Review

Received: 12 November 2015 Revised: 11 January 2016 Accepted article published: 25 January 2016 Published online in Wiley Online Library: 1 March 2016

(wileyonlinelibrary.com) DOI 10.1002/jsfa.7644

Cowpea (Vigna unguiculata L. Walp), a renewed

multipurpose crop for a more sustainable
agri-food system: nutritional advantages
and constraints
Alexandre Gonçalves,a Piebiep Goufo,b Ana Barros,b
Raúl Domínguez-Perles,b Henrique Trindade,b Eduardo A S Rosa,b* Luis
Ferreiraa and Miguel Rodriguesa

Abstract
The growing awareness of the relevance of food composition for human health has increased the interest of the inclusion of
high proportions of fruits and vegetables in diets. To reach the objective of more balanced diets, an increased consumption
of legumes, which constitutes a sustainable source of essential nutrients, particularly low-cost protein, is of special relevance.
However, the consumption of legumes also entails some constraints that need to be addressed to avoid a deleterious impact on
consumers’ wellbeing and health. The value of legumes as a source of nutrients depends on a plethora of factors, including
genetic characteristics, agro-climatic conditions, and postharvest management that modulate the dietary effect of edible
seeds and vegetative material. Thus, more comprehensive information regarding composition, especially their nutritional and
anti-nutritional compounds, digestibility, and alternative processing procedures is essential. These were the challenges to write
this review, which focusses on the nutritional and anti-nutritional composition of Vigna unguiculata L. Walp, an emerging crop all
over the world intended to provide a rational support for the development of valuable foods and feeds of increased commercial
value.
© 2016 Society of Chemical Industry

Keywords: legumes; cowpea; nutritional content; anti-nutrients; digestibility

INTRODUCTION to fulfilling the growing requirements of dietary proteins for

During the last decade, legumes have emerged as an interest-
ing and balanced source of nutrients. In this sense, although
so far soybean has attracted the most attention worldwide,
field pea, common beans, faba bean, cowpea, spring vetch,
and lentils are also cultivated and consumed throughout the
world.1,2
From the most important cultivated legumes, cowpea (Vigna
unguiculata L. Walp) has shown several agronomic, environmen-
tal and economic advantages, contributing to further improve the
diets and incomes of peasant farming across Africa, Asia and South
America.3 – 5 Cowpea has its origin on the southern African region
but has spread and is now cultivated in more than 100 countries
between 40∘ N and 30∘ S latitudes.5 – 7 The environmental advan-
tage of cowpea arises from its ability to grow in semi-arid regions
with low input requirements.3,8 Due to its recognised nutritional
value (high protein and low fat content), which is related to the the
prevention of diverse metabolic and cardiovascular diseases, the
whole above-ground cowpea plant is a multipurpose crop, being
consumed for its leaves, green pods, green beans, mature beans,
or processed into paste or flour and used as a food ingredient.9 – 12
These positive traits have allowed global production to increase
from 1.3 to 7.0 million tons between 1981 and 2013,5 whilst, under
humans.
The occurrence of anti-nutrients such as phytic acid and protease
inhibitors has been stressed as a serious constraint for the dietary
utilisation of cowpea,13 – 15 requiring further development of pro-
cessing alternatives that may allow the production of foods with
lower impact on human health.
In this way, given its nutritional value as well as the reduced
environmental impact of the production systems, intense research
efforts must be redirected to the evaluation of nutrients and
anti-nutrients of cowpea, its digestibility and behaviour during
processing, as well as its potential contribution to human nutrition.
∗ Correspondence to: E Rosa, Centre for the Research and Technology of
Agro-Environmental and Biological Sciences, University of Trás-os-Montes and
Alto Douro, (UTAD-CITAB), Quinta de Prados, 5000–801 Vila Real, Portugal.
E-mail: erosa@utad.pt
a The Animal and Veterinary Research Centre, University of Trás-os-Montes and
Alto Douro, (UTAD-CECAV), Department of Veterinary Sciences, Quinta de
Prados, 5001-801 Vila Real, Portugal
b Centre for the Research and Technology of Agro-Environmental and Biological
Sciences, University of Trás-os-Montes and Alto Douro, (UTAD-CITAB), Quinta
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the current changing climate, cowpea is expected to contribute de Prados, 5000-801 Vila Real, Portugal

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Table 1. Amino acid composition of cowpea above-ground Table 2. Fatty acids composition cowpea seeds
material
Total fat
Seeds Leaves
Degree of ×10−3
Amino acid (g 16 g−1 N) Ref. (g 16 g−1 N) Ref.
Fatty acid unsaturation % g kg−1 DW Ref.
Isoleucine 3.9–4.5 27,31 – 38 9.8–11.1 39
Lauric acid C12:0 0.1–0.2 13–26 19,30,31,50,57
Leucine 7.5–7.8 17.9–19.6
Myristic acid C14:0 0.6–3.1 65–95
Lysine 3.5–7.9 10.3–16.3
Myristoleic acid C14:1 0.5–2.2 65–286
Methionine 1.1–3.5 2.9–4.5
Pentadecanoic acid C15:0 0.1–0.2 18–38
Phenylalanine 4.1–5.7 12.6–14.4
Palmitic acid C16:0 20.6–67.1 4 747–8 949
Tyrosine 3.4–4.5 6.5–9.3
Palmitoleic acid C16:1 0.2–3.6 18–26
Threonine 3.4–4.0 7.8–10.8
Heptadecanoic acid C17:0 0.4–0.8 65–133
Tryptophan 1.1–1.3 2.4–4.1
Stearic acid C18:0 2.9–14.0 780–1 273
Valine 4.5–6.2 11.5–12.8
Oleic acid C18:1 4.4–16.2 1 242–1 710
Aspartate 11.4–13.2 17.0–26.7
Vaccenic acid C18:1 0.2–0.4 16–57
Glutamate 16.5–18.8 24.3–45.3
Linoleic acid C18:2 20.8–40.3 2 821–5 562
Glycine 4.1–4.4 8.5–12.6
Linolenic acid C18:3 9.6–23.1 1 248–3 024
Alanine 4.2–4.5 5.8–9.8
Arachidic acid C20:0 0.1–1.9 195–323
Cysteine 0.6–2.4 1.0–2.9
Paullinic acid C20:1 0.3–0.7 54–95
Serine 4.0–5.2 11.4–11.6
Eicosadienoic acid C20:2 0.0–0.1 5–26
Histidine 2.7–3.4 6.6–8.6
Arachidonic acid C20.4 ≥1.3 –
Arginine 6.8–10.8 16.1–17.3
Behenic acid C22:0 0.1–3.0 324–513
Proline 3.1–6.2 10.4–15.9
Erucic acid C22:1 ≤0.9 ≤117
Information in bold text indicates the most essential amino acids in the cowpea. Docosadienoic acid C22:2 ≥0.7 –
Ligniceric acid C24:0 ≥0.2 –
Total monounsaturated fatty acids 7.5–24.0 1 512–2 291
Total polyunsaturated fatty acids 32.4–64.8 4 074–8 592
NUTRITIONAL COMPOSITION OF COWPEA Total saturated fatty acids 25.1–90.3 6 335–11 350
Macronutrients
Protein and amino acid content DW, dry weight.
With respect to the value of cowpea seeds as a source of proteins,
values in the range 203–394 g kg−1 have been reported, which
with the exception of soybean protein, is close to the content of acids, average 23.3, 35.6 and 55.2%, respectively, suggesting that
the major and under-utilised legumes.1,16 – 23 On the other hand, cowpea meals have the potential to cover human nutritional
cowpea leaves, which constitute an under-utilised vegetative part requirements.11,30
of this legume, compare well with cowpea seeds, with a protein It is important to point out that 5.0–37.0% of the total protein
content ranging from 215 to 437 g kg−1 .24 – 26 in cowpea (mainly globulins) has been reported to be nutritionally
Most of cowpea proteins consist of globulins (vicilins or unavailable.28,47 Nonetheless, bioactive peptides with antioxidant
7S globulins; 44.7–797 g kg−1 protein; 109 g kg−1 seed), fol- activity are successfully obtained from enzymatic proteolysis of
lowed by albumins (150–393 g kg−1 protein; 96 g kg−1 seed), cowpea proteins, indicating an interesting potential to be used as
glutelins (54–248 g kg−1 protein; 21 g kg−1 seed), and prolamins functional food ingredients.11,48,49
(13–131 g kg−1 protein; 18 g kg−1 seed).27 – 30 However, the rela-
tive content of the separate proteins is closely dependent on the
cultivar under consideration.28 Crude fat and fatty acids
Nutritionally, cowpea seeds protein shows a prevalence of glu- Recently, cowpea has been stressed on a low fat content
tamine/glutamic acid (16.5–18.8 g 16 g−1 N), asparagine/aspartic (31–304 g kg−1 ), comparatively to other legumes (chickpea,
acid (11.4–13.2 g 16 g−1 N), and phenylalanine + tyrosine split pea, lentil, green gram and lupine).18,19,22,44 This fact, accord-
(7.5–10.2 g 16 g−1 N), as well as a secondary prevalence of argi- ing to nutritional guidelines, suggests a potential application
nine, leucine, lysine, valine and proline.27,31 – 38 The same ranking in weight restriction diets.1,16,20,21 In addition, the lipid profile
is observed in leaves, even though cowpea leaf protein, based on of cowpea indicates a predominance of triglycerides (41.2% of
their amino acid content, is more than two-fold higher (Table 1) to total fat), followed by phospholipids (25.1% of total fat), mono-
that described for seeds (24.3–45.3, 17.0–26.7, and 19.1–23.7 g glycerides (10.6% of total fat), free fatty acids (7.9% of total fat),
16 g−1 N, for glutamine/glutamic acid, asparagine/aspartic acid, diglycerides (7.8% of total fat), sterols (5.5% of total fat), and
and phenylalanine + tyrosine, respectively).39 hydrocarbons + sterol esters (2.6% of total fat).50,51
When evaluating essential amino acids, diverse reports on With respect to fatty acids, palmitic acid is the predominant
this issue have shown that the limiting amino acids in cow- one (20.5–67.1% of total fatty acids), followed by linoleic
pea are methionine + cysteine, followed by tryptophan and (20.8–40.3% of total fatty acids), linolenic (9.6–30.9%), and
threonine,7,13,20,28,40 – 43 whilst cowpea is an excellent source of stearic acids (2.9–14.0%). The bulk of fatty acids consist of polyun-
lysine.44 – 46 The content of total amino acids in cowpea seed, the saturated fatty acids that range from 40.1 to 78.3% of total
ratio of essential amino acids relative to total amino acids, and the (Table 2).19,30,31,50 – 57 This high level of unsaturated fatty acids is a
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ratio of the essential amino acids relative to non-essential amino nutritionally desirable feature.36

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Nutritional aptitude of cowpea
Most plants of the Fabaceae family are reported to contain
𝛽-sitosterol as the dominant sterol, followed by stigmasterol,19
whilst in cowpea, stigmasterol is present in highest amounts
(40.8–43.3% of total sterols), followed by 𝛽-sitosterol (27.6-39.5%),
campesterol (8.6–15.1%), Δ-5 avenasterol, Δ-7 campesterol,
Δ-7 avenasterol, Δ-7stigmasterol, cholesterol, and brassicast-
erol (0.5–3.4%).50 – 52,58 Triterpenic acids such as oleanolic acid,
maslinic acid, and ursolic acid have also been reported in cowpea,
with contents of up to 3 × 10−3 g kg−1 .19,58
Carbohydrates and dietary fibre
In general, cowpea is characterised by a high proportion of carbo-
hydrates (504–658 g kg−1 ), being recognised as a potential source
of dietary fibre and resistant starches.1,19,59 Dietary fibre content in
cowpea ranges from 160 to 209 g kg−1 .60 In this regard, it has been
reported that pigmented varieties contain more than twice as
much fibre as unpigmented varieties.61 Furthermore, when com-
pared with other legumes, the highest values of starch content
and in vitro starch hydrolysis are usually observed with cowpea
(223–665 g kg−1 ).18,22,23,30,56,60
Eight sugars have been reported in cowpea, namely,
stachyose (17–60 g kg−1 ), sucrose (11–19 g kg−1 ), verbascose
(6–13 g kg−1 ), raffinose (5–10 g kg−1 ), glucose (4–5 g kg−1 ),
fructose (1–2 g kg−1 ), galactose (≤15 g kg−1 ), and maltose
(≤11 g kg−1 ).22,55,59,62 – 66 It is widely known that the oligosac-
charides stachyose, verbascose and raffinose are causative agents
of flatulence in humans, due to the lack of 𝛼-1,6-galactosidase in
the intestinal mucosa.67,68 This constitutes a constraint linked to
cowpea dietary consumption that requires further investigation
on new processing alternatives, contributing to reduce their
content and deleterious effects on the digestive system.
Micronutrients
Vitamins
Apart from a relevant source of essential macronutrients, cow-
pea constitutes an interesting source of micronutrients.18,69 – 72
The major vitamins present in cowpea are those belonging
to the B complex, being reported in the following decreas-
ing order: niacin (7.0–40.0 × 10−3 g kg−1 ) > panthothenic
acid (17.0–22.0 × 10−3 g kg−1 ) > thiamine (2.0–17.0 × 10−3
g kg−1 ) > pyridoxine (2.0–4.0 × 10−3 g kg−1 ) > folic acid
(1.0–4.0 × 10−3 g kg−1 ) > riboflavin (1.0–3.0 × 10−3 g kg−1 ) > biotin
(0.2–0.3 × 10−3 g kg−1 ) > cobalamin (traces).35,65,67,73 – 81 It is
important to mention that the segregation of cowpea varieties
into groups on the basis of the seed coat shows the ‘brown type’
with a higher content from B complex vitamins as compared with
the ‘black type’ and ‘blue-eye type’.82,83 The folic acid pattern has
revealed 5-methyltetrahydrofolate as the predominant folate form
in cowpea.70 This is, however, in contrast with Yarbaeva et al. who
found tetrahydrofolate as the most abundant B vitamin in Tajik
cowpea accessions.84 Concerning vegetative tissues, the partial
characterisation performed so far showed that the total folic acid
content in leaves was up to five-fold higher to those described in
seeds (3.3–20.1 × 10−3 g kg−1 ).85,86
Cowpea appear to be a particularly good source of vitamin C,
with levels in seeds and pods ranging from 52.0 to 554.0 × 10−3
g kg−1 ,33,65,67,69,76,87 – 89 and in leaves from 292.0 to 4100.0 × 10−3
g kg−1 .34,85,90 – 92 It is important to note that the total vitamin
C content in leaves at the maturity stages (4 and 6 weeks) is
mostly in the form of dehydroascorbic acid, an oxidation product
of L-ascorbic acid, which exhibits a lower biological activity.90
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Carotenoids, precursors of vitamin A, are also present in
cowpea contributing to the antioxidant compounds provided
by this legume. Among the carotenoids present in cowpea
seeds, lutein makes up over 70.0% (0.5 × 10−3 g kg−1 ).93 Other
carotenoids present in cowpea are 𝛽-carotene (0.056–0.183,
≤0.156, and 0.1–0.6 × 10−3 g kg−1 in seeds, pods and leaves,
respectively),69,74,76,85,87,92,94 𝛾-carotene (0.1 × 10−3 g kg−1 in
seeds), and cryptoxanthin (<0.1 × 10−3 g kg−1 in seeds).93
Lastly, from the various vitamin E vitamers present in cowpea,
𝛿-tocopherol has been observed with the highest concen-
tration (15.1–109.7 × 10−3 g kg−1 ), followed by 𝛾-tocopherol
(4.3–92.3 × 10−3 g kg−1 ), and 𝛾-tocotrienol (0.7–3.4 × 10−3 g
kg−1 ).27,50,51,95 The vitamin E composition of cowpea seems to
differ significantly from that of most legumes, where 𝛾-tocopherol
dominates.19,72
Mineral content
Studies aimed at assessing the mineral composition of cowpea
seeds have shown a wide variation depending on the specific
cultivars evaluated,32,36,40,46,56,67,96 whilst a gap of information on
the effect of diverse agro-climatic conditions is clearly noticed.
Thus, potassium is the most abundant mineral in seeds and
leaves, with contents ranging from 1.9 to 28.9 g kg−1 . Of partic-
ular importance are the high contents of calcium (383–10 620
and 398.0–8050.0 × 10−3 g kg−1 in seeds and leaves, respectively),
zinc (8.1–118.0 and 3.4–129.1 × 10−3 g kg−1 in seeds and leaves,
respectively), and iron (6.9–218.0 and 16.2–772.9 × 10−3 g kg−1 in
seeds and leaves, respectively).27,30,31,33,34,38,55,61,97 – 100 This mineral
composition emphasises the value of cowpea fulfilling the recom-
mended daily intake in a similar way as other legumes.18,23,37,44,84
Still, there is a dearth of information on the mineral composition
of cowpea pods, an under-exploited material from legumes in gen-
eral, and current data shows lower contents compared with seeds
and leaves.25,74,85,86,101 Thus, as an example, this difference regard-
ing the mineral composition of cowpea pods ralteive to sedes and
leaves has been demonstrated with respect to calcium and iron,
which have been reported in pods in concentrations of 0.60 and
0.04 g kg−1 , on average, respectively.87
PHYTOCHEMICAL COMPOSITION
AND ANTI-NUTRITIONAL FEATURES
The presence of some types of phenolic compounds in legumes
can be nutritionally troublesome since they might interact with
macro- and micronutrients, impairing their proper absorption
throughout the diverse phases of the digestion process.38,97 This is
particularly relevant when concerning proanthocyanidins, which
are often referred as ‘anti-nutrients’. However, other compounds
that affect the bioavailability of nutrients, including phytic acid and
enzyme inhibitors, also represent the anti-nutritional composition
of cowpea.
Proanthocyanidins
The anti-nutritional features of proanthocyanidins are largely
dependent on their conformation. The structural characteristics
of these compounds confer them ability to chelate metals and
bind to proteins, which, in turn, is closely related to the degree
of polymerisation. Thus, proanthocyanidin monomers and dimers
are mostly devoid of binding properties.102 The recent descrip-
tions on the phenolic composition of cowpea with respect to
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anti-nutritional compounds have shown that cowpea is unlikely
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Table 3. Anti-nutritional composition of cowpea seeds

Anti-nutritional factor Units Content (×10−3 ) Refs

Phenolic compounds
Total proanthocyanidins monomers g kg−1 827–3 104 19,25,121,146

Catechin g kg−1 17–297 19,25,121,146

Catechin-3-O-glucoside g catechin kg−1 ≤2 19,25,121,146

Catechin-7-O-glucoside g catechin kg−1 770–2 553 19,25,121,146

Epicatechin g catechin kg−1 ≤11 19,25,121,146

Epicatechin-3-O-glucoside g catechin kg−1 ≤0.3 19,25,121,146

(Epi)afzelechin-3-O-glucoside g procyanidin C1 kg−1 ≤243 19,25,121,146

(Epi)afzelechin-4′ -O-glucoside g procyanidin C1 kg−1 ≤243 19,25,121,146

Total proanthocyanidin dimers g kg−1 40–554 19,25,121,146

Total proanthocyanidin trimers g kg−1 140–633 19,25,121,146

Total proanthocyanidin tetramers g kg−1 120–241 19,25,121,146

Total proanthocyanidin pentamers g kg−1 17–223 19,25,121,146

Total proanthocyanidins hexamers g kg−1 93–769 19,25,121,146

Total proanthocyanidin heptamers g kg−1 ≤90 19,25,121,146

Total proanthocyanidin octamers g kg−1 ≤447 19,25,121,146

>10 proanthocyanidin polymers g kg−1 131–842 19,25,121,146

Enxyme inhibitors
Phytic acid g kg−1 2 640–15 240 13, 30, 31, 36, 40, 61, 62, 65, 98, 114--116

Trypsin inhibitor activity g trypsin inhibited kg−1 3 400–67 080 7,14,27,62,65,114,119,129,146

Chemotrypsin inhibitor activity g chemotrypsin inhibited kg−1 ≤1 600 47,61,121

𝛼-Amylase inhibitor activity g 𝛼-amylase inhibited kg−1 140–8 950 26,34,121

Other compounds
Haemagglutinating activity HUI kg−1 40 000–640 000 24,26,27,47,59,121,145

Cyanogenic glucosides g hydrogen cyanide kg−1 26–800 22,36,101,114

Oxalic acid g kg−1 420–7 700 1,36,77

3, 4-Dihydroxyphenylalanine g kg−1 10 000–23 000 30,65

Total saponins g diosgenin kg−1 2 510–37 160 1,12,91,104

to contain different types and amounts of proanthocyanidins
that might be nutritionally disadvantageous to humans. In this
concern, cowpea proanthocyanidins are mostly constituted by
monomers and low molecular weight oligomers (degree of poly-
merisation 2–4) (Table 3), unlike the typical proanthocyanidins
profile reported for most plant foods where oligomeric and poly-
meric units are dominant.103 – 107 This calls into question the clas-
sification of cowpea proanthocyanidins as ‘anti-nutrients’, which
was further supported by Plahar et al. and Chang et al., who
did not find a relationship between proanthocyanidin content
and mineral absorption/protein digestibility in cowpea.43,102 Addi-
tionally, diverse cowpea varieties have unusually high levels of
(epi)afzelechin glycosides at the expense of (epi)catechins,107
which might be beneficial in terms of improved micronutri-
ent bioaccessibility. On the other hand, additional studies have
pointed out the higher amounts of tannins in cowpea compared
with other legumes,108 as well as the fact that their capacity
for complexing with proteins, polysaccharides and other macro-
molecules contributes to the reduction of protein digestibility.109
Phytic acid
Phytic acid is the principal storage form of phosphorus, although it
is not bioavailable for humans. In the human gut, phytic acid low-
ers the bioavailability of minerals and limits the digestibility of pro-
teins and starch by inhibiting proteases and amylases. Specifically,
phytic acid is known to form complexes with the dietary essential
minerals calcium, zinc, iron and magnesium, making them biolog-
2944
vitamin niacin, potentially contributing to the deficiency disease
pellagra. Furthermore, phytic acid is associated with the devel-
opment of the hard-to-cook defect found in cowpea and other
beans.110 – 112 Still, some potential positive effects of phytic acid
have also been described. In this regard, phytic acid has been
shown to function as an antioxidant, as a hypocholesterolaemic
agent, and as a suppressor of iron-dependent oxidation.113
This phytochemical compound is a characteristic and abundant
constituent of legume seeds. The phytic acid content of cowpea
seeds,13,30,31,36,40,61,62,65,98,114 – 116 pods,87 and leaves91,117 is compa-
rable to that of many other legumes,12,18,22,23,59,64,97,103 ranging
from 2.6 to 15.2 g kg−1 and representing between 29.0 and 52.0%
of total inositol phosphates.18,76,118,119 Most of the phytic acid in
cowpea seeds is located in the cotyledons.120
Several studies indicate that cowpea contains more than 80.0%
of its inositol phosphates in the inositol hexaphosphate form,
while inositol pentaphosphate is the second predominant com-
pound belonging to this class (15.0%). The relative percent-
ages of inositol triphosphate and tetraphosphate are lower than
3.0%.60,103,121 – 123 The phytic acid output from cowpea leaves is
reported to be about two-fold lower in comparison with seeds,
which could indicate higher bioavailability of minerals in cowpea
leaves than in seeds.24
Enzyme inhibitors
Another important concern regarding the anti-nutritional char-
acteristics of cowpea is the presence of enzyme inhibitors. Pro-

ically unavailable for absorption. Phytic acid can also chelate the tease inhibition activities are essential in plant physiology because

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Nutritional aptitude of cowpea
of its contribution to plant resistance to pests. Due to its effi-
ciency, the trypsin inhibitor cowpea gene, CpTi, has been stud-
ied and successfully transferred to tobacco, rice, oil palm and
cotton plants.124 – 127 However, these compounds are also related
with a decreased activity of trypsin, chymotrypsin, and 𝛼-amylase
in the gastrointestinal tract, and have shown a strong variabil-
ity among diverse cultivars. For instance, Marconi et al. assessed
the trypsin and chymotrypsin inhibitor content in 22 cultivated
cowpea varieties from the International Institute of Tropical Agri-
culture germplasm bank and showed that the values changed
from minimums of 9.0 × 106 trypsin inhibited units per kg (TIU
kg−1 ) and 7.1 × 106 chemotrypsin inhibitor units per kg (CIU kg−1 )
to a maximum of 46.7 × 106 TIU kg−1 and 55.8 × 106 CIU kg−1 ,
respectively.128 The inhibition of these enzymes entails a negative
impact on amino acids, peptides, and free sugars availability after
ingestion. The evaluation of the trypsin inhibitor activity of cow-
pea seeds has allowed a negative correlation with the content in
proanthocyanidins to be shown, whilst it seems to be positively
correlated to phytic acid and haemagglutinin contents.129
Other anti-nutrients
Several other factors in cowpea have been related with major neg-
ative physiological effects due to their toxicity once in the blood,
like the presence of haemagglutinins,7,13,14,27,103,119,129 cyanogenic
glucosides,22,36,101,114 oxalic acid,1,36,76 3,4-dihydroxyphenylala-
nine,30,65 and saponins.1,12,91,104 With respect to these compounds,
cowpea does not seem to accumulate quantities that can be toxic
to humans. Nonetheless, decreasing or removing anti-nutrients
that lead to an improved nutritional quality should be pursued
for an efficient utilisation of cowpea in human nutrition.130 In
this regard, legumes in general, and particularly concerning cow-
pea seeds, are heated or hydrated before being consumed by
humans. These processes have been reported to reduce the level
of anti-nutritional factors.15,116
COWPEA DIGESTIBILITY
Cowpea production and its acceptance by consumers might be
affected by the combination of some biotic and abiotic con-
straints that affect the productivity of cowpea, the presence of
anti-nutritional factors such as protease inhibitors, lectin, phytic
acid, tannin, among others, and also the presence of indigestible
compounds inducing flatulence.5,131 Consequently, besides
overcoming the productivity constrains (e.g. viral, bacterial and
parasitic diseases) the human digestibility of cowpea must
be studied through different approaches (e.g. plant breeding,
post-processing methods and others), which in turn might help
improve cowpea nutritional value and, thus, a wider acceptance
by consumers.
Research aimed at breeding and enhancing cowpea cultivars
for early maturity, high yield, resistance to insects and diseases
have been pursued.132,133 The efforts put in by plant breeders
led to several high-yielding cultivars of cowpea being developed.
Nonetheless, the significance of this work may be amplified if these
cultivars are evaluated nutritionally, since new varieties might not
differ in their nutritional value from the traditional varieties.110,130
Cowpea seeds are characterised by a high proportion of car-
bohydrates, with starch being the main component and energy
source. Similar to other legumes, cowpea starch is more slowly
digested than starch from cereals, producing less abrupt changes
in plasma glucose and insulin upon ingestion, which is extremely
J Sci Food Agric 2016; 96: 2941–2951 © 2016 Society of Chemical Industry
www.soci.org
beneficial to human health. However, the long cooking times, the
presence of anti-nutrients, the beany flavour, and the presence of
oligosaccharides causing flatulence are frequent causes limiting
the use of whole cowpea dry seeds as food.130,134,135
The agronomic characterisation of different cowpea culti-
vars is pursued regularly while the chemical, nutritional and
anti-nutritional properties are often overlooked. A negative corre-
lation between the amounts of anti-nutrients described above and
the nutritional value of cowpeas has been shown.110,130 In fact, this
is observed specifically between the quantity of anti-nutritional
factors (phytic acid, tannins, and protease inhibitors) and the pro-
tein and starch digestibility, with negative correlation coefficients
available in some studies of cowpea and also other species.18,136,137
Thus, in breeding programmes, in vitro protein digestibility could
be an important parameter as a fast and reliable tool to evaluate
the nutritional value of food products and sub-products.
As mentioned before, although for human consumption cowpea
is grown primarily for its edible seeds, its leaves and green pods
are also harvested and consumed. Therefore, the nutritional value
of these products should also be considered. The nutritive value
of cowpea grains, leaves and haulms is very high. Cowpea leaves
shows similar nutritional values to those reported for other tropi-
cal leaf vegetables like Solanum nigrum, Amaranthus sp., Cucurbita
moschata and Ipomoea batatas, with the crude protein content of
cowpea ranging from 22.0 to 30.0% in the grain and leaves on a
dry weight basis and from 13.0 to 17.0% in the haulms.138,139 When
comparing cooked cowpea seeds and leaves, the latter presents
seven times more calcium and three times more iron.138 In addi-
tion, although the phosphorus content of cooked cowpea leaves
is 50.0% smaller than in cooked seeds, the phosphorus in leaves
is not present as phytic acid, thus there is a superior bioavailabil-
ity of minerals in cowpea leaves.140 Additionally, cowpea leaves are
good sources of 𝛽-carotene and ascorbic acid.138
As in cowpeas leaves, the nutritional composition and digestibil-
ity of cowpea pods, and the influence of different postharvesting
methods in these parameters, have been rarely studied. Deol and
Bains87 showed that in vitro protein digestibility of green pods
from the dual-purpose cowpea variety ‘CL-367’ (Punjab Agricul-
tural University, Ludhiana, India) picked at the 9th day after flow-
ering is 72.2%. This value is comparable to those reported among
raw cowpea seeds from other varieties, averaging 74.8%.130,141
These numbers, together with the data reporting smaller amounts
of anti-nutrients present in green-tissues, enlightens towards the
nutritional importance of these products as viable sources of pro-
teins and dietary fibre.
Another concern affecting cowpea’s acceptance by consumers
is the flatulence caused by the oligosaccharides stachyose, verbas-
cose and raffinoses, which are ubiquitous in legume seeds.67,68 To
overcome this constraint, food-processing techniques including
dehulling, boiling and soaking, have been evaluated in order
to determine their potential in reducing the fermentability of
indigestible carbohydrates in cowpea and thus reducing the
associated flatulence.131 Complementary to the breeding efforts,
several postharvest processing techniques to improve the nutri-
tional value of cowpea food-products, like the starch and protein
digestibility, have been investigated.
POSTHARVEST MODIFICATION
OF THE NUTRITIONAL VALUE OF COWPEA
The improvement of currently existing postharvest processing
2945
techniques including conventional storage and methods such as
wileyonlinelibrary.com/jsfa
 www.soci.org A Gonçalves et al.

drying, irradiation, dehulling, soaking, germination, fermentation,

Roasting

NS20 ; ↑37

↓73 ; NS37
cooking and roasting, may contribute to nutritionally enhance the

↓59,63
cowpea final products.87

NS37

NS37

NS37

NS37

↓73
↓89
↓95
NA

NA

NA
These processes have been shown to alter in varying degrees
the nutrients (Table 4) and anti-nutrients (Table 5).15 Haemag-

↓30,37,60,84,85,87,94,98,143 ; NS31,60,99 ; ↑60

glutinins, for example, are completely eliminated by boiling and

↓29,31,60,85,87,98 ; NS30,37,94,99,110
autoclaving, while proanthocyanidins and phytic acid are only

↓60,70,71,75,77,78,84 – 86,94,99 ; ↑86

partly affected.63,119 In order to overcome this constraint, gener-
ally, a combination of treatments is applied that allow greater

↓31,94,98 ; NS30,35,37 ; ↑20

Cooking
anti-nutrient losses than single treatments.14

↓59,60,62,63,68,120,146
The maximum reduction in proanthocyanidins has been

↓37,60,87 ; NS30,99

NS30,87,94,99 ; ↑60
described after dehulling, as compared with fermentation or

↓87 ; NS94 ; ↑85

↑30,37,60,87,99

↓85,87 ; NS94
cooking.62,68 Regarding the retention of phenolic compounds, dry

NS31,32,50
heated samples of cowpea showed the highest retention among

↓87,98
dry-heating, soaking and autoclaving treatments.142 When com-

NA
paring three cooking methods, it is observable a higher phytic

↓55,59,60,62,64,95,146
acid reduction in the boiling method, followed by autoclaving and

↓60 ; NS60 ; ↑24,55

Fermentation
microwave cooking.59
A reported optimum-water cooking method did not cause any

↓55,60 ; ↑55

↓95 ; ↑60,95
↓95 ; NS55

↓56 ; ↑56
↓60 ; ↑55
significant change in the mineral and riboflavin contents of cow-

↓55,60

↑55,60

NS92
pea, as compared with the in-excess-water method, which causes

↑20

↑92
↓95
a decrease in these nutrients.31,75

NS41 ; ↑17,18,37,67
↓18 ; NS41 ; ↑17,37
Postharvest processing may allow reducing the concentration

↓18 ; ↑17,33,37,67
NS35,37,41 ; ↑33

↓17,18,41 ; NS37
Germination

↓63,67,146 ; ↑17

↑17,33,67,69,89
of anti-nutrients in cowpea, however this reduction is frequently
accompanied by nutrient losses (Table 4 and Table 5).110,143

↑17,18,41

↑41,69,93
↑18,35,67
Processing method

↓17,41
Nonetheless, when done under controlled conditions, processing

NA

NA
can prevent damage, and even enhance the nutritional quality of
cowpea, causing for instance an improved bioavailability of some

↓59,62,63,66,68,120,146
compounds namely 𝛽-carotene.144 In a similar way, fermentation ↓13 ; NS37,98 ; ↑59
Soaking

can increase the total essential amino acids available.20

↓37,98 ; NS143

↓71,147 ; NS75
↓66,98 ; NS37
The consumption of cowpea is also challenged by the issues
of beany flavour, flatulence and a long cooking time; the latter NS66 ; ↑37
↓37,66
NS98

↑37

↓92

↓92
NA

NA
is either due to a hard-coat or to the hard-to-cook defect result-
ing from storing cowpea at elevated humidity and temperature.145

↓37 ; NS31,75 ; ↑13,18

↓35,75 ; NS35 ; ↑18,35

Steaming has been shown to reduce the beany flavour in cowpea,4
↓13 ; NS31,35,37
↓13,18,31 ; NS37
Dehulling

and fermentation was found to be the most single effective treat-

↓13,37 ; ↑18

ment in removing flatus-causing oligosaccharides, as compared

↓13,18,37
↓18,31,37

with soaking, cooking or germination.55,146

NS32

↓68
NA

NA

NA
NA
The seed coat appears to be the principal anti-cook factor in
cowpea.31 Steaming cowpea for some minutes, followed by drying
↓16,42 ; NS78,145

↓79 – 81 ; NS78

before storage has been reported to reduce the hardness of

Table 4. Effect of processing methodologies on cowpea’s nutrients

Storage

↓47 ; NS78

cowpea, although this process negatively altered the appearance

↓16,42,57
↓58 ; ↑58

of the cooked meal.4 Several alkaline solutions such as ash, ‘kanwa’,

↓57

↓57

↓92

↓92
NA
NA

NA

NA

potash, sodium bicarbonate, sodium hydroxide, lime, or calcium

↓, decrease; ↑, increase; NS, not significant; NA, not available.

hydroxide have also been used, with varying degrees of success,

Irradiation

to overcome the hard-coat and the hard-to-cook phenomenon

in cowpea.120,147,148 These treatments caused losses of vitamins,
↓65,81
↓123
↓65
↓65
↓65

↓65

↑65
↓65
NA
NA

NA

NA

NA

while no deleterious effect has been demonstrated on the protein

content (Table 4 and Table 5).75,78,120,149
Drying

In order to maximise the current processing and utilisation

↓91,92

↓91,92

of cowpea, recent studies have highlighted the effectiveness of

↑91
↑91
↓91

↓91
↓91

↑91

↓95
NA

NA

NA

NA

the response surface methodology and the principal compo-

nent analysis for selecting suitable combinations of processing
Harvesting

methods.17,66,87,90,98
↓24 ; ↑24

↑24,39

↓24,39
NS24

NS24

↓8 6
↓39

↑86
↑90
NA

NA

NA

NA

CONCLUSIONS AND FUTURE PROSPECTS

Oligosaccharides
Carbohydrates

Unravelling the chemical composition and nutritional properties

Amino acids
Compound

Fatty acids

of cowpea has been the subject of several studies. Cowpea has an

Vitamin A

Vitamin C
Vitamin B

Vitamin E
Minerals

Proteins

interesting composition in macro- and micronutrients, but also in

Fibre
Fats
Ash

anti-nutrients. The chemical composition together with the agro-

2946

nomical traits associated to this species (Vigna unguiculata L.), in

wileyonlinelibrary.com/jsfa © 2016 Society of Chemical Industry J Sci Food Agric 2016; 96: 2941–2951
Nutritional aptitude of cowpea www.soci.org

terms of resistance to biotic and abiotic stress, may constitute

Roasting

↓59,63,119
↓43,59,63
a valuable advantage in comparison with other legumes. How-

↓20 ; ↑43
ever, there is the need to update the currently available chemical

NA

NA

NA

NA

NA
↓63

↓63
data. Furthermore, the variability in the levels of cowpea phyto-

↓30,31,59,62,63,68,87,98,110,116,119,120,123,146,148
↓30,31,59,62,63,68,87,98,105,106,119,142,146,148,150
chemicals are dependent on the cultivar and the interaction with
the environment where it is grown, an area of research that has
received little attention.
A substantial amount of work has been done on cowpea seeds

↓60 ; ↑20,78,87,98,99,110,145,148
as a source of nutrients, but the information on cowpea leaves
Cooking

and pods is scarce. In this regard, it has been shown that properly
processed cowpea leaves and pods could provide a sustainable
and equivalent source of protein and vitamins compared with
seeds.
↓14,63,119

↑99,145
↓30,63
↓30,63

Cowpea could contribute valuably to fulfil the basic nutritional

↓150
NA

NA

requirements of humans; however, it provides relatively few calo-

ries, covering, on average, 8.0% of a 2000 kcal daily energy intake.19
↓59,60,62,95,122,146
Fermentation

↓59,62 ; ↑60,146
Since cowpea is usually eaten as a protein complement in cereal-
and tuber-rich diets, this should not have serious nutritional
↓60 ; ↑20

implications. In addition, cowpea flour as a supplement is able

NA

NA

NA
NA

NA

NA

NA

to provide additional vitamin A activity and zinc in cereal-based

↓17,18,63,103,115,116,119,146

weaning foods. Nonetheless, establishing the outcome of cowpea

flour supplementation is an area that requires further research.
Germination

↓18,63,119,146 ; ↑103

Traditionally, food-processing methods are primarily aimed at

NS41 ; ↑17,18,35

making food-products more palatable and improving their flavour.

↓63,103,119

However, new approaches like germination, heating treatment,

Processing method

↑18,41
↓103

soaking, dehulling, fermentation, among others, need to be opti-

NA

NA

NA
↓63

mised to increase the nutritional composition of cowpea seeds

↓13,14,59,62,63,68,98,119,120,146,148

and vegetative materials and their digestibility, whilst decrease the

↓13,59,63,68,98,116,119,120,146,148

anti-nutritional content.
Research studies showed that the accumulation of proteins,
Soaking

proanthocyanidins, haemagglutinins, phenolic compounds, or

NS98,148 ; ↑59,149

phytic acid in cowpea is genetically controlled. Coupled to its high

↓13,14,63,119

adaptability to drought and heat, the nutritional composition

↓14,63

makes cowpea an interesting candidate for genetic improvement

NA

NA

NA
↓13

↓63

programmes, which could ultimately lead to a higher absolute

↓13,18,31,43,68,151

amount of nutrients absorbed after dietary intake. In this sense,

↓13,18,31,68,116
Dehulling

it exists sufficient variability in cowpea germplasm for start-

Table 5. Effect of processing methodologies on cowpea’s anti-nutrients

NS35 ; ↑18

NS43 ; ↑18

ing breeding programmes focused on the nutritional aspects.

Nonetheless, human absorption studies with cowpea are rare, and
NA

NA
NA

NA
↓13

↑13

therefore are in need before such targeted breeding programmes

are initiated.
Storage

↓78,145

↑145

↓150

↓150
NA

NA

NA
NA

NA

↓57

ACKNOWLEDGEMENTS
Irradiation

↓65 ; NS123
↓123 ; ↑65

This work was supported by national funds from FCT - Portuguese

NS, not significant; NA, not available

Foundation for Science and Technology, under the projects

NA

NA

NA

NA
↓65
↓65

↓65
↑65

UID/AGR/04033/2013 and POCI-01-0145-FEDER-006958, and the

↓91,119,142
↓119 ; ↑91

European Project EROLEGUME (Seventh Research Framework

Drying

↓91,119

Programme of the European Union - FP7 research project n∘

NA
NA

NA

NA

NA

NA

↓91

613781). RDP was supported by a Postdoctoral contract from the

Portuguese Foundation for Science and Technology within the
Harvesting

framework of the National Scientific and Technologic System. The

authors declare no conflict of interests.
↑151
NA

NA
NA

NA

NA

NA
NA

NA

NA
Haemagglutinin activity
Chymotrypsin inhibitor

Cyanogenic glucosides

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