Heliyon 5 (2019) e01744
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Heliyon
journal homepage: www.heliyon.com
Review Article
Carbohydrates, proteins, fats and other essential components of food from
native trees in West Africa
Anne Mette Lykke a, Elie Antoine Padonou b, c, *
a
b
c
Department of Bioscience, Aarhus University, Vejlsøvej 25, DK 8600, Silkeborg, Denmark
School of Tropical Forestry, National University of Agriculture, BP 43, Ketou, Benin
Faculty of Agronomic Sciences, University of Abomey-Calavi, Laboratory of Applied Ecology, 01 BP 526, Cotonou, Benin
A R T I C L E I N F O
A B S T R A C T
Keywords:
Agriculture
Food science
Food analysis
Nutrition
Native tree foods contribute to food and nutrition security, health and income generation in sub-Saharan Africa.
However, the specific contribution of native tree foods to nutrition is poorly documented in science and often not
acknowledged in poverty reduction strategies. This review gives an overview on the content of carbohydrates,
proteins, fat, fibers, ash and dry matter of 98 native food tree species from sub-Saharan Africa. Data were grouped
according to the food providing organ (seeds, fruits and leaves). In general, seeds had high content of fat, protein
and dry matter; while leaves had high content of protein and ash. There was no significant difference between the
three organs on the content of fibers and carbohydrate. Some tree foods species were good sources to provide
carbohydrates, proteins, fat, fibers, ash and dry matter.
1. Introduction
In sub-Saharan Africa, it is well-known that indigenous trees traditionally contribute to food and nutrition security, health and income
generation (Hyacinthe et al., 2015; Otori and Mann, 2014; Stadlmayr et
al., 2013). The more specific contribution of food from native trees to
nutrition, however, is poorly documented in science and often not
acknowledged in poverty reduction strategies (Schreckenberg et al.,
2006; Ngome et al., 2017). Therefore, several trees may be considered for
food uses, but their nutritional value is underestimated (FAO, 2013).
Information on the nutrient composition of food is essential to estimate
adequate nutrient intake both at individual and group levels (Joyanes
and Lema, 2006). This information may facilitate the selection of priority
tree food species for domestication programs aimed at improving food
and nutrition security and income generation (Stadlmayr et al., 2013) as
well as for natural resource management and conservation.
Nutritional components generally analysed are carbohydrates, proteins, fat, fibers, ash, vitamins, minerals and dry matter. Carbohydrates
hold a special place in human nutrition providing the largest single
source of energy in the diet and satisfying instinctual desire for sweetness
(Brand-Miller, 2002). A high carbohydrate content is a major source of
readily available energy (Assogbadjo et al., 2012; Bamidele et al., 2015).
Proteins are fundamental elements for metabolism of enzymes, hormones
and many other molecules essential for life. Proteins are composed of 20
amino acids of which nine are essential and need to be provided through
the diet (histidine, isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tryptophan and valine). In early childhood a number of
amino acids, which are not essential in adults, cannot be formed in
adequate amounts. These are conditionally essential, because of the
limited ability of their endogenous formation relative to the magnitude of
demand (arginine, cysteine, glycine, glutamine, histidine, proline and
tyrosine) (Jackson, 2002). There may be disease situations during adult
life whereby a particular amino acid, or group of amino acids, becomes
conditionally essential (Jackson, 2002). Some tree food species are
promising as sources of dietary protein and amino acid supplement for
domestic and industrial use (Djenontin et al., 2009; Igwenyi and Akubugwo, 2010; Lohlum, 2010; Ayessou et al., 2014; Otori and Mann,
2014). Fats are a major source of energy and are the essential fuel for the
brain and growing fetus (Stubbs et al., 2018). They enhance flavour and
palatability of food and make an important contribution to health containing essential fatty acids that cannot be synthesized in the body and
are furthermore required for a range of metabolic and physiological
processes to maintain the structural and functional integrity of cell
membranes (Mann and Skeaff, 2002). High content of fat make certain
oils from native trees a good alternative or supplement to conventional
oil (Bazongo et al., 2014; Niyi, 2014). Fiber has many health benefits and
* Corresponding author.
E-mail address: padonouelie@gmail.com (E.A. Padonou).
https://doi.org/10.1016/j.heliyon.2019.e01744
Received 7 May 2019; Received in revised form 11 May 2019; Accepted 14 May 2019
2405-8440/© 2019 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
A.M. Lykke, E.A. Padonou
Heliyon 5 (2019) e01744
3.2. Seeds
can reduce diabetes (Bazongo et al., 2014) and high blood cholesterol
(Liu et al., 2000). Ash content is obtained by burning away of organic
materials and gives a measure of the contents inorganic minerals (Adubiaro et al., 2011; Amoo et al., 2012; Ayessou et al., 2014). High content
of dry matter in food is an advantage for high shelve-life because it
prevents microbial spoilage and pest attack during storage (Magaia et al.,
2013; Honfo et al., 2014).
The present review gives an overview of the content of carbohydrates,
proteins, fat, fibers, ash and dry matter of 98 native trees food species
from West Africa based on scientific literature.
The PCA performed on the proximate composition of seeds revealed
that the first three axes explained 77% of the variation in proximate
composition (Fig. 1). Fat (71%) and protein (56%) were positively
correlated with axis 1, while carbohydrate (-95%) was negatively
correlated with this axis. The group of species with high content of fat
were Irvingia gabonensis, Telfairia occidentalis, Sclerocarya birrea, Lophira
lanceolata, Balanites aegyptiaca, Pentaclethra macrophylla, Vitellaria paradoxa, Cola millenii. Ricinodendron heudelotii and Annona senegalensis (Appendix 2, Fig. 2). The species with highest content in protein were
Tetracarpidium conophorum, Balanites aegyptiaca, Ricinodendron heudelotii,
Pentaclethra macrophylla, Sphenostylis stenocarpa, Parkia biglobosa, Sclerocarya birrea, Lophira lanceolata, Sterculia africana and Boscia senegalensis
(Appendix 2, Fig. 2). The species with high content of carbohydrate were
Brachystegia nigerica, Diospyros mespiliformis, Saba comorensis, Mucuna
sloanei, Daniellia ogea, Cola pachycarpa, Afrostyrax lepidophyllus, Buchholzia coriacea, Detarium microcarpum and Olax subscorpioides (Appendix
2, Fig. 2).
Ash (72%) and fibers (75%) were positively correlated with axis 2
(Fig. 1). The species with high content of ash were Annona senegalensis,
Cola pachycarpa, Ricinodendron heudelotii, Scorodophloeus zenkeri,
Dichrostachys cinerea, Zanthoxylum zanthoxyloides, Irvingia gabonensis,
Xylopia aethiopica, Tetracarpidium conophorum and Parinari excelsa (Appendix 2, Fig. 2). The species with highest content of fiber were Scorodophloeus zenkeri, Sterculia africana, Acacia macrostachya, Acacia senegal,
Xylopia aethiopica, Dacryodes edulis, Annona senegalensis, Blighia sapida,
Adansonia digitata and Monodora myristica (Appendix 2, Fig. 2).
Dry matter (90.86%) was positively correlated with axis 3. The species with high content of dry matter were Cola acuminata, Saba comorensis, Sphenostylis stenocarpa, Tetracarpidium conophorum, Brachystegia
nigerica, Balanites aegyptiaca, Vitellaria paradoxa, Pentaclethra macrophylla, Sterculia africana and Irvingia gabonensis (Appendix 2, Fig. 2).
2. Methodology
The review was based on literature on tree food species from West
Africa. Species were selected based on floras, plant lists and books
(Akoegninou et al., 2006; Arbonnier, 2002; Matig et al., 2006; Codjia
et al., 2015; Awodoyin et al., 2015) with the following search terms:
“proximate composition”, “carbohydrate”, “protein”, “fat”, “lipid”,
“fiber”, “ash” and “dry matter” as these are the components of proximate
composition. For each variable considered, the reported values were
converted into the same units and synthetized in tables.
Data were grouped according to the food providing organ [seeds, fruit
(all other fruit parts besides of the seeds) and leaves] and analyzed using
ANOVA, Student-Newman-Keuls Test and Principal Component Analysis
(PCA) based on correlation. No data transformation was applied, as data
met the assumption of normality and homoscedasticity based on RyanJoiner test of normality and the Levene test for homogeneity of variances. All the analyses were performed using R statistical software (R
Core Team, 2013).
3. Results
3.1. Proximate composition of the organs
Seeds, fruit and leaves had significantly different proximate composition (Appendix 1). The highest protein content was found in seeds and
leaves, highest fat content in seeds, highest ash content in leaves and
highest dry matter content in seeds. There was no significant difference
between the three organs concerning the content of carbohydrate and
fibers.
3.3. Fruit
A positive correlation was found between dry matter (83%), carbohydrate (82%) and axis 1 (Fig. 3). Fibers (-62%) was negatively correlated with axis 1. Species with high content of dry matter were Saba
comorensis, Saba senegalensis, Dialium guineense, Detarium microcarpum,
Afraegle paniculata, Borassus aethiopum, Bridelia ferruginea, Dennettia tripetala, Adansonia digitata and Canarium schweinfurthii (Appendix 3,
Fig. 4). The Species with high content of carbohydrate were Anisophyllea
Fig. 1. Relation of the specific components of seeds based on PCA analysis (axis 1 vs axis 2 and axis 1 vs axis 3).
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A.M. Lykke, E.A. Padonou
Heliyon 5 (2019) e01744
Fig. 2. Species distribution of the proximate composition of the seeds based on PCA (axis 1 vs axis 2 and axis 1 vs axis 3). Acacmacr: Acacia macrostachya, Acacsene:
Acacia senegal, Adandigi: Adansonia digitata, Afrolepi: Afrostyrax lepidophyllus, Afzeafri: Afzelia africana, Afzebell: Afzelia bella, Annosene: Annona senegalensis, Balaaegy:
Balanites aegyptiaca, Bligsapi: Blighia sapida, Boscsene: Boscia senegalensis, Braceury: Brachystegia eurycoma, Bracnige: Brachystegia nigerica, Buchcori: Buchholzia coriacea, Colaacum: Cola acuminata, Colamill: Cola millenii, Colaniti: Cola nitida, Colapach: Cola pachycarpa, Dacredul: Dacryodes edulis, Daniogea: Daniellia ogea, Danioliv:
Daniellia oliveri, Detamicr: Detarium microcarpum, Dichcine: Dichrostachys cinerea, Diosmesp: Diospyros mespiliformis, Garckola: Garcinia kola, Irvigabo: Irvingia gabonensis, Landtogo: Landolphia togolana, Lannacid: Lannea acida, Lophlanc: Lophira lanceolata, Monomyri: Monodora myristica, Mucusloa: Mucuna sloanei, Olaxsubs: Olax
subscorpioides, Pachglab: Pachira glabra, Parkbigl: Parkia biglobosa, Pentmacr: Pentaclethra macrophylla, Pariexce: Parinari excelsa, Prosafri: Prosopis africana, Riciheud:
Ricinodendron heudelotii, Sabacomo: Saba comorensis, Sclebirr: Sclerocarya birrea, Scorzenk: Scorodophloeus zenkeri, Sphesten: Sphenostylis stenocarpa, Sterafri: Sterculia
africana, Tamaindi: Tamarindus indica, Telfocci: Telfairia occidentalis, Plukcono: Plukenetia conophora, Trecafri: Treculia africana, Tetrcono: Tetracarpidium conophorum,
Vitepara: Vitellaria paradoxa, Xyloaeth: Xylopia aethiopica, Zantzant: Zanthoxylum zanthoxyloides.
Fig. 3. Relation of the nutrition components of fruit based on PCA analysis (axis 1 vs axis 2 and axis 1 vs axis 3).
microcarpum, Treculia africana, Ximenia americana, Cordia sinensis, Grewia
betulaefolia, Dennettia tripetala, Carpolobia lutea, Cola pachycarpa, Tetrapleura tetraptera and Dacryodes edulis (Appendix 3, Fig. 4). The species
with high content of ash were Cola pachycarpa, Ficus sycomorus, Tetrapleura tetraptera, Saba comorensis, Mondia whitei, Sclerocarya birrea, Grewia betulaefolia, Parkia biglobosa, Canarium schweinfurthii and Anisophyllea
laurina (Appendix 3, Fig. 4).
Fat (-84.83%) was negatively correlated with axis 3 (Fig. 3). The
species with high content of fat were Canarium schweinfurthii, Ximenia
laurina, Carpolobia lutea, Dialium guineense, Adansonia digitata, Afraegle
paniculata, Saba senegalensis, Saba comorensis, Chrysophyllum albidum,
Balanites aegyptiaca and Cola pachycarpa (Appendix 3, Fig. 4). Species
with high content of fibers were Parinari curatellifolia, Lannea schimperi,
Sclerocarya birrea, Ficus sycomorus, Bridelia ferruginea, Ximenia Americana,
Vitellaria paradoxa, Gardenia erubescens, Detarium microcarpum and Borassus aethiopum (Appendix 3, Fig. 4).
Protein (74.92%) and ash (76.77%) were positively correlated with
axis 2 (Fig. 3). The species with high content of protein were Detarium
3
A.M. Lykke, E.A. Padonou
Heliyon 5 (2019) e01744
Fig. 4. Species distribution of the proximate composition of fruit based on PCA (axis 1 vs axis 2 and axis 1 vs axis 3). Adandigi: Adansonia digitata, Afrapani: Afraegle
paniculata, Anislaur: Anisophyllea laurina, Annosene: Annona senegalensis, Balaaegy: Balanites aegyptiaca, Boraaeth: Borassus aethiopum, Bridferr: Bridelia ferruginea,
Canaschw: Canarium schweinfurthii, Carplute: Carpolobia lutea, Chryalbi: Chrysophyllum albidum, Colapach: Cola pachycarpa, Cordsine: Cordia sinensis, Dacredul:
Dacryodes edulis, Denntrip: Dennettia tripetala, Detamicr: Detarium microcarpum, Dialguin: Dialium guineense, Ficusyco: Ficus sycomorus, Garckola: Garcinia kola, Garderub, Gardenia erubescens, Grewbetu: Grewia betulaefolia, Irvigabo: Irvingia gabonensis, Landhirs: Landolphia hirsuta, Landowar: Landolphia owariensis, Lannschi: Lannea
schimperi, Mondwhit: Mondia whitei, Paricura: Parinari curatellifolia, Parkbigl: Parkia biglobosa, Sabacomo: Saba comorensis, Sabasene: Saba senegalensis, Sarclati: Sarcocephalus latifolius, Sclebirr: Sclerocarya birrea, Scorzenk: Scorodophloeus zenkeri, Synsdulc: Synsepalum dulcificum, Syzyguin: Syzygium guineense, Tamaindi: Tamarindus
indica, Tetrtetr: Tetrapleura tetraptera, Trecafri: Treculia africana, Vitepara: Vitellaria paradoxa, Vitedoni: Vitex doniana, Ximeamer: Ximenia americana, Zizimaur:
Ziziphus mauritiana.
4. Conclusion
americana, Dacryodes edulis, Saba comorensis, Treculia africana, Saba senegalensis, Bridelia ferruginea, Tetrapleura tetraptera, Ficus sycomorus and
Chrysophyllum albidum (Appendix 3, Fig. 4).
A large number of native species from West Africa have good potentials for food and nutritional supplements. Carbohydrates and fibers
are found in equal amounts in seeds, fruits and leaves. Seeds are
important sources for both protein and fat. Leaves are important sources
for protein and minerals. Fruits generally have lower content of protein,
fat and minerals, but these components are still present.
Species with high amounts of protein and fat in seeds and fruits are
often low in carbohydrates. This is in contrast to leaves, where the species
with highest carbohydrate and fat content often are low in protein. It is
therefore important to focus more specifically on the qualities of each
food source in order to meet the particular local nutritional needs.
The species reviewed in this study are often directly available to local
communities, however they are often neglected in poverty and nutritional strategies. For instance Vitellaria paradoxa is well known for shea
butter production, but several species with higher fat content, such as
Irvingia gabonensis, Sclerocarya birrea and Balanites aegyptiaca, are rarely
use, and could have high potentials for improving food and nutritional
security. Only few of the species are widely known and exported, but
these few species have large economic potentials, eg Vitallaria paradoxa
and Elaeis guineensis. Many other species have similar properties, which
indicates a large unexploited potential from native species in West Africa.
For many species only few chemical analyses have been published
and some of the published data have been found doubtful during the
verification of data for the review. Therefore further analyses are needed
to verify the reported nutritional content and to test for substances that
could have toxic or pathogenic longterm effects that have not been
noticed based on the traditional uses.
Desertification and forest destruction in West Africa is likely to
eliminate nutritionally and economically valuable native species to give
place for less valuable crops, only because the value was not been realized outside particular communities. Greater sustainable use of seeds,
3.4. Leaves
Leave content of carbohydrate (77%), fat (58%), fibers (56%) and ash
(56%) were positively correlated with axis 1 (Fig. 5). The species with
high content of carbohydrate were Tamarindus indica, Vernonia amygdalina, Adansonia digitata, Cissus populnea, Telfairia occidentalis, Lecaniodiscus cupanioides, Gongronema latifolium, Grewia carpinifolia, Pterocarpus
soyauxii and Baphia pubescens (Appendix 4, Fig. 6). The species with high
content of fat were Cissus populnea, Gongronema latifolium, Ceiba pentandra, Vernonia amygdalina, Adansonia digitata, Ficus thonningii, Baphia
pubescens, Pterocarpus mildbraedii, Telfairia occidentalis and Afzelia africana (Appendix 4, Fig. 6). The species with high content of fibers were
Afzelia africana, Ficus thonningii, Sterculia tragacantha, Ceiba pentandra,
Albizia glaberrima, Pterocarpus santalinoides, Vitex doniana, Pterocarpus
mildbraedii, Grewia carpinifolia and Opilia amentacea (Appendix 4, Fig. 6).
The species with high content of ash were Opilia amentacea, Pterocarpus
mildbraedii, Maerua angolensis, Baphia pubescens, Sterculia tragacantha,
Adansonia digitata, Cissus populnea, Ceiba pentandra, Grewia carpinifolia
and Ficus thonningii (Appendix 4, Fig. 6).
Dry matter (67.38%) was positively correlated with axis 2. The species with high content of dry matter were Tamarindus indica, Ceiba pentandra, Grewia carpinifolia, Maerua angolensis, Vernonia amygdalina, Opilia
amentacea, Afzelia africana, Telfairia occidentalis, Pterocarpus soyauxii and
Lecaniodiscus cupanioides (Appendix 4, Fig. 6).
Protein (58.32%) was positively correlated with axis 3 (Fig. 5). The
species with high content of protein were Ficus glumosa, Maerua angolensis, Vitex doniana, Albizia glaberrima, Vernonia amygdalina, Pterocarpus
santalinoides, Blighia unijugata, Gongronema latifolium, Pterocarpus mildbraedii and Telfairia occidentalis (Appendix 4, Fig. 6).
4
A.M. Lykke, E.A. Padonou
Heliyon 5 (2019) e01744
Fig. 5. Relation of the specific components of leaves based on PCA analysis (axis 1 vs axis 2 and axis 1 vs axis 3).
Fig. 6. Species distribution of the proximate composition of leaves based on PCA (axis 1 vs axis 2 and axis 1 vs axis 3). Adandigi: Adansonia digitata, Afzeafri: Afzelia
africana, Albiglab: Albizia glaberrima, Baphpube: Baphia pubescens, Bligunij: Blighia unijugata, Bombbuon: Bombax buonopozense, Ceibpent: Ceiba pentandra, Cisspopu:
Cissus populnea, Ficuglum: Ficus glumosa, Ficuthon: Ficus thonningii, Gonglati: Gongronema latifolium, Grewcarp: Grewia carpinifolia, Hymeulmo: Hymenocardia ulmoides,
Lecacupa: Lecaniodiscus cupanioides, Maerango: Maerua angolensis, Myriarbo: Myrianthus arboreus, Opilamen: Opilia amentacea, Ptermild: Pterocarpus mildbraedii,
Ptersant: Pterocarpus santalinoides, Ptersoya: Pterocarpus soyauxii, Stertrag: Sterculia tragacantha, Tamaindi: Tamarindus indica, Telfocci: Telfairia occidentalis, Vernamyg:
Vernonia amygdalina, Vitedoni: Vitex doniana.
Funding statement
fruits and leaves by local communities could go hand in hand with nature
conservation.
This work was supported by Agropolis, Cariplo and Daniel & Nina
Carasso foundations via the TREEFOOD project.
Declarations
Author contribution statement
Competing interest statement
All authors listed have significantly contributed to the development
and the writing of this article.
The authors declare no conflict of interest.
Additional information
Supplementary content related to this article has been published
5
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Heliyon 5 (2019) e01744
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