Plant Cell, Tissue and Organ Culture (PCTOC) 
https://doi.org/10.1007/s11240-023-02554-y
ORIGINAL ARTICLE
Plant fruit extracts enhance the in vitro propagation of cowpea (Vigna 
unguiculata) on Murashige and Skoog media
Grace Markin1 · John S. Y. Eleblu1,5  · Jacqueline N. Amissah2 · Samuel Reynolds1 · Charline Soraru3 · 
Melanie S. Craze3 · Emma J. Wallington3 · Alison R. Bentley3,4 · Eric Y. Danquah1,2
Received: 22 December 2022 / Accepted: 16 June 2023 
© The Author(s) 2023
Abstract
Cowpea (Vigna unguiculata) is a versatile legume with diverse nutritional and nutraceutical properties that serve as a food 
security and medicinal crop for millions of households across Africa. An efficient protocol was developed to propagate 
shoot tip and cotyledonary node explants from six cowpea breeding accessions in vitro on Murashige and Skoog (MS) basal 
media supplemented with either banana extract, coconut water, orange or tomato juice. Micropropagation performance was 
compared to MS medium supplemented with B5 vitamins. A total of 500 plantlets were obtained in vitro across treatments 
and MS basal media supplemented with tomato juice had the highest micropropagation performance (154 plantlets), followed 
by banana extract (112 plantlets), orange juice (107 plantlets), and coconut water (82 plantlets). Three accessions (AGRAC 
216, TA, and Asontem) were found to be the most amenable to in vitro propagation using plant-derived extracts. Overall, 
this study successfully established that plant-derived extracts can support in vitro cowpea propagation in the absence of 
synthetic plant growth regulators.
Key message 
in vitro propagation of cowpea using plant fruit extracts as cheap substitutes for synthetic plant growth regulators allows for 
large scale production and transformation of cowpeas against environmental stress.
Keywords Cowpea · Micropropagation · Plant-derived extracts · Shoot tip · Cotyledonary node
Introduction
Cowpea (Vigna unguiculata) is an annual, herbaceous, 
Communicated by Sergio J. Ochatt. warm-season crop that belongs to the Fabaceae family 
(Maréchal et al. 1978; OECD 2016). Globally, the total 
*  John S. Y. Eleblu 
 jeleblu@wacci.ug.edu.gh land area estimated for cowpea cultivation is about 12.3 
million hectares with an annual dry grain production of 
1 West Africa Centre for Crop Improvement, College of Basic about 7.2 million metric tons (FAOSTAT 2020a, b). Africa 
and Applied Sciences, University of Ghana, Legon, PMB accounts for 95.2% of the total acreage production, of 
LG 30, Accra, Ghana which Nigeria leads as the world’s leading producer and 
2 Department of Crop Science, College of Agriculture consumer (FAOSTAT 2020a, b). Cowpeas play an essen-
and Consumer Sciences, University of Ghana, Legon, 
P. Box. LG 44, Accra, Ghana tial role in most farming systems as a result of their ability 
3 to curb erosion, fix atmospheric nitrogen, and contribute  National Institute of Agricultural Botany (NIAB), 93 
Lawrence Weaver Rd, Cambridge CB3 0LE, UK to soil fertility via decay of its residues especially for sub-
4 sequent cereal crop rotations (Coba de la Peña 2012; Ron  International Maize and Wheat Improvement Center 
(CIMMYT), Texcoco, Mexico 2015). In terms of health, they are mainly cheap sources 
5 of dietary protein, relatively low in fat and rich in minerals  Biotechnology Centre, College of Basic and Applied 
Sciences, University of Ghana, P. O. Box LG 1195, ACCRA , and vitamins (Carneiro da Silva et al. 2018). Cowpeas can 
Ghana be boiled and eaten as whole meals, or ground into whole 
Vol.:(012 3456789)
 Plant Cell, Tissue and Organ Culture (PCTOC)
or composite flours for the preparation of baby foods, or propagation of clean cowpea planting materials and plant 
used as garnishes (Gómez 2003). Its leaves and green genetic transformation.
pods are used for treatment of diseases such as ulcers and 
measles among several others (Abebe and Alemayehu 
2022). Consequently, this legume has been endorsed as Materials and method
“high-quality proteins” with the sole purpose of decreas-
ing high incidences of nutritional malnutrition to “shift Plant material and culture conditions
the world unto a more sustainable path” (UNDP 2020). In 
Sub-Saharan Africa, millions of people rely on cowpea as Six (6) cowpea accessions (REC 64, Asontem, AGRAC 216, 
a principal component of their daily meals and is widely Tintinwa A, Tintinwa B, Songotra) were obtained from the 
cultivated by small-scale farmers (Enyiukwu et al. 2018; WACCI gene bank, University of Ghana for use in this study.
Langyintuo et al. 2003; Singh et al. 1997). One hundred and seventy-five (175) seeds per accession 
Recently, the supply of cowpea has been unable to meet were surface sterilized with 0.1% mercuric chloride and 
high consumer demands due to an inadequate supply of two drops of Tween-20 for 5 min, then rinsed three times 
clean (specifically disease-free) seeds as planting materials with sterile distilled water. The surface sterilized seeds 
for farmers, as well as limited mass propagation of improved were incubated for 1 week on the germination medium of 
cowpea cultivars for consumers. To curb these problems, Murashige and Skoog basal salts (Phytotechlab) and 30 g/l 
researchers worldwide have identified tissue culture tech- sucrose (Central Drug House) after which the shoot tip and 
nology as an efficient tool to enable the mass propagation cotyledonary node explants were excised and cultured on 
of improved cowpea cultivars and provide clean planting the different propagation media under a laminar flow hood. 
materials for use by farmers (Aragao and Campos 2007; A Completely Randomized Design (6 accessions × 5 treat-
Hussain et al. 2012; Sani et al. 2015; Suman 2017). ments) was used and 5 seeds per treatment were cultured and 
To upscale the production of cowpea using tissue culture, replicated six times.
especially in Sub-Saharan Africa, several challenges need Explants were transferred to different propagation media 
to be addressed, including the high cost of synthetic plant constituted of 100 ml/l of the blended and sieved plant-
growth regulators, delays in product importation, and inac- derived extracts (coconut water, banana extract, orange 
cessibility of these products in local markets. These serve juice, or tomato juice) prepared from fresh fruits in the labo-
as major drawbacks for laboratories with limited resources ratory (Table 1), sucrose, and Murashige and Skoog basal 
especially in developing countries (Datta et al. 2017). medium (Sigma). Murashige and Skoog supplemented with 
To address these barriers, we aimed to develop new proto- B5 vitamins and sucrose served as the control propaga-
cols for reducing costs without compromising on the quality tion medium. All prepared media was solidified with 6 g/l 
of cowpea propagules propagated (Datta et al. 2017; Klerk phytagel and adjusted to pH 5.8 ± 0.5 before autoclaving at 
et al. 2008). We tested the use of readily available, low- 121 °C for 20 min at 15 PSI. All culture conditions were set 
cost plant-derived extracts as substitutes for synthetic plant at a 16-h photoperiod with a temperature of 25 ± 1 °C and 
growth regulators and compared their propagative perfor- light intensity of 3000 Lumens.
mance. Here we report results from the in vitro propaga- After 14 days, well-rooted plantlets were removed from 
tion of six cowpea breeding accessions using shoot tip and culture vessels and any media adhering to them washed off 
cotyledonary node explants cultured on MS media supple- with tap water. They were then placed in 350 cc containers 
mented with four different plant-derived extracts (coconut containing sterile potting soil complete with additional fiber 
water, orange juice, tomato juice, and banana extract). The (Primasta) in the acclimatization chamber, and well misted 
objective of the study was to determine which plant-derived with water before being covered with propagator lids. Over 
extract best supports cowpea in vitro propagation and which the subsequent 2 weeks period, misting was reduced gradu-
genotypes and explant types was best-suited to in vitro prop- ally, and the chamber opened fully before plantlets were 
agation with organic additions. This can be used for mass transferred to the greenhouse in containers with dimensions 
Table 1  Description of plant- Plant-derived extract Degree of maturity Brix Range pH Range of extract
derived extracts
Banana (hybrid cultivar) Fully ripe with no brown 8.0–15.0 4.83–5.36
patches
Tomato (Pectomech) Fully ripe 2.4–3.6 3.97–4.24
Orange (Sweet orange) Fully ripe 7.4–10.3 3.78–5.1
Coconut Semi-matured 4.7–6.1 5.05–5.8
1 3
Plant Cell, Tissue and Organ Culture (PCTOC) 
of 30 cm × 20 cm × 25 cm for full maturation and production in vitro propagation on media supplemented 
of seeds. with plant‑derived extracts
Data collection and analysis Explants of the germinated accessions also responded differ-
ently to the MS media fortified with different plant-derived 
The days to shoot formation, number of shoots per explant, extracts. It was observed that full plantlet regeneration of 
shoot length, number of leaves per explant, days to root for- the various explants on MS medium supplemented with 
mation, root number per explant, root length, plant height, tomato juice and banana juice was obtained on the third day 
and stem girth were recorded daily between the hours of 7:00 of in vitro culture (Fig. 1). At the end of week one, explants 
and 16:00 GMT for two weeks after the culture of explants. cultured on MS medium supplemented with orange had 
All statistical analysis was conducted using the GenStat regenerated into full plantlets whiles those on MS medium 
Analytical Package Twelfth edition. The significant differ- supplemented with coconut lasted 14 days. It was observed 
ences between the observed means per plant-derived extract that injuries caused at excision on the shoot tip explant and 
treatment and genotype were determined using analysis of cotyledonary node explant of the six cowpea accessions 
variance (ANOVA) and Duncan’s Multiple range test and resulted in callus formation amidst the formation of roots 
where significant differences were present, the F-protected for the explants cultured on MS media supplemented with 
least significant difference (F-protected LSD) was used to coconut juice.
separate them.
Acclimatization of in vitro propagated cowpea 
plantlets
Results At the end of the experiment, 500 clean cowpea plantlets 
were fully propagated. These plantlets were placed in accli-
Varying responses of genotypes and explants matization chambers for two weeks. The acclimatization sur-
cultured on different media vival rate was 95%. Total humidity inside the chambers was 
maintained for 1 week after which it was gradually reduced 
The six accessions tested responded differently to in vitro before the transfer of plantlets to the greenhouse where they 
culture on the germination media. At the end of the first produced normal growth (Figs. 2, 3). The table below shows 
week, it was observed that five (AGRAC 216, REC 64, in further detail the genotypes, type of media, and percent-
Asontem, TB, and TA) out of the six accessions were ame- age of cowpea plantlets obtained from each of the explants 
nable to in vitro growth. At the end of two weeks, the rate of (Table 2).  
germination for the cowpea cultivar Songotra was 17.14%. The performance of each cowpea accession was deter-
See germination data in supplementary material. mined by comparing the survival rate of the various explants 
Fig. 1  Image of the cotyledonary node (left) and shoot tip explant (right) of the cowpea accession AGRAC 216 on the third day of culture on 
MS propagation media supplemented with tomato juice
1 3
 Plant Cell, Tissue and Organ Culture (PCTOC)
Fig. 2  Image of the acclimatized plantlets during the first week of acclimatization
Fig. 3  Image of the acclimatized plantlets before and after transfer to the greenhouse
Table 2  Percentage of in vitro Cowpea germplasm Micropropagation medium additive
propagated plantlets produced 
from Cotyledonary node (2A) Tomato juice (%) Banana Orange juice (%) Coconut MSB5 (%)
and Shoot tip (2B) explants extract (%) water (%)
(A)
 AGRAC 86.67 63.33 66.67 40.00 40.00
 TA 66.67 76.67 33.33 36.67 6.67
 TB 16.67 33.33 33.33 16.67 3.33
 ASONTEM 80.00 33.33 23.33 33.33 23.33
 REC064 26.67 3.33 26.67 NR NR
 SONGOTRA NR NA 26.67 NR 6.67
(B)
 AGRAC 100 30.00 60 36.67 30.00
 TA 83.33 46.67 40 56.67 3.33
 TB 13.33 50.00 6.67 20 NR
 ASONTEM 40.00 26.67 33.33 16.67 33.33
 REC064 NR 10.00 NR 6.67 NR
 SONGOTRA NA NR 6.67 10 3.33
in the MS media supplemented with different plant-derived explants were tested for micropropagation in each geno-
extracts against the control. The total survival rate = total type × media treatment. Thus, five (5) explants were cultured 
number of survived explants/ total number of explants cul- per dish and a total of six replicates for each explant. The 
tured) × 100%. NR equals no regeneration. A total of 30 following were observed:
1 3
Plant Cell, Tissue and Organ Culture (PCTOC) 
Overall, the cotyledonary node explant recorded the high- different propagation media respectively (Fig. 4). AGRAC 
est survival rate than the shoot tip explants cultured (see 216 cultured on MS media supplemented with tomato juice 
survived plantlets in supplementary material). In addition, recorded the highest plant height average for shoot tip and 
it is deduced from the analysed data that the shoot tip and TB did the same for cotyledonary node explants on MS 
cotyledonary node explants of AGRAC 216 were the most media supplemented with coconut water. Both explant 
amenable to growth in the MS media supplemented with types, however, recorded Songotra cultured on the control 
varying plant-derived extracts. Collectively, Tintiwa A came medium as having the least plant height average. TB and 
second after AGRAC 216 in terms of the total number of AGRAC 216 were the most efficient in terms of the number 
explants regenerated on the different media for both shoot of roots produced for the different explants type cultured on 
tips and cotyledonary node explants. This was followed by MS media supplemented with banana extract. There was no 
Asontem, Tintiwa B, Songotra, and Rec 064 for shoot tips, direct correlation observed for root number and root length 
with Rec 064 having greater micropropagation than Son- for explants of the various genotypes. The absent bars for the 
gotra for cotyledonary nodes (Table 2). genotypes indicates no regeneration (Fig. 4).
Additionally, significant differences for selected growth Furthermore, significant differences were observed for 
parameters number of leaves, number of shoots, plant height, the performance of the various propagation media based 
root number, and root length were observed between the on selected parameters (Fig. 5). Tomato juice recorded 
explants of the various cowpea accessions cultured on the the least significant difference for days to shoot formation, 
Shoot tip perfomance on banana extract Cotyledonary node performance on banana
propagation medium extract propagation medium
12
20 d
c
18 c 10 bc
16 c bc d c
14 8 bc b c
b c b12 6 bc b b
10 ab ab aa bc a c aa 4 b8 b ab b
6 b b a aa a a
4 c c cd a
d a 2
b
2 a a
b ab ab 0
0 AGRAC 216 Asontem TB TA REC O64-2
AGRAC 216 Asontem TB Rec 064 TA Cowpea lines
Cowpea accessions
Plant height Number of leaves Number of shoots Plant height Number of leaves Number of shoots
Root number Root length Root number Root length
Shoot tip performance on coconut water Performance of the cotyledonary node on 
propagation medium coconut water propagation media
10 c c 12
9 c
8 b 10 b
7 b
6 ab a 8 a a a
5 a a a a a
4 6 b
3 b b 4 ab a
2 a a a a a
1 2
0
AGRAC Asontem TB Rec 064 Songotra TA 0
216 AGRAC 216 Asontem TB TA
Cowpea lines Cowpea lines
Number of leaves Number of shoots Root number Plant height Root length
Fig. 4  Mean values for genotype performances on the different propa- cotyledonary node explants. All measurements in cm, error bars rep-
gation media based on the growth parameters plant height, root num- resent standard error of mean values
ber, root length, shoot number and shoot length of the shoot tip and 
1 3
Parameter means for cowpea lines Parameter means for cowpea lines
Parameter means for cowpea lines Parameter means for cowpea lines
 Plant Cell, Tissue and Organ Culture (PCTOC)
Shoot tip performance on MSB5 propagation Cotyledonary node performance on MSB5 
medium propagation media
8 14
7 b b
b 12
6 b 10 ab
5
a a 8 a4 a a b ca bc a
c
b a
3 6 b ab b
2 a4
b b a
c
1 b2 a
0
-1 AGRAC 216 Asontem Songotra TA
0
AGRAC 216 Asontem TB Songotra TA
-2
Cowpea Lines Cowpea lines
Plant height Root number Plant height Number of leaves Number of shoots Root length
Shoot tip performance on tomato juice Cotyledonary node performance on tomato 
propagation medium juice propagation medium
16 12 c bc
14 bc c 10 ab
12
b b
a
b a
10 8 c a
8 a 6 b bc ac a cc bc
6 b a b a ab b 4 b b b
ab
4 b a a
a a
2 2
0 0
AGRAC 216 Asontem TB TA AGRAC 216 Asontem TB Rec 064 TA
Cowpea lines Cowpea lines
Plant height Number of leaves Number of shoots
Plant height Root number Root length Root number Root length
Performance of the shoot tip on orange Performance of the cotyledonary node on 
juice propagation medium orange juice propagation medium
10
9 c 12c
8 c10 bc
7 abcab
6 c 8 a ab
c b c
5 ca c b b b 6 bc
4 cd cd b bcba
a b3 d 4
b
d a
a a
c c2 c c ab a a bca
1 a
b b 2
0 0
AGRAC 216 Asontem TB Songotra TA AGRAC Asontem TB Songotra REC 064 TA
216
Cowpea lines Cowpea lines
Plant height Number of leaves Number of shoots Plant height Number of leaves
Root number Root length Number of shoots Root number
Fig. 4  (continued)
days to root formation and the highest significant differ- highest difference for shoot number and tomatoes the high-
ence for plant height for both cotyledonary node and est for number of leaves for cotyledonary node explants. 
shoot tip explants. Meanwhile, banana juice displayed the MSB5 media displayed the highest significant difference 
1 3
Parameter means for cowpea lines
Parameter means for cowpea lines Parameter means for cowpea lines
Parameter means for cowpea lines 
Parameter means for cowpea lines Parameter means for cowpea lines 
Plant Cell, Tissue and Organ Culture (PCTOC) 
Media performances for the shoot tip explants 201
12
c d
202
10
8 203
c
6 d c d cd c
c b
b
c b
b b c a d a
b
4 a a d
b
a d a 204bc
bc b b a c2 a a205
0
Tomato Banana Coconut Control Orange 206
Propagaon Media 
Days to shoot formaon Days to root formaon Number of leaves per explan2t07
Plant height Root length Root number (cm)
208
Shoot number
209
Media performance for the cotyledonary node explant
9
d
8
c
7 c cc
6 b b
bc
d c
b c
5 c a c c c a
b b b a b4 aab a b
a a a
d
3 c b
a a
2
1
0
TOMATO CONTROL COCONUT BANANA ORANGE
Propagaon Media
Days to shoot formaon Days to root formaon Number of leaves per explant
Plant height Root length Root number
Shoot number
Fig. 5  Mean values of propagation media performances for the shoot height, e root number, f root length, g shoot number. All measure-
tip and cotyledonary nodes based on the growth parameters a days to ments in cm, error bars represent standard error of mean values
shoot formation, b days to root formation, c number of leaves, d plant 
for root length of the shoot tip explants however the oppo- efficiency for cotyledonary nodes. The absent bars for the 
site is observed for the cotyledonary node explants. MS performance parameter indicates no significant difference. 
media supplemented with orange juice displayed the least Furthermore, the variables on the error bars indicate sig-
significant difference for root length of shoot tips whiles nificant differences in performance for both genotype and 
MS media supplemented with Banana recorded the highest media.
1 3
Parameter means for the propagation media
Parameter means for the propagation media
 Plant Cell, Tissue and Organ Culture (PCTOC)
Discussion supplemented with banana extract came second to pro-
viding the best growth rate for in vitro shoot regenera-
Cowpeas are essential leguminous crops that contribute tion of Celosia spp. It was also deduced from the results 
immensely to the achievement of food security in devel- of the study that MS media supplemented with orange 
oping and underdeveloped countries (UNDP 2020). To juice induced both shoot and root formation for explants 
boost production and optimize the cost associated with in culture. This observation is confirmed by the findings 
the application of tissue culture tehnology for cowpea of Ubalua et al. (2015) who posited efficient shoot and 
improvement via in vitro culture, low-cost tissue culture root formation of cocoyam using orange juice. Finally, MS 
media options that allow the use of plant-derived extracts basal media supplemented with coconut water came forth 
as replacements for synthetic plant growth regulators are as the best plant extract that supported cowpea micropro-
employed (Akter et al. 2007; Datta et al. 2017). pagation due to delayed root formation of the cowpea gen-
The germination and in vitro propagation trends otypes. This observation could be explained by the very 
observed in the present study for the cowpea genotypes high levels of cytokinin and low amounts of auxins pre-
Rec 064, Songotra, and TB could presumably be due to sent in coconut water compared to the other plant-derived 
the general recalcitrance of legumes such as cowpea to extracts (Klerk et al. 2008; Kuraishi and Okumura 1961).
in vitro manipulation (Bakshi and Sahoo 2013; Somers Significant growth rate differences were observed for plant 
et al. 2003). height, root length, shoot number, shoot length and root num-
The cotyledonary node explants were reported to have ber of the various genotypes on the different supplemented 
the highest survival rate than shoot tip explants. This media. Explants' responses to the various in vitro propagation 
observation agrees with the findings of (Adesoye et al. media differed one from the other presumably due to differ-
2010; Chaudhury et  al. 2007; Raji et  al. 2008; Solleti ences in genetic compositions (Brar et al. 1999) and varied 
et al. 2008) who opined that cotyledonary nodes were the absorption rates of nutrients and hormones contained in the 
most efficient and gave the best results for multiple shoot media (Chee 1995; Drew et al. 1993). Gibberellins are respon-
induction. sible for the elongation of plants (Torres 1989) and all the 
All the plant-derived extracts supported the microprop- plant-derived extracts used in this study contain gibberellins 
agation of cowpea and this response could be explained by (Garmendia et al. 2019; Ge et al. 2007, 2008; Khalifah 1966; 
the amounts of vitamins, minerals, and plant growth regu- Radley and Dear 1958; Srivasta and Handa 2005). However, 
lators present in the extracts. Though “undefined” because the significant increase in plant height of shoot tip and coty-
the exact amount of each constituent is unknown and vari- ledonary node explants in the growth media supplemented 
able for each extract, these supplements are reported to with tomato juice and coconut water respectively suggests 
contain plant growth regulators coupled with some nutri- an increased concentration of gibberellins in both juices. The 
ents that support explant growth in plant tissue culture number of roots formed can be attributed to the high amount of 
systems (Caplin and Steward 1948; Saad and Elshahed auxin present in the extracts. The works of Hu et al. (2015) and 
2012). Majority of the time, the amount of auxin to cyto- Klerk et al. (2008)imply a higher number of auxins in bananas, 
kinin in plant culture media determines the type and extent hence the most efficient for the highest root numbers and root 
of organogenesis present in a culture (Saad and Elshahed length for the different explant types. From this study, MS 
2012; Skoog and Miller 1957). The performance of the basal media supplemented with tomato juice, banana extract, 
control media could be explained by the very low concen- or orange juice could be utilized in further cowpea tissue cul-
trations of plant growth regulators present in the medium ture works since they provided optimum growth conditions. 
(Gamborg et al. 1968; Murashige and Skoog 1962). Furthermore, protocols that involved the use of MS basal 
MS basal media supplemented with tomato juice was media supplemented with coconut water could be improved 
determined as the best and most efficient plant-derived further to provide optimum growth conditions for the cow-
extract that supported cowpea in vitro propagation. This pea genotypes in culture. In addition, AGRAC 216, TA, and 
observation agrees with the findings of Ayanlola et al. Asontem proved to have high amenability to growth in MS 
(n.d.) and Norhayati et al. (2011) who reported on the suc- media supplemented with plant-derived extracts. They could 
cessful regeneration of cowpea varieties Ife brown and be optimized for further tissue culture works on cowpea that 
TVU 943 and shoot regeneration of Celosia spp. respec- involves the use of plant-derived extracts.
tively on MS media supplemented with tomato juice. Fur-
thermore, MS media supplemented with banana extract 
was second in terms of shoot formation for the various 
cowpea genotypes. This is in line with the findings of 
Norhayati et al. (2011) who reported that MS basal media 
1 3
Plant Cell, Tissue and Organ Culture (PCTOC) 
Conclusion Open Access This article is licensed under a Creative Commons Attri-
bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long 
Cowpea can undergo successful in vitro propagation on as you give appropriate credit to the original author(s) and the source, 
MS media supplemented with varied plant-derived extracts provide a link to the Creative Commons licence, and indicate if changes 
by optimizing efficient tissue culture media and protocols. were made. The images or other third party material in this article are 
Based on regeneration efficiency in descending order, included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in 
tomato juice, banana extract, orange juice, and coconut the article’s Creative Commons licence and your intended use is not 
water best-supported cowpea growth in vitro. Based on permitted by statutory regulation or exceeds the permitted use, you will 
organ development, tomato juice—promoted early shoot need to obtain permission directly from the copyright holder. To view a 
and root development. Banana extract and orange juice copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
came second and third respectively followed by coconut 
water. Additionally, cowpea genotypes AGRAC 216, TA, 
and Asontem were most amenable to in vitro manipulation References
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