Recommendation of Cowpea Genotypes Based On Adaptability, Stability and Grain Darkening
Recommendation of Cowpea Genotypes Based On Adaptability, Stability and Grain Darkening
Linda Brenna Ribeiro Araújo2*, Francisco Linco de Souza Tomaz3, Leslyene Maria de Freitas4, Cândida
Hermínia Campos de Magalhães Bertini 5, Júlio César DoVale 5
ABSTRACT - The cowpea is a legume that is widely grown in the north-east of Brazil, and which has been gaining ground in other
regions of the country. The main producer is t he state of Ce ará, with a large planted-area, albeit low productivity due to a lack of
producer technology and adapted cultivars. The aim of this study was to i dentify and r ecommend superior genotypes in terms of
adaptability and stability under rainfed and irrigated conditions, in addition to genotypes with reduced grain darkening. To this end, six
experiments were conducted in different districts of Ceará (Crato, Pentecoste, Crateús, Madalena, Bela Cruz and Limoeiro do Norte)
and one laboratory experiment, to evaluate grain darkening. The experimental design of the field trials was of randomised blocks,
with 14 genotypes and 4 re plications. The analysis of variance showed a significant effect from the genotypes and environments
and their interaction, so GGE Biplot analysis was carried out to eva luate adaptability and s tability. To evaluate grain darkening, a
completely randomised design was used in a simple factorial scheme with six previously selected genotypes and five different
storage times (0, 2, 4, 6 and 8 months). There was a significant effect from the genotypes and storage time. Genotype 1 showed the
least darkening, and can be recommended for environments to which it is best adapted (Crato and Crateús). Genotype 9 was considered
the most stable for grain yield, and can be more broadly recommended for the semi-arid region of the state of Ceará.
DOI: 10.5935/1806-6690.20250002
Editor-in-Chief: Eng. Agronomist - manoel.fi[email protected]
*Author for correspondence
Received for publication 20/01/2022; approved on 04/09/2023
*Corresponding author
1
This st udy was ext racted f rom t he doctor’s t hesis o f the le ad a uthor. T he research received the financial s upport of t he Co nselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq)
2
PhD in Agronomy and Plant Science, Federal University of Ceará (UFC), Fortaleza-CE, Brazil, [email protected] (ORCID ID 0000-0002-3554-3908)
3
PhD stu dent in Plant Sc ience, F ederal Rural U niversity o f the Semi-arid Re gion ( UFERSA), Mo ssoró-RN, Br azil, linc [email protected]
(ORCID ID 0000-0002-8696-8914)
4
Undergraduate student of Agronomy, Federal University of Ceará (UFC), Fortaleza-CE, Brazil, [email protected] (ORCID ID 0000-0003-3584-3756)
5
Department o f P lant S cience, F ederal University of Ceará (UFC), Fortaleza-CE, B razil, [email protected] (ORCID ID 0 000-0003-2949-5660),
[email protected] (ORCID ID 0000-0002-3497-9793)
L. B. R. Araújo et al.
Table 1 - Identification, origin and commercial subclass of the genotypes used in the study
Figure 1 - Locations of the value for cultivation and use (VCU) trials in the semi-arid region of the state of Ceará
Table 2 - En vironments of t he t rials a nd their r espective codes, so wing dat e, altitude, ge ographic coordinates, an d ra infall
accumulated during the tests
Grain brightne ss was ana lysed under stora ge Decomposition of th e mean s quare erro r of the
in genotypes s elected for adapt ability an d s tability, interaction i nto simple and complex parts was t hen
using the e xcluded specular re flection method with a estimated usi ng the expression p roposed b y Cruz and
colorimeter (Co loQuest XE , Hun terLab, Un ited Sta tes). Castoldi (1991). The mean va lues we re the n grou ped
The mo isture in the g rains wa s standa rdised at 12% using the Scott-Knott test at 5% probability. The analyses
using the low-t emperature ove n me thod (B razil, 2009) were carried out using the GENES software (Cruz, 2013).
at the Seed Analysis Laboratory (LAS) of the Federal The GGE-Biplot method was used to evaluate the
University of Ceará (U FC). The grain s were pac ked adaptability and stability of the genotypes, separating the
and sealed in 20 μm polyethylene bags. Each package rainfed a nd irriga ted e nvironments. The GG E method
contained 500 g of beans and was stored under ambient considers tw o sou rces of variation (G + G E) w ithout
conditions (25 ºC ± 5 ºC and 55% ± 15% ). The following separating the effect of the genotype and of the interaction
storage times were evaluated: Time I (harvest), Time II (Yan et al., 2007), as shown in the following equation:
(2 months), Time III (4 months), Time IV (6 months)
and Time V (8 months). Yij - - B j = Yi1 j1 + Yi 2 j2 + ijk ( 3)
Data analysis where: Yijk is the mean gra in yield of g enotype i in
environment j; μ is the ove rall mea n; Bj is the effect of
After verifying the normality of the data for grain environment j; γi1 and αj1 are the main scores of genotype i
productivity and hom oscedasticity of the va riances, and environment j, respectively; γi2 and αj2 are the secondary
individual and joint analyses of variance (ANOVA) were scores of genotype i and environment j, respectively; εijk is
carried out. In the individual analyses, the adopted model was: the residue not explained by any of the effects.
Yij = + Gi + B j + ij
1)( The GGE Biplot g raphs we re gene rated by
where: Yij is t he ph enotypic value o f geno type i in the simple dispersion o f γi1 a nd γ i2 for the geno types
block j; μ is the overall mean; Gi is the effect of the ith and αj1 and αj2 for the environments using singular value
genotype; B j is the effect of the jth block; ε ij is the error decomposition, as in the following equation:
associated with the ith genotype in the jth block. Yij - - Bj = 1 i1 j1 + 2 i2 j2 + ijk 4)
(
In order to identify possible genotype x environment where: λ1 and λ2 are the largest eigenvalues for p rincipal
interactions, a joint analysis of variance was carried out as components 1 a nd 2 (P CA1 and PCA2), respectively; ξ1
per the following model: and ξ2 are the e igenvalues of ge notype i for PCA1 and
Yijk = + Gi + A j + Gaij + PCA2, respe ctively; η1 and η2 are the eigenvalues of
( 2)
environment j for PCA1 and PCA2, respectively.
ijk
(1992), and a hea tmap was ge nerated to visual ise under ev aluation. However , t he accuracy is consider ed
the performa nce of the genot ypes in the d ifferent high ( Resende, 2 002), d enoting t he high re liability of the
environments. T he analyses we re carrie d out using recommendation process.
the metan packa ge (Olivoto ; Lúcio, 2020) of the R
The g enotype x e nvironment inte ractions w ere
software (R CORE TEAM, 2017).
complex for grain yield (Figure 2). A different genotype
The assumptions were met for the brightness data, and ranking can be s een in the e nvironments un der ev aluation,
an analysis of variance and mean value test (Scott-Knott) were which were classified as favourable or unfavourable for
carried out for the six most adapted and/or stable genotypes. cultivation based on Annicchiarico (1992). This method helps
The ANOVA was carried out in a simple 6 x 5 factorial scheme to identify stable genotypes, which should have low sensitivity
(6 genotypes x 5 storage times). to unfavourable e nvironments (Pe reira et al., 2009). The
environments Crato, Pentecoste, Crateús and Bela Cruz were
classified as favourable, while Madalena and Limoeiro were
RESULTS AND DISCUSSION unfavourable. In the trials, grain yield ranged from 279.4
to 2250.4 kg ha -1 for Ge notypes 13 (BR S-Tumucumaque)
The joint analysis of variance showed a significant and 4 (Bico-de-ouro 1-5-24), respectively.
difference between the genotypes (p < 0.05) and
environments (p < 0.01), as wel l as fo r the genotype x Complex genotype x environment interactions are
environment interaction (p < 0.01) (Table 3). This shows more challenging for breeders (Evangelista et al., 2021)
that the genotypes had different behaviours for grain yield, as they make broader recommendations difficult. Due
and that this varia ble was a lso influenced by the grow th to these interactions, the a nalyses of a daptability and
environments, i n addition to the re being an int eraction stability ar e even more i mportant for sele cting and
between these factors. Similar results were obtained when recommending th e eva luated s trains m ore pre cisely
evaluating the yield of cowpea genotypes in different (Cruz et al., 2021). The GGE B iplot method becomes
environments in Bra zil and in other areas of prod uction highly rele vant in thes e c ases, as it af fords greater
(Abiriga et al., 2020; Araméndis-Tatis; Cardona-Ayala; precision when making the selection (Cruz et al., 2020).
Espitia-Camacho, 202 1; Cruz et al., 2020; Melo et al., The two principal components generated by the GGE
2020; Sousa et al., 2017; Tomaz et al., 2022). analyses (PC1 and PC2) explained 89.47% and 86.94% of
The significant G x E i nteraction explains the the total variation in the grain yield data for the irrigated and
evaluations of adaptability and s tability (Araméndiz-Tatis; rainfed environments, respectively (Figure 3). This shows
Cardona-Ayala; Espi tia-Camacho, 2021), espe cially that the bip lots repr esented th e existi ng interactio ns well,
when it com es to a com plex tr ait such a s grai n yield the components becoming even more representative as the
(Abiriga et al., 2020). F urthermore, the predominance of cultivation systems separated. Th e first two components
the complex part of the G x E interaction confirms that the capture less info rmation as th e environmen ts under
behaviour of the strains varied greatly in the environments evaluation increase (Tomaz et al., 2022).
Table 3 - Summary of the joint variance analysis, the simple and complex parts of the genotype x environment interaction, and accuracy
for grain yield (kg ha-1) in 14 cowpea genotypes evaluated in six locations in the state of Ceará
Figure 2 - Heatmap, grouping of average yield values (kg ha -1), and the c lassification as per Annicchiarico (1992) for the different
environments
Source: prepared by the author. Gen: genotypes. Mea n values followed by the same uppercase letter on a line and/or the same lowercase letter in a
column are part of the same grouping by Scott-Knott test at 5% probability
Figure 3 - GGE Biplot for the trials in the irrigated and rainfed environments
Source: prepared by the author. A and E: Genotype ranking; B and F: Mean vs. stability; C and G: Who wins where; D an d H: Di scrimination vs.
representation. A, B, C and D: Irrigated trials. E, F, G and H: Rainfed trials
In the GGE Biplot (G + GE), the first principal The GGE Bi plot method is also able to class ify
component (PC1) correlates grain yield with the effect the e nvironments under analysis based on their
of th e ge notype, while t he second compone nt (PC2) representativeness and ability to discr iminate b etween
summarises the sour ces of variation that lead to lar ge genotypes. The length of the environment vector in relation
differences i n the yi eld of t hese genot ypes between to PC1 is lin ked to its d iscriminating ability (Cruz et al.,
locations, repr esenting t he genot ype x envi ronment 2020; Yan et al., 2000; Yan; T inker, 2006). Figures 3-D
interaction (Cruz et al., 2020; Yan et al., 2000). and 3-H show that Crateús (E3) and Pentecoste (E2) were
therefore the mo st discriminating amo ng th ose evalua ted
In Figure 3, graphs A and E sh ow the ranking
for ir rigated and rain fed c ultivation, re spectively. Th ese
of the genotypes, where the arrow in the centre of the
environments were also t he most r epresentative, d ue
concentric ci rcles m arks th e sa me distance between
to t he s maller a ngle formed wi th t he a xis of th e a verage
the origin and the longest vector of the environments,
environment (Tomaz et al., 2022). Environments that form
representing the ideal genotype. This genotype would
acute angles to each other show a positive correlation, while
have high gr ain yield in all the environments under
environments whose axes form obtuse angles are negatively
evaluation (Melo et al., 2020; Y an; T inker, 2006).
correlated (Yan; Tinker, 2006). These results corroborate the
As such, individuals lo cated clo sest to this ideoty pe,
classification made b y Annicchiarico, since the correlation
such as Genotypes 4 and 10 for irrigated en vironments
between favourable and unfavourable environments, such as
and Genotypes 9 and 2 f or rainfed environments, ha d the
E1 with E5 or E2 with E6, was small or non-existent. Melo
highest grain pr oductivity a nd sta bility i n t hese locations.
et al. (2020) and Tomaz et al. (2022) al so fo und p ositive
The different results for environments with different water
and negative correlations between environments in cowpea
regimes highlight the importance of separating the analyses
cultivation in the state of Ceará when using the GGE Biplot
of adaptability and stability for these production systems.
method. It is, therefore, essential to id entify genotypes for
Graphs B a nd F (F igure 3 ) a re entitled ‘M ean vs . each condition due t o the significant differences between
Stability’ and show that genotypes wit h greater projection the environments under evaluation. Furthermore, the design
on the PC2 axis had lower stability albeit higher productivity of genetic improvement programs for the cowpea must take
in the envir onments clo sest to their p osition. The arro w this into account, developing strategies that enable efficient
indicates the coordinate of the average environment, which recommendations to be made.
is the point of greatest stability (Cruz et al., 2020).
There was a significant difference in grain
Genotype 10 was s table when i rrigated, but did not darkening u nder storage b etween the s ix g enotypes
have the highest average, while genotype 13 stood out in the selected for their adaptability and stability (1, 2, 4, 8, 9 and 10)
Bela Cruzand Crateús environments (greater adaptability), but (p < 0.01). Storage times were also significant (p < 0.01),
did not have the same performance in the Crato environment, albeit wi th no inte raction. T here was a reduc tion in t he
showing less stability. Genotype 10 was also considered stable brightness (L*) o f all the ge notypes over time; however,
when evaluated in the state of Rio de Janeiro under different Genotype 1 showed less darkening compared to the other
soil and climate conditions (Cruz et al., 2020). genotypes, with no difference in brightness between
Under rainfed conditions for example, Genotype six and eight mo nths of storage, and di ffering from the
8 sho wed st able b ehaviour in ea ch of th e thre e others stati stically in the final evaluation (Figure 4). In
environments, but also did n ot s tand ou t i n te rms o f the colorimetric analysis, brightness is the most important
productivity. St able genotypes s how s imilar behaviour variable for detecting grain darkening, as it evaluates the
over a range of environments, while adapted genotypes lightness of the colour of the seed coat, ranging from black
benefit under specific conditions. In each case, only to white (Ribeiro; Jost; Cargnelutti Filho, 2004).
those that meet the objectives of the program should be
Genotypes 2, 4, 8, 9 a nd 10 show ed a reduc tion of
analysed and selected (Carvalho et al., 2016).
more tha n 21 .5% in grain brig htness wit h sto rage, while
The GGE Biplot analysis is able to de limit mega Genotype 1 showed a reduction of only 16.3%. The genotype
environments in whic h th e patte rn of the genotype x that da rkened the most after e ight m onths of s torage wa s
environment interactio n is sim ilar, representin g simple Genotype 2, w ith a 25 .8% reduction in brightness. Ribeiro,
interactions or inte ractions with s maller changes in Jost an d C argnelutti F ilho ( 2004) gi ve a n ide al b rightness
genotype ranking (Carvalho et al., 2016). F igure 3- C of greater than 53 for carioca beans, and state that this value
shows th e for mation of two mega environments, w ith should not change with time or the environmental conditions;
Genotypes 4 a nd 13 located at t he ver tices, c lassifying however, they h ighlight the p ossibility of this cha nge. The
them as more responsive to these locations. In Figure 3-G, brightness of the genotypes selected here ranged f rom 52.9
Genotypes 9 and 3 were the most ada pted to the three to 54.6 at the time of harvest (Time I); by the end of the eight
mega environments that were formed. months of storage, the brightness ranged from 41.5 to 45.7.
Figure 4 - Boxplot of grain brightness (L*) in selected cowpea genotypes under storage
Source: prepared by the author. Time I: harvest; Time II: 2 months; Time 3: 4 months; Time 4: 6 months; Time 5: 8 months
Genotype 1 had the high est average in the Cra to Genotypes 1, 4, 8 and 10 are the most adapted to irrigated
environment and the third highest average in Crateús, both environments, and Genotypes 1, 2, 7 and 9 to locations with
under irrigated conditions. It was unstable in each of t he rainfed cultivation;
environments under evaluation, but can be recommended
2. Genotype 1, in addition to its good productive performance,
as an alternative cultivar with less grain darkening in the
shows le ss gr ain dark ening und er sto rage, a nd c an be
locations to which it adapted best. Genotype 9, considered
recommended to prod ucers a nd rese llers wh o stock the
stable, had a highe r grain yi eld than the controls in each
product, so as to guarantee better post-harvest quality.
of the e valuated en vironments, a nd was s econd only to
Genotype 1 in terms of less darkening. Cruz et al. (2020)
also recommended the genotype for its stability.
For t hese cu ltivars to b ecome a vailable on t he
ACKNOWLEDGEMENTS
market, it is important they be launched and pub licised, The aut hors would like to th ank E mbrapa Me io-
and the f armers and seed p roducers en couraged. In t he Norte for providing the seeds for the trials, and the Conselho
seed distribution program for farmers in the state of Ceará, Nacional de Desenvolvimento Científico e Tecnológico
only tw o cultivars ( Pujante a nd I PA 207 Miranda) we re (CNPq) for the doctoral scholarship granted to the lead author.
available last year due to a lack of offers from the bidders
(Ceará, 202 1). The distribut ion of see ds from s table or
adapted cultivars in certain locations in the state can help
increase the average productivity of the cowpea in Ceará, REFERENCES
of 329 kg ha-1 (EMBRAPA, 2020).
ABIRIGA, F . et al. Ha rnessing genotype-by-environment
interaction to determine adaptability of advanced cowpea lines
to multiple environments in Uganda. Journal of Plant Breeding
CONCLUSIONS and Crop Science, v. 12, n. 2, p. 131-145, 2020.
ABREU, H. K. A. et al. Adaptability and stability of cowpea
1. Genotype 9 can be recommended for the semi-arid region genotypes v ia R EML/BLUP and gge bipl ot. Bioscience
of the state of Ceará as it is stable and has good grain yield. Journal, v. 35, n. 4, p. 1071-1082, 2019.
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