />
closely linked to the vrs1 locus (row-type gene) indicates multiple origins of sixrowed<br />
barley (Hordeum vulgare L.) Theor. Appl. Genet. 104:54-60.<br />
21. Ubisch, G. von. 1916. Beitrag zu einer Faktorenanalyse von Gerste. Z.<br />
Indukt. Abstammungs. Vererbungsl. 17:120-152.<br />
22. Woodward, R.W. 1949. The inheritance of fertility in the lateral florets of the<br />
four barley groups. Agron. J. 41:317-322.<br />
Prepared:<br />
T.E. Haus. 1975. BGN 5:106.<br />
Revised:<br />
J.D. Franckowiak and U. Lundqvist. 1997. BGN 26:49-50.<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:192-194.<br />
194
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 7<br />
Locus name: Naked caryopsis 1<br />
Locus symbol: nud1<br />
BGS 7, Naked caryopsis 1, nud1<br />
Previous nomenclature and gene symbolization:<br />
Naked caryopsis = k (14).<br />
Naked caryopsis = s (21).<br />
Naked caryopsis = n (6, 9).<br />
Hulless = h (10).<br />
Inheritance:<br />
Monofactorial recessive (6, 14, 19).<br />
Located in chromosome 7HL [1L] (3, 11, 12, 14, 20), near the centromere (3, 11),<br />
about 9.6 cM proximal from the lks2 (short awn 2) locus (15), about 10.5 cM<br />
proximal from the dsp1 (dense spike 1) locus (15, 16), in bin 7H-07 about 13.1<br />
cM distal from RFLP marker MWG808 (2), co-segregating with AFLP markers<br />
KT3 and KT7 and SCAR marker sKT7 (7), about 0.06 cM distal from SCAR<br />
marker sTK3 and the same distance proximal from sTK9 (17).<br />
Description:<br />
The lemma and palea do not adhere to the caryopsis and the grain will thresh<br />
free of the hull at maturity. The naked caryopsis trait is expressed in all<br />
environments (16). The naked lines fail to produce a cementing substance<br />
present in covered lines (4). The nud1.a mutant depressed the expression by 10<br />
to 20% of other traits such as plant height, seed weight (1, 8) and altered malt<br />
quality parameters (8). The nud1.a gene is often associated with the dsp1.a<br />
(dense spike 1) gene in Japanese cultivars (16). Allele IV of the marker sKT7<br />
near the nud1 locus was the only one found in naked barley cultivars (18);<br />
however, the geographic distribution for haplotypes of allele IV suggest migration<br />
of naked types toward eastern Asia (18).<br />
Origin of mutant:<br />
In an unknown cultivar, but its origin was monophyletic probably in southwestern<br />
Iran (18), widespread in cultivated barley in Asia.<br />
Mutational events:<br />
nud1.a in Himalaya (CIho 1312) (21); nud1.b in Haisa (Mut 4129), nud1.c (Mut<br />
3041/62) in Ackermann's Donaria (PI 161974) (13).<br />
Mutant used for description and seed stocks:<br />
nud1.a in Himalaya (GSHO 115), nud1.a from Sermo (CIho 7776) in Betzes (PI<br />
129430)*7 (CIho 16559, GP 37), nud1.a from Sermo in Compana (CIho 5438)*7<br />
(CIho 16185, GP 41), nud1.a from Sermo in Decap (CIho 3351)*7 (CIho 16563,<br />
GP 45) (5); nud1.a from Stamm (PI 194555) in Betzes*7 (CIho 16566, GP 48),<br />
nud1.a from Stamm in Compana*7 (CIho 16183, GP 50), nud1.a from Stamm*7<br />
in Freja (CIho 7130)*7 (CIho 16568, GP 52) (5); nud1.a from R.I. Wolfe's Multiple<br />
Recessive Marker Stock in Bowman (PI 483237)*8 (GSHO 1847).<br />
References:<br />
1. Choo, T-M., K.M. Ho, and R.A. Martin. 2001. Genetic analysis of a hulless X<br />
covered cross of barley using doubled-haploid lines. Crop Sci. 41:1021-1026.<br />
2. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
195
Barley Genetics Newsletter (2007) 37: 188-301<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
3. Fedak, G., T. Tsuchiya, and S.B. Helgason. 1972. Use of monotelotrisomics<br />
for linkage mapping in barley. Can. J. Genet. Cytol. 14:949-957.<br />
4. Gaines, R.L., D.B. Bechtel, and Y. Pomeranz. 1985. A microscopic study on<br />
the development of a layer in barley that causes hull-caryopsis adherence.<br />
Cereal Chem. 62:35–40.<br />
5. Hockett, E.A. 1981. Registration of hulless and hulless short-awned spring<br />
barley germplasm (Reg. nos. GP 35 to 52). Crop Sci. 21:146-147.<br />
6. Hor, K.S. 1924. Interrelations of genetic factors in barley. Genetics 9:151-180.<br />
7. Kikuchi, S., S. Taketa, M. Ichii, and S. Kawasaki. 2003. Efficient fine mapping<br />
of the naked caryopsis gene (nud) by HEGS (high efficiency genome<br />
scanning)/AFLP in barley. Theor. Appl. Genet. 108:73-78.<br />
8. McGuire, C.F., and E.A. Hockett. 1981. Effect of awn length and naked<br />
caryopsis on malting quality of Betzes barley. Crop Sci. 21:18-21.<br />
9. Miyake, K., and Y. Imai. 1922. [Genetic studies in barley. 1.] Bot. Mag., Tokyo<br />
36:25-38. [In Japanese.]<br />
10. Neatby, K.W. 1926. Inheritance of quantitative and other characters in a<br />
barley cross. Sci. Agric. 7:77-84.<br />
11. Persson, G. 1969. An attempt to find suitable genetic markers for dense ear<br />
loci in barley I. Hereditas 62:25-96.<br />
12. Robertson, D.W. 1937. Inheritance in barley. II. Genetics 22:443-451.<br />
13. Scholz, F. 1955. Mutationsversuche an Kulturpflanzen. IV. Kulturpflanze<br />
3:69-89.<br />
14. So, M., S. Ogura, and Y. Imai. 1919. [A linkage group in barley.] Nogaku<br />
Kaiho 208:1093-1117. [In Japanese.]<br />
15. Takahashi, R., J. Hayashi, T. Konishi, and I. Moriya. 1975. Linkage analysis<br />
of barley mutants. BGN 5:56-60.<br />
16. Takahashi, R., J. Yamamoto, S. Yasuda, and Y. Itano. 1953. Inheritance and<br />
linkage studies in barley. Ber. Ohara Inst. landw. Forsch. 10:29-52.<br />
17. Taketa, S., T. Awayama, S. Amano, Y. Sakurai, and M. Ichii. 2006. Highresolution<br />
mapping of the nud locus controlling the naked caryopsis in barley.<br />
Plant Breed. 125:337-342.<br />
18. Taketa, S., S. Kikuchi, T. Awayama, S. Yamamoto, M. Ichii, and S. Kawasaki.<br />
2004. Monophyletic origin of naked barley inferred from molecular analyses of a<br />
marker closely linked to the naked caryopsis gene (nud). Theor. Appl. Genet.<br />
108:1236-1242.<br />
19. Tschermak, E. von. 1901. Über Züchtung neuer Getreiderassen mittelst<br />
künstlicher Kreuzung. Kritisch-historische Betrachtungen. Zeitschrift für das<br />
landwirtschaftliche Versuchswesen Oesterreich 4:1029-1060.<br />
20. Tsuchiya, T., and R.J. Singh. 1973. Further information on telotrisomic<br />
analysis in barley. Barley Genet. Newsl. 3:75-78.<br />
21. Ubisch, G. von. 1921. Beitrag zu einer Faktorenanalyse von Gerste. III. Z.<br />
Indukt. Abstammungs. Vererbungsl. 25:198-200.<br />
Prepared:<br />
T. Tsuchiya and T.E. Haus. 1971. BGN 1:110.<br />
Revised:<br />
J.D. Franckowiak and T. Konishi. 1997. BGN 26:51-52.<br />
J.D. Franckowiak. 2007. BGN 37:195-196.<br />
196
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 10<br />
Locus name: Short awn 2<br />
Locus symbol: lks2<br />
BGS 10, Short awn 2, lks2<br />
Previous nomenclature and gene symbolization:<br />
Short awn = a (15, 16).<br />
Short awn = lk (14).<br />
Short awn 2 = lk1 (7).<br />
Short awn 2 = lk2 (11).<br />
Short awn 4 = lk4 (2, 5).<br />
Inheritance:<br />
Monofactorial recessive (6, 7, 12).<br />
Located in chromosome 7HL [1L] (6, 13), estimates range from 7.9 to 10.5 cM<br />
distal from the nud1 (naked caryopsis 1) locus (3, 12, 13), about 2.8 cM distal<br />
from molecular marker WG541 in bin 7H-05 (8), about 8.6 cM proximal from<br />
RFLP marker WG380B in bin 7H-08 (1).<br />
Description:<br />
Awns of both central and lateral spikelets are reduced to about 3/5 of the long<br />
awned type. Texture of the short awn is finer and more flexible than that of the<br />
long awn, especially in non-uzu genotypes (13, 14). The awn length of<br />
heterozygotes in some crosses is shorter that of the normal parent. Other plant<br />
characteristics are apparently unaltered by the lks2.b gene.<br />
Origin of mutant:<br />
Spontaneous occurrence in some cultivars distributed in China, Japan, Korea,<br />
and Nepal (5, 10, 12, 14).<br />
Mutational events:<br />
lks2.b in many cultivars of Oriental origin, often associated with the dsp1.a<br />
(dense spike 1) gene (6, 12, 14); a possible mutant in Morex (CIho 15773) (9,<br />
10).<br />
Mutant used for description and seed stocks:<br />
lks2.b in Honen 6 (OUJ469, PI 307495, GSHO 566) (14); lks2.b from Sermo<br />
(CIho 7776) in Betzes (PI 129430)*7 (CIho 16558, GP 36), lks2.b from Sermo in<br />
Compana (CIho 5438)*7 (CIho 16188, GP 40), lks2.b from Sermo in Decap (CIho<br />
3351)*7 (CIho 16562, GP 44) (4); lks2.b from R.I. Wolfe's Multiple Recessive<br />
Stock in Bowman (PI 483237)*9 (GSHO 1850).<br />
References:<br />
1. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
2. Eslick, R.F., and E.A. Hockett. 1967. Allelism for awn length, lk2, in barley<br />
(Hordeum species). Crop Sci. 7:266-267.<br />
3. Eslick, R.F., and E.A. Hockett. 1972. Recombination values of four genes on<br />
chromosome 1. BGN 2:123-126.<br />
4. Hockett, E.A. 1981. Registration of hulless and hulless short-awned spring<br />
barley germplasm (Reg. nos. GP 35 to 52). Crop Sci. 21:146-147.<br />
5. Litzenberger, S.C., and J.M. Green. 1951. Inheritance of awns in barley.<br />
Agron. J. 43:117-123.<br />
197
Barley Genetics Newsletter (2007) 37: 188-301<br />
6. Miyake, K., and Y. Imai. 1922. [Genetic studies in barley. 1.] Bot. Mag., Tokyo<br />
36:25-38. [In Japanese.]<br />
7. Myler, J.L. 1942. Awn inheritance in barley. J. Agric. Res. 65:405-412.<br />
8. Pozzi, C., P. Faccioli, V. Terzi, A.M. Stanca, S. Cerioli, P. Castiglioni, R. Fink,<br />
R. Capone, K.J. Müller, G. Bossinger, W. Rohde, and F. Salamini. 2000.<br />
Genetics of mutations affecting the development of a barley floral bract. Genetics<br />
154:1335-1346.<br />
9. Ramage, T. 1984. A semi-dominant short awn mutant in Morex. Barley Genet.<br />
Newsl. 14:19-20.<br />
10. Ramage, T., and J.L.A. Eckhoff. 1985. Assignment of mutants in Morex to<br />
chromosomes. Barley Genet. Newsl. 15:22-25.<br />
11. Robertson, D.W., G.A. Wiebe, and F.R. Immer. 1941. A summary of linkage<br />
studies in barley. J. Am. Soc. Agron. 33:47-64.<br />
12. So, M., S. Ogura, and Y. Imai. 1919. [A linkage group in barley.] Nogaku<br />
Kaiho 208:1093-1117. [In Japanese.]<br />
13. Takahashi, R., J. Hayashi, T. Konishi, and I. Moriya. 1975. Linkage analysis<br />
of barley mutants. Barley Genet. Newsl. 5:56-60.<br />
14. Takahashi, R., J. Yamamoto, S. Yasuda, and Y. Itano. 1953. Inheritance and<br />
linkage studies in barley. Ber. Ohara Inst. landw Forsch. 10:29-52.<br />
15. Takezaki, Y. 1927. [On the genetical formulae of the length of spikes and<br />
awns in barley, with reference to the computation of the valency of the heredity<br />
factors.] Rep. Agric. Exp. Sta., Tokyo 46:1-43. [In Japanese.]<br />
16. Ubisch, G. von. 1921. Beitrag zu einer Faktorenanalyse von Gerste. III. Z.<br />
Indukt. Abstammungs. Vererbungsl. 25:198-200.<br />
Prepared:<br />
R. Takahashi. 1972. BGN 2:176.<br />
Revised:<br />
R. Takahashi and T. Tsuchiya. 1973. BGN 3:119.<br />
J.D. Franckowiak and T. Konishi. 1997. BGN 26:54-55.<br />
J.D. Franckowiak 2007. BGN 37:197-198.<br />
198
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 22, Reaction to Schizaphis graminum 1, Rsg1<br />
Stock number: BGS 22<br />
Locus name: Reaction to Schizaphis graminum 1 (greenbug)<br />
Locus symbol: Rsg1<br />
Previous nomenclature and gene symbolization:<br />
Greenbug resistance = Grb (8).<br />
Resistance to Schizaphis graminum Rondani (greenbug) = Rsg,,a (3).<br />
Inheritance:<br />
Monofactorial dominant (2, 3, 9).<br />
Located in chromosome 7H [1] (4).<br />
Description:<br />
Resistant seedlings infested with greenbugs (aphids) are not killed, while<br />
susceptible seedlings are killed, eight weeks after infestation by the buildup of<br />
the greenbug population (2, 3, 4). The resistance provided by Post 90 (PI<br />
549081), having the Rsg1.a gene, to most S. graminum biotypes was commonly<br />
2 to 3 readings on a 1 to 9 scale (7). Accessions with the Rsg1.a gene conferred<br />
resistance to most, but not all greenbug populations (5).<br />
Origin of mutant:<br />
Natural occurrence in Bozu Omugi (OUJ028, PI 87181), Derbent (PI 76504), and<br />
Kearney (PI 539126, CIho 7580) (1, 3).<br />
Mutational events:<br />
Rsg1.a in Bozu Omugi, Derbent, Kearney, Dobaku (PI 87817), and CIho 5087<br />
(PI 82683) (3, 7).<br />
Mutant used for description and seed stocks:<br />
Rsg1.a in Bozu Omugi (GSHO 1317); Rsg1.a in Post 90 (PI 549081) from Will ()<br />
(5).<br />
References:<br />
1. Atkins, I.M., and R.G. Dahms. 1945. Reaction of small-grain varieties to<br />
greenbug attack. <strong>US</strong>DA Tech. Bull. 901.<br />
2. Gardenshire, J.H. 1965. Inheritance and linkage studies on greenbug<br />
resistance in barley (Hordeum vulgare L.). Crop Sci. 5:28-29.<br />
3. Gardenshire, J.H., and H.L. Chada. 1961. Inheritance of greenbug resistance<br />
in barley. Crop Sci. 1:349-352.<br />
4. Gardenshire, J.H., N.A. Tuleen, and K.W. Stewart. 1973. Trisomic analysis of<br />
greenbug resistance in barley, Hordeum vulgare L. Crop Sci. 13:684-685.<br />
5. Mornhinweg, D.W., L.H. Edwards, L.H. Smith, G.H. Morgan, J.A. Webster,<br />
D.R. Porter, and B.F. Carver. 2004. Registration of ‘Post 90’ barley. Crop Sci.<br />
44:2263.<br />
6. Porter, D.R., J.D. Burd, and D.W. Mornhinweg. 2007. Differentiating greenbug<br />
resistance genes in barley. Euphytica 153:11-14.<br />
7. Porter, D.R., and D.W. Mornhinweg. 2004. Characterization of greenbug<br />
resistance in barley. Plant Breed. 123:493-494.<br />
8. Robertson, D.W., G.A. Wiebe, and R.G. Shands. 1955. A summary of linkage<br />
studies in cultivated barley, Hordeum species: Supplement II, 1947-1953. Agron.<br />
J. 47:418-425.<br />
9. Smith, O.D., A.M. Schlehuber, and B.C. Curtis. 1962. Inheritance studies of<br />
greenbug (Toxoptera graminum Rond.) resistance in four varieties of winter<br />
barley. Crop Sci. 2:489-491.<br />
Prepared:<br />
199
Barley Genetics Newsletter (2007) 37: 188-301<br />
J.G. Moseman. 1976. BGN 6:119.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:68.<br />
J.D. Franckowiak. 2007. BGN 37:199-200.<br />
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BGS 32, Reaction to Puccinia hordei 9, Rph9<br />
Stock number: BGS 32<br />
Locus name: Reaction to Puccinia hordei 9 (barley leaf rust)<br />
Locus symbol: Rph9<br />
Previous nomenclature and gene symbolization:<br />
Resistance to Puccinia hordei Otth 9 = Pa9 (3, 7, 8).<br />
Resistance to Puccinia hordei Otth 9 = Pa9 (2).<br />
Resistance to Puccinia hordei 12 = Rph12 (1, 2, 9).<br />
Inheritance:<br />
Monofactorial dominant (1, 5, 6).<br />
Located in chromosome 5HL [7L] (5), about 26.1 cM distal from the raw1<br />
(smooth awn 1) locus (5), in bin 5H-11 about 9.3 cM proximal from esterase 9<br />
(Est9) and about 22.5 cM proximal from STS marker ABC155 (1), about 29.2 cM<br />
distal from the var1 (variegated 1) locus (1).<br />
Description:<br />
Seedling reaction types range from 0; or necrotic fleck to 23- or reduced pustule<br />
size (4, 6), but 0; reactions are more common with the Rph9.z allele, formerly<br />
Rph12.z (1, 2, 5). The resistant reaction of the Rph9.i allele is temperature<br />
sensitive and is inactivated above 20°C (3). Heterozygotes show an intermediate<br />
reaction to pathogenic isolates of Puccinia hordei (5). The original cultivar<br />
‘Trumpf’ was also marketed in the United Kingdom as ‘Triumph’.<br />
Origin of mutant:<br />
Natural occurrence in Abyssinian (Hor 2596, CIho 1234) (3, 7); natural<br />
occurrence in Hordeum vulgare subsp. spontaneum, but transferred to the<br />
cultivar Trumpf (Triumph, PI 548762, GSHO 1590) (2, 9).<br />
Mutational events:<br />
Rph9.i in Abyssinian (3, 7); Rph9.z in Trumpf (2, 9).<br />
Mutant used for description and seed stocks:<br />
Rph9.i in Abyssinian (GSHO 1601); Rph9.i in Bowman (PI 483237)*8 (GSHO<br />
1866); Rph12.z in Trumpf (GSHO 1590); Rph12.z in Bowman (PI 483237)*9<br />
(GSHO 2145).<br />
References:<br />
1. Borovkova, I.G., Y. Jin, and B.J. Steffenson. 1998. Chromosomal location and<br />
genetic relationship of leaf rust resistance genes Rph9 and Rph12 in barley.<br />
Phytopathology 88:76-80.<br />
2. Clifford, B.C. 1985. Barley leaf rust. p. 173-205. In W.R. Bushnell and A.P.<br />
Roelfs (eds.) The Cereal Rusts, Vol. II. Academic Press, New York.<br />
3. Clifford, B.C., and A.C.C. Udeogalanya. 1976. Hypersensitive resistance of<br />
barley to brown rust (Puccinia hordei Otth). p. 27-29. In Proc. 4th Eur. Medit.<br />
Cereal Rusts Conf., Interlaken, Switzerland.<br />
4. Golan, T., Y. Anikster, J.G. Moseman, and I. Wahl. 1978. A new virulent strain<br />
of Puccinia hordei. Euphytica 27:185-189.<br />
5. Jin, Y., G.D. Stadler, J.D. Franckowiak, and B.J. Steffenson. 1993. Linkage<br />
between leaf rust resistance genes and morphological markers in barley.<br />
Phytopathology 83:230-233.<br />
6. Reinhold, M., and E.L. Sharp. 1982. Virulence types of Puccinia hordei from<br />
North America, North Africa and the Middle East. Plant. Dis. 66:1009-1011.<br />
7. Tan, B.H. 1977. A new gene for resistance to Puccinia hordei in certain<br />
Ethiopian barleys. Cereal Rust Bull. 5:39-43.<br />
201
Barley Genetics Newsletter (2007) 37: 188-301<br />
8. Udeogalanya, A.C.C., and B.C. Clifford. 1976. Genetical, physiological and<br />
pathological relationships of resistance to Puccinia hordei and P. striiformis in<br />
Hordeum vulgare. Trans. Br. Mycol. Soc. 71:279-287.<br />
9. Walther, U. 1987. Inheritance of resistance to Puccinia hordei Otth in the<br />
spring barley variety Trumpf. Cereal Rusts Powdery Mildews Bull. 15:20-26.<br />
Prepared:<br />
J.D. Franckowiak and Y. Jin. 1997. BGN 26:81.<br />
J.D. Franckowiak and Y. Jin. 1997. BGN 26:281 as BGS 333, Reaction to<br />
Puccinia hordei 12, Rph12.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:201-202.<br />
202
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Stock number: BGS 41<br />
Locus name: Brachytic 7<br />
Locus symbol: brh7<br />
BGS 41, Brachytic 7, brh7<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-w = brh.w (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 5HS [7S] (1), approximately 4.6 cM proximal from SSR<br />
marker Bmac0113 in bin 5H-04 (1).<br />
Description:<br />
Plants are about 5/6 of normal height and awns are about 3/4 of normal length.<br />
The rachis internodes are slightly shorter than normal for Bowman. The seedling<br />
leaf of brh7 plants is short and wide and leaf blades are wider than those of<br />
normal sibs. The Bowman line with brh7 showed less lodging than Bowman.<br />
Although the kernels of brh7 plants seem plumper and more globose shaped<br />
than those from normal sibs, the primary difference is a 10 to 15% reduction in<br />
kernel length. Kernel weights and grain yields of the brh7 line are slightly lower<br />
than those of normal Bowman (1, 2).<br />
Origin of mutant:<br />
An induced mutant in Volla (PI 280423) (4).<br />
Mutational events:<br />
brh7.w in Volla (7101, DWS1211) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh7.w in Volla (GSHO 1687); brh7.w in Bowman (PI 483237)*7 (GSHO 1943).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: semidwarf genes.<br />
A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Gaul, H. 1986. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:81.<br />
Revised:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:203.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 44<br />
Locus name: Brachytic 16<br />
Locus symbol: brh16<br />
BGS 44, Brachytic 16, brh16<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-v = brh.v (2).<br />
Inheritance:<br />
Monofactorial recessive (2, 3).<br />
Located in chromosome 7HL [1L] (1), approximately 7.4 cM proximal from SSR<br />
marker Bmag0135 in bin 7H-13 (1).<br />
Description:<br />
Plants are less than 2/3 of normal height and awns are about 3/4 of normal<br />
length in the Bowman backcross-derived line. The peduncle is about 2/3 normal<br />
length. The rachis internodes are slightly shorter than normal. The tip of the spike<br />
has a fascinated appearance because spikelets are very close together. The<br />
seed yield of the Bowman line with brh16 was less than 1/3 of Bowman’s yield.<br />
Since kernels per spikes and kernel size were not reduced, much of the yield<br />
loss was probably associated with reduced tillering (1). The original introduction<br />
(HE 2816) contained two dwarf mutants, but only brh16.v gene was isolated in<br />
the Bowman backcross-derived line.<br />
Origin of mutant:<br />
Probably an ethyl methanesulphonate induced mutant in Korál (PI 467778) (4).<br />
Mutational events:<br />
brh16.v in HE 2816 (DWS1176) from a cross between two semidwarf mutants (3,<br />
4).<br />
Mutant used for description and seed stocks:<br />
brh16.v in HE 2816/Bowman (GSHO 1686); brh16.v in Bowman (PI 483237)*7<br />
(GSHO 2177).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
3. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: semidwarf genes.<br />
A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
4. Váša, M. 1986. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:204.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 60<br />
Locus name: Liguleless 1<br />
Locus symbol: lig1<br />
BGS 60, Liguleless 1, lig1<br />
Previous nomenclature and gene symbolization:<br />
Ligule and auricle less = al (9).<br />
Liguleless = li (8).<br />
Exauriculum = aur-a (1).<br />
Inheritance:<br />
Monofactorial recessive (9).<br />
Located in chromosome 2HL (6, 9, 10); about 25.1 cM distal from the mtt4<br />
(mottled leaf 4) locus (2); and near AFLP marker E3633-1 in subgroup 21 of the<br />
Proctor/Nudinka map (7).<br />
Description:<br />
The ligule and auricle of all leaves are absent, and the leaf blades are erect<br />
along the stem. Liguleless plants can be identified visually at all stages of growth<br />
(9). Reverse mutation of some mutants is possible (4). The fine structure analysis<br />
of the lig1 locus conducted by Konishi (5) showed that some mutants can<br />
recombine. Bowman backcross-derived lines with lig1 gene are similar in<br />
agronomic traits and maturity to Bowman (2).<br />
Origin of mutant:<br />
A spontaneous mutant in an unknown cultivar, Muyoji (liguleless) (8).<br />
Mutational events:<br />
lig1.my as Muyoji (OUL007) (9); lig1.ky in Koyo (PI 190819), lig1.a1 (OUM001),<br />
lig1.a2 in Akashinriki (PI 467400, OUJ659); lig1.c1, lig1.c2, lig1.c3, lig1.c4 in<br />
Chikurin Ibaraki 1 (OUJ030, CIho 7370) (5); aur-a.1 (lig1.b1) (NGB 114359), aura.2<br />
(lig1.b2) (NGB 114360), aur-a.7 (lig1.b7) (NGB 114365), aur-a.8 (lig1.b8)<br />
(NGB 114366), aur-a.9 (lig1.b9) (NGB 114367) in Bonus (PI 189763), aur-a.3<br />
(lig1.b3) (NGB 114361), aur-a.4 (lig1.b4) (NGB 114362), aur-a.5 (lig1.b5) (NGB<br />
114363), aur-a.6 (lig1.b6) (NGB 114364), aur-a.10 (lig1.b10) (NGB 114368) in<br />
Foma (CIho 11333) (5); aur-a.11 (NGB 114369), aur-a.12 (NGB 114370, NGB<br />
114371) in Kristina, aur-a.13 (NGB 114372), aur-a.14 (NGB 114373) in Bonus,<br />
aur-a.15 (NGB 119377) in Golf (PI 488529) (6); lig1.2 in Bonus, found in eli-2<br />
(eligulum-2) (NGB 115389) stock as the second mutant (2).<br />
Mutant used for description and seed stocks:<br />
lig1.my as Muyoji (GSHO 6); lig1.my in Bowman (PI 483237)*8 (GSHO 1930);<br />
lig1.2 in Bowman*5 (2).<br />
References:<br />
1. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Hayashi, J., T. Konishi, I. Moriya, and R. Takahashi. 1984. Inheritance and<br />
linkage studies in barley. VI. Ten mutant genes located on chromosomes 1 to 7,<br />
except 3. Ber. Ohara Inst. landw. Biol., Okayama Univ. 18:227-250.<br />
4. Konishi, T. 1975. Reverse mutation at the ligule-less locus (li) of barley. BGN<br />
5:21-23.<br />
5. Konishi, T. 1981. Reverse mutation and interallelic recombination at the liguleless<br />
locus in barley. p. 838-845. In M.J.C. Ascher, R.P. Ellis, A.M. Hayter, and<br />
R.N.H. Whitehouse. (eds.) Barley Genetics. IV. Proc. Fourth Int. Barley Genet.<br />
Symp. Edinburgh. Edinburgh University Press.<br />
6. Lundqvist, U. (Unpublished).<br />
205
Barley Genetics Newsletter (2007) 37: 188-301<br />
7. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
8. Robertson, D.W., G.A. Wiebe, and R.G. Shands. 1955. A summary of linkage<br />
studies in barley: Supplement II, 1947-1953. Agron. J. 47:418-425.<br />
9. Takahashi, R., J. Yamamoto, S. Yasuda, and Y. Itano. 1953. Inheritance and<br />
linkage studies in barley. Ber. Ohara Inst. landw. Forsch. 10:29-52.<br />
10. Woodward, R.W. 1957. Linkages in barley. Agron. J. 49:28-32.<br />
Prepared:<br />
T. Tsuchiya and T.E. Haus. 1971. BGN 1:120.<br />
Revised:<br />
J.D. Franckowiak, U. Lundqvist, T. Konishi. 1997. BGN26:96.<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:205-206.<br />
206
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 79<br />
Locus name: White streak 7<br />
Locus symbol: wst7<br />
BGS 79, White streak 7, wst7<br />
Previous nomenclature and gene symbolization:<br />
Ribbon grass = rb (6).<br />
White streak-k = wst,,k (10).<br />
White streak-B = wst,,B (8).<br />
Inheritance:<br />
Monofactorial recessive (2, 11).<br />
Located in chromosome 2HL (5, 7, 8,9), about 22.0 cM distal from the gpa1<br />
(grandpa 1) locus (2, 9), over 29.4 cM distal from the lig1 (liguleless 1) locus (8),<br />
in bin 2H-15 about 6.1 cM from RFLP marker MWG949A (1).<br />
Description:<br />
Vertical white streaks of variable width and number develop in the leaf blades of<br />
young secondary tillers. Fewer white streaks and fewer tillers with white streaks<br />
occur as environmental conditions become warm. White streaks can be found<br />
until near maturity, but they are difficult to observe after heading under field<br />
conditions. Often the lower or first leaves on early tillers have more and wider<br />
streaks. The mutant has no apparent affect on agronomic traits in the Bowman<br />
backcross-derived line (4).<br />
Origin of mutant:<br />
A spontaneous mutant isolated by Robertson (6, 11).<br />
Mutational events:<br />
wst7.k in an unknown cultivar (2, 11).<br />
Mutant used for description and seed stocks:<br />
wst7.k in an unknown cultivar (GSHO 247); wst7.k from R.I. Wolfe's Multiple<br />
Recessive Marker Stock in Bowman (PI 483237)*7 (GSHO 1935).<br />
References:<br />
1. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
2. Doney, D.L. 1961. An inheritance and linkage study of barley with special<br />
emphasis on purple pigmentation or the auricle. M.S. Thesis. Utah State Univ.,<br />
Logan.<br />
3. Franckowiak, J.D. 1996. Coordinator's report: Chromosome 2. Barley Genet.<br />
Newsl. 25:88-90.<br />
4. Franckowiak, J.D. (Unpublished).<br />
5. Kasha, K.J. 1982. Coordinator's report: Chromosome 6. Barley Genet. Newsl.<br />
12:90-92.<br />
6. Robertson, D.W., G.A. Wiebe, and R.G. Shands. 1947. A summary of linkage<br />
studies in barley: Supplement I, 1940-1946. J. Am. Soc. Agron. 39:464-473.<br />
7. Schondelmaier, J., G. Fischbeck, and A. Jahoor. 1993. Linkage studies<br />
between morphological and RFLP markers in the barley genome. Barley Genet.<br />
Newsl. 22:57-62.<br />
8. Shin, J.S., S. Chao, L. Corpuz, and T.K. Blake. 1990. A partial map of the<br />
barley genome incorporating restriction fragment length polymorphism,<br />
207
Barley Genetics Newsletter (2007) 37: 188-301<br />
polymerase chain reaction, isozyme, and morphological marker loci. Genome<br />
23:803-810.<br />
9. Walker, G.W.R. 1974. Linkage data for rb and mt3. Barley Genet. Newsl. 4:90-<br />
91.<br />
10. Wolfe, R.I., and J.D. Franckowiak. 1991. Multiple dominant and recessive<br />
genetic marker stocks in spring barley. Barley Genet. Newsl. 20:117-121.<br />
11. Woodward, R.W. 1957. Linkages in barley. Agron. J. 49:28-32.<br />
Prepared:<br />
J.D. Franckowiak and R.I. Wolfe. 1997. BGN 26:117.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:207-208.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 82<br />
Locus name: Zeocriton 1<br />
Locus symbol: Zeo1<br />
BGS 82, Zeocriton 1, Zeo1<br />
Previous nomenclature and gene symbolization:<br />
"Kurz und dicht" = Knd (6).<br />
Inheritance:<br />
Monofactorial incomplete dominant (5).<br />
Located in chromosome 2HL, about 9.2 cM distal from the lig1 (liguleless 1)<br />
locus (4), in bin 2H-13 about 7.3 cM distal from RFLP marker cnx1 (1).<br />
Description:<br />
Plants heterozygous for Zeo1 have short culms, compact spikes, and wide<br />
kernels. Homozygotes have shorter culms (short peduncle), very compact<br />
spikes, large outer glumes with long awns, and reduced fertility. Generally, the<br />
spike emerges from the side of the sheath in homozygotes. Although the name<br />
zeocriton is used for this gene, this gene is not from Spratt, the dense ear type<br />
described by Engledow (2).<br />
Origin of mutant:<br />
An X-ray induced mutant in Donaria (PI 161974) (5).<br />
Mutational events:<br />
Zeo1.a in Donaria (Mut 2657) (5); Zeo1.b, received as "Kurz und dicht" and<br />
placed in R.I. Wolfe's Multiple Dominant Marker Stock (GSHO 1614), was<br />
probably derived from Mut 2657 (3, 6).<br />
Mutant used for description and seed stocks:<br />
Zeo1.a in Donaria (GSHO 1613); Zeo1.a in Bowman (PI 483237)*5 (GSHO<br />
1931); Zeo1.b in Bowman*9 (GSHO 1932).<br />
References:<br />
1. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szücs, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
2. Engledow, F.L. 1924. Inheritance in barley. III. The awn and the lateral floret<br />
(cont'd): fluctuation: a linkage: multiple allelomorphs. J. Genet. 14:49-87.<br />
3. Franckowiak, J.D. 1992. Allelism tests among selected semidwarf barleys.<br />
Barley Genet. Newsl. 21:17-23.<br />
4. Luna Villafaña, A., and J.D. Franckowiak. 1995. (Unpublished).<br />
5. Scholz, F., and O. Lehmann. 1958. Die Gaterslebener Mutanten der<br />
Saatgerste in Beziehung zur Formenmannigfaltigkeit der Art Hordeum vulgare<br />
L.s.l. I. Kulturpflanze 6:123-166.<br />
6. Wolfe, R.I. (Unpublished).<br />
Prepared:<br />
J.D. Franckowiak and R.I. Wolfe. 1997. BGN 26:120.<br />
J.D. Franckowiak. 2007. BGN 37:209.<br />
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Stock number: BGS 85<br />
Locus name: Yellow streak 4<br />
Locus symbol: yst4<br />
BGS 85, Yellow streak 4, yst4<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Located in chromosome 2HL, near the vrs1 (six-rowed spike 1) locus (2), in bin<br />
2H-07 near RFLP marker CDO537 (3).<br />
Description:<br />
Plants have a yellow-green color with numerous, vertical yellow streaks in the<br />
leaves. The yellow-green color is retained until maturity, but the yellow streaks<br />
may be difficult to observe after heading. Plant vigor and height are reduced,<br />
heading is delayed, and seed yields are low.<br />
Origin of mutant:<br />
A sodium azide induced mutant in Glenn (CIho 15769) (1).<br />
Mutational events:<br />
yst4.d in Glenn (DWS1059) (2).<br />
Mutant used for description and seed stocks:<br />
yst4.d in Glenn (GSHO 2502); yst4.d in Bowman (PI 483237)*7 (GSHO 1922).<br />
References:<br />
1. Faue, A.C. 1987. Chemical mutagenesis as a breeding tool for barley. M.S.<br />
Thesis. North Dakota State Univ., Fargo.<br />
2. Faue, A.C., A.E. Foster, and J.D. Franckowiak. 1989. Allelism testing of an<br />
induced yellow streak mutant with the three known yellow streak mutants. Barley<br />
Genet. Newsl. 19:15-16.<br />
3. Kleinhofs, A. 2002. Integrating molecular and morphological/physiological<br />
marker maps. Coordinator’s Report. Barley Genet. Newsl. 32:152-159.<br />
Prepared:<br />
J.D. Franckowiak. 1997. BGN 26:123.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:210.<br />
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Stock number: BGS 87<br />
Locus name: Chlorina seedling 14<br />
Locus symbol: fch14<br />
BGS 87, Chlorina seedling 14, fch14<br />
Previous nomenclature and gene symbolization:<br />
Chlorina seedling 14 = f14 (2).<br />
Inheritance:<br />
Monofactorial recessive (2, 4).<br />
Located in chromosome 2HL (2), probably between the vrs1 (six-rowed spike 1)<br />
and the ant2 (anthocyanin-less 2) loci (2), likely in bin 2H-11 (3, 4).<br />
Description:<br />
Seedlings have a pale yellow-green color. The leaves gradually become greener<br />
starting at the tip of the leaf blade, and mutant plants are indistinguishable in<br />
color from normal sibs at heading (2). When grown in the field, plants produce<br />
slightly thinner kernels with about a 10% reduction in kernel weight (1).<br />
Origin of mutant:<br />
A spontaneous mutant in Shyri (Lignee 640//Kober/Teran 78) from Ecuador (2).<br />
Mutational events:<br />
fch14.w in Shyri (2, 5).<br />
Mutant used for description and seed stocks:<br />
fch14.w in Shyri (GSHO 1739); fch14.w in Bowman (PI 483237)*6 (GSHO 1911).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived<br />
lines of spring barley. Barley Genet. Newsl. 24:63-70.<br />
3. Kleinhofs, A. 2006. Integrating molecular and morphological/physiological<br />
marker maps. Barley Genet. Newsl. 36:66-82.<br />
4. Kudrna, D., A. Kleinhofs, A. Kilian, and J. Soule. 1996. Integrating visual<br />
markers with the Steptoe x Morex RFLP map. p. 343. In A.E. Slinkard, G.J.<br />
Scoles, and B.G. Rossnagel (eds.) Proc. Fifth Int. Oat Conf. & Seventh Int.<br />
Barley Genet. Symp., Saskatoon. Univ. of Saskatchewan, Saskatoon.<br />
5. Rivendeneira, M. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak. 1997. BGN 26:125.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:211.<br />
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BGS 88, Reaction to Puccinia hordei 2, Rph2<br />
Stock number: BGS 88<br />
Locus name: Reaction to Puccinia hordei 2 (barley leaf rust)<br />
Locus symbol: Rph2<br />
Previous nomenclature and gene symbolization:<br />
Resistance to Puccinia anomala Rostr = Pa (2).<br />
Resistance to Puccinia hordei Otth 2 = Pa2 (9, 15).<br />
Resistance to Puccinia hordei A = A (8, 9).<br />
Inheritance:<br />
Monofactorial incomplete dominant (2, 14).<br />
Located in chromosome 5HS [7S] (1), not in the long arm (10), distal from the<br />
secondary constriction (1), in bin 5H-04 about 3.5 cM proximal from RFLP<br />
marker CDO749 (1).<br />
Description:<br />
The seedling reaction type is 0 n - 1 c with race 4 culture 57-19 (2); heterozygotes<br />
have reaction types ranging from 1 to 3, depending on parents. Responses will<br />
vary for homozygotes and heterozygotes when different rust cultures are tested<br />
(8).<br />
Origin of mutant:<br />
Natural occurrence in Peruvian (CIho 935) and several other cultivars (2, 4, 6,<br />
12, 15, 16).<br />
Mutational events:<br />
Rph2.b in Peruvian (4, 12); Rph2.j in Batna (CIho 3391) (7, 12); Rph2.k in<br />
Weider (No 22, PI 39398) (2, 11, 15); Rph2.l in Juliaca (PI 39151) (3, 12);<br />
Rph2.m in Kwan (PI 39367, GSHO 1392) ( 2, 4, 12); Rph2.n in Chilean D (PI<br />
48136) (4, 14); an allele at the Rph2 locus is present in Purple Nepal (CIho<br />
1373), Modia (CIho 2483), Morocco (CIho 4975), Barley 305 (CIho 6015), Marco<br />
(PI 94877) (2); Austral (CIho 6358) (4, 6, 7, 12); Marocaine 079 (CIho 8334) (6);<br />
Q21861 (PI 584766), TR306 (1, 13); accessions with a second Rph gene<br />
besides the Rph2 allele include Carre 180 (CIho 3390), CIho 14077 (12); Ricardo<br />
(PI 45492) (2, 14, 16); Ariana (CIho 14081) (11, 12, 16); Quinn (PI 39401) (8, 9);<br />
Bolivia (PI 36360) (2, 8, 9); Reka 1 (CIho 5051) (4, 6, 7, 12); tentative Rph2 allele<br />
symbols are Rph2.q in Quinn, Rph2.r in Bolivia (GSHO 1598), Rph2.s in Ricardo,<br />
Rph2.t in Reka 1 (GSHO 1594), and Rph2.u in Ariana based on differential<br />
reactions and different cultivar origins (5, 8, 9, 12); Rph2.y from HJ198*3/HS2310<br />
(PI 531841, GSHO 1595) (3).<br />
Mutant used for description and seed stocks:<br />
Rph2.b in Peruvian (GSHO 1593); Rph2.b in Bowman (PI 483237)*3 (GSHO<br />
2320); Rph2.t from Rika 1 in Bowman*8 (GSHO 2321).<br />
References<br />
1. Borovkova, I.G., Y. Jin, B.J. Steffenson, A. Kilian, T.K. Blake, and A. Kleinhofs.<br />
1997. Identification and mapping of a leaf rust resistance gene in the barley line<br />
Q21861. Genome 40:236-241.<br />
2. Henderson, M.T. 1945. Studies of the sources of resistance and inheritance of<br />
reaction to leaf rust, Puccinia anomala Rostr., in barley. Ph.D. Thesis. Univ. of<br />
Minnesota, St. Paul.<br />
3. Jin, Y., G.H. Cui, B.J. Steffenson, and J.D. Franckowiak. 1996. New leaf rust<br />
resistance genes in barley and their allelic and linkage relationships with other<br />
Rph genes. Phytopathology 86:887-890.<br />
212
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4. Levine, M.N., and W.J. Cherewick. 1952. Studies on dwarf leaf rust of barley.<br />
U.S. Dept. Agr. Tech. Bull.1056. 17p.<br />
5. Moseman, J.G., and L.W. Greeley. 1965. New physiological strains of Puccinia<br />
hordei among physiological races identified in the United States from 1959<br />
through 1964. Plant Dis. Rep. 49:575-578.<br />
6. Moseman, J.G., and C.W. Roane. 1959. Physiologic races of barley leaf rust<br />
(Puccinia hordei) isolated in the United States from 1952 to 1958. Plant Dis. Rep.<br />
43:1000-1003.<br />
7. Reinhold, M., and E.L. Sharp. 1982. Virulence types of Puccinia hordei from<br />
North America, North Africa and the Middle East. Plant. Dis. 66:1009-1011.<br />
8. Roane, C.W. 1962. Inheritance of reaction to Puccinia hordei in barley. I.<br />
Genes for resistance among North American race differentiating varieties.<br />
Phytopathology 52:1288-1295.<br />
9. Roane, C.W., and T.M. Starling. 1967. Inheritance of reaction to Puccinia<br />
hordei in barley. II. Gene symbols for loci in different cultivars. Phytopathology<br />
57:66-68.<br />
10. Roane, C.W., and T.M. Starling. 1989. Linkage studies with genes<br />
conditioning leaf rust reaction in barley. Barley Newsl. 33:190-192.<br />
11. Sharp, E.L., and M. Reinhold. 1982. Resistance gene sources to Puccinia<br />
hordei in barley. Plant Dis. 66:1012-1013.<br />
12. Starling, T.M. 1955. Sources, inheritance, and linkage relationships of<br />
resistance to race 4 of leaf rust (Puccinia hordei Otth), race 9 of powdery mildew<br />
(Erysiphe graminis hordei El. Marchal), and certain agronomic characters in<br />
barley. Iowa State Coll. J. Sci. 30:438-439.<br />
13. Steffenson, B.J., and Y. Jin. 1997. A multi-allelic series at the Rph2 locus for<br />
leaf rust resistance in barley. Cereal Rusts Powdery Mildews Bull. (in press).<br />
14. Tan, B.H. 1977. Evaluation host differentials of Puccinia hordei. Cereal Rust<br />
Bull. 5:17-23.<br />
15. Watson, I.A., and F.C. Butler. 1947. Resistance to barley leaf rust (Puccinia<br />
anomala Rostr.). Linnean Soc. New South Wales, Proc. 72:379-386.<br />
16. Zloten, R.R. 1952. The inheritance of reaction to leaf rust in barley. M.S.<br />
Thesis. Univ. of Manitoba, Winnipeg.<br />
Prepared:<br />
Y. Jin and J.D. Franckowiak. 1997. BGN 26:126-127.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:212-213.<br />
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BGS 96, Reaction to Puccinia hordei 15, Rph15<br />
Stock number: BGS 96<br />
Locus name: Reaction to Puccinia hordei 15 (barley leaf rust)<br />
Locus symbol: Rph15<br />
Previous nomenclature and gene symbolization:<br />
Rph16 = Reaction to Puccinia hordei 16 (6, 9).<br />
Inheritance:<br />
Monofactorial dominant (1, 2).<br />
Located in chromosome 2HS (1, 6); over 32.3 cM proximal from the vrs1 (sixrowed<br />
spike 1) locus (1); in bin 2H-6 near molecular markers MWG874 (6) and<br />
MWG2133 (9); cosegregation with AFLP marker P13M40 (9); about 25.2cM<br />
distal from the centromere (9); about 14 cM proximal from the Eam1 (Early<br />
maturity 1) locus (3).<br />
Description:<br />
The seedling reaction to most isolates of Puccinia hordei is a relatively large<br />
necrotic fleck, hypersensitive reaction (1). The seedling infection type of<br />
heterozygotes is indistinguishable from that of homozygous resistant seedlings.<br />
Alleles at this locus were found in six of the first seven Rph genes from Hordeum<br />
vulgare subsp spontaneum evaluated in Bowman backcross-derived lines (1, 2).<br />
The Rph15 locus is likely allelic to Rph16 based on the failure to recover<br />
susceptible recombinants (9). Only one of the 350 leaf rust isolates (90-3 from<br />
Israel) was found to be virulent on Rph15 lines (4, 9). Resistance to isolate 90-3<br />
was observed in progeny from a cross between a line with Rph15 to another<br />
source of leaf rust resistance (8). Rph15 represents one of the most effective leaf<br />
rust resistance genes reported in Hordeum vulgare (9).<br />
Origin of mutant:<br />
Natural occurrence in accession PI 355447 of Hordeum vulgare subsp<br />
spontaneum, but isolated in a selection that contained one Rph gene from the<br />
original accession crossed to Bowman (PI 483237) (1, 7).<br />
Mutational events:<br />
Rph15.ad in PI 355447 (1, 2, 5), PI 354937, PI 391024, PI 391069, PI 391089,<br />
and PI 466245 (1, 2); Rph15.ae from HS084 (6, 9); PI 466245 has at least two<br />
genes for leaf rust resistance (7).<br />
Mutant used for description and seed stocks:<br />
Rph15.ad in selection from a cross to Bowman (GSHO 1586); Rph15.ad in<br />
Bowman*8 (GSHO 2330).<br />
References:<br />
1. Chicaiza, O. 1996. Genetic control of leaf rust in barley. Ph.D. dissertation,<br />
North Dakota State Univ., Fargo.<br />
2. Chicaiza, O., J.D. Franckowiak, and B.J. Steffenson. 1996. New sources of<br />
resistance to leaf rust in barley. pp. 706-708. In A.E. Slinkard, G.J. Scoles, and<br />
B.G. Rossnagel (eds.). Proc. Fifth Int. Oat Conf. & Seventh Int. Barley Genet.<br />
Symp., Saskatoon. Univ. of Saskatchewan, Saskatoon.<br />
3. Falk, A.B. (Personal communications).<br />
4. Fetch, T.G. Jr., B.J. Steffenson, and Y. Jin. 1998. Worldwide virulence of<br />
Puccinia hordei on barley. Phytopathology 88:S28.<br />
5. Franckowiak, J.D., Y. Jin, and B.J. Steffenson. 1997. Recommended allele<br />
symbols for leaf rust resistance genes in barley. Barley Genet. Newsl. 27:36-44.<br />
6. Ivandic, V.,U. Walther, and A. Graner. 1998. Molecular mapping of a new gene<br />
214
Barley Genetics Newsletter (2007) 37: 188-301<br />
in wild barley conferring complete resistance to leaf rust (Puccinia hordei Otth).<br />
Theor. Appl. Genet. 97:1235-1239.<br />
7. Jin, Y., and B.J. Steffenson. 1994. Inheritance of resistance to Puccinia hordei<br />
in cultivated and wild barley. J. Hered. 85:451-454.<br />
8. Sun, Y., J.D. Franckowiak, and S.M. Neate. 2005. Reactions of barley lines to<br />
leaf rust, caused by Puccinia hordei. Proceedings of the 18th North American<br />
Barley Researchers Workshop, July 17-20, 2005, Red Deer, Alberta, Canada<br />
http://www1.agric.gov.ab.ca/$department/deptdocs.nsf/all/fcd10135#Reactions<br />
9. Weerasena, J.S., B.J. Steffenson, and A.B. Falk. 2004. Conversion of an<br />
amplified fragment length polymorphism marker into a co-dominant marker in the<br />
mapping of the Rph15 gene conferring resistance to barley leaf rust, Puccinia<br />
hordei Otth. Theor. Appl. Genet. 108:712-719.<br />
Prepared:<br />
J.D. Franckowiak and O. Chicaiza. 1998. BGN 28:29.<br />
Revised:<br />
J.D. Franckowiak. 2005. BGN 35:186.<br />
J.D. Franckowiak. 2007. BGN 37:214-215.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 98<br />
Locus name: Early maturity 6<br />
Locus symbol: Eam6<br />
BGS 98, Early maturity 6, Eam6<br />
Previous nomenclature and gene symbolization:<br />
Early heading = Ea (9).<br />
Early maturity 6 = Ea6 (7).<br />
Inheritance:<br />
Monofactorial dominant (9).<br />
Located in chromosome 2HS, about 13.5 cM proximal from the vrs1 (six-rowed<br />
spike 1) locus (9), near the gsh5 (glossy sheath 5) locus based on linkage drag<br />
(1, 2), near molecular marker ABC167b in bin 2H-08 (5, 8).<br />
Description:<br />
Alleles at the Eam6 locus alter the timing of floral initiation when barley is grown<br />
under long-day conditions. In temperate climates, the Eam6.h gene induces<br />
spring barley to head two to five days earlier than plants with the recessive allele<br />
(1, 5). A much stronger response to long photoperiods is associated with the<br />
Eam1 gene. Tohno-oka et al. (8) reported that Eam6 gene from Morex (CIho<br />
15773) is effective when the photoperiod is 13 hours or longer and that the Eam1<br />
gene from Steptoe (CIho 15229) induces early heading when the photoperiod is<br />
14 hours or longer. In North Dakota, plants with both the Eam1 and Eam6 genes<br />
head one to two days earlier than those with only the Eam1 gene (1). The<br />
factors, Eam1 and Eam6, for early heading were studied possibly by Yasuda (10)<br />
and named “A” and “B”, respectively. A QTL for long-day photoperiod response<br />
in North American two-rowed and six-rowed barleys in the Eam6 region of 2H<br />
was reported by Moralejo et al. (6) and Horsley et al. (3), respectively. Eam6 may<br />
interact with other maturity genes because a QTL for early heading was detected<br />
in 2HS under both short- and long-day environments in the Harrington/Morex<br />
mapping population (4).<br />
Origin of mutant:<br />
Natural occurrence in many spring, six-rowed barley, represented by the cultivar<br />
Morex (CIho 15773) (8).<br />
Mutational events:<br />
Eam6.h in an unknown cultivar (8), possibly Trebi (CIho 936) (1); Eam6.h in<br />
Morex (4, 5, 8).<br />
Mutant used for description and seed stocks:<br />
Eam6.h in Morex (CIho 15773, GSHO 2492); Eam6.h from Nordic (CIho 15216)<br />
in Bowman (PI 483237) (1).<br />
References:<br />
1. Franckowiak, J.D., and G. Yu (Unpublished).<br />
2. Franckowiak, J.D., and U. Lundqvist. 1997. BGS 355, glossy sheath 5, gsh5.<br />
Barley Genet. Newsl. 26:300-301.<br />
3. Horsley, R.D., D. Schmierer, C. Maier, D. Kudrna, C.A. Urrea, B.J. Steffenson,<br />
P.B. Schwarz, J.D. Franckowiak, M.J. Green, B. Zhang, and A. Kleinhofs. 2006.<br />
Identification of QTLs associated with Fusarium head blight resistance in barley<br />
accession CIho 4196. Crop Sci. 46:145-156.<br />
4. Krasheninnik, N. 2005. Genetic association of Fusarium head blight resistance<br />
and morphological traits in barley. Ph.D. Thesis. North Dakota state Univ., Fargo,<br />
ND.<br />
216
Barley Genetics Newsletter (2007) 37: 188-301<br />
5. Marquez-Cedillo, L.A., P.M. Hayes, A. Kleinhofs, W.G. Legge, B.G.<br />
Rossnagel, K. Sato, S.E. Ullrich, and D. M. Wesenberg. 2001. QTL analysis of<br />
agronomic traits in barley based on the doubled haploid progeny of two elite<br />
North American varieties representing different germplasm groups. Theor. Appl.<br />
Genet. 103:625-637.<br />
6. Moralejo, M., J.S. Swanston, P. Muñoz, D. Prada, M. Elía, J.R. Russell, L.<br />
Ramsay, L. Cistué, P. Codesal, A.M. Casa, I. Romagosa, W. Powell, and J.L.<br />
Molina-Cano. 2004. Use of new EST markers to elucidate the genetic differences<br />
in grain protein content between European and North American two-rowed<br />
malting barleys. Theor. Appl. Genet. 110:116-125.<br />
7. Robertson, D.W., G.A. Wiebe, R.G. Shands, and A. Hagberg. 1965. A<br />
summary of linkage studies in cultivated barley, Hordeum species: Supplement<br />
III, 1954-1963. Crop Sci. 5:33-43.<br />
8. Tohno-oka, T., M. Ishit, R. Kanatani, H. Takahashi, and K. Takeda. 2000.<br />
Genetic analysis of photoperiodic response of barley in different daylength<br />
conditions. p. 239-241. In S. Logue (ed.) Barley Genetics VIII. Volume III. Proc.<br />
Eighth Int. Barley Genet. Symp., Adelaide. Dept. Plant Science, Waite Campus,<br />
Adelaide University, Glen Osmond, South Australia.<br />
9. Woodward, R.W. 1957. Linkages in barley. Agron. J. 49:28-32.<br />
10. Yasuda, S. 1958. (Genetic analysis of the response to short photoperiod in a<br />
barley cross by means of the partitioning method.) Nogaku Kenkyu 46:54-62 [In<br />
Japanese].<br />
Prepared:<br />
J.D. Franckowiak and T. Konishi. 2002. BGN 32:86-87.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:216-217.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 100, Slender dwarf 4, sld4<br />
Stock number: BGS 100<br />
Locus name: Slender dwarf 4<br />
Locus symbol: sld4<br />
Previous nomenclature and gene symbolization:<br />
Slender dwarf d = sld.d (2).<br />
Inheritance:<br />
Monofactorial recessive (5, 6).<br />
Located in chromosome 7HS [5S] (4), near AFLP marker E 4134-2 in subgroup 6<br />
of the Proctor/Nudinka map (4).<br />
Description:<br />
Plants with the sld4.d gene have reduced vigor and are light green in color during<br />
early stages of growth (6). The sld4.d mutant is apparently very environmentally<br />
sensitive in the Bowman derived line. Plants can vary from less than 1/2 to 3/4 of<br />
normal height and heading can be delayed over 10 days in certain environments.<br />
The number of fertile spikelets per spike varies from 2/3 normal to near normal.<br />
Depending on the delay in heading, kernels vary from very thin to near normal.<br />
Grain yield of the Bowman backcross-derived line can vary from very low to<br />
nearly normal (1).<br />
Origin of mutant:<br />
A neutron induced mutant in Two-row Glacier (5). (Glacier is available as CIho<br />
6976.)<br />
Mutational events:<br />
sld4.d in Two-row Glacier (80-T-5899-2-13, DWS1368) (2, 3, 5).<br />
Mutant used for description and seed stocks:<br />
sld4.d in Two-row Glacier (GSHO 2479); sld4.d in Bowman (PI 483237)*7<br />
(GSHO 1880).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Franckowiak, J.D. 1999. Coordinator’s report: Semidwarf genes. Barley Genet.<br />
Newsl. 29:74-79.<br />
3. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
4. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
5. Ramage, R.T., and P. Curtis. 1981. A light green, dwarf mutant located on<br />
chromosome 2. Barley Genet. Newsl. 11:37-38.<br />
6. Ramage, R.T., and R.A. Ronstadt-Smith. 1983. Location of a light green dwarf<br />
mutant on chromosome 2. Barley Genet. Newsl. 13:62-64.<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:89.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:218.<br />
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BGS 101, Absent lower laterals 1, als1<br />
Stock number: BGS 101<br />
Locus name: Absent lower laterals 1<br />
Locus symbol: als1<br />
Previous nomenclature and gene symbolization:<br />
Absent lower laterals = als (2).<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Located in chromosome 3HL (2, 3, 5, 6), about 31.2 cM distal from the uzu1 (uzu<br />
1) locus (2), about 39.7 cM proximal from the cur2 (curly 2) locus (3), and near<br />
AFLP marker E4234-11 in subgroup 28 of the Proctor/Nudinka map (4).<br />
Description:<br />
Lateral spikelets at the base of the spike fail to develop or are partially<br />
developed. Tillers are large, coarse, and stiff, and only 1 or 2 tillers are produced<br />
in the six-rowed stock. The plants resemble those of the (cul2) uniculm 2 mutant<br />
(2). Plants of the Bowman backcross-derived line commonly produce 3 to 5 tillers<br />
with short spikes; and seed yields are very low (1).<br />
Origin of mutant:<br />
A gamma-ray induced mutant in Montcalm (CIho 7149) (2).<br />
Mutational events:<br />
als1.a in Montcalm (Alb Acc 281) (2).<br />
Mutant used for description and seed stocks:<br />
als1.a in Montcalm (GSHO 1065); als1.a in Bowman (PI 483237)*7 (GSHO<br />
1990).<br />
References:<br />
1. Babb, S., and G.J. Muehlbauer. 2003. Genetic and morphological<br />
characterization of the barley uniculm 2 (cul2) mutant. Theor. Appl. Genet.<br />
106:846-857.<br />
2. Kasha, K.J., and G.W.R. Walker. 1960. Several recent barley mutants and<br />
their linkages. Can. J. Genet. Cytol. 2:397-415.<br />
3. Konishi, T., J. Hayashi, I. Moriya, and R. Takahashi. 1984. Inheritance and<br />
linkage studies in barley. VII. Location of six new mutant genes on chromosome<br />
3. Ber. Ohara Inst. landw. Biol., Okayama Univ. 18:251-264.<br />
4. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
5. Shahla, A., and T. Tsuchiya. 1983. Telotrisomic analysis in Triplo 3S in barley.<br />
Barley Genet. Newsl. 13:25.<br />
6. Singh, R.J., and T. Tsuchiya. 1974. Further information on telotrisomic<br />
analysis in barley. Barley Genet. Newsl. 4:66-69.<br />
Prepared:<br />
T. Tsuchiya and T.E. Haus. 1971. BGN 1:123.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:135.<br />
J.D. Franckowiak. 2007. BGN 37:219.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 102, Uzu 1, uzu1<br />
Stock number: BGS 102<br />
Locus name: Uzu 1 (semi-brachytic)<br />
Locus symbol: uzu1<br />
Previous nomenclature and gene symbolization:<br />
Normal vs uzu = h (12).<br />
Uzu = u (4).<br />
Uzu (semi-brachytic) = uz (11).<br />
Uzu 2 = uz2 (3, 13, 15).<br />
Uzu 3 = uz3 (3, 13, 15).<br />
Hordeum vulgare BR-insensitive 1 = HvBRI1 (1).<br />
Inheritance:<br />
Monofactorial recessive (4, 7, 9, 11).<br />
Located in chromosome 3HL (5, 6, 11), about 17.6 cM proximal from the alm1<br />
(albino lemma 1) locus (10), in bin 3H-06 near cDNA marker, C1271 (1).<br />
Description:<br />
The uzu1.a gene has pleiotropic effects on the elongation of the coleoptile, leaf,<br />
culm, rachis internode, awn, glume, and kernel (8, 9, 11). These organs are often<br />
reduced in length and increased in width. Changes in organ length are<br />
temperature sensitive, but heading date and maturity are unaltered. The<br />
coleoptile of uzu plants shows a prominent projection or hook near the apex.<br />
Sometimes the coleoptile of the mutant shows a V-shaped notch on the side<br />
opposite from the projection. Thus, the apex of the coleoptile has two notches,<br />
one on each side (9, 13, 14). The temperature sensitive reduction in culm length<br />
of uzu1.a plants ranges from less than 15% in cool environments to over 75% in<br />
warm ones. Chono et al. (1) reported that the uzu1.a or HVBRI1 gene is caused<br />
by a mutation that changed a highly conserved residue of the kinase domain of<br />
BRI1 (Arabidopsis BR-insensitive 1) (brassinosteroids) receptor protein from His-<br />
857 to Arg-857.<br />
Origin of mutant:<br />
Natural occurrence in many cultivars of Japanese origin (8, 9).<br />
Mutational events:<br />
uzu1.a in many Japanese cultivars (9, 13, 15); uzu1.b (092AR) in Aramir (PI<br />
467781) (2).<br />
Mutant used for description and seed stocks:<br />
uzu1.a in Baitori 11 (OUJ371, PI 182624, GSHO 1300); uzu1.a in Bowman (PI<br />
483237)*7 (GSHO 1963).<br />
References:<br />
1. Chono, M., I. Honda, H. Zeniya, K. Yoneyama, D. Saisho, K. Takeda, S.<br />
Takatsuto, T. Hoshino and Y. Watanabe. 2003. A semidwarf phenotype of barley<br />
uzu results from a nucleotide substitution in the gene encoding a putative<br />
brassinosteroid receptor. Plant Physiol. 133:1209-1219.<br />
2. Gruszka, D., J. Zbieszczyk, M. Kwasniewski, I. Szarejko and M. Maluszynski.<br />
2006. A new allele in a uzu gene encoding brassinosteroid receptor. Barley<br />
Genet. Newsl. 36:1-2.<br />
3. Leonard, W.H., H.O. Mann, and L. Powers. 1957. Partitioning method of<br />
genetic analysis applied to plant height inheritance in barley. Colorado Agric.<br />
Expt. St. Tech. Bull. 60:1-24.<br />
4. Miyake, K., and Y. Imai. 1922. [Genetic studies in barley. 1.] Bot. Mag., Tokyo<br />
220
Barley Genetics Newsletter (2007) 37: 188-301<br />
36:25-38. [In Japanese.]<br />
5. Singh, R.J., A. Shahla, and T. Tsuchiya. 1982. Telotrisomic analysis of three<br />
genes with newly obtained telotrisomic, Triplo 3S, in barley. BGN 12:42-44.<br />
6. Singh, R.J., and T. Tsuchiya. 1974. Further information on telotrisomic<br />
analysis in barley. Barley Genet. Newsl. 4:66-69.<br />
7. So, M., S. Ogura, and Y. Imai. 1919. [A linkage group in barley.] J. Sci. Agric.<br />
Soc. Jpn. 208:1093-1117. [In Japanese.]<br />
8. Takahashi, R. 1942. Studies on the classification and the geographical<br />
distribution of the Japanese barley varieties. I. Significance of the bimodal curve<br />
of the coleoptile length. Ber. Ohara Inst. landw. Forsch. 9:71-90.<br />
9. Takahashi, R. 1951. Studies on the classification and geographical distribution<br />
of the Japanese barley varieties. II. Correlative inheritance of some quantitative<br />
characters with the ear types. Ber. Ohara Inst. landw. Forsch. 9:383-398.<br />
10. Takahashi, R., and J. Hayashi. 1959. Linkage study of albino lemma<br />
character in barley. Ber. Ohara Inst. landw. Biol., Okayama Univ. 11:132-140.<br />
11. Takahashi, R., and J. Yamamoto. 1951. Studies on the classification and<br />
geographical distribution of the Japanese barley varieties. III. On the linkage<br />
relation and the origin of the "uzu" or semi-brachytic character in barley. Ber.<br />
Ohara Inst. landw. Forsch. 9:399-410.<br />
12. Takezaki, Y. 1927. On the genetical formulae of the length of spikes and<br />
awns in barley, with reference to the computation of the valency of the heredity<br />
factors. Rep. Agric. Exp. Sta., Tokyo 46:1-42.<br />
13. Tsuchiya, T. 1972. Genetics of uz, uz2 and uz3 for semi-brachytic mutations<br />
in barley. Barley Genet. Newsl. 2:87-90.<br />
14. Tsuchiya, T. 1976. Allelism testing in barley. II. Allelic relationships of three<br />
uzu genes. Crop Sci. 16:496-499.<br />
15. Tsuchiya, T. 1981. Further results on the allelic relationships of three uzu<br />
genes in barley. J. Hered. 72:455-458.<br />
Prepared:<br />
T. Tsuchiya and T.E. Haus. 1971. BGN 1:124.<br />
Revised:<br />
T. Tsuchiya. 1984. BGN 14:92.<br />
J.D. Franckowiak and T. Konishi. 1997. BGN 26:136-137.<br />
J.D. Franckowiak. 2007. BGN 37:220-221.<br />
221
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 108<br />
Locus name: Albino lemma 1<br />
Locus symbol: alm1<br />
BGS 108, Albino lemma 1, alm1<br />
Previous nomenclature and gene symbolization:<br />
Albino lemma = al (9).<br />
Eburatum = ebu-a (3).<br />
Inheritance:<br />
Monofactorial recessive (9).<br />
Located in chromosome 3HS (9), about 16.5 cM distal from the uzu1 (uzu 1)<br />
locus (2, 5, 6, 7, 8, 9), in bin 3H-04 about 4.8 cM proximal from RFLP marker<br />
MWG844B (1).<br />
Description:<br />
The lemma and palea are white in color and mostly devoid of chlorophyll, but<br />
they terminate into green tips with green awns. The basal part of lower leaf<br />
sheaths and stem nodes are devoid of chlorophyll. Ligules and joints between<br />
the leaf sheath and blade are white in color (9, 10). Plant vigor is reduced slightly<br />
and maturity is delayed in the Bowman backcross-derived line.<br />
Origin of mutant:<br />
Spontaneous occurrence in an unknown cultivar (Russia 82) (OUU086, NSL<br />
43389) (9).<br />
Mutational events:<br />
alm1.a in Russia 82 (9); alm1.b in Liberty (CIho 9549) (2); alm1.c (Mut 966/61) in<br />
Proctor (PI 280420) (4); ebu-a.1 (NGB 115236), -a.2 (NGB 115237), -a.3 (NGB<br />
115238) in Foma (CIho 11333) (3, 10); ebu-a.4 (NGB 115239), -a.5 (NGB<br />
115240) in Foma (6).<br />
Mutant used for description and seed stocks:<br />
alm1.a in Russia 82 (GSHO 270); alm1.a in Bowman (PI 483237)*8 (GSHO<br />
1953).<br />
References:<br />
1. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
2. Eslick, R.F., W.L. McProud. 1974. Positioning of the male sterile 5 (msg5) on<br />
chromosome 3. Barley Genet. Newsl. 4:16-23.<br />
3. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
4. Häuser, H., and G. Fischbeck. 1972. Translocations and genetic analysis of<br />
other mutants. Barley Genet. Newsl. 2:28-29.<br />
5. Konishi, T., J. Hayashi, I. Moriya, and R. Takahashi. 1984. Inheritance and<br />
linkage studies in barley. VII. Location of six new mutant genes on chromosome<br />
3. Ber. Ohara Inst. landw. Biol., Okayama Univ. 18:251-264.<br />
6. Lundqvist, U. (Unpublished).<br />
7. Moriya, I., and R. Takahashi. 1980. Linkage studies of three barley mutants.<br />
Barley Genet. Newsl. 10:47-51.<br />
8. Nonaka, S. 1973. A new type of cultivar, Mitake, with very few in number, but<br />
222
Barley Genetics Newsletter (2007) 37: 188-301<br />
thick and stiff culms. Barley Genet. Newsl. 3:45-47.<br />
9. Takahashi, R., and J. Hayashi. 1959. Linkage study of albino lemma character<br />
in barley. Ber. Ohara Inst. landw. Biol., Okayama Univ. 11:132-140.<br />
10. Tsuchiya, T. 1973. Allelism testing between established marker stocks and<br />
Swedish mutants. Barley Genet. Newsl. 3:67-68.<br />
Prepared:<br />
T. Tsuchiya and T.E. Haus. 1971. BGN 1:130.<br />
Revised:<br />
T. Tsuchiya. 1980. BGN 10:111.<br />
J.D. Franckowiak and U. Lundqvist. 1997. BGN 26:143.<br />
J.D. Franckowiak and U. Lundqvist. 2007. BGN 37:222-223.<br />
223
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 122, Reaction to Puccinia hordei 5, Rph5<br />
Stock number: BGS 122<br />
Locus name: Reaction to Puccinia hordei 5 (barley leaf rust)<br />
Locus symbol: Rph5<br />
Previous nomenclature and gene symbolization:<br />
Resistance to Puccinia hordei Otth 5 = Pa5 (6, 7).<br />
Resistance to Puccinia hordei B = B (5).<br />
Resistance to Puccinia hordei Otth = X (5, 6).<br />
Resistance to Puccinia hordei Otth = Pax (6).<br />
Resistance to Puccinia hordei 6 = Rph6.f (11).<br />
Inheritance:<br />
Monofactorial incomplete dominant (3, 5, 6).<br />
Located in chromosome 3HS (4, 11); about 7.0 cM distal from Rph7 (11), about<br />
0.5 cM proximal from RFLP marker CDO549 (11), about 2.5 cM distal from RFLP<br />
marker MWG2021 (4).<br />
Description:<br />
Rph5.e in Magnif 102 showed a seedling infection type of 0 - 0; c with race 4<br />
culture 57-19, and Rph6.f from Bolivia had a 0; n - 1 c seedling infection type with<br />
race 4 culture 57-19. (3). Heterozygotes frequently show an intermediate<br />
response (type 2 or 3 reaction) to inoculation with pathogenic races, and<br />
incomplete dominance is observed in segregating progenies (3, 5, 6). Zhong et<br />
al. (11) demonstrated that Rph5.e is allelic to the Rph6.f gene extracted from<br />
Bolivia (PI 36360). Rph6.f was identified as a monofactorial dominant, but an<br />
allele at the Rph2 (reaction to Puccinia hordei 2) locus is present in the original<br />
cultivar Bolivia (PI 36360) (5, 6, 8).<br />
Origin of mutant:<br />
Natural occurrence in Quinn (PI 39401) (6, 10); natural occurrence in Bolivia (PI<br />
36360) (2, 5).<br />
Mutational events:<br />
Rph5.e in Magnif 102 (PI 337140) (10), Rph5.f (formerly Rph6.f) in Bolivia (11),<br />
Rph5.ai in Quinn along with Rph2.q (5, 6).<br />
Mutant used for description and seed stocks:<br />
Rph5.e in Malteria Heda*4/Quinn (Magnif 102, GSHO 1597) (10); Rph5.e in<br />
Bowman (PI 483237)*8 (GSHO 1865); Rph5.f in Bowman*8 (GSHO 2323);<br />
Rph6.f in Bolivia (GSHO 1598); Rph6.f (without an Rph2 allele) in Bowman (PI<br />
483237)*4 (GSHO 2323) (1).<br />
References:<br />
1. Chicaiza, O., J.D. Franckowiak, and B.J. Steffenson. 1996. Backcross-derived<br />
lines of barley differing for leaf rust resistance genes. pp. 198-200. In G.H.J.<br />
Kema, R.E. Niks, and R.A. Daamen (eds.) Proc. 9th Eur. & Medit. Cereal Rusts<br />
& Mildews Conf. Drukkerij Ponsen en Looijen B.V., Wageningen.<br />
2. Henderson, M.T. 1945. Studies of the sources of resistance and inheritance of<br />
reaction to leaf rust, Puccinia anomala Rostr., in barley. Ph.D. Thesis. Univ. of<br />
Minnesota, St. Paul.<br />
3. Jin, Y., G.H. Cui, B.J. Steffenson, and J.D. Franckowiak. 1996. New leaf rust<br />
resistance genes in barley and their allelic and linkage relationships with other<br />
Rph genes. Phytopathology 86:887-890.<br />
4. Mammadov, J.A., J.C. Zwonitzer, R.M. Biyashev, C.A. Griffey, Y. Jin, B.J.<br />
Steffenson, and M.A. Saghai Maroof. 2003. Molecular mapping of leaf rust<br />
224
Barley Genetics Newsletter (2007) 37: 188-301<br />
resistance gene Rph5 in barley. Crop Sci. 43:388-393.<br />
5. Roane, C.W. 1962. Inheritance of reaction to Puccinia hordei in barley. I.<br />
Genes for resistance among North American race differentiating varieties.<br />
Phytopathology 52:1288-1295.<br />
6. Roane, C.W., and T.M. Starling. 1967. Inheritance of reaction to Puccinia<br />
hordei in barley. II. Gene symbols for loci in differential cultivars. Phytopathology<br />
57:66-68.<br />
7. Roane, C.W., and T.M. Starling. 1970. Inheritance of reaction to Puccinia<br />
hordei in barley. III. Genes in the cultivars Cebada Capa and Franger.<br />
Phytopathology 60:788-790.<br />
8. Starling, T.M. 1955. Sources, inheritance, and linkage relationships of<br />
resistance to race 4 of leaf rust (Puccinia hordei Otth), race 9 of powdery mildew<br />
(Erysiphe graminis hordei El. Marchal), and certain agronomic characters in<br />
barley. Iowa State Coll. J. Sci. 30:438-439.<br />
9. Starling, T.M. 1956. Sources, inheritance, and linkage relationships of<br />
resistance to race 4 of leaf rust (Puccinia hordei Otth), race 9 of powdery mildew<br />
(Erysiphe graminis hordei El. Marchal), and certain agronomic characters in<br />
barley. Iowa State Coll. J. Sci. 30:438-439.<br />
10. Yahyaoui, A.H., and E.L. Sharp. 1987. Virulence spectrum of Puccinia hordei<br />
in North Africa and the Middle East. Plant Dis. 71:597-598.<br />
11. Zhong, S., R.J. Effertz, Y. Jin, J.D. Franckowiak, and B.J. Steffenson. 2003.<br />
Molecular mapping of the leaf rust resistance gene Rph6 in barley and its linkage<br />
relationships with Rph5 and Rph7. Phytopathology 93:604-609.<br />
Prepared:<br />
C.W. Roane. 1976. BGN 6:122.<br />
J.D. Franckowiak and Y. Jin. 1997. BGN 26:501, as BGS 575, Reaction to<br />
Puccinia hordei 6, Rph6.<br />
Revised:<br />
J.D. Franckowiak and Y. Jin. 1997. BGN 26:157.<br />
J.D. Franckowiak and B.J. Steffenson. 2005. BGN 35:188.<br />
J.D. Franckowiak. 2007. BGN 37:224-225.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 130<br />
Locus name: Early maturity 10<br />
Locus symbol: eam10<br />
BGS 130, Early maturity 10, eam10<br />
Previous nomenclature and gene symbolization:<br />
Early maturity sp = easp (8).<br />
Inheritance:<br />
Monofactorial recessive (8).<br />
Located in chromosome 3HL (8); about 2.0 ± 5.8 cM from the Est1-Est4<br />
(esterase 1, esterase 4) locus (8); about 5.8 cM distal from RFLP marker<br />
Xmwg546 (1).<br />
Description:<br />
In winter nurseries at Cuidad Obregón, Sonora, Mexico and Davis, California,<br />
<strong>US</strong>A, plants of Super Precoz 2H head about 11 days earlier than lines with the<br />
genes eam7.g or eam8.k for photoperiod insensitivity from Atsel and Sv Mari,<br />
respectively (8). The eam10.m gene appears to suppress expression of the<br />
eam7.g and eam8.k genes (8). Plants expressing eam10.m become chlorotic<br />
(yellow green) under photothermal stress. Zeaxanthin increases at the expense<br />
of chlorophyll and other pigments (7). The chlorotic appearance is similar to that<br />
observed in plants homozygous for other recessive genes for early maturity<br />
(eam7, eam8, and eam9) (2, 5, 7). Plants in the Bowman eam10.m line head two<br />
days earlier than Bowman under long days and are slightly shorter (5).<br />
Origin of mutant:<br />
Present in Super Precoz 2H (PI 527381) from Russia (7), but originating probably<br />
as an induced mutant in MC20 (3, 4, 7).<br />
Mutational events:<br />
eam10.m in Super Precoz 2H plus a dominant maturity enhancer ( 4, 5, 7);<br />
eam10.m in Amber Nude without the enhancer (4).<br />
Mutant used for description and seed stocks:<br />
eam10.m in Super Precoz 2H (GSHO 2504); eam10.m in Amber Nude (GSHO<br />
2505); eam10.m from Super Precoz in Bowman (PI 483237)*5.<br />
References:<br />
1. Börner, A., G.H. Buck-Sorlin, P.M. Hayes, S. Malyshev, and V. Korzun. 2002.<br />
Molecular mapping of major genes and quantitative trait loci determining<br />
flowering time in response to photoperiod in barley. Plant Breed. 121:129-132.<br />
2. Dormling, I., and Å. Gustafsson. 1969. Phytotron cultivation of early barley<br />
mutants. Theor. Appl. Genet. 39:51-61.<br />
3. Favret, E.A. 1972. El mejoramiento de las plantas por induccíon de<br />
mutaciones en latinoamerica. p. 49-59. In Induced Mutations and Plant<br />
Improvement. Int. Atomic Energy Agency, Vienna.<br />
4. Favret, E.A., and G.S. Ryan. 1966. New useful mutants in plant breeding. p.<br />
49-61. In Mutations in Plant Breeding. Int. Atomic Energy Agency, Vienna.<br />
5. Franckowiak, J.D. (Unpublished).<br />
6. Gallagher, L.W. (Unpublished).<br />
7. Gallagher, L.W., A.A. Hafez, S.S. Goyal, and D.W. Rains. 1994. Nuclear<br />
mutations affecting chloroplastic pigments of photoperiod-insensitive barley.<br />
Plant Breed. 113:65-70.<br />
8. Gallagher, L.W., K.M. Soliman, and H. Vivar. 1991. Interactions among loci<br />
conferring photoperiod insensitivity for heading time in spring barley. Crop Sci.<br />
226
Barley Genetics Newsletter (2007) 37: 188-301<br />
31:256-261.<br />
Prepared:<br />
L.W. Gallagher and J.D. Franckowiak. 1997. BGN 26:166.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:226-227.<br />
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BGS 136, Reaction to Puccinia hordei 7, Rph7<br />
Stock number: BGS 136<br />
Locus name: Reaction to Puccinia hordei 7 (barley leaf rust)<br />
Locus symbol: Rph7<br />
Previous nomenclature and gene symbolization:<br />
Resistance to Puccinia hordei Otth y = Pay (7).<br />
Resistance to Puccinia hordei Otth 5 = Pa5 (10).<br />
Resistance to Puccinia hordei Otth 7 = Pa7 (8).<br />
Inheritance:<br />
Monofactorial dominant (7, 10).<br />
Located in chromosome 3HS (14, 15), linkage to markers in the centromeric<br />
region was reported (11), about 24.0 cM from the ant17 (proanthocyanidin-free<br />
17) locus (5), in bin 3H-01 about 1.3 cM distal from RFLP marker cMWG691 (6),<br />
about 3.2 cM from receptor-like kinase gene Hv3Lrk (2), about 7.0 cM proximal<br />
from Rph5 locus (16).<br />
Description:<br />
The seedling reaction type is 0; n - 1 c (4, 11). Temperature studies show that<br />
resistance conferred by the Rph7.g gene is not expressed well above 20ºC (4,<br />
15). Cebada Capa is indistinguishable from the cultivar Forrajera Klein (possibly<br />
identical to PI 331904) (1). The Rph7 regions from Morex (rph7) and Cebada<br />
Capa (Rph7) were sequenced and compared to similar regions from 39 other<br />
cultivars. The data suggest that a large amount of haplotype variability exists in<br />
the cultivated barley gene pool and indicate rapid and recent divergence at this<br />
locus (12).<br />
Origin of mutant:<br />
Natural occurrence in Cebada Capa (PI 53911) (7, 8, 10).<br />
Mutational events:<br />
Rph7.g in Cebada Capa (7, 8, 10); Rph7.g in France 7 and France 21 (7);<br />
Rph7.g in Dabat, Gondar (PI 199964), and La Estanzuela (9, 13, 16); Rph7.ac in<br />
Tu17a, a Bowman backcross-derived line from Tunisia 17 (3).<br />
Mutant used for description and seed stocks:<br />
Rph7.g in Cebada Capa (GSHO 1318); Rph7.g in Bowman (PI 483237)*8<br />
(GSHO 1994).<br />
References:<br />
1. Arias, G. (Personal communications).<br />
2. Brunner, S., B. Keller, and C. Feuillet. 2000. Molecular mapping of the Rph7.g<br />
leaf rust resistance gene in barley (Hordeum vulgare L.). Theor. Appl. Genet.<br />
101:783-788.<br />
3. Chicaiza, O., J.D. Franckowiak, and B.J. Steffenson. 1996. New sources of<br />
resistance to leaf rust in barley. p. 706-708. In A.E. Slinkard, G.J. Scoles, and<br />
B.G. Rossnagel (eds.) Proc. Fifth Int. Oat Conf. & Seventh Int. Barley Genet.<br />
Symp., Saskatoon. Univ. of Saskatchewan, Saskatoon.<br />
4. Clifford, B.C., and H. W. Roderick. 1981. Detection of cryptic resistance of<br />
barley to Puccinia hordei. Trans. Br. Mycol. Soc. 76:17-24.<br />
5. Falk, D.E. 1985. Genetic studies with proanthocyanidin-free barley. Barley<br />
Genet. Newsl. 15:27-30.<br />
6. Graner, A., S. Streng, A. Drescher, Y. Jin, I. Borovkova, and B.J Steffenson.<br />
2000. Molecular mapping of the leaf rust resistance gene Rph7 in barley. Plant<br />
Breed. 119:389-392.<br />
228
Barley Genetics Newsletter (2007) 37: 188-301<br />
7. Johnson, R. 1968. The genetics of resistance of some barley varieties to<br />
Puccinia hordei. p. 160-162. In Proc. Eur. Medit. Cereal Rust Conf., Oeiras,<br />
Portugal.<br />
8. Nover, I., and C.O. Lehmann. 1974. Resistenzeigenschaften im Gersten- und<br />
Weizensortiment Gatersleben. 18. Prüfung von Sommergersten auf ihr Verhalten<br />
gegen Zwergrost (Puccinia hordei Otth). Kulturpflanze 22:25-43.<br />
9. Parlevliet, J.E. 1976. The genetics of seedling resistance to leaf rust, Puccinia<br />
hordei Otth, in some spring barley cultivars. Euphytica 25:249-254.<br />
10. Roane, C.W., and T.M. Starling. 1970. Inheritance of reaction to Puccinia<br />
hordei in barley. III. Genes in the cultivars Cebada Capa and Franger.<br />
Phytopathology 60:788-790.<br />
11. Roane, C.W., and T.M. Starling. 1989. Linkage studies with genes<br />
conditioning leaf rust reaction in barley. Barley Newsl. 33:190-192.<br />
12. Scherrer, B., E. Isidore, P. Klein, J-S. Kim, A. Bellec, B. Chalhoub, B. Keller,<br />
and C. Feuillet. 2005. Large intraspecific haplotype variability at the Rph7 locus<br />
results from rapid and recent divergence in the barley genome. Plant Cell<br />
17:361-374.<br />
13. Tan, B.H. 1978. Verifying the genetic relationships between three leaf rust<br />
resistance genes in barley. Euphytica 27:317-323.<br />
14. Tuleen. N.A., and M.E. McDonald. 1971. Location of genes Pa and Pa5.<br />
Barley Newsl. 15:106-107.<br />
15. Udeogalanya, A.C.C., and B.C. Clifford. 1976. Genetical, physiological and<br />
pathological relationships of resistance to Puccinia hordei and P. striiformis in<br />
Hordeum vulgare. Trans. Br. Mycol. Soc. 71:279-287.<br />
16. Zhong, S., R.J. Effertz, Y. Jin, J.D. Franckowiak, and B.J. Steffenson. 2003.<br />
Molecular mapping of the leaf rust resistance gene Rph6 in barley and its linkage<br />
relationships with Rph5 and Rph7. Phytopathology 93:604-609.<br />
Prepared:<br />
J.D. Franckowiak and Y. Jin. 1997. BGN 26:173.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:228-229.<br />
229
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 142<br />
Locus name: Brachytic 8<br />
Locus symbol: brh8<br />
BGS 142, Brachytic 8, brh8<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-ad = brh.ad (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 5, 6).<br />
Located in chromosome 3HS (4), near the btr1 (non-brittle rachis 1) locus based<br />
on linkage drag (4), about 26.3 cM proximal from SSR marker HVM60 in bin 3H-<br />
08 (1).<br />
Description:<br />
In the Bowman backcross-derived line, brh8 plants are 3/4 to 5/6 of normal<br />
height and awns are 2/3 to 3/4 of normal length. The peduncle is 3/4 normal<br />
length. The seedling leaf of brh8 plants is shorter and wider than those of normal<br />
sibs and the leaf blades are slightly wider. Kernels of brh8 plants are shorter than<br />
that of normal sibs and their weights are nearly15% lower. Heading dates are 2<br />
or 3 days later, spikes have 3 to 4 more kernels, and rachis internodes are about<br />
20% shorter. Grain yield is nearly normal (1, 2).<br />
Origin of mutant:<br />
Probably a sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494,<br />
NGB 14667) (6).<br />
Mutational events:<br />
brh8.ad in Birgitta (17:16:1, DWS1008) (5, 6).<br />
Mutant used for description and seed stocks:<br />
brh8.ad in Birgitta (GSHO 1671); brh8.ad in Bowman (PI 483237)*8 (GSHO<br />
1944).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived<br />
lines of spring barley. Barley Genet. Newsl. 24:63-70.<br />
5. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
6. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:92.<br />
Revised:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:230.<br />
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Stock number: BGS 148<br />
Locus name: Brachytic 14<br />
Locus symbol: brh14<br />
BGS 148, Brachytic 14, brh14<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-q = brh.q (4).<br />
Inheritance:<br />
Monofactorial recessive (4, 5).<br />
Located in chromosome 3HL (2), approximately 24.9 cM proximal from SSR<br />
marker Bmac0029 in bin 3H-15 (2).<br />
Description:<br />
Plants are about 2/3 normal height and awns, peduncles are about 2/3 normal<br />
length, and rachis internodes are about 7/8 normal length (2, 6, 7). Seedling<br />
leaves of brh14.q plants are relatively short, but they do respond to gibberellic<br />
acid treatment (1). Leaf blades are about 3/4 normal length. The kernels of brh14<br />
plants are slightly shorter and smaller than those of normal sibs, but there are<br />
slightly more kernels per spike. However, the grain yields of the brh14 line to<br />
average 1/3 to 1/4 of those for Bowman reduced because tillering was reduced.<br />
Plants show an erect growth habit (2, 3). Failure of the internode below the<br />
peduncle to elongate was observed in double dwarfs involving brh14.q in the<br />
Akashinriki genetic background (7).<br />
Origin of mutant:<br />
An ethyl methanesulfonate induced mutant in Akashinriki (OUJ659, PI 467400)<br />
(6, 7).<br />
Mutational events:<br />
brh14.q in Akashinriki (OUM131, dw-d, DWS1035, GSHO 1682) (4, 5, 6, 7).<br />
Mutant used for description and seed stocks:<br />
brh14.q in Akashinriki (GSHO 1682); brh14.q in Bowman (PI 483237)*6 (GSHO<br />
2175).<br />
References:<br />
1. Börner, A. 1996. GA response in semidwarf barley. Barley Genet. Newsl.<br />
25:24-26.<br />
2. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
3. Franckowiak, J.D. (Unpublished).<br />
4. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
5. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
6. Konishi, T. 1976. The nature and characteristics of EMS-induced dwarf<br />
mutants in barley. p. 181-189. In H. Gaul (ed.). Barley Genetics III. Proc. Third<br />
Int. Barley Genet. Symp., Garching, 1975. Verlag Karl Thiemig, München.<br />
7. Konishi, T. 1977. Effects of induced dwarf genes on agronomic characters in<br />
barley. p. 21-38. In Use of dwarf mutations. Gamma-Field Symposium No. 16.<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:231.<br />
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BGS 149, Reaction to Puccinia coronata var. hordei 1, Rpc1<br />
Stock number: BGS 149<br />
Locus name: Reaction to Puccinia coronata var. hordei 1<br />
Locus symbol: Rpc1<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial dominant (3).<br />
Located in chromosome 3H centromeric region (1), approximately 2.5 cM from<br />
RAPD marker OPO08-700 (1).<br />
Description:<br />
Crown rust of barley was identified as a new disease of barley in North America<br />
(2). In seedling tests, resistant cultivars exhibited necrotic or chlorotic flecks (0; to<br />
; infection types) at infection sites and no sporulation (3). Adult plant reactions of<br />
Hor2596 were resistant to moderately resistant (3). Hor 2596 is one of the<br />
differential lines for barley leaf rust (caused by Puccinia hordei), see BGS 032,<br />
Rph9.i (reaction to Puccinia hordei 9). The F1 plants from the Bowman/Hor2596<br />
cross exhibited slightly higher infection types (1,2 reaction) than the resistant<br />
parent (3).<br />
Origin of mutant:<br />
Natural occurrence in Abyssinian (Hor 2596, CIho 1234) (3).<br />
Mutational events:<br />
Rpc1.a in Hor 2596 (3).<br />
Mutant used for description and seed stocks:<br />
Rpc1.a in Hor 2596 (GSHO 1601) (3).<br />
References:<br />
1. Agrama, H.A., L. Dahleen, M. Wentz, Y, Jin, and B. Steffenson. 2004.<br />
Molecular mapping of the crown rust resistance gene Rpc1 in barley.<br />
Phytopathology 94:858-861.<br />
2. Jin, Y., and B. Steffenson. 1999. Puccinia coronata var. hordei nov.:<br />
morphology and pathogenicity. Mycologia 91:877-884.<br />
3. Jin, Y., and B. Steffenson. 2002. Sources and genetics of crown rust<br />
resistance in barley. Phytopathology 92:1064-1067.<br />
Prepared:<br />
Y. Jin and J.D. Franckowiak. 2007. BGS 37:232.<br />
232
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 155<br />
Locus name: Glossy leaf 1<br />
Locus symbol: glf1<br />
BGS 155, Glossy leaf 1, glf1<br />
Previous nomenclature and gene symbolization:<br />
Waxless bloom on leaves = w1 (11).<br />
Glossy = gl (9).<br />
Glossy leaves = gl (16).<br />
Glossy leaf = gl (15).<br />
Glossy seedling 2 = gl2 (3, 9).<br />
Eceriferum-zh = cer-zh (4).<br />
Inheritance:<br />
Monofactorial recessive (9).<br />
Located in chromosome 4HL (3, 9, 12, 14), about 7.5 cM distal from the lbi2 (long<br />
basal rachis internode 2) locus (1), and about 4.8 cM distal from the Mlg (Reg2,<br />
reaction to Erysiphe graminis 2) locus (1).<br />
Description:<br />
Surface wax coating on the leaf blade appears absent from the seedling stage to<br />
near maturity, and leaves have a shiny appearance (wax code ++ ++ -) (4).<br />
Plants are semidwarf, relatively weak, and late in heading. The stock in the<br />
Bonus is highly sterile (4), but the Bowman backcross-derived line has nearly<br />
complete fertility. The lack of surface waxes reduces the ability of growing germ<br />
tube of certain fungi to find the stomata openings (10).<br />
Origin of mutant:<br />
A radiation induced mutant in Himalaya (CIho 1312) (9, 13), an X-ray induced<br />
mutant in Bonus (PI 189763) (4).<br />
Mutational events:<br />
glf1.a, glf1.b (gl2, GSHO 22) in Himalaya (13); glf1.f in 34-119-1, glf1.g in II-34-<br />
199-7-2 (GSHO 89) (2); cer-zh.54 (NGB 110938) in Bonus (4, 5); cer-zh.266<br />
(NGB 111153), -zh.308 (NGB 111195), -zh.357 (NGB 111244, NGB 117254), -<br />
zh.366 (NGB 111253), -zh.432 (NGB 111320), -zh.433 (NGB 111321, NGB<br />
117256) in Foma (CIho 11333) (5, 8); cer-zh.325 (NGB 111212) in Foma (5); cerzh.373<br />
(NGB 111260) in Foma (6); cer-zh.865 (NGB 111753) in Bonus (7).<br />
Mutant used for description and seed stocks:<br />
glf1.a in Himalaya (GSHO 98); cer-zh.54 in Bonus (GSHO 455) is used for<br />
allelism tests; glf1.a in Bowman (PI 483237)*8 (GSHO 2015).<br />
References:<br />
1. Forster, B.P. 1993. Coordinator's report: Chromosome 4. Barley Genet. Newsl.<br />
22:75-77.<br />
2. Haus, T.E., and T. Tsuchiya. 1972. Allelic relationships among glossy seedling<br />
mutants. Barley Genet. Newsl. 2:79-80.<br />
3. Immer, F.R., and M.T. Henderson. 1943. Linkage studies in barley. Genetics<br />
28:419-440.<br />
4. Lundqvist, U., and D. von Wettstein. 1962. Induction of eceriferum mutants in<br />
barley by ionizing radiations and chemical mutagens. Hereditas 48:342-362.<br />
5. Lundqvist, U., and D. von Wettstein. 1971. Stock list for the eceriferum<br />
mutants. Barley Genet. Newsl. 1:97-102.<br />
6. Lundqvist, U., and D. von Wettstein. 1973. Stock list for the eceriferum<br />
mutants II. Barley Genet. Newsl. 3:110-112.<br />
233
Barley Genetics Newsletter (2007) 37: 188-301<br />
7. Lundqvist, U., and D. von Wettstein. 1977. Stock list for the eceriferum<br />
mutants IV. Barley Genet. Newsl. 5:92-96.<br />
8. Lundqvist, U., P. von Wettstein-Knowles, and D. von Wettstein. 1968.<br />
Induction of eceriferum mutants in barley by ionizing radiations and chemical<br />
mutagens. II. Hereditas 59:473-504.<br />
9. Robertson, D.W., and O.H. Coleman. 1942. Location of glossy and yellow leaf<br />
seedlings in two linkage groups. J. Am. Soc. Agron. 34:1028-1034.<br />
10. Rubiales, D., M.C. Ramirez, T.L.W. Carver, and R.E. Niks. 2001. Abnormal<br />
germling development by brown rust and powdery mildew on cer barley mutants.<br />
Hereditas 135:271-276.<br />
11. Smith, L., 1951. Cytology and genetics of barley. Bot. Rev. 17:1-355.<br />
12. Tsuchiya, T. 1981. Revised linkage maps of barley. 1981. Barley Genet.<br />
Newsl. 11:96-98.<br />
13. Tsuchiya, T., and T.E. Haus. 1973. Allelism testing in barley. I. Analysis of<br />
ten mapped genes. J. Hered. 64:282-284.<br />
14. Tsuchiya, T., and R.J. Singh. 1982. Chromosome mapping in barley by<br />
means of telotrisomic analysis. Theor. Appl. Genet. 61:201-208.<br />
15. Walker, G.W.R., J. Dietrich, R. Miller, and K. Kasha. 1963. Recent barley<br />
mutants and their linkages II. Genetic data for further mutants. Can. J. Genet.<br />
Cytol. 5:200-219.<br />
16. Woodward, R.W. 1957. Linkages in barley. Agron. J. 49:28-32.<br />
Prepared:<br />
T.E. Haus and T. Tsuchiya. 1971. BGN 1:141 as BGS 155, Glossy seedling, gl;<br />
and BGN 1:145 as BGS 159, Glossy seedling 2, gl2.<br />
U. Lundqvist. 1975. BGN 5:144 as BGS 426, Eceriferum-zh, cer-zh.<br />
Revised:<br />
T. Tsuchiya. 1980. BGN 10:114 as BGS 155, Glossy seedling, gl; and BGN<br />
10:116 as BGS 159, Glossy seedling 2, gl2.<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:181-182.<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:233-234.<br />
234
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 157<br />
Locus name: Brachytic 2<br />
Locus symbol: brh2<br />
BGS 157, Brachytic 2, brh2<br />
Previous nomenclature and gene symbolization:<br />
Brachytic 2 = br2 (9).<br />
Breviaristatum-l = ari-l (4, 5).<br />
Inheritance:<br />
Monofactorial recessive (8).<br />
Located in chromosome 4HL (8), about 1.5 cM proximal from the glf3 (glossy leaf<br />
3) locus (3, 8), over 22.8 cM proximal from the Kap1 (hooded lemma 1) locus (8),<br />
near AFLP marker E4140-7 in subgroup 38-40 of the Proctor/Nudinka map (7),<br />
and about 15.9 cM distal from SSR marker Bmag0353 near the boundary<br />
between bins 4H-06 and 4H-07 (2).<br />
Description:<br />
Plant height and vigor are reduced to about 2/3 normal; the awn is less than 1/4<br />
normal length; the spike is semi-compact; and the leaf, kernel, glume and glume<br />
awn, rachilla, and coleoptile are shorter than in the original cultivar. Auricles are<br />
well developed and larger than those of the original cultivar (9). In the Bowman<br />
backcross-derived lines, the peduncle is about 1/2 normal length, kernel weights<br />
are slightly over 2/3 normal, yield is about 1/2 normal; however, rachis internode<br />
lengths are normal (2). The ari-l.3 allele at the brh2 locus is sensitive to<br />
gibberellic acid treatment (1).<br />
Origin of mutant:<br />
An X-ray induced mutant in Svanhals (PI 5474) (9).<br />
Mutational events:<br />
brh2.b in Svanhals (Kmut 28, OUM283) (8); ari-l.3 (NGB 115848) in Bonus (PI<br />
189763) (5); ari-l.132 (NGB 115942) in Foma (CIho 11333) (6); ari-l.135 (NGB<br />
115945), -l.145 (NGB 115956), -l.214 (NGB 116023), -l.237 (NGB 116047) in<br />
Foma, -l.257 (NGB 116066) in Kristina (NGB 1500) (5).<br />
Mutant used for description and seed stocks:<br />
brh2.b in Svanhals (GSHO 573); ari-l.3 in Bonus (GSHO 1660); brh2.b in<br />
Bowman (PI 483237)*7 (GSHO 2016); ari-l.3 in Bowman*7 (GSHO 2017).<br />
References:<br />
1. Börner, A. 1996. GA response in semidwarf barley. Barley Genet. Newsl.<br />
25:24-26.<br />
2. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
3. Forster, B.P. 1993. Coordinator's report: Chromosome 4. BGN 22:75-77.<br />
4. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
5. Kucera, J., U. Lundqvist, and Å. Gustafsson. 1975. Inheritance of<br />
breviaristatum mutants in barley. Hereditas 80:263-278.<br />
6. Lundqvist, U. (Unpublished).<br />
7. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
8. Takahashi, R., J. Hayashi, and I. Moriya. 1971. Linkage studies in barley.<br />
235
Barley Genetics Newsletter (2007) 37: 188-301<br />
Barley Genet. Newsl. 1:51-58.<br />
9. Tsuchiya, T. 1962. Radiation breeding in two-rowed barley. Seiken Ziho 14:21-<br />
34.<br />
Prepared:<br />
T.E. Haus and T. Tsuchiya. 1971. BGN 1:143.<br />
Revised:<br />
T. Tsuchiya. 1980. BGN 10:115.<br />
J.D. Franckowiak and U. Lundqvist. 1997. BGN 26:184.<br />
J.D. Franckowiak and L. S. Dahleen. 2007. BGN 37:235-236.<br />
236
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 178<br />
Locus name: Intermedium spike-c<br />
Locus symbol: int-c<br />
BGS 178, Intermedium spike-c, int-c<br />
Previous nomenclature and gene symbolization:<br />
Intensifer for Z = W (22).<br />
Infertile intermedium = i (12, 20, 21).<br />
Allelic series I h , I, i (12, 23).<br />
Intermedium spike-c = int-c (6, 7, 17, 18).<br />
Six-rowed spike 5 = v5 (24).<br />
Inheritance:<br />
Monofactorial recessive (4, 5, 21, 24).<br />
Located in chromosome 4HS (3, 5, 18, 21, 24), about 13.1 cM proximal from the<br />
fch9 (chlorina seedling 9) locus, about 14.5 cM distal from the Kap1 (hooded<br />
lemma 1) locus (2, 3, 4, 5, 11), and about 3.5 cM from AFLP marker E4143-5 in<br />
subgroup 8 of the Proctor/Nudinka map (19).<br />
Description:<br />
Alleles at the int-c (v5) locus alter the size of lateral spikelets. The lemma apex of<br />
lateral kernels is rounded or weakly pointed, awnless or short-awned (1, 9, 16).<br />
Lower lateral spikelets may develop poorly in some int-c mutants (4), while seed<br />
development may occur in all lateral spikelets of others (6, 15). Variability in<br />
lateral spikelet development exists among the int-c mutants and environmental<br />
conditions can alter expressivity. The Int-c.a (formerly I) allele in six-rowed barley<br />
increases the size of lateral spikelets, while the int-c.b (formerly i) allele in tworowed<br />
barley prevents anther development in lateral spikelets (9, 22). The int-c.5<br />
mutant in Bonus produces fertile stamens in lateral spikelets (9). In the presence<br />
of the Int-c.h (formerly I h ) allele of Mortoni, lateral spikelets are male fertile and<br />
may occasionally set seed (8, 12). Spikes of vrs5.n (v5) plants appear similar to<br />
those of six-rowed barley, but lateral spikelets are smaller (less than half the size<br />
of the central spikelets) and broader (3, 4).<br />
Origin of mutant:<br />
Natural occurrence in many two-rowed barley cultivars; an X-ray induced mutant<br />
in Gamma 4 (3, 5).<br />
Mutational events:<br />
int-c.b (i) in two-rowed barley (23); Int-c.h (I h ) in Mortoni (CIho 2210, GSHO 72)<br />
(8, 12); vrs5.n (v5) in Gamma 4 (38X-197, OUM338) (3, 5, 14); int-c.5 (NGB<br />
115423) in Bonus (PI 189763) (15, 18); int-c.7 (NGB 115425), -c.62 (NGB<br />
116835), -c.63 (NGB 115481) in Bonus, -c.13 (NGB 115431), -c.15 (NGB<br />
115433), -c.16 (NGB 115434), -c.18 (NGB 115436), -c.25 (NGB 115443), -c.29<br />
(NGB 115447) in Foma (CIho 11333), -c.33 (NGB 115451), -c.38 (NGB 115456),<br />
-c.45 (NGB 115463), -c.48 (NGB 115466), -c.49 (NGB 115467), -c.53 (NGB<br />
115471), -c.56 (NGB 115474), -c.60 (NGB 115478) in Kristina (NGB 1500) (15);<br />
int-c.70 (NGB 115488), -c.76 (NGB 115494), -c.78 (NGB 115496), -c.84 (NGB<br />
115502) in Bonus, -c.95 (NGB 115513) in Hege (NGB 13692) (13).<br />
Mutant used for description and seed stocks:<br />
vrs5.n in Gamma 4 (GSHO 776); int-c.b in Hordeum distichon var. nigrinudum<br />
(GSHO 988); int-c.5 in Bonus (GSHO 1765); int-c.b from Compana (CIho 5438)<br />
in Bonneville (CIho 7248)*6 (CIho 16176) (10); vrs5.n in Bowman (PI 483237)*6<br />
(GSHO 2002); int-c.5 in Bowman*6 (GSHO 2003).<br />
237
Barley Genetics Newsletter (2007) 37: 188-301<br />
References:<br />
1. Åberg, E., and G.A. Wiebe. 1945. Irregular barley, Hordeum irregulare sp. nov.<br />
J. Washington Acad. Sci. 35:161-164.<br />
2. Forster, B.P. 1993. Coordinator's report: Chromosome 4. Barley Genet. Newsl.<br />
22:75-77.<br />
3. Fukuyama, T. 1982. Locating a six-rowed gene v5 on chromosome 4 in barley.<br />
Barley Genet. Newsl. 12:29-31.<br />
4. Fukuyama, T., J. Hayashi, and R. Takahashi. 1975. Genetic and linkage<br />
studies of the five types of induced 'six-row' mutants. Barley Genet. Newsl. 5:12-<br />
13.<br />
5. Fukuyama, T., R. Takahashi, and J. Hayashi. 1982. Genetic studies on the<br />
induced six-rowed mutants in barley. Ber. Ohara Inst. landw. Biol., Okayama<br />
Univ. 18:99-113.<br />
6. Gustafsson, Å., and U. Lundqvist. 1980. Hexastichon and intermedium<br />
mutants in barley. Hereditas 92:229-236.<br />
7. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
8. Gymer, P.T. 1977. Probable allelism of Ii and int-c genes. Barley Genet.<br />
Newsl. 7:34-35.<br />
9. Gymer, P.T. 1978. The genetics of the six-row/two row character. Barley<br />
Genet. Newsl. 8:44-46.<br />
10. Hockett, E.A. 1985. Registration of two- and six-rowed isogenic Bonneville<br />
barley germplasm. Crop Sci. 25:201.<br />
11. Jensen, J. 1987. Linkage map of barley chromosome 4. p. 189-199. In S.<br />
Yasuda and T. Konishi (eds.) Barley Genetics V. Proc. Fifth Int. Barley Genet.<br />
Symp., Okayama. 1986. Sanyo Press Co., Okayama.<br />
12. Leonard, W.H. 1942. Inheritance of fertility in the lateral spikelets of barley.<br />
Genetics 27:299-316.<br />
13. Lundqvist, U. (Unpublished).<br />
14. Lundqvist, U. 1991. Coordinator's report: Ear morphology genes. Barley<br />
Genet. Newsl. 20:85-86.<br />
15. Lundqvist, U., and A. Lundqvist. 1988. Induced intermedium mutants in<br />
barley: origin, morphology and inheritance. Hereditas 108:13-26.<br />
16. Nötzel, H. 1952. Genetische Untersuchungen an röntgeninduzierten<br />
Gerstenmutanten. Kühn-Archiv 66:72-132.<br />
17. Nybom, N. 1954. Mutation types in barley. Acta Agric. Scand. 4:430-456.<br />
18. Persson, G. 1969. An attempt to find suitable genetic markers for dense ear<br />
loci in barley I. Hereditas 62:25-96.<br />
19. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration<br />
of a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
20. Robertson, D.W. 1929. Linkage studies in barley. Genetics 14:1-36.<br />
21. Robertson, D.W. 1933. Inheritance in barley. Genetics 18:148-158.<br />
22. Ubisch, G. von. 1916. Beitrag zu einer Faktorenanalyse von Gerste. Z.<br />
Indukt. Abstammungs. Vererbungsl. 17:120-152.<br />
23. Woodward, R.W. 1947. The I h , I, i allels in Hordeum deficiens genotypes of<br />
barley. J. Am. Soc. Agron. 39:474-482.<br />
24. Woodward, R.W. 1957. Linkages in barley. Agron. J. 49:28-32.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:200-201.<br />
238
Barley Genetics Newsletter (2007) 37: 188-301<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:237-239.<br />
239
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 179<br />
Locus name: Hairy leaf sheath 1<br />
Locus symbol: Hsh1<br />
BGS 179, Hairy leaf sheath 1, Hsh1<br />
Previous nomenclature and gene symbolization:<br />
Hairy leaf sheath = Hs (7).<br />
Inheritance:<br />
Monofactorial dominant (4, 5, 6).<br />
Located in chromosome 4HL (6), over 8.7 cM proximal from the yhd1 (yellow<br />
head 1) locus, and over 22.5 cM distal from the mlo (reaction to Erysiphe<br />
graminis hordei-o) locus (3, 5), in bin 4H-12 about 1.1 cM proximal from RFLP<br />
marker HVM067 (2).<br />
Description:<br />
Short hairs (1 to 3 mm) are scattered or in rows on leaf sheaths of the basal part<br />
of the plant. The density of hairs varies considerably among cultivars and with<br />
changes in growing conditions. With few exceptions, no hairs are observed on<br />
the sheath of upper leaves (4, 5). Heterozygotes and smooth awned cultivars<br />
seem to have fewer hairs.<br />
Origin of mutant:<br />
Natural occurrence in a few cultivars and in some accessions of Hordeum<br />
vulgare subsp spontaneum (1, 5, 6).<br />
Mutational events:<br />
Hsh1.a introduced into cultivated barley from its wild progenitor (5).<br />
Mutant used for description and seed stocks:<br />
Hsh1.a in Kimugi (OUL012, GSHO 986) (5, 6); Hsh1.a from R.I. Wolfe's Multiple<br />
Dominant Marker Stock in Bowman (PI 483237)*10 (GSHO 2026).<br />
References:<br />
1. Cauderon, A. 1951. Étude de I'hérédité de trois couples de caractères<br />
morphologiques chez l'orge cultivée. Ann. Amélior. Plant. 1:9-19.<br />
2. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
3. Hayashi, J., and R. Takahashi. 1986. Location of hs for hairy sheath and yh for<br />
yellow head character on barley chromosome 4. Barley Genet. Newsl. 16:24-27.<br />
4. Patterson, F.L., and R.G. Shands. 1957. Independent inheritance of four<br />
characters in barley. Agron. J. 49:218-219.<br />
5. Takahashi, R., and J. Hayashi. 1966. Inheritance and linkage studies in barley.<br />
II. Assignment of several new mutant genes to their respective linkage groups by<br />
the trisomic method of analysis. Ber. Ohara Inst. landw. Biol., Okayama Univ.<br />
13:185-198.<br />
6. Takahashi, R., J. Hayashi, and S. Yasuda. 1957. [Four genes in linkage in<br />
barley, which are inherited independently of the markers in the known seven<br />
linkage groups in barley.] Nogaku Kenkyu 45:1-10. [In Japanese.]<br />
7. Takahashi, R., S. Yasuda, J. Yamamoto, and I. Shiojiri. 1953. [Physiology and<br />
genetics of ear emergence in barley and wheat. II. Genic analysis of growth-habit<br />
in two spring barleys.] Nogaku Kenkyu 40:157-168. [In Japanese.]<br />
Prepared:<br />
240
Barley Genetics Newsletter (2007) 37: 188-301<br />
R. Takahashi. 1972. BGN 2:184 as BGS 158.<br />
Revised:<br />
J.D. Franckowiak and T. Konishi. 1997. BGN 26:202.<br />
J.D. Franckowiak. 2007. BGN 37:240-241.<br />
241
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 185<br />
Locus name: Brachytic 5<br />
Locus symbol: brh5<br />
BGS 185, Brachytic 5, brh5<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-m = brh.m (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 5).<br />
Located in chromosome 4HS (4), near the int-c (intermedium spike-c) locus (4),<br />
about 13.0 cM proximal from SSR marker Bmac0310 near the boundary between<br />
bins 4H-06 and 4H-07 (1).<br />
Description:<br />
Plants are about 3/4 normal height and awns are about 3/4 of normal length.<br />
Peduncles are less than 2/3 normal length. Seedling leaves of brh5 plants are<br />
relatively short. The kernels of brh5 plants are shorter than those of normal sibs<br />
and weigh about 30% less. Plants lodge easily and the grain yield is about 1/2<br />
normal (1, 2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(6).<br />
Mutational events:<br />
brh5.m in Birgitta (17:18:2, DWS1010) (5, 6).<br />
Mutant used for description and seed stocks:<br />
brh5.m in Birgitta (GSHO 1678); brh5.m in Bowman (PI 483237)*7 (GSHO 2001).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived<br />
lines of spring barley. Barley Genet. Newsl. 24:63-70.<br />
5. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
6. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:100.<br />
Revised:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:242.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 186<br />
Locus name: Slender dwarf 3<br />
Locus symbol: sld3<br />
BGS 186, Slender dwarf 3, sld3<br />
Previous nomenclature and gene symbolization:<br />
Anthocyanin-free = ant-567 (5).<br />
Proanthocyanidin-free 17.567 = ant17.567 (4).<br />
Slender dwarf e = sld.e (3).<br />
Inheritance:<br />
Monofactorial recessive (1).<br />
Located in chromosome 4HS, based on linkage drag with the int-c (intermedium<br />
spike-c) locus (2).<br />
Description:<br />
Plants show reduced vigor and are about 3/4 normal height. The number of<br />
spikelets per spike is about 3/4 that of normal sibs and kernels are slightly<br />
smaller. Rachis internodes can be slightly longer and grain yields are about 3/4<br />
normal (1). The mutant gene sld3.e was isolated as a second mutant in the stock<br />
ant17.567 (proanthocyanidin-free 17) (1). The Bowman backcross-derived line<br />
for sld3.e does not show a reduction in anthocyanin pigmentation or the large<br />
reduction in kernel size (1).<br />
Origin of mutant:<br />
A sodium azide induced mutant isolated with ant-567 in Manker (CIho 15549) (5).<br />
Mutational events:<br />
sld3.e in ant17.567 (DWS1050) (1).<br />
Mutant used for description and seed stocks:<br />
sld3.e in Bowman/ant17.567 (GSHO 2480); sld3.e in Bowman (PI 483237)*7<br />
(GSHO 1998).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived<br />
lines of spring barley. Barley Genet. Newsl. 24:63-70.<br />
3. Franckowiak, J.D. 1999. Coordinator’s report: Semidwarf genes. Barley Genet.<br />
Newsl. 29:74-79.<br />
4. Jende-Strid, B. 1988. Coordinator's report: Anthocyanin genes. Stock list of ant<br />
mutants kept at the Carlsberg Laboratory. Barley Genet. Newsl. 18:74-79.<br />
5. Ullrich, S.E. 1983. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:101.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:243.<br />
243
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 187<br />
Locus name: Brachytic 9<br />
Locus symbol: brh9<br />
BGS 187, Brachytic 9, brh9<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-k = brh.k (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 4HS (1), about 11.7 cM distal from SRR marker<br />
Bmac0310 in bin 4H-06 (1).<br />
Description:<br />
Culms and peduncles are about 3/4 normal length and awns are 3/4 to 5/6 of<br />
normal length. Rachis internodes are slightly shorter than those of normal sibs.<br />
Seedling leaves of brh9 plants are relatively short. The kernels of brh9 plants are<br />
shorter and kernel weight are about 20% lower than those of normal sibs. Grain<br />
yields averaged less than 1/2 normal (1, 2); however, plants appeared nearly<br />
normal when grown in Dundee, Scotland (2). The brh9.k gene was found to nonallelic<br />
at brh5 locus, which is located in the same region of 4HS (1).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(5).<br />
Mutational events:<br />
brh9.k in Birgitta (17:14:4, DWS1006) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh9.k in Birgitta (GSHO 1676); brh9.k in Bowman (PI 483237)*6 (GSHO 2170).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:244.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 203, Black lemma and pericarp 1, Blp1<br />
Stock number: BGS 203<br />
Locus name: Black lemma and pericarp 1<br />
Locus symbol: Blp1<br />
Previous nomenclature and gene symbolization:<br />
Black lemma and caryopsis = B (6).<br />
Black pericarp = Bk (1).<br />
Black lemma and pericarp = B (7).<br />
Inheritance:<br />
Monofactorial dominant (1, 4, 6).<br />
Located in chromosome 1HL [5L] (3, 5), about 16.0 cM proximal from the trd1<br />
(third outer glume 1) locus (3), in bin 1H-13 about 8.8 cM proximal from RFLP<br />
marker ABC261 (2).<br />
Description:<br />
Black pigmentation of the lemma and pericarp develops slightly before<br />
maturation of the spike. Pigmented organs may include all parts of the spike,<br />
awns, the upper portion of the stem, and upper leaves. The intensity of<br />
pigmentation associated with each of the dominant alleles at the Blp1 locus is<br />
characteristic of that allele, and is relatively stable over environments (7). Black<br />
seed is produced by melanin-like pigment in the pericarp (1). Woodward (7)<br />
reports that the dominance ranking of alleles at the Blp1 locus is related to the<br />
intensity of black pigmentation they confer, with the Blp1.b (B) allele conferring<br />
extreme black pigmentation. The Blp1.mb (B mb ) allele is associated with medium<br />
black and a reduced distribution pattern; and the Blp1.g (B g ) allele is associated<br />
with light black or gray coloration (7, 8).<br />
Origin of mutant:<br />
Natural occurrence in several cultivars (6, 7).<br />
Mutational events:<br />
Blp1.b (B) in Hordeum distichon var nigrinudum No 1 (7); Blp1.mb (B mb ) in CIho<br />
2970 (GSHO 226) (7); Blp1.g (B g ) in Blackhull (CIho 878, GSHO 199) and Black<br />
Smyrna (CIho 191, GSHO 222) (7).<br />
Mutant used for description and seed stocks:<br />
Blp1.b in Hordeum distichon var nigrinudum No 1 (GSHO 988); Blp1.b from R.I.<br />
Wolfe's Multiple Dominant Marker Stock (GSHO 1580) in Bowman (PI 483237)*8<br />
(GSHO 2054).<br />
References:<br />
1. Buckley, G.F.H. 1930. Inheritance in barley with special reference to the color<br />
of the caryopsis and lemma. Sci. Agric. 10:460-492.<br />
2. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
3. Ivanova, K.V. 1937. A new character in barley "third outer glume" — Its<br />
inheritance and linkage with color of the flowering glumes. Bull. Appl. Bot.,<br />
Genet., & Pl. Breed. II. 7:339-353.<br />
4. Robertson, D.W. 1929. Linkage studies in barley. Genetics 14:1-36.<br />
5. Robertson, D.W., G.A. Wiebe, R.G. Shands, and A. Hagberg. 1965. A<br />
summary of linkage studies in cultivated barley, Hordeum species: Supplement<br />
245
Barley Genetics Newsletter (2007) 37: 188-301<br />
III, 1954-1963. Crop Sci. 5:33-43.<br />
6. Tschermak, E. von. 1901. Über Züchtung neuer Getreiderassen mittelst<br />
künstlicher Kreuzung. Kritisch-historische Betrachtungen. Zeitschrift für das<br />
landwirtschaftliche Versuchswesen Oesterreich 4:1029-1060.<br />
7. Woodward, R.W. 1941. Inheritance of melanin-like pigment in the glumes and<br />
caryopses of barley. J. Agric. Res. 63:21-28.<br />
8. Woodward, R.W. 1942. Linkage relationships between the allelomorphic<br />
series, B, B mb , B g , and Atat factors in barley. J. Amer. Soc. Agron. 34:659-661.<br />
Prepared:<br />
T.E. Haus and T. Tsuchiya. 1971. BGN 1:148.<br />
Revised:<br />
J.D. Franckowiak and U. Lundqvist. 1997. BGN 26:209.<br />
J.D. Franckowiak. 2007: BGN 37:245-246.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 214<br />
Locus name: Early maturity 8<br />
Locus symbol: eam8<br />
BGS 214, Early maturity 8, eam8<br />
Previous nomenclature and gene symbolization:<br />
Early heading k = eak (26).<br />
Early maturity-a = ea-a (8, 21).<br />
Praematurum-a = mat-a (3, 8, 13, 14, 26).<br />
Erectoides-o = ert-o (8, 18).<br />
Inheritance:<br />
Monofactorial recessive (3, 7).<br />
Located in chromosome 1HL [5L] (21), about 11.4 cM distal from the trd1 (third<br />
outer glume 1) locus and 20.9 cM distal from the Blp1 (black lemma and pericarp<br />
1) locus (21, 24).<br />
Description:<br />
Early heading is associated with decreased culm length, spike length, kernels per<br />
spike, and grain yield (16, 24, 26). When grown in the fall at Kurashiki, Japan,<br />
plants head about 20 days earlier than the standard mid-season cultivar,<br />
Akashinriki, because they are day-length neutral or photoperiod insensitive (26).<br />
Day-length neutrality is observed in early heading mutants isolated from spring<br />
barley in Sweden (2, 9). Under controlled environmental conditions, number of<br />
days to heading does not change as photoperiod is altered (2, 10). All mat-a<br />
induced mutants are characterized by yellowish-green seedlings at an early<br />
stage of development under controlled environmental conditions (1). Other eam8<br />
mutants show a similar response by becoming yellow green under specific<br />
growing conditions, 8 to 12 hours of illumination at low temperatures (below<br />
10°C) plus high temperature (20°C or higher) during the dark period (5, 21, 24).<br />
The color change is caused by photothermal stress, which increases the<br />
zeaxanthin content at the expense of chlorophyll and other pigments (5, 19, 24).<br />
The mutant stock mat-a.8 was released as the cultivar Mari (9, 11). When grown<br />
under 12 h days, the levels of phytochrome B (phyB) decreases in light-grown<br />
BMDR-1 plants, containing an allele at the eam8 locus, compared to normal<br />
plants (12). The instability of phyB content was reported to be responsible for<br />
photoperiod insensitivity of eam8 mutants (12). Under continuous light and with<br />
far-red light treatment for seven days, most differences in heading date between<br />
BMDR-1 and BMDR-8 (Shabet) are eliminated (19).<br />
Origin of mutant:<br />
An X-ray induced mutant in Maja (PI 184884, NGB 8815) (6, 7, 10); natural<br />
occurrence in Kinai 5 (OUJ493) and Kagoshima Gold (OUJ219) (21, 25).<br />
Mutational events:<br />
ert-o.16 (NGB 112618) in Maja (6); eam8.k in Kagoshima Gold, Kinai 5 (CIho<br />
11560), and Kindoku (OUU332) (21, 22, 25); mat-a.8 (NGB 1491, NGB 4694,<br />
NGB 14656, NGB 110008), -a.11 (NGB 110011), -a.12 (NGB 110012) in Bonus<br />
(PI 189763) (7, 14); mat-a.27 (NGB 110027), -a.45 (NGB 110045), -a.46 (NGB<br />
110046), -a.48 (NGB 110048), -a.62 (NGB 110062) in Bonus, -a.110 (NGB<br />
110110), -a.130 (NGB 110130), -a.153 (NGB 110153), -a.221 (NGB 110221), -<br />
a.238 (NGB 110238), -a.255 (NGB 110255), -a.272 (NGB 110272), -a.274 (NGB<br />
110274), -a.287 (NGB 110287), -a.289(NGB 110289), -a.294 (NGB 110294), -<br />
a.325 (NGB 110325), -a.338 (NGB 110338), -a.370 (NGB 110370), -a.384 (NGB<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
110384), -a.390 (NGB 110390),-a.404 (NGB 110404), -a.406 (NGB 110406), -<br />
a.407 (NGB 110407) in Foma (CIho 11333), -a.509 (NGB 110509), -a.641 (NGB<br />
110641), -a.703 (NGB 110703), -a.733 (NGB 110733),in Kristina (NGB 1500), -<br />
a.753 (NGB 110753), -a.796 (NGB 110796), -a.797 (NGB 110797), -a.813 (NGB<br />
110813), -a.832 (NGB 110832), -a.903 (NGB 116858), -a.909 (NGB 117440), -<br />
a.921 (NGB 117452) in Bonus, -a.961 (NGB 117492), -a.970 (NGB 117501), -<br />
a.976 (NGB 117507), -a.984 (NGB 117515), -a.1011 (NGB 117542), in Sv<br />
79353, -a.1032 (NGB 117563), -a.1033 (NGB 117564), -a.1034 (NGB 117565), -<br />
a.1035 (NGB 117566), -a.1036 (NGB 117567), -a.1037 (NGB 117568), -a.1039<br />
(NGB 117570), -a.1040 (NGB 117571), -a.1041 (NGB 117572), -a.1042 (NGB<br />
117573), -a.1043 (NGB 117574), -a.1044 (NGB 117575), -a.1045 (NGB 117576),<br />
-a.1046 (NGB 117577), -a.1047 (NGB 117578), -a.1048 (NGB 117579), -a.1049<br />
(NGB 117580) in Sv Vg74233 (13); mat-a.1050 (NGB 117581), -a.1051 (NGB<br />
117582), -a.1052 (NGB 117583), -a.1053 (NGB 117584), -a.1054 (NGB 117585),<br />
-a.1055 (NGB 117586), -a.1056 (NGB 117587), -a.1057 (NGB 117588), -a.1058<br />
(NGB 117589), -a.1059 (NGB 117590), -a.1060 (NGB 117591), -a.1061 (NGB<br />
117592), -a.1062 (NGB 117593), -a.1063 (NGB 117594), -a.1064 (NGB 117595),<br />
-a.1065 (NGB 117596), -a.1067 (NGB 117598), -a.1069 (NGB 117600), -a.1070<br />
(NGB 117601), -a.1071 (NGB 117602), -a.1072 (NGB 117603), -a.1073 (NGB<br />
117604), -a.1074 (NGB 117605) in Sv Vg74233 (15); eam8.q (Ea8), eam8.r<br />
(Ea9), eam8.s (Ea10), eam8.t (Ea16) in Chikurin Ibaraki 1 (OUJ069, CIho 7370,<br />
GSHO 783) (23); eam8.u (Mut 2571) in Donaria (PI 161974) (5, 17); eam8.v in<br />
Munsing (CIho 6009, GSHO 636) (4, 19, 20); eam8.w in Early Russian (CIho<br />
13839) (4), BMDR-1 (eam8.y) from the original mutant in a dwarf line<br />
backcrossed to Shabet (CIho 13827) (19).<br />
Mutant used for description and seed stocks:<br />
eam8.k in Kinai 5 (OUJ439, GSHO 765); ert-o.16 in Maja (GSHO 489); eam8.k in<br />
Bonus*5 (25); mat-a.8 in Tochigi Golden*5 (25); eam8.u in Munsing/7*Titan<br />
(CIho 16526) (20); eam8.k in Bowman (PI 483237)*7 (GSHO 2063); ert-o.16 in<br />
Bowman*7 (GSHO 2064).<br />
References:<br />
1. Dormling, I., and Å. Gustafsson. 1969. Phytotron cultivation of early barley<br />
mutants. Theor. Appl. Genet. 39:51-61.<br />
2. Dormling, I., Å. Gustafsson, H.R. Jung, and D. von Wettstein. 1966. Phytotron<br />
cultivation of Svalöf's Bonus barley and its mutant Svalöf's Mari. Hereditas<br />
56:221-237.<br />
3. Favret, E.A., and J.H. Frecha. 1967. Allelism test of genes for earliness. Barley<br />
Newsl. 10:121.<br />
4. Gallagher, L.W. (Unpublished).<br />
5. Gallagher, L.W., A.A. Hafez, S.S. Goyal, and D.W. Rains. 1994. Nuclear<br />
mutations affecting chloroplastic pigments of photoperiod-insensitive barley.<br />
Plant Breed. 113:65-70.<br />
6. Gustafsson, Å. 1947. Mutations in agricultural plants. Hereditas 33:1-100.<br />
7. Gustafsson, Å., A. Hagberg, and U. Lundqvist. 1960. The induction of early<br />
mutants in Bonus barley. Hereditas 46:675-699.<br />
8. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
9. Gustafsson, Å., A. Hagberg, G. Persson and K. Wiklund. 1971. Induced<br />
mutations and barley improvement. Theor. Appl. Genet. 41:239-248.<br />
10. Gustafsson, Å., and U. Lundqvist. 1976. Controlled environment and short-<br />
248
Barley Genetics Newsletter (2007) 37: 188-301<br />
day tolerance in barley mutants. p. 45-53. In Induced Mutants in Cross-breeding.<br />
Proc. Advisory Group, Vienna. 1975. Int. Atomic Energy Agency, Vienna.<br />
11. Hagberg, A. 1961. [Svalöfs original Mari barley.] Aktuellt från Svalöf.<br />
Allmänna Svenska Utsädesaktiebolaget. p. 13-16. [In Swedish.]<br />
12. Hanumappa, M., L.H. Pratt, M.-M. Cordonnier-Pratt, and G.F. Deitzer. . 1999.<br />
A photoperiod-insensitive barley line contains a light-labile phytochrome B. Plant<br />
Physiol. 119:1033-1040.<br />
13. Lundqvist, U. 1991. Swedish mutation research in barley with plant breeding<br />
aspects. A historical review. p. 135-148. In Plant Mutation Breeding for Crop<br />
Improvement. Proc. Int. Symp. Vienna, 1990. Int. Atomic Energy Agency, Vienna.<br />
14. Lundqvist, U. 1992. Coordinator's report: Earliness genes. Barley Genet.<br />
Newsl. 21:127-129.<br />
15. Lundqvist, U. (Unpublished).<br />
16. Mellish, D.R., B.L. Harvey, and B.G. Rossnagel. 1978. The effect of a gene<br />
for earliness in 2-row barley. Barley Newsl. 22:76.<br />
17. Mettin, D. 1961. Mutationsversuche an Kulturpflanzen. XII. Über das<br />
genetische Verhalten von frühreifen Gerstenmutanten. Züchter 31:83-89.<br />
18. Persson, G., and A. Hagberg. 1969. Induced variation in a quantitative<br />
character in barley. Morphology and cytogenetics of erectoides mutants.<br />
Hereditas 61:115-178.<br />
19. Principe, J.M., W.R. Hruschka, B. Thomas, and G.F. Deitzer. 1992. Protein<br />
differences between two isogenic cultivars of barley (Hordeum vulgare L.) that<br />
differ in sensitivity to photoperiod and far-red light. Plant Physiol. 98:1444-1450.<br />
20. Smail, V.W., R.F. Eslick, and E.A. Hockett. 1986. Isogenic heading date<br />
effects on yield component development in 'Titan' barley. Crop Sci. 26:1023-<br />
1029.<br />
21. Takahashi, R., and S. Yasuda. 1971. Genetics of earliness and growth habit<br />
in barley. p. 388-408. In R.A. Nilan (ed.) Barley Genetics II. Proc. Second Int.<br />
Barley Genet. Symp., Pullman, WA, 1969. Washington State Univ. Press,<br />
Pullman.<br />
22. Takahashi, R., S. Yasuda, J. Hayashi, T. Fukuyama, I. Moriya, and T.<br />
Konishi. 1983. Catalogue of barley germplasm preserved in Okayama University.<br />
Inst. Agric. Biol. Sci., Okayama Univ., Kurashiki, Japan. 217 p.<br />
23. Ukai, Y., and A. Yamashita. 1981. Early mutants of barley induced by ionizing<br />
radiation and chemicals. p. 846-854. In M.J.C. Asher, R.P. Ellis, A.M. Hayter, and<br />
R.N.H. Whitehouse (eds.) Barley Genetics IV. Proc. Fourth Int. Barley Genet.<br />
Symp., Edinburgh. Edinburgh Univ. Press, Edinburgh.<br />
24. Yasuda, S. 1977. Linkage of the earliness gene eak and its pleiotropic effects<br />
under different growth conditions. Ber. Ohara Inst. landw. Biol., Okayama Univ.<br />
17:15-28.<br />
25. Yasuda, S. 1978. Effects of the very early gene, eak, on yield and its<br />
components in barley. BGN 8:125-127.<br />
26. Yasuda, S., T. Konishi, and H. Shimoyama. 1965. [Varietal difference in<br />
yellowing of barleys under a certain controlled condition of temperature and<br />
photoperiod, and its mode of inheritance.] Nogaku Kenkyu 51:53-65. [In<br />
Japanese.]<br />
Prepared:<br />
S. Yasuda. 1972. BGN 2:198.<br />
Revised:<br />
J.D. Franckowiak, U. Lundqvist, T. Konishi, and L.W. Gallagher. 1997. BGN<br />
26:213-215.<br />
249
Barley Genetics Newsletter (2007) 37: 188-301<br />
J.D. Franckowiak and U. Lundqvist. 2007. BGN 37:247-250.<br />
250
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 222<br />
Locus name: Necrotic leaf spot 1<br />
Locus symbol: nec1<br />
BGS 222, Necrotic leaf spot 1, nec1<br />
Previous nomenclature and gene symbolization:<br />
Mutant no. 10 (2).<br />
Parkland spot = sp,,b (1).<br />
Inheritance:<br />
Monofactorial recessive (2, 4).<br />
Located in chromosome 1HL [5L] (1, 2, 4), near the centromere (1), about 34.5<br />
cM proximal from the wst5 (white streak 5) locus (3, 5), about 10.0 cM distal from<br />
the msg1 (male sterile genetic 1) locus (4, 6), in bin 5H-09 near EST marker<br />
BF630384 (7).<br />
Description:<br />
Small black-brown spots develop on all light-exposed parts of the plant starting<br />
near the leaf tip at the three-leaf stage (1, 2). The spots are oval (the longest<br />
dimension is parallel to the leaf veins) and generally less than 1 to 2 mm in size.<br />
The spots are concentrated in awn and the most distal parts of the leaf blade, but<br />
may occur on all plant parts (2, 4). The nec1 locus is an orthologue of<br />
Arabidopsis necrotic mutant HLM1 that encodes the cyclic nucleotide-gated ion<br />
channel 4 (7).<br />
Origin of mutant:<br />
A mutant induced by combined treatment with gamma-rays and diethyl sulfate of<br />
Carlsberg II (CIho 10114, NGB 5085) (2).<br />
Mutational events:<br />
nec1.a in Carlsberg II (Mutant no 10) (2, 3); sp,,b (GSHO 1284) in Parkland<br />
(CIho 10001) (1, 4); a mutant in Morex (CIho 15773) (6); FN085 and FN370 in<br />
Steptoe (CIho 15229) (7); FN338 in Morex (CIho 15773) (7).<br />
Mutant used for description and seed stocks:<br />
nec1.a in Carlsberg II (GSHO 989); nec1.a from R.I. Wolfe's Chromosome 5<br />
Marker Stock in Bowman (PI 483237)*7 (GSHO 2052).<br />
References:<br />
1. Fedak, G., T. Tsuchiya, and S.B. Helgason. 1972. Use of monotelotrisomics<br />
for linkage mapping in barley. Can. J. Genet. Cytol. 14:949-957.<br />
2. Jensen, J. 1971. Mapping of 10 mutant genes for necrotic spotting in barley by<br />
means of translocation. p. 213-219. In R.A. Nilan (ed.) Barley Genetics II. Proc.<br />
Second Int. Barley Genet. Symp., Pullman, WA, 1969. Washington State Univ.<br />
Press, Pullman.<br />
3. Jensen, J. 1992. Coordinator's report: Chromosome 5. Barley Genet. Newsl.<br />
21:89-92.<br />
4. Jensen, J., and J.H. Jørgensen. 1973. Locating some genes on barley<br />
chromosome 5. Barley Genet. Newsl. 3:25-27.<br />
5. Nielsen, G., H. Johansen, and J. Jensen. 1983. Localization on barley<br />
chromosome 5 of the locus Pgd2 coding for phosphogluconate dehydrogenase.<br />
Barley Genet. Newsl. 13:57-59.<br />
6. Ramage, T., and J.L.A. Eckhoff. 1985. Assignment of mutants in Morex to<br />
chromosomes. Barley Genet. Newsl. 15:22-25.<br />
7. Rostoks, N., D. Schmierer, S. Mudie, T. Drader, R. Brueggeman, D. Caldwell,<br />
R. Waugh, and A. Kleinhofs. 2006. Barley necrotic locus nec1 encodes the cyclic<br />
251
Barley Genetics Newsletter (2007) 37: 188-301<br />
nucleotide-gated ion channel 4 homologous to the Arabidopsis Hlm1. Mol. Gen.<br />
Genomics 275:159-168.<br />
Prepared:<br />
J. Jensen. 1981. BGN 11:101.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:220.<br />
J.D. Franckowiak. 2007. BGN 37:251-252.<br />
252
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 253<br />
Locus name: Uniculm 2<br />
Locus symbol: cul2<br />
BGS 253, Uniculm 2, cul2<br />
Previous nomenclature and gene symbolization:<br />
Uniculm 2 = uc2 (8).<br />
Inheritance:<br />
Monofactorial recessive (8).<br />
Located in chromosome 6HL (4, 6), about 1.3 cM distal from the gsh4 (glossy<br />
sheath 4) locus (3, 5), about 11.4 cM from the msg36 (male sterile genetic 36)<br />
locus (3, 5), about 2.2 cM proximal from the rob1 (orange lemma 1) locus (3, 4,<br />
5), about 8.8 cM from RFLP markers cMWG679 and ABG458 (1), and about 6.2<br />
cM from AFLP marker E4343-10 in subgroup 54 of the Proctor/Nudinka map (7).<br />
Description:<br />
The cul2 plants have a single elongated culm (stem), the stem is much greater in<br />
diameter than normal, and plants are usually earlier than normal (8). The cul2<br />
plants initiate vegetative axillary meristems, but tillers fail to develop (1). Irregular<br />
placement of some spikelets and male fertility in lateral spikelets occur in the<br />
original stock (5) and in the Bowman backcross-derived line (1). Yield of uniculm<br />
plants is not restored when grown under high plant populations (2). Double<br />
mutant combinations with most other mutants that affect tiller number resulted in<br />
a uniculm vegetative phenotype (1).<br />
Origin of mutant:<br />
A thermal neutron induced mutant in Kindred (CIho 6969) (8).<br />
Mutational events:<br />
cul2.b in Kindred (GBC379) (5), cul2.k (unc k ) in an unknown cultivar from the<br />
Max-Planck-Institut für Züchtungsforschung (7).<br />
Mutant used for description and seed stocks:<br />
cul2.b in Kindred (GSHO 531, CIho 115530); cul2.b in Bowman (PI 483237)*4<br />
(GSHO 2074); cul2.b plus rob1.a from sel 79Cal in Bowman*8 (GSHO 2075).<br />
References:<br />
1. Babb, S., and G.J. Muehlbauer. 2003. Genetic and morphological<br />
characterization of the barley uniculm2 (cul2) mutant. Theor. Appl. Genet.<br />
106:846–857.<br />
2. Dofing, S.M. 1996. Near-isogenic analysis of uniculm and conventional-tillering<br />
barley lines. p. 617-619. In A.E. Slinkard, G.J. Scoles, and B.G. Rossnagel (eds.)<br />
Proc. Fifth Int. Oat Conf. & Seventh Int. Barley Genet. Symp., Saskatoon. Univ.<br />
of Saskatchewan, Saskatoon.<br />
3. Falk, D.E., M.J. Swartz, and K.J. Kasha. 1980. Linkage data with genes near<br />
the centromere of barley chromosome 6. Barley Genet. Newsl. 10:13-16.<br />
4. Hayashi, J., T. Konishi, I. Moriya, and R. Takahashi. 1984. Inheritance and<br />
linkage studies in barley. VI. Ten mutant genes located on chromosomes 1 to 7,<br />
except 3. Ber. Ohara Inst. landw. Biol., Okayama Univ. 18:227-250.<br />
5. Kasha, K.G., D.E. Falk, and A. Ho-Tsai. 1978. Linkage data with genes on<br />
chromosome 6. Barley Genet. Newsl. 8:61-65.<br />
6. Kirby, E.J. 1973. Effect of temperature on ear abnormalities in uniculm barley.<br />
J. Exp. Bot. 24:935-947.<br />
7. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
253
Barley Genetics Newsletter (2007) 37: 188-301<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
8. Shands, R.G. 1963. Inheritance and linkage of orange lemma and uniculm<br />
characters. Barley Newsl. 6:35-36.<br />
Prepared:<br />
C.R. Burnham. 1971. BGN 1:156.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:234.<br />
J.D. Franckowiak. 2007. BGN 37:253-254.<br />
254
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 254<br />
Locus name: Orange lemma 1<br />
Locus symbol: rob1<br />
BGS 254, Orange lemma 1, rob1<br />
Previous nomenclature and gene symbolization:<br />
Orange lemma = pl (14).<br />
Orange lemma = br (1, 2).<br />
Orange lemma = o (15).<br />
Robiginosum-o = rob-o (6).<br />
Inheritance:<br />
Monofactorial recessive (1, 2, 14, 15).<br />
Located in chromosome 6HS (4, 5, 17, 18), about 10.8 cM proximal from the<br />
msg36 (male sterile genetic 36) locus (5, 9), and about 2.2 cM distal from the<br />
cul2 (uniculm 2) locus (5, 7, 9), in bin 6H-06 near RFLP marker HVM031 (3).<br />
Description:<br />
The lemma, palea, and rachis have an orange pigmentation that is present in<br />
immature spikes, can be observed at heading, and is retained in mature grain<br />
and spikes (2, 15). The orange pigmentation is visible at the base of sheath of<br />
seedlings and in exposed nodes after jointing. Internodes have a layer of orange<br />
tissue and stems have an orange color as the straw dries. The mutant stock for<br />
rob1.f (OUM189) has a lighter orange lemma color than that in other mutants at<br />
the rob1 locus (10). The Bowman backcross-derived line with the rob1 gene had<br />
slightly lower acid-detergent lignin (ADL) content than Bowman (13), but it was<br />
also more susceptible to common root rot, caused by Bipolaris sorokiniana (11).<br />
Origin of mutant:<br />
A spontaneous mutant in CIho 5649 (15).<br />
Mutational events:<br />
rob1.a in CIho 5649 (GBC340, GSHO 707) (8, 15); rob1.b (OUM185), rob1.c<br />
(OUM186), rob1.d (OUM187), rob1.e (OUM188), rob1.f (OUM189) in Akashinriki<br />
(OUJ659, PI 467400) (10); rob1.1 (NGB 115071, NGB 119367), rob1.2 (NGB<br />
115072, NGB 119368) in Bonus (PI 189763), rob1.3 (NGB 115073, NGB<br />
119369), rob1.4 (NGB 115074, NGB 119370), rob1.5 (NGB 115075, NGB<br />
119371), rob1.6 (NGB 115076, 119372) in Foma (CIho 11333), rob1.7 (NGB<br />
115077, NGB 119373) in Kristina (NGB 1500) (12); rob1.g (200A12/8/2) from<br />
Emir (CIho 11790) isolated following a cross to Hordeum bulbosum (16).<br />
Mutant used for description and seed stocks:<br />
rob1.a in CIho 5649 (GSHO 707); rob1.a in Bowman (PI 483237)*8 (GSHO<br />
2069).<br />
References:<br />
1. Bauman, A. 1926. Barley with orange lemmas. Bull. Appl. Bot. & Pl. Breed.<br />
16:181-186.<br />
2. Buckley, G.F.H. 1930. Inheritance in barley with special reference to the color<br />
of caryopsis and lemma. Sci. Agric. 10:460-492.<br />
3. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch,<br />
S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda,<br />
M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe<br />
Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl.<br />
Genet. 103:415-424.<br />
4. Falk, D.E. 1994. Creation of a marked telo 6S trisomic for chromosome 6.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Barley Genet. Newsl. 23:32.<br />
5. Falk, D.E., M.J. Swartz, and K.J. Kasha. 1980. Linkage data with genes near<br />
the centromere of barley chromosome 6. Barley Genet. Newsl. 10:13-16.<br />
6. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
7. Hayashi, J., T. Konishi, I. Moriya, and R. Takahashi. 1984. Inheritance and<br />
linkage studies in barley. VI. Ten mutant genes located on chromosomes 1 to 7,<br />
except 3. Ber. Ohara Inst. landw. Biol., Okayama Univ. 18:227-250.<br />
8. Ivanova, K.V. 1937. A new character in barley "third outer glume" — Its<br />
inheritance and linkage with color of the flowering glumes. Bull. Appl. Bot.,<br />
Genet. & Pl. Breed. II. 7:339-353.<br />
9. Kasha, K.G., D.E. Falk, and A. Ho-Tsai. 1978. Linkage data with genes on<br />
chromosome 6. Barley Genet. Newsl. 8:61-65.<br />
10. Konishi, T. (Unpublished).<br />
11. Kutcher, H.R., K.L. Bailey, B.G. Rossnagel, and J.D. Franckowiak. 1996.<br />
Linked morphological and molecular markers associated with common root rot<br />
reaction in barley. Can. J. Plant Sci. 76:879-883.<br />
12. Lundqvist. U. (Unpublished).<br />
13. Meyer, D.W., J.D. Franckowiak, and R.D. Nudell. 2006. Forage quality of<br />
barley hay. Agronomy Abstracts 2006.<br />
14. Miyake, K., and Y. Imai. 1922. [Genetic studies in barley. 1.] Bot. Mag.,<br />
Tokyo 36:25-38. [In Japanese.]<br />
15. Myler, J.L., and E.H. Stanford. 1942. Color inheritance in barley. J. Am. Soc.<br />
Agron. 34:427-436.<br />
16. Pickering, R.A. 2003. (Personal communications).<br />
17. Ramage, R.T., C.R. Burnham, and A. Hagberg. 1961. A summary of<br />
translocation studies in barley. Crop Sci. 1:277-279.<br />
18. Shahla, A., J.W. Shim, and T. Tsuchiya. 1983. Association of the gene o for<br />
orange lemma with the short arm of chromosome 6 (6S) in barley. BGN 13:83-<br />
84.<br />
Prepared:<br />
C.R. Burnham. 1971. BGN 1:157.<br />
Revised:<br />
J.D. Franckowiak, T. Konishi, and U. Lundqvist. 1997. BGN 26:235-236.<br />
J.D. Franckowiak and U. Lundqvist. 2007. BGN 37:255-256.<br />
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Stock number: BGS 266<br />
Locus name: Erectoides-e<br />
Locus symbol: ert-e<br />
BGS 266, Erectoides-e, ert-e<br />
Previous nomenclature and gene symbolization:<br />
Erectoides-17 = ert-17 (2).<br />
Dense spike = la (4).<br />
Dense spike 9 = l9 (4).<br />
Inheritance:<br />
Monofactorial recessive (2, 4, 8).<br />
Located in chromosome 6HL (3, 7, 8), about 27.2 cM distal from the xnt5 (xantha<br />
seedling 5) locus (5), over 26.9 cM distal from the Aat2 (aspartate<br />
aminotransferase 2) locus (11).<br />
Description:<br />
Spikes are very compact with rachis internode length values from 1.2 to 1.5 mm.<br />
Plants are about 2/3 normal height. Partial fertility and reduced vigor are noted<br />
among ert-e mutants. The peduncle is very short and spikes often emerge from<br />
the side of the flag sheath (7, 9). A large deficiency of mutant plants is frequently<br />
noted in segregating populations (7). Spike density decreases greatly when<br />
plants are treated with GA3 as the flag leaf emerges (10). The mutant ert-e.17 is<br />
allelic to mutant dsp9.i (dense spike 9, see BGS 258) (1).<br />
Origin of mutant:<br />
An X-ray induced mutant in Bonus (PI 189763) (2).<br />
Mutational events:<br />
ert-e.17 (NGB 112619 ), -e.65 (NGB 112664) in Bonus (2); ert-e.94 (NGB<br />
112693), -e.143 (NGB 112742) in Bonus, -e.331 (NGB 112846), -e.396 (NGB<br />
114150) in Foma (CIho 11333) (9); dsp9.i (OUM113) in Akashinriki (4); dsp9.j<br />
(OUM106), dsp9.k (OUM107), dsp9.l (OUM115), dsp9.m (OUM118) in<br />
Akashinriki (6).<br />
Mutant used for description and seed stocks:<br />
ert-e.17 in Bonus (GSHO 477); ert-e.17 in Bowman (PI 483237)*6 (GSHO 2091);<br />
dsp9.i in Akashinriki (GSHO 1774); dsp9.i in Bowman (PI 483237)*7 (GSHO<br />
2090).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Hagberg, A., Å. Gustafsson, and L. Ehrenberg. 1958. Sparsely contra densely<br />
ionizing radiations and the origin of erectoid mutants in barley. Hereditas 44:523-<br />
530.<br />
3. Hagberg, A., G. Persson, and A. Wiberg. 1963. Induced mutations in the<br />
improvement of self-pollinated crops. p. 105-124. In E. Åkerberg and A. Hagberg<br />
(eds.) Recent Plant Breeding Research. Svalöf 1946-1961. Almqvist & Wiksell,<br />
Stockholm.<br />
4. Konishi, T. 1973. Genetic analyses of EMS-induced mutants in barley. Barley<br />
Genet. Newsl. 3:28-31.<br />
5. Konishi, T. 1978. New linkage data on chromosome 6 of barley. Barley Genet.<br />
Newsl. 8:71-72.<br />
6. Konishi, T. (Unpublished).<br />
7. Persson, G. 1969. An attempt to find suitable genetic markers for the dense<br />
ear loci in barley I. Hereditas 62:25-96.<br />
257
Barley Genetics Newsletter (2007) 37: 188-301<br />
8. Persson, G., and A. Hagberg. 1962. Linkage studies with the erectoides loci.<br />
Barley Newsl. 5:46-47.<br />
9. Persson, G., and A. Hagberg. 1969. Induced variation in a quantitative<br />
character in barley. Morphology and cytogenetics of erectoides mutants.<br />
Hereditas 61:115-178.<br />
10. Stoy, V., and A. Hagberg. 1967. Effects of growth regulators on ear density<br />
mutants in barley. Hereditas 58:359-384.<br />
11. Yoshimi, R., and T. Konishi. 1995. Linkage analysis of several isozyme loci in<br />
barley. Barley Genet. Newsl. 24:35-37.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:246.<br />
Revised as BGS 258:<br />
T. Konishi and J.D. Franckowiak. 1997. BGS 258, Dense spike 9, dsp9. BGN<br />
26:239.<br />
Revised:<br />
J.D. Franckowiak and U. Lundqvist. 2007. BGN 37:257-258.<br />
258
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 306<br />
Locus name: Variegated 1<br />
Locus symbol: var1<br />
BGS 306, Variegated 1, var1<br />
Previous nomenclature and gene symbolization:<br />
Variegated = va (4).<br />
Inheritance:<br />
Monofactorial recessive (4).<br />
Located in chromosome 5HL [7L] (4), about 4.6 cM proximal from the raw1<br />
(smooth awn 1) locus (3. 4), in bin 5H-09 about 29.2 cM proximal from the Rph9<br />
(reaction to Puccinia hordei 9) locus (1).<br />
Description:<br />
Narrow white stripes develop on young leaves, but they are not as well defined<br />
than those of white streak 7 (wst7). White stripes are visible on the foliage and<br />
stems of older plants (3). When sown in plots, selections homozygous for the<br />
var1.a gene have a whitish cast until heading (2). The Bowman backcrossderived<br />
line for var1 shows slightly delayed heading and may be slightly shorter<br />
(2).<br />
Origin of mutant:<br />
A gamma-ray induced mutant in Montcalm (CIho 7149) (4).<br />
Mutational events:<br />
var1.a in Montcalm (Alb Acc 311) (4).<br />
Mutant used for description and seed stocks:<br />
var1.a in Montcalm (GSHO 1278); var1.a in Bowman (PI 483237)*7 (GSHO<br />
2121).<br />
References:<br />
1. Borovkova, I.G., Y. Jin, and B.J. Steffenson. 1998. Chromosomal location and<br />
genetic relationship of leaf rust resistance genes Rph9 and Rph12 in barley.<br />
Phytopathology 88:76-80.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Jensen, J. 1981. Construction of a barley chromosome 7 linkage map. p. 927-<br />
939. In M.J.C. Asher, R.P. Ellis, A.M. Hayter, and R.N.H. Whitehouse (eds.)<br />
Barley Genetics IV. Proc. Fourth Int. Barley Genet. Symp., Edinburgh. Edinburgh<br />
Univ. Press, Edinburgh.<br />
4. Walker, G.W.R., J. Dietrich, R. Miller, and K. Kasha. 1963. Recent barley<br />
mutants and their linkages II. Genetic data for further mutants. Can. J. Genet.<br />
Cytol. 5:200-219.<br />
Prepared:<br />
T.E. Haus and T. Tsuchiya. 1971. BGN 1:165.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:257.<br />
J.D. Franckowiak. 2007. BGN 37:259.<br />
259
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 348<br />
Locus name: Early maturity 5<br />
Locus symbol: Eam5<br />
BGS 348, Early maturity 5, Eam5<br />
Previous nomenclature and gene symbolization:<br />
Early maturity = Ea (5, 8).<br />
Early maturity 3 = Ea3 (2, 3).<br />
Early maturity 5 = Ea5 (4).<br />
Early maturity 8 = Ea8 (6).<br />
Inheritance:<br />
Monofactorial dominant (9).<br />
Located in chromosome 5HL [7L] (2), very close to the raw1 (smooth awn 1)<br />
locus (1, 8, 9).<br />
Description:<br />
Plants with the Eam5 gene head 3 to 10 days earlier than normal sibs under<br />
short-day conditions (1, 5). Early heading is commonly associated a shorter<br />
stature compared to normal sibs. The slight reduction in height is also observed<br />
under long-day conditions. Peduncles and rachis internodes are slightly<br />
shortened (1). The Eam5 gene appears to be the common early maturity gene<br />
present in winter sown spring barley cultivars used in China and Japan; and it is<br />
present in the ICARDA/CIMMYT barley lines developed in Mexico. Complex<br />
interactions with other genes conditioning photoperiod response have been<br />
observed (1, 9). Takahashi and Yasuda (7) classified plants that were about 10<br />
days earlier than normal spring barley as having the Sgh2.II (spring growth of<br />
habit 2, grade 2) gene, but an earliness factor closely linked to the rough awn<br />
gene was earlier identified in spring barley (8).<br />
Origin of mutant:<br />
Natural occurrence in Indian cultivars (2, 3) and isolated from ICARDA/CIMMYT<br />
selection CMB85-533-H-1Y-1B-0Y-5B (Higuerilla*2/Gobernadora) (1).<br />
Mutational events:<br />
Eam5.x in CMB85-533 (1), Eam5.x in a number of Chinese cultivars planted in<br />
the fall (9).<br />
Mutant used for description and seed stocks:<br />
Eam5.x in CMB85-533; Eam5.x in Bowman (PI 483237)*6 (GSHO 3424).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Jain, K.B.L. 1961. Genetic studies in barley. III. Linkage relations of some<br />
plant characters. Indian J. Genet. Plant Breed. 21:23-33.<br />
3. Murty, G.S., and K.B.L. Jain. 1960. Genetic studies in barley. II. Inheritance of<br />
fertility of lateral florets and certain other characters. J. Indian Botan. Soc.<br />
39:281-308.<br />
4. Nilan, R.A. 1964. The cytology and genetics of barley, 1951-1962. Monogr.<br />
Suppl. 3, Res. Stud. Vol. 32, No. 1. Washington State Univ. Press, Pullman.<br />
5. Robertson, D.W., G.A. Wiebe, and F.R. Immer. 1941. A summary of linkage<br />
studies in barley. J. Am. Soc. Agron. 33:47-64.<br />
6. Robertson, D.W., G.A. Wiebe, R.G. Shands, and A. Hagberg. 1965. A<br />
summary of linkage studies in cultivated barley, Hordeum species: Supplement<br />
III, 1954-1963. Crop Sci. 5:33-43.<br />
260
Barley Genetics Newsletter (2007) 37: 188-301<br />
7. Takahashi, R., and S. Yasuda. 1971. Genetics of earliness and growth habit in<br />
barley. p. 388-408. In R.A. Nilan (ed.) Barley Genetics II. Proc. Second Int.<br />
Barley Genet. Symp., Pullman, WA, 1969. Washington State Univ. Press,<br />
Pullman.<br />
8. Wexelsen, H. 1934. Quantitative inheritance and linkage in barley. Hereditas<br />
18:307-348.<br />
9. Yu, G. 2006. Development of early maturing two-rowed malting barley with<br />
Fusarium head blight resistance. Ph.D. Thesis. North Dakota State University,<br />
Fargo.<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:109.<br />
Revised:<br />
J.D. Franckowiak and G. Yu. 2007. BGN 37:260-261.<br />
261
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 349<br />
Locus name: Brachytic 4<br />
Locus symbol: brh4<br />
BGS 349, Brachytic 4, brh4<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-j = brh.j 4, 5).<br />
Inheritance:<br />
Monofactorial recessive (4, 5).<br />
Located in chromosome 2HL (3), about 14.1 cM distal from SSR marker<br />
EBmac0850 in bin 2H-08 (1).<br />
Description:<br />
Seedling leaves of brh4 plants are short and wide compared to those of<br />
Bowman. Plants are 3/4 to 5/6 normal height and awns are about 3/4 of normal<br />
length. Plants have a rather erect growth habit. Peduncle length is about 3/4<br />
normal and rachis internodes are slightly shortened. Heading is delayed by about<br />
2 days and the fertile spikelet number is increased by over 3, but these effects<br />
could be caused by pleiotropism or linkage drag with the Eam6 (early maturity 6)<br />
locus. The kernels of brh4 plants are slightly shorter (8.3 vs. 9.4 mm), more<br />
globose shaped, and slightly smaller (46 vs. 56 mg) than those of Bowman. The<br />
yield reduction was non-significant in comparisons between the brh4.j Bowman<br />
line and Bowman (1, 2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(6).<br />
Mutational events:<br />
brh4.j in Birgitta (17:13:6, DWS1005) (5, 6).<br />
Mutant used for description and seed stocks:<br />
brh4.j in Birgitta (GSHO 1675); brh4.j in Bowman (PI 483237)*7 (GSHO 2130).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived<br />
lines of spring barley. Barley Genet. Newsl. 24:63-70.<br />
4. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
5. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
6. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak. 2002. BGN 32:110.<br />
Revised:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:262.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 350<br />
Locus name: Brachytic 6<br />
Locus symbol: brh6<br />
BGS 350, Brachytic 6, brh6<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-r = brh.r (3).<br />
Brachytic-s = brh.s (3).<br />
Inheritance:<br />
Monofactorial recessive (4, 5).<br />
Located in chromosome 5HS [7S] (4), about 12.0 cM distal from SSR marker<br />
Bmag0394 in bin 5H-03 (1).<br />
Description:<br />
Plants of the Bowman backcross-derived line are 2/3 to 3/4 normal height and<br />
awns are 1/2 to 2/3 normal length. The seedling leaf of brh6 plants is shorter than<br />
that of normal sibs (1, 2). Peduncles and leaf blades are 2/3 normal length and<br />
the grain is nearly normal (1). However, kernels are nearly 20% lighter than those<br />
of Bowman with both decreased length and width (1, 2). Although grain yield of<br />
the near-isogenic line for brh6 was lower than those of tall Akashinriki, the brh6<br />
line was considered a high yielding dwarf (7).<br />
Origin of mutant:<br />
An ethyl methanesulfonate induced mutant in Akashinriki (OUJ659, PI 467400)<br />
(6, 7).<br />
Mutational events:<br />
brh6.r in Akashinriki (OUM133, dw-h, DWS1036, GSHO 1683), brh6.s in<br />
Akashinriki (OUM135, DWS1037, GSHO 1684) (3, 5, 6).<br />
Mutant used for description and seed stocks:<br />
brh6.r in Akashinriki (GSHO 1683); brh6.r in Bowman (PI 483237)*7 (GSHO<br />
2132).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D. 1995. Notes on linkage drag in Bowman backcross derived<br />
lines of spring barley. Barley Genet. Newsl. 24:63-70.<br />
5. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
6. Konishi, T. 1976. The nature and characteristics of EMS-induced dwarf<br />
mutants in barley. p. 181-189. In H. Gaul (ed.). Barley Genetics III. Proc. Third<br />
Int. Barley Genet. Symp., Garching, 1975. Verlag Karl Thiemig, München.<br />
7. Konishi, T. 1977. Effects of induced dwarf genes on agronomic characters in<br />
barley. p. 21-38. In Use of dwarf mutations. Gamma-Field Symposium No. 16.<br />
Prepared:<br />
J.D. Franckowiak and T. Konishi. 2002. BGN 32:111.<br />
Revised:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:263.<br />
263
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 377, Shrunken endosperm genetic 1, seg1<br />
Stock number: BGS 377<br />
Locus name: Shrunken endosperm genetic 1<br />
Locus symbol: seg1<br />
Previous nomenclature and gene symbolization:<br />
Shrunken endosperm = se1 (4).<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Located in chromosome 7HL [1L] (3), linked to the msg23 (male sterile genetic<br />
23) locus (5).<br />
Description:<br />
Seed is long and thin and the 100-kernel weight is about 33% of normal. Good<br />
stands can be established in the field if optimum environmental conditions prevail<br />
during germination and emergence (3, 5). This mutant is associated with an<br />
increase in percentage lysine in the protein (5). Tannins are not deposited in<br />
seg1 chalazal cell central vacuoles, but rather appeared to cause cytoplasmic<br />
disorganization and cell death (1). Light microscopy revealed that seg1 mutants<br />
exhibited premature termination of grain filling because of the necrosis and<br />
crushing of the chalazal and nucellar projection of the pericarp early during grain<br />
filling (2).<br />
Origin of mutant:<br />
A spontaneous mutant in Betzes (PI 129430) (3).<br />
Mutational events:<br />
seg1.a in Betzes (3, 4).<br />
Mutant used for description and seed stocks:<br />
seg1.a in Betzes (GSHO 750); seg1.a in Bowman (PI 483237)*7 (GSHO 1852).<br />
References:<br />
1. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1984. Development of tannin<br />
vacuoles in chalazal and seed coat of barley in relation to early chalazal necrosis<br />
in the seg1 mutant. Planta 161:540-549.<br />
2. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1985. Anatomy of immature<br />
grains of eight material effect shrunken endosperm barley mutants. Amer. J. Bot.<br />
72:248-256.<br />
3. Jarvi, A.J. 1970. Shrunken endosperm mutants in barley, Hordeum vulgare.<br />
Ph.D. Thesis. Montana State Univ., Bozeman.<br />
4. Jarvi, A.J., and R.F. Eslick. 1971. BGS 377, Normal vs. shrunken endosperm,<br />
se1. Barley Genet. Newsl. 1:190.<br />
5. Jarvi, A.J., and R.F. Eslick. 1975. Shrunken endosperm mutants in barley.<br />
Crop Sci. 15:363-366.<br />
Prepared:<br />
A.J. Jarvi and R.F. Eslick. 1971. BGN 1:190.<br />
Revised:<br />
R.F. Eslick. 1976. BGN 6:135.<br />
T. Tsuchiya. 1980. BGN 10:124.<br />
J.D. Franckowiak. 1997. BGN 26:325.<br />
J.D. Franckowiak. 2007. BGN 37:264.<br />
264
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 379, Shrunken endosperm genetic 3, seg3<br />
Stock number: BGS 379<br />
Locus name: Shrunken endosperm genetic 3<br />
Locus symbol: seg3<br />
Previous nomenclature and gene symbolization:<br />
Shrunken endosperm = se3 (5).<br />
Proanthocyanidin-free 17 = ant17 (3).<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Located in chromosome 3HS (1, 6), over 30.8 cM from the centromere (6).<br />
Description:<br />
Seed size is reduced to about 33% of normal when grown under field conditions.<br />
Seeds are long and thin similar to those from seg1 plants; seeds are viable and<br />
good stand establishment is possible (6). Light microscopy revealed that seg3<br />
mutants exhibited premature termination of grain filling because of the necrosis<br />
and crushing of the chalazal and nucellar projection of the pericarp early during<br />
grain filling (2). The mutant ant17.148 is an allele at the seg3 locus (3); thus, all<br />
mutants at the proanthocyanidin-free 17 (ant17) locus might be alleles at the<br />
shrunken endosperm genetic 3 locus. Alleles at the seg3 locus that have been<br />
examined in the Bowman genetic background showed a variable reduction in<br />
kernel weight: ant17.148 and seg3.c about 1/3 normal and ant17.567 about 3/4<br />
normal (3). The seg3 locus was named before the ant17 locus, but many more<br />
mutants were identified at the ant17 locus. Therefore, see BGS 599 for a<br />
complete listing of ant17 mutants.<br />
Origin of mutant:<br />
A spontaneous mutant in Compana (PI 539111) (4).<br />
Mutational events:<br />
seg3.c in Compana (4, 5), ant17.148 (Galant, NGB 13698) in Triumph (PI<br />
268180, NGB 13678) (3).<br />
Mutant used for description and seed stocks:<br />
seg3.c in Compana (GSHO 752); seg3.c in Bowman (PI 483237)*7 (GSHO<br />
1957), ant17.148 in Bowman*4 (GSHO 1973).<br />
References:<br />
1. Boyd, P.W., and D. E. Falk. 1990. (Personal communications).<br />
2. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1985. Anatomy of immature<br />
grains of eight material effect shrunken endosperm barley mutants. Amer. J. Bot.<br />
72:248-256.<br />
3. Franckowiak, J.D. (Personal communications).<br />
4. Jarvi, A.J. 1970. Shrunken endosperm mutants in barley, Hordeum vulgare.<br />
Ph.D. Thesis. Montana State Univ., Bozeman.<br />
5. Jarvi, A.J., and R.F. Eslick. 1971. BGS 379, Normal vs. shrunken endosperm,<br />
se3. Barley Genet. Newsl. 1:191.<br />
6. Jarvi, A.J., and R.F. Eslick. 1975. Shrunken endosperm mutants in barley.<br />
Crop Sci. 15:363-366.<br />
Prepared:<br />
A.J. Jarvi and R.F. Eslick. 1971. BGN 1:191.<br />
B. Jende-Strid. 1999. BGN 29:88-89, as BGS 599, proanthocyanidin-free 17,<br />
ant17.<br />
Revised:<br />
265
Barley Genetics Newsletter (2007) 37: 188-301<br />
R.F. Eslick. 1976. BGN 6:137.<br />
T. Tsuchiya. 1980. BGN 10:126.<br />
J.D. Franckowiak. 1997. BGN 26:327.<br />
J.D. Franckowiak and U. Lundqvist. 2007. BGN 37:265-266.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 380, Shrunken endosperm genetic 4, seg4<br />
Stock number: BGS 380<br />
Locus name: Shrunken endosperm genetic 4<br />
Locus symbol: seg4<br />
Previous nomenclature and gene symbolization:<br />
Shrunken endosperm = se4 (3).<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Located in chromosome 7HL [1L] (2), over 34.0 cM from the centromere (4).<br />
Description:<br />
Under field conditions in the original mutant, seed size is reduced to about 38%<br />
of normal and seed set is about 50% of normal. Stand establishment is poor<br />
under field conditions (2, 4). Endosperms of seg4 were characterized by<br />
progressively distorted, disorganized growth, but the quantity of endosperm<br />
tissue at maturity varied from severely reduced to near normal (1). The stock<br />
described in the 1997 revision was incorrect and was in fact a mixture with<br />
GSHO 755 (BGS 382, shrunken endosperm xenia 1, sex1), which was identified<br />
by Dr. Marion Röder, IPK, Gatersleben.<br />
Origin of mutant:<br />
A spontaneous mutant in Compana (PI 539111) (2).<br />
Mutational events:<br />
seg4.d in Compana (2, 3).<br />
Mutant used for description and seed stocks:<br />
seg4.d in Compana (GSHO 753).<br />
References:<br />
1. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1985. Anatomy of immature<br />
grains of eight material effect shrunken endosperm barley mutants. Amer. J. Bot.<br />
72:248-256.<br />
2. Jarvi, A.J. 1970. Shrunken endosperm mutants in barley, Hordeum vulgare.<br />
Ph.D. Thesis. Montana State Univ., Bozeman.<br />
3. Jarvi, A.J., and R.F. Eslick. 1971. BGS 380, Normal vs. shrunken endosperm,<br />
se4. Barley Genet. Newsl. 1:192.<br />
4. Jarvi, A.J., and R.F. Eslick. 1975. Shrunken endosperm mutants in barley.<br />
Crop Sci. 15:363-366.<br />
Prepared:<br />
A.J. Jarvi and R.F. Eslick. 1971. BGN 1:192.<br />
Revised:<br />
R.F. Eslick. 1976. BGN 6:138.<br />
T. Tsuchiya. 1980. BGN 10:127.<br />
J.D. Franckowiak. 1997. BGN 26:328.<br />
J.D. Franckowiak. 2007. BGN 37:267.<br />
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BGS 396, Shrunken endosperm genetic 6, seg6<br />
Stock number: BGS 396<br />
Locus name: Shrunken endosperm genetic 6<br />
Locus symbol: seg6<br />
Previous nomenclature and gene symbolization:<br />
Shrunken endosperm = se6 (3).<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Located in chromosome 3HL (4).<br />
Description:<br />
Seed size is reduced, but the degree reduction is affected by environment. Seed<br />
weights of 25, 50, and 75% of normal are reported for plants grown in the field in<br />
Arizona, in the field in Montana, and in the greenhouse in Arizona, <strong>US</strong>A,<br />
respectively (4). Pollen mother cell meiosis and pollen fertility are normal. Seed<br />
from seg6.g plants can be used to establish stands under field conditions (4).<br />
Light microscopy revealed that seg6 mutants exhibited premature termination of<br />
grain filling because of the necrosis and crushing of the chalazal and nucellar<br />
projection of the pericarp early during grain filling (1).<br />
Origin of mutant:<br />
A spontaneous mutant in Ingrid (CIho 10083) (3).<br />
Mutational events:<br />
seg6.f in an unknown hybrid, seg6.g in Ingrid (3).<br />
Mutant used for description and seed stocks:<br />
seg6.g in Ingrid (GSHO 2467); seg6.g in Bowman (PI 483237)*4 (GSHO 1975).<br />
References:<br />
1. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1985. Anatomy of immature<br />
grains of eight material effect shrunken endosperm barley mutants. Amer. J. Bot.<br />
72:248-256.<br />
2. Ramage, R.T. 1983. Chromosome location of shrunken endosperm mutants<br />
seg6g and seg8k. Barley Genet. Newsl. 13:64-65.<br />
3. Ramage, R.T., and R.F. Eslick. 1975. BGS 396, Shrunken endosperm, xenia,<br />
se6. Barley Genet. Newsl. 5:114.<br />
4. Ramage, R.T., and J.F. Scheuring. 1976. Shrunken endosperm mutants seg6<br />
and seg7. Barley Genet. Newsl. 6:59-60.<br />
Prepared:<br />
R.T. Ramage and R.F. Eslick. 1975. BGN 5:114.<br />
Revised:<br />
R.T. Ramage and R.F. Eslick. 1976. BGN 6:141.<br />
T. Tsuchiya. 1980. BGN 10:130.<br />
R.T. Ramage. 1983. BGN 13:115.<br />
J.D. Franckowiak. 1997. BGN 26:344.<br />
J.D. Franckowiak. 2007. BGN 37:268.<br />
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BGS 397, Shrunken endosperm genetic 7, seg7<br />
Stock number: BGS 397<br />
Locus name: Shrunken endosperm genetic 7<br />
Locus symbol: seg7<br />
Previous nomenclature and gene symbolization:<br />
Shrunken endosperm = se7 (4).<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Location is unknown.<br />
Description:<br />
Seed size is reduced less than other seg mutants, but the degree reduction is<br />
affected by environment. Seed weights of 40, 75, and 90% of normal are<br />
reported for plants grown in the field in Arizona, in the field in Montana, and in the<br />
greenhouse in Arizona, <strong>US</strong>A, respectively (4). Pollen mother cell meiosis and<br />
pollen fertility are normal. Seed from seg7.h plants can be used to establish<br />
stands under field conditions (4). The shrunken endosperm trait was very difficult<br />
to detect in the crosses to Bowman (2). Light microscopy revealed that seg7<br />
mutants exhibited premature termination of grain filling because of the necrosis<br />
and crushing of the chalazal and nucellar projection of the pericarp early during<br />
grain filling (1).<br />
Origin of mutant:<br />
A spontaneous mutant in Ingrid (CIho 10083) (3).<br />
Mutational events:<br />
seg7.h in Ingrid (4).<br />
Mutant used for description and seed stocks:<br />
seg7.h in Ingrid (GSHO 2468); seg7.h in Bowman (PI 483237)*3 (GSHO 2352).<br />
References:<br />
1. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1985. Anatomy of immature<br />
grains of eight material effect shrunken endosperm barley mutants. Amer. J. Bot.<br />
72:248-256.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Ramage, R.T., and R.F. Eslick. 1975. BGS 397, Shrunken endosperm, xenia,<br />
se7. Barley Genet. Newsl. 5:115.<br />
4. Ramage, R.T., and J.F. Scheuring. 1976. Shrunken endosperm mutants seg6<br />
and seg7. Barley Genet. Newsl. 6:59-60.<br />
Prepared:<br />
R.T. Ramage and R.F. Eslick. 1975. BGN 5:115.<br />
Revised:<br />
R.T. Ramage and R.F. Eslick. 1976. BGN 6:142.<br />
T. Tsuchiya. 1980. BGN 10:131.<br />
J.D. Franckowiak. 1997. BGN 26:345.<br />
J.D. Franckowiak. 2007. BGN 37:269.<br />
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Stock number: BGS 437<br />
Locus name: Eceriferum-zt<br />
Locus symbol: cer-zt<br />
BGS 437, Eceriferum-zt, cer-zt<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Location in chromosome 2HS (1), in bin 2H-01 about 16.8 distal from SSR<br />
marker Bmac0134 (1).<br />
Description:<br />
Surface wax coating on the spike appears slightly reduced (wax code + ++ ++)<br />
(3). The reduction in the surface wax seemed greater in plants selected from the<br />
backcrosses to Bowman (wax code +/- ++ ++).<br />
Origin of mutant:<br />
An ethyl methanesulfonate and neutron induced mutant in Foma (CIho 11333)<br />
(2).<br />
Mutational events:<br />
cer-zt.389 (NGB 111276), -zt.479 (NGB 111367) in Foma (2).<br />
Mutant used for description and seed stocks:<br />
cer-zt.389 in Foma (GSHO 1527); cer-zt.389 in Bowman (PI 483237)*2 (GSHO<br />
2205).<br />
References:<br />
1. Dahleen, L.S., and J.D. Franckowiak. 2006. SSR linkages to eight additional<br />
morphological marker traits. Barley Genet. Newsl. 36:12-16.<br />
2. Lundqvist, U. (Unpublished).<br />
3. Lundqvist, U., and D. von Wettstein. 1971. Stock list for the eceriferum<br />
mutants. Barley Genet. Newsl. 1:97-102.<br />
Prepared:<br />
U. Lundqvist. 1975. BGN 5:155.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:389.<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:270.<br />
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Stock number: BGS 449<br />
Locus name: Eceriferum-yf<br />
Locus symbol: cer-yf<br />
BGS 449, Eceriferum-yf, cer-yf<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Location is unknown.<br />
Description:<br />
Surface wax coating on the leaf blade is reduced (wax code ++ ++ +) (3). In the<br />
Bowman backcross-derived line, mutant plants have pale green leaves, heading<br />
is delayed by several days, and plants are slightly shorter (1).<br />
Origin of mutant:<br />
A neutron induced mutant in Bonus (PI 189763) (2).<br />
Mutational events:<br />
cer-yf.652 (NGB 111540), -yf.804 (NGB 111692) in Bonus (3).<br />
Mutant used for description and seed stocks:<br />
cer-yf.652 in Bonus (GSHO 1539); cer-yf.652 in Bowman (PI 483237)*3 (GSHO<br />
2212).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Lundqvist, U. (Unpublished).<br />
3. Lundqvist, U., and D. von Wettstein. 1973. Stock list for the eceriferum<br />
mutants II. Barley Genet. Newsl. 3:110-112.<br />
Prepared.<br />
U. Lundqvist. 1975. BGN 5:167.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:401.<br />
Revised:<br />
U. Lundqvist. 2007. BGN 37:271.<br />
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BGS 455, Shrunken endosperm genetic 8, seg8<br />
Stock number: BGS 455<br />
Locus name: Shrunken endosperm genetic 8<br />
Locus symbol: seg8<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Located in chromosome 7H [1] (4).<br />
Description:<br />
Seed size is reduced and maturity is delayed. Seed weights of 24, 23, and 27%<br />
of normal are reported for plants grown in the field in Arizona, in the field in<br />
Montana, and in the greenhouse in Arizona, <strong>US</strong>A, respectively (4). Pollen mother<br />
cell meiosis and pollen fertility are normal. Seed from seg8.k plants can be used<br />
to establish stands under field conditions (4). Endosperms of seg8 developed as<br />
two-filled lateral lobes with no central endosperm lobe, resulting in a distinct<br />
dorsal crease (1).<br />
Origin of mutant:<br />
A spontaneous mutant in 60Ab1810-53 (CIho 15686) (3).<br />
Mutational events:<br />
seg8.k in 60Ab1810-53 (3, 4).<br />
Mutant used for description and seed stocks:<br />
seg8.k in 60Ab1810-53 (GSHO 2469); seg8.k in Bowman (PI 483237)*5 (GSHO<br />
1854).<br />
References:<br />
1. Felker, F.C., D.M. Peterson, and O.E. Nelson. 1985. Anatomy of immature<br />
grains of eight material effect shrunken endosperm barley mutants. Amer. J. Bot.<br />
72:248-256.<br />
2. Ramage, R.T. 1983. Chromosome location of shrunken endosperm mutants<br />
seg6g and seg8k. Barley Genet. Newsl. 13:64-65.<br />
3. Ramage, R.T., and C.L. Crandall. 1981. BGS 453, Shrunken endosperm,<br />
seg8. Barley Genet. Newsl. 11:103.<br />
4. Ramage, R.T., and C.L. Crandall. 1981. Shrunken endosperm mutant seg8.<br />
Barley Genet. Newsl. 11:34.<br />
Prepared:<br />
R.T. Ramage and C.L. Crandall. 1981. BGN 11:103 as BGS 453.<br />
Revised:<br />
R.T. Ramage. 1983. BGN 13:116 as BGS 453.<br />
T. Tsuchiya. 1983. BGN 13:117. BGS number changed to BGS 455.<br />
J.D. Franckowiak. 1997. BGN 26:405.<br />
J.D. Franckowiak. 2007. BGN 37:272.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 474<br />
Locus name: Laxatum-a<br />
Locus symbol: lax-a<br />
BGS 474, Laxatum-a, lax-a<br />
Previous nomenclature and gene symbolization:<br />
Laxatum-01 = lax-01 (3, 6, 11).<br />
Laxatum-a = lax-a 01 (12).<br />
Inheritance:<br />
Monofactorial recessive (7, 11).<br />
Located in chromosome 5HL [7L] (7, 10), about 2.4 cM proximal from the ari-e<br />
(breviaristatum-e) locus (5, 14), and about 3.1 cM from the ert-g (erectoides-g)<br />
locus (5, 12, 13).<br />
Description:<br />
Florets have five anthers with two developing from transformed lodicules (3, 15);<br />
however, the extra anthers are deficient in having two rather than four<br />
microsporangia (1). The grain is thin and angular and caryopses are exposed<br />
between the lemma and palea. The awn has a very wide base, without a distinct<br />
notch in the lemma attachment region. Rachis internodes are 13% longer than<br />
normal. Tillers arise at oblique angles giving isolated plants an appearance of a<br />
tufty growth habit (6). Treatment of leaves after tillering with GA3 increases rachis<br />
internode length (15).<br />
Origin of mutant:<br />
A gamma-ray induced mutant in Bonus (PI 189763) (2, 6, 9).<br />
Mutational events:<br />
lax-a.01 (NGB 116334), -a.4 (NGB 116338), -a.8 (NGB 116342), -a.20 (NGB<br />
116354), -a.37 (NGB 116372), -a.39 (NGB 116374), -a.54 (NGB 116388) in<br />
Bonus (6, 9); lax-a.92 (NGB 116425, NGB 116426) in Bonus (9); lax-a.208 (NGB<br />
116435, NGB 116436), -a.218 (NGB 116446), -a.222 (NGB 116450), -a.229<br />
(NGB 116457, NGB 116458), -a.249, -a.256 (NGB 116483), -a.278 (NGB<br />
116503), -a.286 (NGB 116510) in Foma (CIho 11333) (6, 8); -a.353 (NGB<br />
116559, NGB 116560), -a.369 (NGB 116578, 116579), -a.373 (NGB 116583), -<br />
a.398 (NGB 116608), -a.405 (NGB 116613), -a.406 (NGB 116614) in Kristina<br />
(NGB 14661) (7); -a.413 (NGB 116621, NGB 116622), -a.434 (NGB 116647), -<br />
a.437 (NGB 116650), -a.444 (NGB 116658, NGB 116659), -a.448 (NGB<br />
116664), -a.450 (NGB 116667, NGB 116668), -a.455 (NGB 116674, NGB<br />
116675), -a.472 (NGB 116695) in Bonus (8); a lax-a mutant (Mut 2100/61) in<br />
Proctor (PI 280420) (4).<br />
Mutant used for description and seed stocks:<br />
lax-a.8 in Bonus (GSHO 1775); lax-a.8 in Bowman (PI 483237)*7 (GSHO 2103).<br />
References:<br />
1. Bossinger, G., W. Rohde W, U. Lundqvist U, and F. Salamini F. 1992.<br />
Genetics of barley development: mutant phenotypes and molecular aspects. p.<br />
231-264. In P.R. Shewry (ed.) Barley: Genetics, Biochemistry, Molecular Biology<br />
and Biotechnology. CAB International, Oxford.<br />
2. Ehrenberg, L., Å. Gustafsson, and U. Lundqvist. 1961. Viable mutants induced<br />
in barley by ionizing radiations and chemical mutagens. Hereditas 47:257-278.<br />
3. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
273
Barley Genetics Newsletter (2007) 37: 188-301<br />
4. Häuser, H., and G. Fischbeck. 1972. Translocations and genetic analysis of<br />
other mutants. BGN 2:28-29.<br />
5. Jensen, J. 1981. Construction of a barley chromosome 7 linkage map. p. 927-<br />
939. In M.J.C. Asher, R.P. Ellis, A.M. Hayter, and R.N.H. Whitehouse (eds.)<br />
Barley Genetics IV. Proc. Fourth Int. Barley Genet. Symp., Edinburgh. Edinburgh<br />
Univ. Press, Edinburgh.<br />
6. Larsson, H.E.B. 1985. Morphological analysis of laxatum barley mutants.<br />
Hereditas 103:239-253.<br />
7. Larsson, H.E.B. 1985. Linkage studies with genetic markers and some<br />
laxatum barley mutants. Hereditas 103:269-279.<br />
8. Lundqvist, U. (Unpublished).<br />
9. Persson, G. 1969. An attempt to find suitable genetic markers for the dense<br />
ear loci in barley II. Hereditas 63:1-28.<br />
10. Persson, G., and A. Hagberg. 1965. Localization of nine induced mutations in<br />
the barley chromosomes. Barley Newsl. 8:52-54.<br />
11. Persson, G., and A. Hagberg. 1969. Induced variation in a quantitative<br />
character in barley. Morphology and cytogenetics of erectoides mutants.<br />
Hereditas 61:115-178.<br />
12. Søgaard, B. 1974. Three-point tests on chromosome 1 and 7. BGN 4:70-73.<br />
13. Søgaard, B. 1977. The localization of eceriferum loci in barley. IV. Three<br />
point tests of genes on chromosome 7 in barley. Carlsberg Res. Commun. 42:35-<br />
43.<br />
14. Stoy, V., and A. Hagberg. 1967. Effects of growth regulators on ear density<br />
mutants in barley. Hereditas 58:359-384.<br />
15. Wettstein, D. von, Å. Gustafsson, and L. Ehrenberg. 1959.<br />
Mutationsforschung und Züchtung. p. 7-50. In Arbeitsgemeinschaft für Forschung<br />
des Landes Nordrhein-Westfalen, Heft 73. Westdeutscher Verlag Köln und<br />
Opladen.<br />
Prepared:<br />
H.E.B. Larsson and U. Lundqvist. 1986. BGN 16:57.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:421-422.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:273-274.<br />
274
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BGS 516, Reaction to Septoria passerinii 2, Rsp2<br />
Stock number: BGS 516<br />
Locus name: Reaction to Septoria passerinii 2<br />
Locus symbol: Rsp2<br />
Previous nomenclature and gene symbolization:<br />
Resistance to Septoria passerinii Sacc = Sep2 (1, 2).<br />
Inheritance:<br />
Monofactorial incomplete dominant (2).<br />
Location in chromosome 1HS [5S] (3), about 3.9 cM from the Rsp3 (reaction to<br />
Septoria passerinii 3) locus (2), cosegregation with SCAR marker E-ACT/M-CAA-<br />
170a and close to the Rsp3 locus (3), about 17.6 cM proximal from marker RFLP<br />
Act8 (3).<br />
Description:<br />
The Rsp2.b gene conditions a high level of resistance to a single spore culture of<br />
Septoria passerinii isolated in Minnesota, <strong>US</strong>A. Pycnidia are observed in some,<br />
but not all lesions, on all F1 plants (2).<br />
Origin of mutant:<br />
Natural occurrence in accession CIho 4780 (PI 70837) (2).<br />
Mutational events:<br />
Rsp2.b in PI 70837.<br />
Mutant used for description and seed stocks:<br />
Rsp2.b in PI 70837 (GSHO 2511).<br />
References:<br />
1. Moseman, J.G. 1972. Report on genes for resistance to pests. Barley Genet.<br />
Newsl. 2:145-147.<br />
2. Rasmusson, D.C., and W.E. Rogers. 1963. Inheritance of resistance to<br />
septoria in barley. Crop Sci. 3:161-163.<br />
3. Zhong, S., H. Toubia-Rahme, B.J. Steffenson, and K.P. Smith. 2006.<br />
Molecular mapping and marker-assisted selection of genes for septoria speckled<br />
leaf blotch resistance in barley. Phytopathology 96:993-999.<br />
Prepared:<br />
D.C. Rasmusson. 1988. BGN 18:85 as BGS 466.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:442.<br />
J.D. Franckowiak. 2007. BGN 37:275.<br />
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BGS 517, Reaction to Septoria passerinii 3, Rsp3<br />
Stock number: BGS 517<br />
Locus name: Reaction to Septoria passerinii 3<br />
Locus symbol: Rsp3<br />
Previous nomenclature and gene symbolization:<br />
Resistance to Septoria passerinii Sacc = Sep3 (1, 2).<br />
Inheritance:<br />
Monofactorial incomplete dominant (2).<br />
Location in chromosome 1HS [5S] (3), about 3.9 cM from the Rsp2 (reaction to<br />
Septoria passerinii 2) locus (2), cosegregation with SCAR marker E-ACT/M-CAA-<br />
170a and close to the Rsp2 locus (3), about 17.6 cM proximal from marker RFLP<br />
Act8 (3).<br />
Description:<br />
The Rsp3.c gene conditions a high level of resistance to a single spore culture of<br />
Septoria passerinii isolated in Minnesota, <strong>US</strong>A. Infection occurs on F1 seedlings,<br />
but is limited to a few lesions (2).<br />
Origin of mutant:<br />
Natural occurrence in selection II-51-43 from a Feebar/Kindred cross (CIho<br />
10644) (2).<br />
Mutational events:<br />
Rsp3.c in CIho 10644.<br />
Mutant used for description and seed stocks:<br />
Rsp3.c in CIho 10644 (GSHO 2512).<br />
References:<br />
1. Moseman, J.G. 1972. Report on genes for resistance to pests. Barley Genet.<br />
Newsl. 2:145-147.<br />
2. Rasmusson, D.C., and W.E. Rogers. 1963. Inheritance of resistance to<br />
septoria in barley. Crop Sci. 3:161-163.<br />
3. Zhong, S., H. Toubia-Rahme, B.J. Steffenson, and K.P. Smith. 2006.<br />
Molecular mapping and marker-assisted selection of genes for septoria speckled<br />
leaf blotch resistance in barley. Phytopathology 96:993-999.<br />
Prepared:<br />
D.C. Rasmusson. 1988. BGN 18:86 as BGS 467.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:443.<br />
J.D. Franckowiak. 2007. BGN 37:276.<br />
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Stock number: BGS 518<br />
Locus name: Semidwarf 1<br />
Locus symbol: sdw1<br />
BGS 518, Semidwarf 1, sdw1<br />
Previous nomenclature and gene symbolization:<br />
Denso dwarf = denso (5, 12).<br />
Inheritance:<br />
Monofactorial recessive (5, 13), although some F1's tend to be intermediate in<br />
height compared to their parents (1, 8).<br />
Location in chromosome 3HL (2, 9), probably proximal from the gsh2 (glossy<br />
sheath 2) locus, near RFLP marker PSR170 (9), in bin 3H-11 (7), near RFLP<br />
marker R1545 (16).<br />
Description:<br />
Plants homozygous for the sdw1.a gene range from 10 to 30 cm shorter than<br />
normal sibs, with expression partial dependent on environment (1, 12, 14). Spike<br />
length is variable, but fully as long as normal barley. The stock used for<br />
description of the sdw1.a gene, M21, has the short straw and long spike of the<br />
original 'Jotun Mutant' as well as a large culm diameter from its parent 'Vantage'<br />
(1, 14). The semidwarf mutants, 'Diamant' and ‘Abed Denso’, are alleles at the<br />
sdw1 locus (5, 10). Alleles at the sdw1 locus are associated with semiprostrate<br />
juvenile growth (5, 12), delayed maturity (4, 6, 12, 15), and reduced malt quality<br />
(4, 6, 12). The sdw1 mutants are GA sensitive (3, 16), and they are very likely<br />
mutants in an orthologue of the rice sd1 gene (16), which encodes a GA-oxidase<br />
that produces lower levels of GA and therefore causes the dwarf phenotype (11).<br />
The original cultivar ‘Trumpf’ was also marketed in the United Kingdom as<br />
‘Triumph’.<br />
Origin of mutant:<br />
An X-ray induced mutant in the Norwegian cultivar Jotun (PI 467357) isolated as<br />
Jotun 22 by Knut Mikaelsen (1, 8).<br />
Mutational events:<br />
sdw1.a in Jotun (66/86, GSHO 1414) (14); sdw1.c (denso) in Abed Denso (PI<br />
361639) (5); sdw1.d (Diamant) in Valticky (5); sdw1.e (Risø 9265) in Bomi (PI<br />
43371) (5).<br />
Mutant used for description and seed stocks:<br />
sdw1.a in M21 (CIho 15481, GSHO 2513) from the cross Jotun Mutant/Kindred//<br />
Vantage (13); sdw1.d in Trumpf (Triumph, PI 548762, GSHO 2465); sdw1.a from<br />
a Jotun derivative in Bowman (PI 483237)*7 (GSHO 1978); sdw1.d from Trumpf<br />
in Bowman*4 (GSHO 1979).<br />
References:<br />
1. Ali, M.A.M., O. Okiror, and D.C. Rasmusson. 1978. Performance of semidwarf<br />
barley. Crop Sci. 18:418-422.<br />
2. Barau, U.M., K.J. Chambers, W.T.B. Thomas, C.A. Hackett, V. Lea, P. Jack,<br />
B.P. Forster, R. Waugh, and W. Powell. 1993. Molecular mapping of genes<br />
determining height, time to heading, and growth habit in barley (Hordeum<br />
vulgare). Genome 36:1080-1087.<br />
3. Boulger, M.C., R.G. Sears, and W.E. Kronstad. 1982. An investigation of the<br />
association between dwarfing sources and gibberellic acid response in barley. p.<br />
550-553. In M.J.C. Asher, R.P. Ellis, A.M. Hayter, and R.N.H. Whitehouse (eds.)<br />
277
Barley Genetics Newsletter (2007) 37: 188-301<br />
Barley Genetics IV. Proc. Fourth Int. Barley Genet. Symp., Edinburgh. Edinburgh<br />
Univ. Press, Edinburgh.<br />
4. Foster, A.E., and A.P. Thompson. 1987. Effects of a semidwarf gene from<br />
Jotun on agronomic and quality traits of barley. p. 979-982. In S. Yasuda and T.<br />
Konishi (eds.) Barley Genetics V., Proc. Fifth Int. Barley Genet. Symp.,<br />
Okayama, 1986. Sanyo Press Co., Okayama.<br />
5. Haahr, V., and D. von Wettstein. 1976. Studies of an induced, high-yielding<br />
dwarf-mutant of spring barley. p. 215-218. In H. Gaul (ed.) Barley Genetics III.,<br />
Proc. Third Int. Barley Genet. Symp., Garching, 1975. Verlag Karl Thiemig,<br />
München.<br />
6. Hellewell, K.B., D.C. Rasmusson, M. Gallo-Meagher. 2000. Enhancing yield of<br />
semidwarf barley. Crop Sci. 40:352-358.<br />
7. Kleinhofs, A. 2006. Integrating molecular and morphological/physiological<br />
marker maps. Barley Genet. Newsl. 36:66-82.<br />
8. Lambert, J.W., and M. Shafi. 1959. Inheritance and heritability of height in<br />
three barley crosses. Barley Newsl. 3:7-8. (Abstr.)<br />
9. Laurie, D.A., N. Pratchett, C. Romero, E. Simpson, and J.W. Snape. 1993.<br />
Assignment of the denso dwarfing gene to the long arm of chromosome 3 (3H) of<br />
barley by use of RFLP markers. Plant Breed. 111:198-203.<br />
10. Mickelson, H.R., and D.C. Rasmusson. 1994. Genes for short stature in<br />
barley. Crop Sci. 34:1180-1183.<br />
11. Murai, M., T. Komazaki, and S. Sato. 2004. Effects of sd1 and Ur1 (Undulate<br />
rachis – 1) on lodging resistance and related traits in rice. Breed. Science 54:<br />
333-340.<br />
12. Powell, W., P.D.J. Caligari, W.T.B. Thomas, and J.L. Jinks. 1985. The effects<br />
of major genes on quantitatively varying characters in barley. 2. The denso and<br />
day length response loci. Heredity 54:349-352.<br />
13. Powell, W., P.D.J. Caligari, W.T.B. Thomas, and J.L. Jinks. 1991. The effects<br />
of major genes on quantitatively varying characters in barley. 4. The GPert and<br />
denso loci and quality characters. Heredity 66:381-389.<br />
14. Rasmusson, D.C., E.E. Banttari, and J.W. Lambert. 1973. Registration of<br />
M21 and M22 semidwarf barley. Crop Sci. 13:777.<br />
15. Yin, X., P.C. Struik, F.A. van Eeuwijk, P. Stam, and J. Tang. 2005. QTL<br />
analysis and QTL-based prediction of flowering phenology in recombinant inbred<br />
lines of barley. J. Exp. Bot. 56 (413):967-976.<br />
16. Zhang, J., X. Yang, P. Moolhuijzen, C. Li, M. Bellgard, R. Lance, and R.<br />
Appels. 2005. Towards isolation of the barley green revolution gene.<br />
Proceedings of Australian Barley Technical Symposium 2005.<br />
http://www.cdesign.com.au/proceedings_abts2005/posters%20(pdf)/poster_li.pdf.<br />
Prepared:<br />
D.C. Rasmusson. 1988. BGN 18:87 as BGS 468.<br />
Revised:<br />
J.D. Franckowiak. 1997. BGN 26:444-445.<br />
J.D. Franckowiak. 2007. BGN 37:277-278.<br />
278
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 546<br />
Locus name: Intermedium spike-k<br />
Locus symbol: int-k<br />
BGS 546, Intermedium spike-k, int-k<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Located in chromosome 7H [1] (2) in the centromeric region closely linked to<br />
markers Bmag0217 and Bmac0162 in bins 6 to 7 (2).<br />
Description:<br />
The spike is short and dense in the original mutant. Lateral spikelets are<br />
enlarged and the apex is pointed, and they occasionally have a short awn. Seed<br />
set does not occur in lateral spikelets and the central spikelets are semi-sterile<br />
(3). Plants of the original stock have a dense coating of surface waxes. In the<br />
Bowman backcross-derived line, plants are small and weak (about 1/2 normal<br />
height) and have short spikes (1/2 normal), reduced awn length (3/4 normal), and<br />
very poor seed set. Awns of plants in the derived line are semi-rough, but F1<br />
hybrids with Bowman have semismooth awns (1).<br />
Origin of mutant:<br />
An ethyl methanesulfonate induced mutant in Kristina (NGB 1500) (3).<br />
Mutational events:<br />
int-k.47 in Kristina (3).<br />
Mutant used for description and seed stocks:<br />
in-k.47 in Kristina (GSHO 1770, NGB 115465); int-k.47 in Bowman (PI<br />
483237)*6.<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Dahleen, L.S., and J.D. Franckowiak. 2006. SSR Linkages to Eight Additional<br />
Morphological Marker Traits. Barley Genet. Newsl. 36:12-16.<br />
3. Lundqvist, U., and A. Lundqvist. 1988. Induced intermedium mutants in barley:<br />
origin, morphology and inheritance. Hereditas 108:13-26.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:472.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:279.<br />
279
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 547<br />
Locus name: Intermedium spike-m<br />
Locus symbol: int-m<br />
BGS 547, Intermedium spike-m, int-m<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (4).<br />
Location is unknown.<br />
Description:<br />
The spike is very short and has irregular rachis internode lengths. Lateral<br />
spikelets are enlarged and pointed, but they do not set seed. Spikelet density at<br />
the base of the spike is increased. Rachis internodes at the tip of the spike are<br />
very short, and the spike appears to have two or three fused or fasciated terminal<br />
spikelets. Tillering of int-m plants is increased (1, 4) and heading is slightly earlier<br />
(4).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Bonus (PI 189763) (3).<br />
Mutational events:<br />
int-m.85 (NGB 115503) in Bonus (3); int-m.la (GSHO 1773) in Lamont (PI<br />
512036) (2).<br />
Mutant used for description and seed stocks:<br />
int-m.85 in Bonus (GSHO 1772); int-m.85 in Bowman (PI 483237)*7 (GSHO<br />
2273); int-m.la in Bowman (PI 483237)*5 (GSHO 2274).<br />
References:<br />
1. Babb, S., and G.J. Muehlbauer. 2003. Genetic and morphological<br />
characterization of the barley uniculm 2 (cul2) mutant. Theor. Appl. Genet.<br />
106:846-857.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Lundqvist, U. (Unpublished).<br />
4. Lundqvist, U., and A. Lundqvist. 1988. Induced intermedium mutants in barley:<br />
origin, morphology and inheritance. Hereditas 108:13-26.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:473.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:280.<br />
280
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 566<br />
Locus name: Erectoides-t<br />
Locus symbol: ert-t<br />
BGS 566, Erectoides-t, ert-t<br />
Previous nomenclature and gene symbolization:<br />
Erectoides-55 = ert-55 (7).<br />
Brachytic 4 = br4 (10).<br />
Brachytic-g = brh.g (3).<br />
Brachytic 3 = brh3 (4).<br />
Brachytic-i = brh.i (3).<br />
Brachytic-y = brh.y (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 5, 7, 9).<br />
Located in chromosome 2HS (2), approximately 11.4 cM distal from SSR marker<br />
Bmac0134 (2), near the boundary between bins 2H-01 and 2H-02 (2).<br />
Description:<br />
Spikes are semicompact, rachis internode length is about 2.7 mm in the original<br />
mutant, and culm length is about 2/3 of normal. These phenotypic traits plus<br />
short awns are inherited together (9). Based on general appearance of the<br />
plants, ert-t can be placed in the brachytic class of semidwarf mutants (3, 10).<br />
Awns are about 2/3 normal length and curled or coiled near their tips. The ertt.55<br />
mutant has a short seedling leaves and is sensitive to gibberellic acid<br />
treatment (1). In the Bowman backcross-derived lines, peduncles are about 2/3<br />
normal, rachis internodes are slightly short, and lodging is reduced. Kernels are<br />
slightly lighter and yields are about 1/2 normal (2).<br />
Origin of mutant:<br />
An X-ray induced mutant in Bonus (PI 189763) (7).<br />
Mutational events:<br />
ert-t.55 in Bonus (NGB 112654) (7); brh3.g (17:10:1, DWS1002), brh3.h (17:11:3,<br />
DWS1003), brh3.i (17:12:1, DWS1004) in Birgitta (NGB 1494, NGB 14667) (2, 3,<br />
4, 8); brh3.y (10001, DWS1230, GSHO 1688) in Bido (PI 399485) (2, 3, 6).<br />
Mutant used for description and seed stocks:<br />
ert-t.55 in Bonus (GSHO 494); ert-t.55 in Bowman (PI 483237)*7 (GSHO 2257);<br />
brh3.g in Birgitta (GSHO 1672); brh3.g in Bowman*7 (GSHO 2167); brh3.y in<br />
Bowman*6 (GSHO 2178).<br />
References:<br />
1. Börner, A. 1996. GA response in semidwarf barley. Barley Genet. Newsl.<br />
25:24-26.<br />
2. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
BGN 24:56-59.<br />
4. Franckowiak. 2002. BGS 631, Brachytic 3, brh3. BGN 32:132.<br />
5. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
6. Gaul, H. 1986. (Personal communications).<br />
7. Hagberg, A., Å. Gustafsson, and L. Ehrenberg. 1958. Sparsely contra densely<br />
ionizing radiations and the origin of erectoid mutants in barley. Hereditas 44:523-<br />
281
Barley Genetics Newsletter (2007) 37: 188-301<br />
530.<br />
8. Lehmann, L.C. 1985. (Personal communications).<br />
9. Persson, G., and A. Hagberg. 1969. Induced variation in a quantitative<br />
character in barley. Morphology and cytogenetics of erectoides mutants.<br />
Hereditas 61:115-178.<br />
10. Tsuchiya, T. 1976. Allelism testing of genes between brachytic and<br />
erectoides mutants. Barley Genet. Newsl. 6:79-81.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:492.<br />
J.D. Franckowiak. 2002. BGS 631, Brachytic 3, brh3. BGN 32:132.<br />
Revised:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:281-282.<br />
282
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 577, Reaction to Schizaphis graminum 2, Rsg2<br />
Stock number: BGS 577<br />
Locus name: Reaction to Schizaphis graminum 2 (greenbug)<br />
Locus symbol: Rsg2<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial dominant (1).<br />
Location is unknown.<br />
Description:<br />
Resistant seedlings infested with greenbugs (aphids) are not killed or severely<br />
stunted by a buildup of the greenbug population, but susceptible seedlings are<br />
killed or severely stunted (1, 4). The resistance provided by PI 426756 (4 to 5<br />
readings on a 1 to 9 scale) to most S. graminum biotypes was less effective than<br />
that provided the Rsg1.a gene in Post 90 (PI 549081) (2 to 3 readings) (3). PI<br />
426756 was confirmed to provide resistance (2 to 3 readings) to the TX1 isolate<br />
of S. graminum, which produces a susceptible reaction (9 reading) on Post 90<br />
(2).<br />
Origin of mutant:<br />
Natural occurrence in Joa (PI 426756) (1, 4).<br />
Mutational events:<br />
Rsg2.b in PI 426756 (1).<br />
Mutant used for description and seed stocks:<br />
Rsg2.b in PI 426756 (GSHO 2513).<br />
References:<br />
1. Merkle, O.G., J.A. Webster, and G.H. Mogen. 1987. Inheritance of a second<br />
source of greenbug resistance in barley. Crop Sci. 27:241-243.<br />
2. Porter, D.R., J.D. Burd, and D.W. Mornhinweg. 2007. Differentiating greenbug<br />
resistance genes in barley. Euphytica 153:11-14.<br />
3. Porter, D.R., and D.W. Mornhinweg. 2004. Characterization of greenbug<br />
resistance in barley. Plant Breed. 123:493-494.<br />
4. Webster, J.A., and K.J. Starks. 1984. Sources of resistance in barley to two<br />
biotypes of greenbug Schizaphis graminum (Rondani), Homoptera: Aphididae.<br />
Protect. Ecol. 6:51-55.<br />
Prepared:<br />
J.D. Franckowiak. 1997. BGN 26:503.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:283.<br />
283
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 586<br />
Locus name: Bracteatum-d<br />
Locus symbol: bra-d<br />
BGS 586, Bracteatum-d, bra-d<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (1).<br />
Located in chromosome 1HL [5L] (5), about 4.1 cM from AFLP marker E3634-7<br />
(5), probably in bin 1H-14 based on the association with trd1 (third outer glume 1)<br />
(2, 5).<br />
Description:<br />
The characteristic trait of this mutant is the presence of a bract (third outer<br />
glume) outside the two empty glumes of the central spikelet. The bract<br />
subtending the lowest spikelet is always the largest, embracing in some cases<br />
about one-half the spike. Bracts become progressively smaller toward the tip of<br />
the spike. Mutants have elongated basal rachis internodes (3, 4). Pozzi et al. (5)<br />
suggested that bra-d.7 is allelic to trd1 (third outer glume 1) or near the trd1<br />
locus. Allelism studies demonstrated that bra-d.7 is not an allele at the trd1 locus<br />
(3).<br />
Origin of mutant:<br />
An ethylene imine induced mutant in Foma (CIho 11333) (3).<br />
Mutational events:<br />
bra-d.7 (NGB 114310) in Foma (3).<br />
Mutant used for description and seed stocks:<br />
bra-d.7 in Foma (GSHO 1696); bra-d.7 in Bowman (PI 483237)*3 (GSHO 2185).<br />
References:<br />
1. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
2. Kleinhofs, A. 2006. Integrating molecular and morphological/physiological<br />
marker maps. Barley Genet. Newsl. 36:66-82.<br />
3. Lundqvist, U. (Unpublished).<br />
4. Nybom, N. 1954. Mutation types in barley. Acta Agric. Scand. 4:430-456.<br />
5. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 1997. BGN 26:513.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:284.<br />
284
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 593<br />
Locus name: Awned palea 1<br />
Locus symbol: adp1<br />
BGS 593, Awned palea 1, adp1<br />
Previous nomenclature and gene symbolization:<br />
Awned palea = adp (1).<br />
Inheritance:<br />
Monofactorial recessive (1, 2).<br />
Located in chromosome 3HL (2), about 5.8 cM distal from AFLP marker E3634-8<br />
in subgroup 27 of the Proctor/Nudinka map (2).<br />
Description:<br />
This mutant was isolated as a partially female sterile plant with abnormal spikes.<br />
The palea is elongated to form two awns (2), which are derived from two fused<br />
bracts that form the palea (3). Pistils are often transformed into leafy buds and<br />
result in low female fertility and greatly reduced seed set. Two of the anthers<br />
appear normal and the third is deformed to some extent (1). Pollen fertility is<br />
good (1).<br />
Origin of mutant:<br />
A spontaneous mutant in an inbred line (1).<br />
Mutational events:<br />
adp1.a in an unknown inbred line (1).<br />
Mutant used for description and seed stocks:<br />
adp1.a in a selection, with the eog1.a (elongated outer glume 1) gene from<br />
Svalöfs Guldkorn 91 [AHOR 226, a mutant of Gull (CIho 1145, GSHO 466)] (1),<br />
crossed to the unknown line (GSHO 1618); adp1.a in Bowman*5 (GSHO 1950).<br />
References:<br />
1. Ahokas, H. 1977. A mutant of barley: Awned palea. Barley Genet. Newsl. 7:8-<br />
10.<br />
2. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
3. Williams, R.F. 1975. The Shoot Apex and Leaf Growth. Cambridge University<br />
Press, Cambridge.<br />
Prepared:<br />
J.D. Franckowiak. 1998. BGN 28:34.<br />
Revised:<br />
J.D. Franckowiak. 2007. BGN 37:285.<br />
285
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 599, Proanthocyanidin-free 17, ant17<br />
Stock number: BGS 599<br />
Locus name: Proanthocyanidin-free 17<br />
Locus symbol: ant17<br />
Previous nomenclature and symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (4, 5).<br />
Located in chromosome 3HS (1), it has been shown that ant17.148 is an allele at<br />
the seg3 (shrunken endosperm genetic 3, see BGS 379) locus (2).<br />
Description:<br />
Under normal growing conditions no anthocyanin pigmentation is observed in the<br />
mutant plants. The testa layers of the grain of the ant17 mutants lack<br />
proanthocyanidins and catechins, but accumulate homoeriodictyol and<br />
chrysoeriol (7, 10). A full length cDNA clone from barley, coding for a protein<br />
consisting of 377 amino acids (42 kDa), has been isolated. It shows a homology<br />
of 71% to the flavanone-3-hydroxylase enzyme protein from Antirrhinum majus<br />
(12). It is likely that the ant17 gene codes for one subunit and the ant22 gene for<br />
the other subunit of the dimeric flavanone 3-hydroxylase enzyme, which<br />
catalyzes the conversion of flavanones into dihydroflavanols (7, 12). The mutant<br />
stock ant17.148 was released as cultivar Galant (11). Alleles at the ant17 locus<br />
that have been examined in the Bowman genetic background showed a variable<br />
reduction in kernel weight: ant17.148 and seg3.c about 1/3 normal and ant17.567<br />
about 3/4 normal (2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Nordal (NGB 13680) (3).<br />
Mutational events:<br />
ant17.103, 17.104, 17.105, 17.139, 17.140, 17.142, 17.143, 17.145 in Nordal (4);<br />
ant17.107 in Alf (NGB13682) (4); ant17.147, 17.148 (Galant) (NGB 13698),<br />
17.150, 17.151, 17.153, 17.154, 17.180, 17.185 in Triumph (PI 268180, NGB<br />
13678) (4); ant17.352 in Triumph (5); ant17.160 in Gula Abed (NGB 13681) (4);<br />
ant17.165, 17.167, 17.169, 17.171, 17.174, 17.182 in Ark Royal (PI 447006) (4);<br />
ant17.192, 17.193 in Georgie (PI 447012, NGB 13683) (4); ant17.199 in Secobra<br />
4681 (4); ant17.200 in Secobra 4681 (5); ant17.208 in Hege 876 (4); ant17.210,<br />
17.211, 17.217 in Hege 802 (4); ant 17.216 in Hege 802 (5); ant17.220, 17.221,<br />
17.224, in Secobra 4743 (NGB 13679) (4); ant17.227 in Ca 59995 (5); ant17.231<br />
in Tron (4); ant17.237, 17.239, 17.241, 17.242, 17.247, 17.249 in Gunhild (PI<br />
464655, NGB 13690) (4); ant17.243, 17.246 in Gunhild (5); ant17.250, 17.251,<br />
17.252, 17.253, 17.255 in Tokak (PI 264251) (4); ant17.267, 17.268, 17.269 in<br />
Secobra 18193 (NGB 13684) (4); ant17.270 in Secobra 18193 (5); ant17.280 in<br />
Hege 550/75 (NGB 13692) (9); ant17.288, 17.289, 17.290 in Hege 550/75 (4);<br />
ant17.293, 17.294, 17.295, 17.296 in Bonus (PI 189763) (4); ant17.297, 17.298,<br />
17.300, 17.301, 17.307 in Ca 41507 (4); ant17.306, 17.340 in Ca 41507 (5);<br />
ant17.316 in Ca 33787 (NGB 13693) (5); ant17.318, 17.321, 17.326 in Harry (PI<br />
491575) (5); ant17.331 in Hege A2/A4 (5); ant17.335, 17.336, 17.338 in<br />
Ackermann 724/5/7 (5); ant17.359 in Hege15/74-1A (5); ant17.370 in Ackermann<br />
72/440 (5); ant17.372, 17.413, 17.414, 17.417, 17.418, 17.419, 17.444 in Kaya<br />
(5); ant17.375 in Fanette (6); ant17.379, 17.382, 17.383, 17.386, 17.387, 17.388,<br />
17.389, 17.390, 17.391, 17.464, 17.465 in Irene (5); ant17.405 in Odin (6);<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
ant17.408 in KMJ 326 (5); ant17.410, 17.447 in Catrin (5); ant17.421 in VBS<br />
18707 (5); ant17.422, 17.423, 17.424, 17.426 in NZ 3789 (5); ant17.432 in NZ<br />
1836-3 (5); ant17.438, 17.439 in NZ 732.01 (5); ant17.440 in Nordal (5);<br />
ant17.450 in Ca 601427 (5); ant17.453, 17.455, 17.457, 17.458 in Ackermann<br />
1734/5 (5); ant17.462 in Pamela (5); ant17.469, 17.470 in Grit (PI 548764, NGB<br />
13685) (5); ant17.475 in Zenit (PI 564447, NGB 13686) (5); ant17.476 in Zenit<br />
(6); ant17.480 in Secobra 9709 (5); ant17.501 in Advance (CIho 15804) (4);<br />
ant17.504 in Karla (CIho 15860) (4); ant17.506, 17.507, 17.508, 17.509 in OR<br />
9114 (4); ant17.515, 17.516, 17.518 in WA9037-75 (4); ant17.520 in WA9044-75<br />
(4); ant17.530 in Morex (CIho15773) (4); ant17.537, 17.595, 17.619, 17.620 in<br />
Advance (5); ant17.560, 17.561, 17.563, 17.565, 17.567 in Manker (CIho 15549)<br />
(5); ant17.597 in Morex (6); ant17.598 in Morex (5); ant17.600 in S 80351 (5);<br />
ant17.601 in Moravian 111 (CIho 15812) (5); ant17.604 in Harrington (6);<br />
ant17.612 in Andre‚ (PI 469107) (5); ant17.624 in Klages (CIho 15478) (5);<br />
ant17.625 in Robust (M36, PI 476976) (5); ant17.630 in Azure (CIho 15865) (13);<br />
ant17.636, 17.658 in Cougbar (PI 496400) (13); ant17.637 in 8892-78 (13);<br />
ant17.661 in Crest (PI 561409) (13); ant17.1502, 17.1505, 17.1519 in Amagi-Nijo<br />
(4); ant17.1510, 17.1511 in Haruna- Nijo (4); ant17.1515 in Nirakei 61 (4);<br />
ant17.1537 in Nirakei 62 (5); ant17.1544 in Nirakei 63 (5); ant17.1534 in<br />
Nirasaki-Nijo 14 (5); ant17.2022, 17.2067 in Natasha (PI 592171) (6); ant17.2084<br />
in Hege 694/82 (9); ant17.2106 in Ca 708912 (8); ant17.5019 in Sonja (PI<br />
302047) (9); ant17.5024 in Ackermann 72/27/4 (6); ant17.5028 in Trigger (PI<br />
473541) (9); ant17.5034 in Kaskade (9); ant17.5035, 17.5036, 17.5037 in Video<br />
(6); ant17.5038, 17.5039, 17.5040, 17.5042 in Sonja (6); ant17.5044 in<br />
Ackermann 27/220/8 (6).<br />
Mutant used for description and seed stock:<br />
ant17.139 in Nordal (NGB 13697); ant17.148 (Galant) in Triumph (NGB 13698,<br />
GSHO 1628); ant17.148 in Bowman (PI 483237)*4 (GSHO 1973); ant17.567 in<br />
Manker (GSHO 1629); ant17.567 in Bowman*5 (GSHO 1974), seg3.c in<br />
Bowman (PI 483237)*7 (GSHO 1957).<br />
References:<br />
1. Boyd, P.W., and D. E. Falk. 1990. (Personal communications).<br />
2. Franckowiak, J.D. (Personal communications).<br />
3. Jende-Strid, B. 1978. Mutation frequencies obtained after sodium azide<br />
treatments in different barley varieties. Barley Genet. Newsl. 8:55-57.<br />
4. Jende-Strid, B. 1984. Coordinator's report: Anthocyanin genes. Barley Genet.<br />
Newsl. 14:76-79.<br />
5. Jende-Strid, B. 1988. Coordinator's report: Anthocyanin genes. Stock list of ant<br />
mutants kept at the Carlsberg Laboratory. Barley Genet. Newsl. 18:74-79.<br />
6. Jende-Strid, B. 1991. Coordinator's report: Anthocyanin genes. Barley Genet.<br />
Newsl. 20:87-88.<br />
7. Jende-Strid, B. 1993. Genetic control of flavonoid biosynthesis in barley.<br />
Hereditas 119:187-204.<br />
8. Jende-Strid, B. 1993. Coordinator's report: Anthocyanin genes. Barley Genet.<br />
Newsl. 22:136-137.<br />
9. Jende-Strid, B. 1995. Coordinator's report: Anthocyanin genes Barley Genet.<br />
Newsl. 24:162-165.<br />
10. Jende-Strid, B., and K.N. Kristiansen. 1987. Genetics of flavonoid<br />
biosynthesis in barley. p. 445-453. In: S. Yasuda and T. Konishi (eds.) Barley<br />
Genetics V. Proc. Fifth Int. Barley Genet. Symp., Okayama 1986. Sanyo Press<br />
Co., Okayama.<br />
287
Barley Genetics Newsletter (2007) 37: 188-301<br />
11. Larsen, J., S. Ullrich, J. Ingversen, A. E. Nielsen, J.S. Gochan, and J. Clanay.<br />
1987. Breeding and malting behaviour of two different proanthocyanidin-free<br />
barley gene sources. p. 767-772. In S. Yasuda and T. Konishi (eds.) Barley<br />
Genetics V. Proc. Fifth Int. Barley Genet. Symp., Okayama. 1986. Sanyo Press<br />
Co., Okayama.<br />
12. Meldgaard, M. 1992. Expression of chalcone synthase, dihydroflavonol<br />
reductase, and flavanone 3-hydroxylase in mutants in barley deficient in<br />
anthocyanin and proanthocyanidin biosynthesis. Theor. Appl. Genet. 83:695-706.<br />
13. Ullrich, S., and J. Cochran. 1998. (Personal communications).<br />
Prepared:<br />
B. Jende-Strid. 1999. BGN 29:88-89.<br />
Revised:<br />
B. Jende-Strid and U. Lundqvist. 2007. BGN 37:286-288.<br />
288
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 617<br />
Locus name: Uniculme 4<br />
Locus symbol: cul4<br />
BGS 617, Uniculme 4, cul4<br />
Previous nomenclature and gene symbolization:<br />
Uniculme-5 = uc-5 (3).<br />
Inheritance:<br />
Monofactorial recessive (3).<br />
Located in chromosome 3HL (5), near AFLP marker E4143-4 in subgroup 32 of<br />
the Proctor/Nudinka map (5).<br />
Description:<br />
Plants produce 1 to 4 tillers that are twisted and have slightly bowed culm<br />
internodes. All secondary tillers are shorter than the primary tiller and have a<br />
curly appearance. Often secondary tillers are trapped at the base of the primary<br />
tiller (2, 4). Compared to normal sibs, cul4 plants have peduncles that are slightly<br />
to 50% longer. Rachis internodes are slightly elongated, and kernels are slightly<br />
longer. Plant height varies from 2/3 normal to slightly taller than Bowman. The<br />
mutant cul4.15 exhibits the most variation in height over environments (2). Under<br />
greenhouse conditions, Bowman line for cul4.5 developed only two axillary tillers,<br />
and it was uniculm when combined with the cul2.b (uniculm 2) gene (1).<br />
Origin of mutant:<br />
An ethylene oxide induced mutant in Bonus (PI 189763) (4).<br />
Mutational events:<br />
cul4.3 in Bonus (GSHO 2495, NGB 115062), cul4.5 in Bonus (NGB 115063),<br />
cul4.15 (NGB 115064) in Foma (CIho 11333), cul4.16 in Bonus (NGB 115065)<br />
(4).<br />
Mutant used for description and seed stocks:<br />
cul4.5 in Bonus (GSHO 2493, NGB 115063); cul4.5 in Bowman (PI 483237)*7<br />
(GSHO 2361).<br />
References:<br />
1. Babb, S., and G.J. Muehlbauer. 2003. Genetic and morphological<br />
characterization of the barley uniculm2 (cul2) mutant. Theor. Appl. Genet.<br />
106:846–857.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
4. Lundqvist, U. (Unpublished).<br />
5. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
Prepared:<br />
J.D. Franckowiak and U. Lundqvist. 2002. BGN 32:118.<br />
Revised:<br />
J.D. Franckowiak and U. Lundqvist. 2007. BGN 37:289.<br />
289
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 623<br />
Locus name: Eligulum-a<br />
Locus symbol: eli-a<br />
BGS623, Eligulum-a, eli-a<br />
Previous nomenclature and gene symbolization:<br />
Eligulum-a = lig-a (2).<br />
Eligulum-3 = eli-3 (4).<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Location is unknown.<br />
Description:<br />
Plants do not have ligules in the junction between the sheath and leaf blade,<br />
auricles are rudimentary and asymetrically displaced. Plants are about 2/3 of<br />
normal height and have very wide leaves (3, 4). The peduncle is short and spike<br />
emergence from the sheath of the flag leaf is poor. Spikes have a compact<br />
arrangement of spikelets and are extremely compacted near the tip (1, 3). The<br />
culm breaks very easily just below the nodes. The Bowman backcross-derived<br />
lines have glume awns that are nearly twice as long as those of Bowman, but the<br />
lemma awns are about 2/3 of normal (1).<br />
Origin of mutant:<br />
An ethylene imine induced mutant in Foma (CIho 11333) (2, 3).<br />
Mutational events:<br />
eli-a.2 (NGB 115389), eli-a.3 (NGB 115390), -a.7 (NGB 115392), -a.9 (NGB<br />
115393), -a.10 (NGB 115394) in Foma (3); eli-a.11 (NGB 115395), -a.14 (NGB<br />
115397) in Kristina (NGB 1500); eli-a.15 (NGB 115398), -a.16 (NGB 151399) in<br />
Bonus (PI 189763 (4), -a.216 (FN216) in Steptoe (CIho 15229) (1, 3).<br />
Mutant used for description and seed stocks:<br />
eli-a.3 in Foma (NGB 115390); eli-a.3 in Bowman (PI 483237)*3.<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
3. Kleinhofs, A. (Unpublished).<br />
4. Lundqvist, U. (Unpublished).<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 2002. BGN 32:126.<br />
Revised:<br />
J.D. Franckowiak and A. Kleinhofs. 2005. BGN 35:192.<br />
Revised:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:290.<br />
290
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 633<br />
Locus name: Many noded dwarf 6<br />
Locus symbol: mnd6<br />
BGS 633, Many noded dwarf 6, mnd6<br />
Previous nomenclature and gene symbolization:<br />
Densinodosum-6 = den-6 (3, 4).<br />
Inheritance:<br />
Monofactorial recessive (4).<br />
Located in chromosome 5HL [7L] (5), near AFLP marker E3743-3 in subgroup 65<br />
of the Proctor/Nudinka map (5).<br />
Description:<br />
Plants with the mnd6.6 gene are about 2/3 normal height and have many<br />
elongated internodes in each culm (1, 4). The number of elongated internodes<br />
can be up to 20 in the original stock when grown in Sweden. Kernels are thin and<br />
small (4). The number of tillers per plant is increased compared to normal sibs.<br />
Peduncles are very short, about 1/3 normal length, and awns are about 1/2<br />
normal length. Spikes are shorter with slightly over half the kernel number of<br />
Bowman. The Bowman backcross-derived line has 9 to 10 elongated internodes<br />
per tiller. Kernels of the Bowman mnd6 line are thinner and about 2/3 of normal<br />
weight (2). The grain yields of the mnd6 line are about 3/4 normal (2).<br />
Origin of mutant:<br />
An ethylene imine induced mutant in Bonus (PI 189763) (4).<br />
Mutational events:<br />
mnd6.6 in Bonus (NGB 114514) (4); mnd6.8 in Bonus (NGB 114516) (4, 5).<br />
Mutant used for description and seed stocks:<br />
mnd6.6 in Bonus (GSHO 1713), mnd6.6 in Bowman (PI 483237)*7 (GSHO<br />
2235).<br />
References:<br />
1. Bossinger, G., U. Lundqvist, W. Rohde, and F. Salamini. 1992. Genetics of<br />
plant developm ent in barley. p. 989-1017. In L. Munck, K. Kirkegaard, and B.<br />
Jensen (eds.). Barley Genetics VI. Proc. Sixth Int. Barley Genet. Symp.,<br />
Helsingborg, 1991. Munksgaard Int. Publ., Copenhagen.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson. 1969. A proposed<br />
system of symbols for the collection of barley mutants at Svalöv. Hereditas<br />
62:409-414.<br />
4. Lundqvist, U. (Unpublished).<br />
5. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
Prepared:<br />
U. Lundqvist and J. D. Franckowiak. 2002. BGN 32:134.<br />
Revised:<br />
U. Lundqvist and J. D. Franckowiak. 2007. BGN 37:291.<br />
291
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 636<br />
Locus name: Tip sterile 2<br />
Locus symbol: tst2<br />
BGS 636, Tip sterile 2, tst2<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (4).<br />
Location is unknown.<br />
Description:<br />
Spikes of tst2.b plants are 1/4 to 1/2 of normal length because seed set fails in<br />
the upper portion of the spike. Slow or poor development of the spike reduces<br />
both the number of rachis internodes and number of fertile spikelets (1, 4). Most<br />
spikes of the Bowman backcross-derived line set less than 10 seeds. Plants are<br />
shorter than are normal sibs because peduncles fail to elongate normally. Both<br />
rachis internode length and awn length are reduced in tst2 plants (1).<br />
Origin of mutant:<br />
An X-ray induced mutant in Donaria (PI 161974) (3, 4).<br />
Mutational events:<br />
tst2.b in Donaria (Mut. 2249, DWS1337) (2, 3).<br />
Mutant used for description and seed stocks:<br />
tst2.b in Donaria (GSHO 1781); tst2.b in Bowman (PI 483237)*5 (GSHO 2280).<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
3. Scholz, F. 1956. Mutationsversuche an Kulturpflanzen. V. Die Vererbung<br />
zweier sich variabel manifestierender Übergangsmerkmale von bespelzter zu<br />
nackter Gerste bei röntgeninduzierten Mutanten. Kulturpflanze 4:228-246.<br />
4. Scholz, F., and O. Lehmann. 1958. Die Gaterslebener Mutanten der<br />
Saatgerste in Beziehung zur Formenmannigfaltigkeit der Art Hordeum vulgare<br />
L.s.l.I. Kulturpflanze 6:123-166.<br />
Prepared:<br />
J.D. Franckowiak and U. Lundqvist. 2002. BGN 32:137.<br />
Revised:<br />
J.D. Franckowiak. 2005. BGN 35:193. (Locus symbol was changed from lin2.)<br />
J.D. Franckowiak. 2007. BGN 37:292.<br />
292
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 653<br />
Locus name: Brachytic 10<br />
Locus symbol: brh10<br />
BGS 653, Brachytic 10, brh10<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-l = brh.l (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 2HS (1), approximately 12.9 cM distal from SSR marker<br />
Bmac0850 in bin 2H-08 (1).<br />
Description:<br />
Plants are about 3/4 normal height and peduncles are over 3/4 normal length.<br />
Awns are about 3/4 of normal length. Rachis internodes are slightly shorter than<br />
those of normal sibs, but the number of fertile rachis nodes is increased by over<br />
2. Seedling leaves of brh10 plants are relatively short. Kernels of the Bowman<br />
brh10 line are shorter (7.3 vs. 9.6 mm) and about 20% lighter than those of<br />
Bowman. Plants have and erect growth habit and grain yields averaged 20% less<br />
than those of Bowman (1, 2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(5).<br />
Mutational events:<br />
brh10.l in Birgitta (17:15:2, DWS1007) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh10.l in Birgitta (GSHO 1677); brh10.l in Bowman (PI 483237)*7 (GSHO 2171).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:293.<br />
293
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 654<br />
Locus name: Brachytic 11<br />
Locus symbol: brh11<br />
BGS 654, Brachytic 11, brh11<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-n = brh.n (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 5HS [7S] (1), about 6.7 cM proximal from SSR marker<br />
Bmac0113 in bin 5H-04 (1).<br />
Description:<br />
Plants are 2/3 to 3/4 normal height and peduncles are 3/4 to 5/6 normal length.<br />
The length of the rachis internodes is about 3/4 as long as those of normal sibs.<br />
Seedling leaves of brh11 plants are relatively short. Kernels of the Bowman<br />
brh11 line are shorter (7.2 vs. 9.6 mm) and about 25% lighter than those of<br />
Bowman. Plants have an erect growth habit and grain yields of the brh11 line<br />
averaged less than 1/2 of those for Bowman (1, 2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(5).<br />
Mutational events:<br />
brh11.n in Birgitta (17:19:2, DWS1011) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh11.n in Birgitta (GSHO 1679); brh11.n in Bowman (PI 483237)*6 (GSHO<br />
2172).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:294.<br />
294
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 655<br />
Locus name: Brachytic 12<br />
Locus symbol: brh12<br />
BGS 655, Brachytic 12, brh12<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-o = brh.o (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 5HS [7S] (1), approximately 13.5 cM distal from SSR<br />
marker Bmag0387 in bin 5H-03 (1).<br />
Description:<br />
Plants are 2/3 to 3/4 of normal height. Awns and peduncles are about 3/4 normal<br />
length. The length of the rachis internodes is about 3/4 of normal sibs. Seedling<br />
leaves of brh12 plants are relatively short. Kernels of the Bowman brh12 line are<br />
shorter (7.9 vs. 9.6 mm) and about 20% lighter than those of Bowman. Grain<br />
yields of the brh12 line averaged slightly more than 1/2 of those for Bowman (1,<br />
2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(5).<br />
Mutational events:<br />
brh12.o in Birgitta (17:20:2, DWS1012) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh12.o in Birgitta (GSHO 1680); brh12.o in Bowman (PI 483237)*7 (GSHO<br />
2173).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:295.<br />
295
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 656<br />
Locus name: Brachytic 13<br />
Locus symbol: brh13<br />
BGS 656, Brachytic 13, brh13<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-p = brh.p (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 5HS [7S] (1), approximately 8.7 cM distal from SSR<br />
marker Bmag0387 in bin 5H-03 (1).<br />
Description:<br />
Plants are about 2/3 normal height and awns are about 1/2 normal length.<br />
Peduncles and leaf blades are about 2/3 and 3/4 normal length, respectively. The<br />
length of the rachis internodes is about 3/4 of that of Bowman. The spikelets at<br />
the tip of the spike are close together giving a fascinated appearance. Seedling<br />
leaves of brh13 plants are relatively short. Plants lodge relatively easily. Kernels<br />
of the Bowman brh13 line are about the same size as those of Bowman, but<br />
kernel weights are about 20% less. The brh13 plants have and erect growth habit<br />
and their grain yields are about 1/2 of those Bowman (1, 2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Birgitta (NSGC 1870, NGB 1494, NGB 14667)<br />
(5).<br />
Mutational events:<br />
brh13.p in Birgitta (18:02:4, DWS1013) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh13.p in Birgitta (GSHO 1681); brh13.p in Bowman (PI 483237)*6 (GSHO<br />
2174).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Lehmann, L.C. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:296.<br />
296
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 657<br />
Locus name: Brachytic 15<br />
Locus symbol: brh15<br />
BGS 657, Brachytic 15, brh15<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-u = brh.u (3).<br />
Inheritance:<br />
Monofactorial recessive (2, 3).<br />
Location is unknown (1).<br />
Description:<br />
Plants have numerous tillers with small leaves, spikes, and kernels. Prior to<br />
heading plants appear to be grassy culms similar to those produced by the sld2<br />
(slender dwarf 2) and sld4 (slender dwarf 4) mutants, but heading is not<br />
drastically delayed. Culms and peduncles are about 1/2 normal length. Awns and<br />
rachis internodes are slightly shorter than those of normal sibs. Leaf blades are<br />
narrow and about 1/2 normal length. Mutant plants headed 2 to 3 days later than<br />
normal sibs. No lodging was observed. Spikes of brh15 plants had nearly 4 fewer<br />
kernels than those of Bowman. Kernels of the Bowman brh15 line are slightly<br />
shorter (8.6 vs. 9.6 mm), thinner (3.4 vs. 3.8 mm), and about 30% lighter than<br />
those of Bowman. The grain yield of the brh15 line averaged about 2/3 of that<br />
recorded for Bowman (1, 2).<br />
Origin of mutant:<br />
A N-methyl-N-nitrosourea induced mutant in Julia (PI 339811) (5, 6).<br />
Mutational events:<br />
brh15.u in Julia (409 JK, DWS1156) (4, 6).<br />
Mutant used for description and seed stocks:<br />
brh15.u in 409 JK/Bowman (GSHO 1685); brh15.u in Bowman (PI 483237)*5<br />
(GSHO 2176).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Micke, A. and M. Maluszynski, M. 1984. List of semi-dwarf cereal stocks. In<br />
Semi-dwarf Cereal Mutants and Their Use in Cross-breeding II. IAEA-TECDOC-<br />
307. IAEA, Vienna.<br />
6. Szarejko I., M. Maluszynski, M. Nawrot, and B. Skawinska-Zydron, 1988.<br />
Semi-dwarf mutants and heterosis in barley. II. Interaction between several<br />
mutant genes responsible for dwarfism in barley. p. 241-246. In Semi-dwarf<br />
Cereal Mutants and their Use in Cross-breeding III. IAEA- TECDOC-455, IAEA,<br />
Vienna.<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:297.<br />
297
Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 658<br />
Locus name: Brachytic 17<br />
Locus symbol: brh17<br />
BGS 658, Brachytic 17, brh17<br />
Previous nomenclature and gene symbolization:<br />
Semidwarf mutant = Mo4 (5).<br />
Brachytic-ab = brh.ab (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 5HS [7L] (1), approximately 11.6 cM proximal from SSR<br />
marker Bmag0387 in bin 5H-03 (1).<br />
Description:<br />
Plants are about 3/4 normal height and awns are 5/6 of normal length. Peduncles<br />
are slightly shortened. Rachis internodes are about 20% shorter than those of<br />
normal sibs. Seedling leaves of brh17 plants are relatively short. Kernels of the<br />
Bowman brh17 line are shorter (7.7 vs. 9.6 mm) and about 20% lighter than<br />
those of Bowman. Lodging is reduced in the backcross-derived line and grain<br />
yields averaged slightly less than those of Bowman (1, 2).<br />
Origin of mutant:<br />
A sodium azide induced mutant in Morex (CIho 15773) (6).<br />
Mutational events:<br />
brh17.ab in Morex (Wa14355-83, Mo4, DWS1260) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh17.ab in Morex (GSHO 1669); brh17.ab in Bowman (PI 483237)*6 (GSHO<br />
2181).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Nedel, J.L., S.E. Ullrich, J.A. Clancy, and W.L. Pan. 1993. Barley semidwarf<br />
and standard isotype yield and malting quality response to nitrogen. Crop Sci.<br />
33:258-263.<br />
6. Ullrich, S.E., and Aydin, A. 1988. Mutation breeding for semi-dwarfism in<br />
barley. p. 135-144. In Semi-dwarf Cereal Mutants and Their Use in Crossbreeding<br />
III. IAEA-TECDOC-455. IAEA, Vienna.<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:298.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 659<br />
Locus name: Brachytic 18<br />
Locus symbol: brh18<br />
BGS 659, Brachytic 18, brh18<br />
Previous nomenclature and gene symbolization:<br />
Brachytic-ac = brh.ac (3).<br />
Inheritance:<br />
Monofactorial recessive (3, 4).<br />
Located in chromosome 5HS [7L] (1), approximately 9.2 cM distal from SSR<br />
marker Bmac0163 in bin 5H-01(1).<br />
Description:<br />
Plants are about 2/3 normal height and awns are less than 2/3 of normal length.<br />
Peduncles are slightly coiled and about 5/6 the length of those of normal sibs.<br />
Rachis internodes are about 20% shorter than those of Bowman. Seedling<br />
leaves of brh18 plants are relatively short. Kernels of brh18 plants are similar in<br />
weight to those of Bowman, but slightly shorter. Lodging is reduced, but grain<br />
yields averaged slight more than 1/2 of those for Bowman (1, 2).<br />
Origin of mutant:<br />
An induced mutant backcrossed into Triumph (CIho 11612, GSHO 2465) (5).<br />
Mutational events:<br />
brh18.ac in mo6/4*Triumph (402B, DWS1277) (4, 5).<br />
Mutant used for description and seed stocks:<br />
brh18.ac in Mo6/4*Triumph (GSHO 1670); brh18.ac in Bowman (PI 483237)*6<br />
(GSHO 2182).<br />
References:<br />
1. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak. 2005. Characterization<br />
and molecular mapping of genes determining semidwarfism in barley. J. Hered.<br />
96:654-662.<br />
2. Franckowiak, J.D. (Unpublished).<br />
3. Franckowiak, J.D. 1995. The brachytic class of semidwarf mutants in barley.<br />
Barley Genet. Newsl. 24:56-59.<br />
4. Franckowiak, J.D., and A. Pecio. 1992. Coordinator’s report: Semidwarf<br />
genes. A listing of genetic stocks. Barley Genet. Newsl. 21:116-127.<br />
5. Falk, D. 1985. (Personal communications).<br />
Prepared:<br />
J.D. Franckowiak and L.S. Dahleen. 2007. BGN 37:299.<br />
299
Barley Genetics Newsletter (2007) 37: 188-301<br />
BGS 660, Narrow leafed dwarf 2, nld2<br />
Stock number: BGS 660<br />
Locus name: Narrow leafed dwarf 2<br />
Locus symbol: nld2<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (1, 2).<br />
Location is unknown.<br />
Description:<br />
Mutant plants have narrow, dark green leaves, which are erect with welldeveloped<br />
midribs. Auricles degenerate to tiny projections, but ligules are<br />
normal. Stem internodes are short, and the upper ones are curved. Spikelets are<br />
relatively narrow and small, and seed set may be low. Kernels of the Bowman<br />
nld2 line are thinner (3.2 vs. 3.8 mm) and about 35% lighter than those of<br />
Bowman (1). Plants are 1/2 to 1/3 of normal height, the spike commonly emerges<br />
from the side of the sheath before anthesis. Awns of the nld2.b line are similar in<br />
length to those of Bowman. The nld2.b Bowman line is more vigorous than nld1.a<br />
in Christchurch, New Zealand and in North Dakota greenhouse nurseries, but<br />
nld1.a was more vigor in the Dundee, Scotland nursery. Seed yields are<br />
generally less than 20% of those of Bowman (1).<br />
Origin of mutant:<br />
A fast neutron induced mutant in Steptoe (CIho 15229) (2).<br />
Mutational events:<br />
nld2.b in Steptoe (2).<br />
Mutant used for description and seed stocks:<br />
nld2.b in Steptoe; nld2.b in Bowman (PI 483237)*6.<br />
References:<br />
1. Franckowiak, J.D. (Unpublished).<br />
2. Kleinhofs, A. (Unpublished).<br />
Prepared:<br />
J.D. Franckowiak and A. Kleinhofs. 2007. BGN 37:300.<br />
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Barley Genetics Newsletter (2007) 37: 188-301<br />
Stock number: BGS 661<br />
Locus name: Double seed 1<br />
Locus symbol: dub1<br />
BGS 661, Double seed 1, dub1<br />
Previous nomenclature and gene symbolization:<br />
None.<br />
Inheritance:<br />
Monofactorial recessive (2).<br />
Located in chromosome 5HL [7L] (2), near AFLP marker E4038-4 in subgroups<br />
66 to 67 of the Proctor/Nudinka map (2).<br />
Description:<br />
The modification of the top of spike is distinctive and occurs on all tillers. The tip<br />
of the spike is compacted and a few spikelets form two and three fertile florets<br />
adjacent to each other. The double spikelets have fused lemmas and paleas<br />
often enclose the part of two, occasionally more, flowers: six anthers and two<br />
ovaries (1). The tip of the spike appears phenotypically similar to those of int-m<br />
(intermedium spike-m) mutants (1).<br />
Origin of mutant:<br />
An X-ray and ferrisulfate induced mutant in Bonus (PI 189763) (1).<br />
Mutational events:<br />
dub1.1 (NGB 114331), dub1.2 (NGB 114332) in Bonus (1); dub1.3 (NGB<br />
114333), dub1.7 (NGB 114337), dub1.8 (NGB 114338), dub1.9 (NGB 114339),<br />
dub1.10 (NGB114340), dub1.11 (NGB 114341), dub1.12 (NGB 114342) in Foma<br />
(CIho 11333) (1); dub1.18a (NGB 114345), dub1.18b (NGB 114346, 114347) in<br />
Kristina (NGB 1500) (1); dub1.19 (NGB 114348), dub1.20 (NGB 114349,<br />
114350) in Bonus (1).<br />
Mutant used for description and seed stocks:<br />
dub1.1 (NGB 114331) in Bonus (2).<br />
References:<br />
1. Lundqvist, U. (Unpublished).<br />
2. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F. Salamini. 2003. Integration of<br />
a barley (Hordeum vulgare) molecular linkage map with the position of genetic<br />
loci hosting 29 developmental mutants. Heredity 90:390-396.<br />
Prepared:<br />
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:301.<br />
301