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Comparative Wood Anatomy of Atraphaxis taxa in
Turkey
Funda Erşen Bak a,* and Derya Cesur b
The wood anatomy of four Atraphaxis taxa that have natural distribution in
Turkey—Atraphaxis billardieri Jaub. & Spach, Atraphaxis billardieri subsp.
tournefortii (Jaup. & Spach) Lovelius, Atraphaxis spinosa L., and endemic
Atraphaxis grandiflora (Willd.)—were compared in this study. The wood
samples were sectioned according to standard techniques. Samples were
macerated with Schultze’s method. Tangential and radial vessel
diameters, intervessel pit diameters, vessel wall thickness, vessel
elements length, dimensions of libriform fibres (lengths, widths, cell wall
thickness and, lumen diameter), and uniseriate and biseriate ray heights
were measured, and the number of vessels per mm2, number of rays per
mm, and number of vessels per group were counted. The qualitative
features such as growth rings, vessel grouping, presence of helical
thickening and storied structure, vestured pits, type of perforation plate,
and arrangement of axial parenchyma were determined. These four
species of Atraphaxis shrubs differ in some wood characteristics such as
growth rings, vessel grouping, vestured pits, height and density of rays,
number of vessels per mm2, and the dimensions of the vessel.
Keywords: Wood Anatomy; Atraphaxis; Turkey
Contact information: a: Department of Forest Faculty, Artvin Çoruh University, Artvin, Turkey;
b: Graduate School of Natural and Applied Sciences, Artvin Çoruh University, Artvin, Turkey;
* Corresponding author: fundaersenbak@artvin.edu.tr
INTRODUCTION
The genus Atraphaxis L. (Polygonaceae) comprises 36 woody species across
Southern Europe, Northern Africa, and predominately Asia (Siberia, China, and Mongolia)
(Schuster et al. 2011; Yurtseva et al. 2014; 2016; Tavakkoli et al. 2015). There have been
many taxonomic studies (morphologic, palynology, and phylogenetic) on the genus
(Pavlov 1970; Haraldson 1978; Ronse Decraene and Akeroyd 1988; Ronse Decraene et al.
2000; Sanchez et al. 2011; Schuster et al. 2011; Sun and Zhang 2012; Tavakkoli et al.
2015; Yurtseva et al. 2016). However, few studies have been reported on the wood
anatomy of Atraphaxis genus (Jansen et al. 2001; Carlquist 2003; 2010). Jansen et al.
(2001) has mentioned A. spinosa (from Turkey) briefly in their study that vestured pits
were investigated, and their taxonomic and evolutionary significance was evaluated.
Similarly, species of A. pungens (M. Bieb.) Jaub. & Spach and A. frutescens (L.) Ewersm.
were partially mentioned in studies where wood diversity of Polygonaceae (Carlquist 2003)
was examined and Caryophyllales (Carlquist 2010) was evaluated in terms of ontogenetic,
phylogenetic, and ecologic factors.
According to floral characteristics (the numbers of tepal, stamen, style, and achene
type), the genus Atraphaxis is divided into two subgenera Euatraphaxis and Tragopyrum
(Pavlov 1970) or two sections Atraphaxis and Tragopyrum (Rechinger and SchimanCzeika 1968). The genus Atraphaxis is represented by 5 species in Turkey, one of which
Erşen Bak & Cesur (2021). “Atraphaxis anatomy,” BioResources 16(1), 835-845.
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is endemic (A. grandiflora Willd.). Four taxa (A. grandiflora Willd., A. angustifolia Jaub.
& Spach, A. billardieri Jaub. & Spach, and A. billardieri Jaub. & Spach subsp. tournefortii
(Jaub. & Spach) Lovelius) are indicated in the subgen. Tragopyrum, while one species (A.
spinosa L.) is categorized in the subgen. Atraphaxis (Cullen 1967). Although the only
Turkish record is the type specimen, the presence of Atraphaxis angustifolia in Turkey is
doubtful (Cullen 1967).
The goal of the present study is to describe and compare the anatomical
characteristics of wood in Atraphaxis billardieri, A. billardieri subsp. tournefortii, A.
grandiflora, and A. spinosa.
EXPERIMENTAL
Materials
All studied Atraphaxis taxa spread in arid areas and especially dry hillsides and
rocky slopes. A. spinosa is an erect shrub up to 1 m, A. grandiflora (˂ 30 cm) and taxa of
A. billardieri (≤ 60 cm) are low shrubs. All of them are deciduous. Plant samples were
collected from natural habitats in Turkey (Table 1).
Table 1. The Collection Data of Studied Samples
Taxon
Atraphaxis billardieri
Atraphaxis billardieri
subsp. tournefortii
Atraphaxis grandiflora
Atraphaxis spinosa
Locality
Antalya, Elmalı, Avlan Gölü, 1055m
363403N, 295657 E
Artvin, 400 m
411233N, 420203 E
Erzincan, Refahiye, 1750 m
395401N, 384532 E
Iğdır, Aralık, 823 m
395215N, 443019 E
Voucher Number
Erşen Bak 300
Erşen Bak 301
Erşen Bak 302
Erşen Bak & Cesur 9
Methods
Wood samples were obtained from the stem section and were boiled, and stored in
equal amounts of ethanol, glycerin, and distilled water (Merev 1998). Wood materials were
sectioned with a sliding microtome and cryostat. The sections were stained with safranine
O and alcian blue (Ruzin 1999; Ives 2001). Macerations were prepared using Schultze’s
method (Normand 1972). On sections, tangential and radial vessel diameters, intervessel
pit diameters, vessel wall thickness and, uniseriate and biseriate ray heights were measured,
and the number of vessels per mm2, the number of rays per mm, and the number of vessels
per group were counted. The vessel elements length, libriform fibre dimensions (lengths
and widths, the thickness of cell walls, lumen diameter) were measured on macerated
specimens. The qualitative features such as growth rings, vessel grouping, presence of
helical thickening and storied structure, vestured pits, type of perforation plate, and the
arrangement of axial parenchyma were determined. Also, vulnerability ratio (vessel
diameter/ numbers of vessels per mm2) and mesomorphy value (vulnerability ratio x vessel
element length) were calculated by using vessel member characteristics (Carlquist 1977,
1988). The average value for each feature was based on 30 measurements and/or counts.
Vessel diameter was measured on the basis of the lumen and was taken at the widest point.
Vessel element length, including its tails, was obtained from macerations (Baas et al. 1983;
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Carlquist 1988). Measurements and microphotographs were taken with an Olympus BX 53
microscope with DP73 (Tokyo, Japan). Helical thickening and vestured pits were examined
in detail using a Zeiss/Evo LS10 scanning electron microscope (Cambridge, UK). All wood
terms used conform to the usage of the International Association of Wood Anatomists
Committee on Nomenclature (IAWA Committee 1989).
RESULTS AND DISCUSSION
The wood anatomical properties of the Atraphaxis species are listed below. The
mean values of selected quantitative data are listed in Table 2.
Atraphaxis billardieri
Wood ring porous, growth ring distinct (marked by thick-walled, radially flattened
latewood fibres and vascular tracheids) (Fig. 1a). Vessels ca. 146 to 156/mm2, mostly
solitary (63%) and in usually tangential multiples of 2 to 3 (4 to 7), or in small radial groups
or in small clusters (Fig. 1a), mean number of vessels per group 1.65 (2 to 7); rounded in
cross-section, earlywood vessels tangential diameter 30 to 80 µm and radial diameter 30 to
83 µm, latewood vessels tangential diameter 10 to 26 µm and radial diameter 10 to 36 µm,
walls 2.28 to 5.54 µm in thickness. Narrow vessels intergrading with vascular tracheids in
latewood. Vessel member length 125 to 202 µm. Perforations simple in horizontal to
oblique end walls. Intervessel pits vestured (Fig. 2c), alternate, rounded or oval, 3.68 to
5.98 µm in diameter, with slit-like or coalescent apertures. Earlywood (±) and latewood
(+) vessels with helical thickenings, vessel members storied. Axial parenchyma scanty
paratracheal or vasicentric. Libriform fibres 314 to 491 µm long, 10.31 to 21.36 µm wide,
thin- to thick-walls (2.52 to 5.29 µm), simple pits on radial and tangential walls. Rays 1 to
6 / mm, mostly uniseriate (45 to 230 µm, 2 to 9 cells high) and biseriate (80 to 195 µm),
very rarely multiseriate (3 cells); heterocellular. Silica bodies were observed and darkstaining deposits in ray and axial parenchyma cells.
Atraphaxis billardieri subsp. tournefortii
Wood ring porous, growth ring distinct (marked by thick-walled, radially flattened
latewood fibres and vascular tracheids) (Figs. 1b, 1e). Vessels ca. 108 to 168 / mm2, solitary
(44%) and in usually tangential (predominantly at the beginning of growth ring) to radial
multiples of 2 to 4 (8 to 10), or in small clusters (Fig. 1e), mean number of vessels per
group 2.5 (2 to 10); rounded in cross-section, earlywood vessels tangential diameter 25 to
80 µm and radial diameter 35 to 85 µm, latewood vessels tangential diameter 15 to 35 µm
and radial diameter 10 to 35 µm, walls 3.26 to 9.01 µm in thickness. Narrow vessels
intergrading with vascular tracheids in latewood. Vessel member length 115 to 195 µm.
Perforations simple in horizontal to oblique end walls. Intervessel pits vestured (Fig. 2b),
alternate, rounded or oval, 4.8 to 7.96 µm in diameter, with slit-like or coalescent apertures.
Earlywood (±) and latewood (+) vessels with helical thickenings, vessel members storied.
Axial parenchyma scanty paratracheal or vasicentric. Libriform fibres 210 to 430 µm long,
11.62 to 23.24 µm wide, thin- to thick-walls (2.49 to 4.15 µm), simple pits on radial and
tangential walls. Rays 1 to 5 / mm, mostly uniseriate (45 to 325 µm, 2 to 11 cells high) and
biseriate (90 to 525 µm), very rarely multiseriate (3 cells); heterocellular. Silica bodies
(Fig. 2j) and dark-staining deposits were observed in ray and axial parenchyma cells.
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Atraphaxis grandiflora
Growth rings distinct. Wood diffuse to semi-ring porous (Figs. 1c, 1f). Vessels ca.
140 to 181 / mm2, in tangential and radial group of 2 to 4, or in clusters (5 to 9), or solitary
(48%) (Fig. 1f), mean number of vessels per group 2.1 (2 to 9); rounded to angular in crosssection, earlywood vessels tangential diameter 28 to 70 µm and radial diameter 29 to 70
µm, latewood vessels tangential diameter 10 to 23 µm and radial diameter 8 to 29 µm,
walls 2.28 to 4.96 µm in thickness. Vasicentric vascular tracheids present adjacent to in
vessel grouping. Vessel member length 105 to 220 µm. Perforations simple in horizontal
to oblique end walls. Intervessel pits vestured (Fig. 2a), alternate, rounded or oval, 3.84 to
7.03 µm in diameter, with slit-like or coalescent apertures (Fig. 2f). Wide (rare and very
slightly) and narrow (+) vessels with helical thickenings (Fig. 2f), vessel members storied
(Fig. 2h). Axial parenchyma scanty paratracheal or vasicentric. Libriform fibres 275 to 550
µm long, 11.18 to 45.32 µm wide, thin- to thick-walls (2.12 to 8.65 µm), simple pits on
radial and tangential walls. Rays 1 to 5 / mm, uniseriate (40 to 325 µm, 2 to 10 cells high)
and biseriate (130 to 430 µm), rarely multiseriate; heterocellular (Fig. 2i).
Table 2. Some Wood Anatomical Characteristics of Atraphaxis taxa
A. billardieri
A. grandiflora
A. spinosa
subsp. tournefortii
ETD
52.28
45.83
41.49
53.23
ERD
56.38
58.83
49.74
74.09
LTD
18.34
20.65
15.44
19.50
LRD
20.84
23.55
16.32
23.95
VF
150
128
155
185
VG
1.65
2.5
2.1
2.5
VEL
155.93
159.50
156.50
109.08
LFL
400.68
348.50
396.50
322.33
LFW
15.61
17.18
22.10
14.82
UH
115.31 / 4
118.95 / 4
130.11 / 4
127.92 / 6
BH
129.25
178.67
191.47
282.01
U
4
4
3
8
V
0.23
0.26
0.18
0.20
ME
36.71
41.42
28.74
21.44
ETD= earlywood vessels tangential diameter, m; ERD= earlywood vessels radial diameter, m;
LTD= latewood vessels tangential diameter, m; LRD= latewood vessels radial diameter, m; VF=
number of vessels per mm2;VG= mean number of vessels per group; VEL= vessel element length,
m; LFL= libriform fibre length, m; LFW= libriform fibre width, m; UH= uniseriate ray height, m
/ cell; BH= biseriate ray height, m; U= number of rays / mm; V= vulnerability ratio (mean tangential
vessel diameter / number of vessels per mm2); ME= mesomophy value (vulnerability ratio x vessel
element length).
Characteristics
A. billardieri
Atraphaxis spinosa
Growth rings distinct. Wood diffuse to semi-ring porous (Figs 1d, 1g). Vessels ca.
165 to 229 / mm2, in radial group of 2 to 4 (7 to 9), or in clusters, or solitary (42%) (Fig.
1g), mean number of vessels per group 2.5 (2 to 9); rounded to angular in cross-section,
vessels of two distinct diameter classes (Fig. 2g), earlywood vessels tangential diameter 32
to 87 µm and radial diameter 42 to 115 µm, latewood vessels tangential diameter 10 to 37
µm and radial diameter 10 to 40 µm, walls 1.96 to 3.99 µm thick. Vasicentric vascular
tracheids present adjacent to in vessel grouping. Vessel member length 75 to 137 µm.
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Fig. 1. Transverse sections. a: Atraphaxis billardieri, b, e: Atraphaxis billardieri subsp. tournefortii,
c, f: Atraphaxis grandiflora, d, g: Atraphaxis spinosa. a, b: Wood ring porous and growth ring
boundaries distinct, c, d: Wood diffuse to semi ring porous and growth ring boundaries distinct, e:
Vessels in tangential to radial multiples or small clusters, scanty paratracheal or vasicentric axial
parenchyma, f: Vessels in tangential and radial group, or in clusters, g: Vessels in radial multiples
of 2 to 4 or in clusters, axial parenchyma scanty paratracheal or vasicentric. Scale bars a-c= 200
µm, e-g = 100 µm.
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Fig. 2. Some qualitative characteristics. a, f, h: Atraphaxis grandiflora, b, ı, j: Atraphaxis billardieri
subsp. tournefortii, c: Atraphaxis billardieri, d, e, g: Atraphaxis spinosa. a-d: Vestured pits on vessel
walls, e: Helical thickening in earlywood vessel elements, f: Helical thickening in vessel elements
(left) and coalescent pit apertures (right), g: Two distinct vessel diameter classes in ring boundaries,
h: Vessel elements storied, i: Heterocellular uniseriate, biseriate and multiseriate rays; j: silica
bodies (arrow) in ray cells. Scale bars a, c, d=2 µm; b, e, f= 10 µm; g=50 µm; h, i= 100 µm.
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Perforations simple in horizontal to oblique end walls. Intervessel pits vestured
(Fig. 2d), alternate, rounded or oval, 4.99 to 6.23 µm in diameter, with slit-like or
coalescent apertures. Vessels with helical thickenings (Fig. 2e), vessel members storied.
Axial parenchyma scanty paratracheal or vasicentric, and scanty marginal (1 to 2 seriate,
initial). Libriform fibres 175 to 400 µm long, 8.3 to 21.58 µm wide, thin- to thick-walls
(2.49 to 5.81 µm), simple pits on radial and tangential walls. Rays 5 to 11 / mm, uniseriate
(58 to 225 µm, 3 to 13 cells high) and biseriate (110 to 470 µm), rarely multiseriate;
heterocellular.
Although growth ring boundaries were distinct in all taxa, it was hard to distinguish
latewood vessels in Atraphaxis spinosa. Narrow vessels and wide vessels were observed
mixed in growth rings, especially in the growth ring initial. Vessel dimorphism, in which
vessel diameter tends to be bimodal distribution (nearly), occurs in the Atraphaxis spinosa
(Figs. 1d, 1g, 2g). Vines and xerophytes often have vessels of two distinct diameter classes
(Baas and Schweingruber 1987; Carlquist 1988).
The mean numbers of vessels per group for these species are difficult to obtain
because earlywood vessels can be solitary or in small groups, while very narrow latewood
vessel groups (include vascular tracheids) can arrange wider patches. It is very difficult to
distinguish between very narrow vessels and vascular tracheids in cross-section. Vascular
tracheids occur at the end of the growth rings in all studied Atraphaxis taxa, and
intergrading vessels (Figs. 1e, 1f). Carlquist (1986; 1988) reported vascular tracheids are
formed at the end of the growth rings and do not tend to occur intermixed with vessels. In
addition, Baas and Zhang (1986) and IAWA Committee (1989) reported they also formed
around latewood vessel groups (in a vasisentric position). Because vascular tracheids that
lack perforation plate are resistant to embolism, these cells protect the cambium during the
dry season, in regions with water stress conditions at the end of the growing season.
Vascular tracheids that provide safety to the conductive system are tend to be seen in
drought-deciduous species.
A. spinosa has the shortest and the narrowest fibers among the studied taxa. There
were shorter vessel elements and more numerous vessels in A. spinosa than the other
Atraphaxis taxa (Table 2). According to Baas (1973), Baas et al. (1983), Baas and Carlquist
(1985), and Wheeler and Baas (1991), vessel elements length and vessel diameter decrease
from mesic vegetation towards xeric vegetation, and numbers of vessels in the unit area
and vessels grouping rate increase. The calculated mesomorphy values are low for A.
spinosa (21.44), A. grandiflora (28.74), A. billardieri (36.71), and A. billardieri subsp
tournefortii (41.42). A mesomorphy value of less than 75 indicates xeromorphy (Carlquist
1977; 1988). A. spinosa has more xeromorphic characteristics compared to the others
(Table 2). This species is spread in more arid fields. This value was calculated as 4.99 for
A. pungens depending on vessel member features given in Carlquist (2003). Vessel
dimorphism, vessel grouping, more numerous vessel, short vessel elements, and the
presence of vascular tracheids are for safety against air embolism and provide increased
safety rather than efficiency in the conductive system in drought taxa (Baas et al. 1983;
Carlquist 1988)
While the diagnostic value of vestured pits is limited (Nair 1998; Jansen et al.
1998), they can be employed in wood identification (Bailey 1933; Ohtani and Ishida 1976;
Merev 2003). The presence of vestured pits has been reported in Polygonaceae for some
genera and species (Jansen et al. 1998, 2001; Carlquist 2003). Jansen et al. (2001) and
Carlquist (2003; 2010) detected vestured pits in A. spinosa (from Turkey), and A. pungens
and A. frutescens, respectively. In the present study, vestured pits were photographed with
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SEM for A. spinosa, A. grandiflora, A. billardieri, and A. billardieri subsp. tournefortii
(Figs. 2a to 2d). The vestured pits examined in this study not only differ among the studied
taxa but also are different from the species examined in Carlquist’s studies (2003; 2010).
Therefore, these findings emphasize the potential significance of vestured pits in the
identification of Atraphaxis wood.
Carlquist (2003) stated that the width of multiseriate rays is two cells for most
members of Polygonaceae. In this study, multiseriate rays (≥ 3) are very rare. While
biseriate rays are as common as uniseriate rays in A. spinosa, uniseriate rays are more
common in A. billardieri, A. billardieri subsp. tournefortii, and A. grandiflora. The ray
heights (uniseriate and biseriate) of A. spinosa and A. grandiflora are greater than
subspecies of A. billardieri (Table 2). In A. spinosa and A. grandiflora, the mean height of
uniseriate rays is 127.92 and 130.11, respectively. In A. spinosa the mean height of biseriate
rays was found to be longest among studied taxa. For Atraphaxis pungens, the mean height
of uniseriate rays was 73, and the mean height of biseriate rays was 259 in Carlquist's study
(2003). The number of rays per mm in A. spinosa more numerous than the other taxa. Ray
frequency is more useful in wood identification than ray height because the latter varies
depending on ontogeny (Carlquist 1988).
In earlier studies, storied narrow vessels and associated axial parenchyma strands
were recorded for Calligonum arborescens, Antigonon leptopus, Dedeckera eurekensis,
Eriogonum deserticola, E. heermannii, Gymnnopodium antigonoides, Polygonum
lapathifolium (Carlquist 2003), and Calligonum polygonoides (Erşen Bak and Cesur 2020)
in Polygonaceae. Carlquist (2003) has not reported a storied structure for Atraphaxis
pungens, although this feature is relatively common in Polygonaceae. In this study, vessels
are storied all studied Atraphaxis taxa, as shown for Atraphaxis grandiflora (Fig. 2h).
Helical thickenings in earlywood and/or latewood vessels are reported for
Atraphaxis taxa (Figs. 2e, 2f). The presence of helical thickenings in vessels is reported for
Eriogonum fasciculatum, E. giganteum (Carlquist and Hoekman 1985), Atraphaxis
pungens, Bilderdykia multiflora, Eriogonum giganteum, E. heermannii, and
Muehlenbeckia astonii (Carlquist 2003) in Polygonaceae. Grooves interconnecting
(coalescent) pit apertures (Fig. 2f), which can be confused with helical thickening, were
observed especially in earlywood or wide vessel walls of studied Atraphaxis woods. These
grooves have been reported in vessels of Antigonon leptopus, Calligonum arborescens,
Coccoloba laurifolia, Eriogonum fasciculatum, E. kennedyi, and Polygonum lapathifolium
in Polygonaceae (Carlquist 2003).
CONCLUSIONS
1. These four species of Atraphaxis shrubs differ in some wood characteristics such as
growth rings, vessel grouping, vestured pits, height and density of rays, number of
vessels per mm2, and the dimensions of the vessel.
2. The properties such as vessel dimorphism, vessel grouping, vessel density, short vessel
member, presence of helical thickening, and tracheids, which provide safety in the water
conduction, are common in drought-deciduous species as Atraphaxis.
3. Vestured pits may be an important characteristic in identifying Atraphaxis wood.
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ACKNOWLEDGEMENTS
This study was financed by the Artvin Çoruh University Scientific Research
Projects Unit with the project numbered as 2013.F10.01.02. The authors thank Dilek
Öztekin and İbrahim Onkaş for assistance in obtaining the samples.
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PEER-REVIEWED ARTICLE
bioresources.com
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Article submitted: October 13, 2020; Peer review completed: November 21, 2020;
Revised version received and accepted: December 4, 2020; Published: December 10,
2020.
DOI: 10.15376/biores.16.1.835-845
Erşen Bak & Cesur (2021). “Atraphaxis anatomy,” BioResources 16(1), 835-845.
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