Morphological Plasticity of Economical Traits
in Pigeonpea Genotypes Grown in South Africa
Maletsema Alina Mofokeng1, Zaid Bello1, Kingstone Mashingaidze1 and Abe Shegro Gerrano2,3*
1Agricultural
Research Council-Grain Crops, Private Bag X 1251, Potchefstroom, 2520, South
Africa
2Agricultural
Research Council - Vegetables, Industrial and Medicinal Plants, Private Bag X
293, Pretoria, 0001. South Africa
3Food
Security and Safety Focus Area, Faculty of Natural and Agricultural Sciences,
North-West University, Mmabatho, South Africa
*For correspondence:
agerrano@arc.agric.za; ORCID: http://orcid.org/0000-0001-7472-8246
Received
09 May 2022; Accepted16 July 2022; Published 25 August 2022
Abstract
Pigeonpea (Cajunus cajan) is a leguminous crops cultivated in tropical and
sub-tropical region of the world. This crop is one of underutilized and future
food security plant species grown southern Africa. The objective of the study
was to assess morphological variability among nineteen tested pigeonpea
genotypes using multivariate analysis. The experimental trial was conducted at
Mafikeng and Nelspruit sites located in Northwest and Mpumalanga Provinces of
South Africa. A randomized complete block design with three replications at all
sites. Data were recorded on quantitative and qualitative traits and analysed
using univariate (ANOVA), and multivariate analysis tools. Significant genotype
effect was observed for plant height (PH), pod bearing (PDB) and seed number
per pod (SNP) among the studied genotypes. Seed yield (SY) was positively
correlated with seed number per pod (SNT), seed number per plant (SNP) and pod
weight (PWT), whereas PBD was negatively associated with hundred seed weight
(HSW). Principal component analysis (PCA) revealed five significant principal
components (PCs), which accounted for 84.70% of phenotypic variation among the
studied genotypes. The Shannon Weaver diversity indices ranged from 0.98–1.00,
indicating the presence of variation among the qualitative traits measured. The
clustering analysis grouped genotypes into three main groups, with ICEAP00554,
ICEAP000979-1, ICEAP00540, and Karatu-1 being the most diverse and singletons.
Hence, use of multivariate analyses revealed the existence of morphological
variation among the test pigeonpea genotypes for breeding population. These
identified genotypes could be used as potential parental lines in a pigeonpea
breeding programme for direct production and development of new high yielding
varieties in the country. © 2022 Friends Science Publishers
Keywords: Agro-morphology;
Characterization; Pigeonpea; Similarity; Variation
Introduction
Pigeonpea (Cajunus cajan) is a diploid (2n = 2x = 22) legume (Maesen 1990)
grown in tropical and subtropical regions of the world. It is underutilised
crop species, despite its contribution to food and nutritional security (Lin-Qi
2014). The crop improves the fertility of the soil through atmospheric nitrogen
fixation and increase production and productivity of the crop (Adebowale and
Maliki 2011; Choudhary et al. 2013; Saidia
et al. 2019). Pigeonpea can be
intercropped with cereal-based cropping system (Lin-Qi 2014). It is sources of
macro- and micronutrients, vitamins and phytochemical compositions (Saxena et al. 2010; Gerrano et al. 2022). It is considered as
climate smart crop in tropical and sub-tropical regions. It has the ability to
withstand drought and give good economic benefits when planted under dryland
farming conditions and sustain the livelihood of resource poor rural
populations in tropical and sub-tropical regions of the African continent
including South Africa. Furthermore, the crop helps in protecting the
environment from soil erosion and degradation, improve the fertility of the
soil, increase crop production and productivity at marginal crop lands towards
soil and food security strategies. It is climate smart crop that adapt to the
current climate change, which is tolerant to heat, drought, diseases and insect
pests (Odeny 2007).
The seed of the crop can
be eaten as a green vegetable and dry pulse and is an important source of
nutritional components (Faris et al.
1987; Choudhary et al. 2013). The
green pods and foliage of the plant can be used as animal feed (Mallikarjuna et al. 2011). The crop is cultivated by
the resource poor small scale farmers with the low input agriculture in South
Africa. Hence, identification of potential candidate genotypes and development
of improved cultivars for increased production and productivity of the crop is
important.
For an efficient
evaluation and utilisation of the plant genetic resources, understanding and
knowledge of genetic diversity, genetic background information, collection and
classification are important and the basis for crop improvement programs (Khan et al. 2014; Syafii et al. 2015), which is elucidated through different marker systems
such as agro-morphological, biochemical and molecular markers. Among these,
morphological characterisation is considered as the initial step for designing
breeding programs (Smith and Smith 1989; Khan et al. 2014), it is influenced by the growing environmental
condition unlike that of DNA-based markers. Yohane et al. (2020) reported the existence of widest variability among
test pigeonpea based on their morphological performance in Malawi. Assessing
genetic variability helps to study heterosis (Virk et al. 2003), selection of transgressive breeding segregants and
genes of novelty, and has a role in collection and maintenance of germplasm for
crop improvement (Duran et al. 2009)
in the gene bank for future use.
Potential parental lines
were identified for improvement (Malik et
al. 2014; Syed et al. 2019)
through using multivariate statistical analysis. It is effective statistical
tool for studying the differences and similarities between and within the
genotypes (Peeters and Martinelli 1989; Kovacic 1994; Rachovska et al. 2002; Mondal 2003; Mohammadi and
Prasanna 2003; Ajmal et al. 2013;
Immad et al. 2018), which can help to
generate new breeding population in the breeding programme. The knowledge and
understanding of the crop species and their respective descriptors are
necessary for informed breeding strategy (Upadhyaya et al. 2007; Gbaguidi et al.
2018). The assessment of phenotypic plasticity using morphological characters
in pigeonpea is importance in order to determine the existing variability in
the population which will finally enable the identification and selection of
potential and superior lines of the genotypes for production and breeding.
Therefore, the objective of the current study was to assess the variability and
their interrelationship in pigeonpea genotypes using morphological traits.
Materials and Methods
Plant material and trial
sites
The 19 pigeonpea genotypes were
evaluated on the field during the summer season of 2019–2020. The origins of
the pigeonpea genotypes used in the study are listed in Table 1.
The experimental trials
were conducted at the North West University research farm at Mafikeng (25°48ʹ
S, 45°38ʹ E; 1012 m. a. s. l.) in North West Province and the Agricultural
Research Council – Tropical and Subtropical research station in Nelspruit
(25.49°89’ S, 31.35°37’
E; 670 m. a. s. l.) in Mpumalanga Provinces during 2019/2020 cropping season in
South Africa. Pigeonpea is widely grown predominantly in this two Province in
South Africa and have extreme variations in agro-climatic conditions. The soils
on the North West University farm belongs to the Hutton series, with sandy loam
and a yellow sand alternating (Molope 1987; Kasirivu et al. 2011), while the Nelspruit research station field consisted
of sandy loam soil. During the season, Mafikeng received a summer total
rainfall, with a mean of 571 mm during the cropping season. The mean maximum temperature
is 37°C, while the mean minimum temperature is 9°C during this cropping season.
The field in Nelspruit is characterised by mean maximum temperature of 28°C.
The mean minimum temperature is12.5°C with
a precipitation of about 796 mm during the cropping season in 2019/2020.
Trial design and management
The trials were laid out in a
randomized complete block design replicated three times with a plot consisting
of two rows of 4 m length. The inter- and intra-row spacing’s were 90 and 60
cm, respectively. The experiment was conducted during summer cropping season in
rainfed condition based on the farmers practice. Weeding was done manually. No
fertilizer was applied to simulate low input cropping system in the region
(Gerrano et al. 2015).
Data collection
Data were recorded according to
standard descriptor list for pigeonpea (IBPGR 1994). Data were recorded from
three randomly selected plants in the middle of each row per replications. The
qualitative data recorded included base flower colour, second flower colour,
vigour at 50% flowering, pod form, seed colour pattern, seed shape, and pattern
of streaks. The list of quantitative traits studied and their data collection
method is presented in Table 2.
Statistical data analysis
The recorded quantitative data
were analysed using analysis of variance (ANOVA), principal component analysis
(PCA), and Pearson correlations. The qualitative data were analysed using
frequencies, spearman correlations, and Shannon Weaver Table 1: Pigeonpea germplasm used in the study
Entry
number |
Genotype
Name |
Origin/source |
1 |
ICEAP 01147 |
Kenya |
2 |
ICEAP 01154-2 |
Kenya |
3 |
ICEAP 01150-1 |
Kenya |
4 |
ICEAP 01179 |
Kenya |
5 |
ICEAP 00979-1 |
Kenya |
6 |
ICEAP 01172-2-4 |
Kenya |
7 |
ICEAP 01159 |
Kenya |
8 |
ICEAP 01544-2 |
Kenya |
9 |
ICEAP 00540 |
Kenya |
10 |
ICEAP 00554 |
Kenya |
11 |
ICEAP 00557 |
Kenya |
12 |
ICEAP 00850 |
Kenya |
13 |
Ilonga 14-M1 |
Tanzania |
14 |
Mali |
Tanzania |
15 |
Ilonga 14-M2 |
Tanzania |
16 |
Karatu-1 |
Tanzania |
17 |
Kiboko |
Tanzania |
18 |
Komboa |
Tanzania |
19 |
Tumia |
Tanzania |
Table 2: A list of economical
traits measured, abbreviations and definitions
Trait |
Abbreviation |
Measurement/definitions |
Plant
height (cm) |
PHT |
Height of a plant from the
base of the stem to the tip of the plant at harvest |
Days
to 50% flowering |
DFF |
Number of days from planting
until 50% of the plants have flowered in a plot |
Pod
bearing (cm) |
PDB |
Distance from lowest to the
top most of the plant |
Leaf
length mm) |
LFL |
Length from the tip of the
leaf to the leaf petiole |
Leaf
width (mm) |
LFW |
Length in the middle of the
leaf from one tip to the other tip |
Branch
number |
BRN |
Number of branches per plant |
Stem
diameter (cm) |
STD |
Diameter of plant stem |
Pod
length (mm) |
PDL |
Length of the pod from bottom
end to top end at harvest |
Pod
width (mm) |
PDW |
Length at the centre of the
pod from one end to the other end/diameter |
100
Seed weight (g) |
HSW |
Weight of 100 seed picked
randomly for each genotype |
Pod
weight (g) |
PWT |
Weight of dry pods harvested
from each genotype |
Seed
number per pod |
SNT |
Number of seeds in a pod (average
of 10 pods) |
Seed
number per plant |
SNP |
A number of seeds produced by
a single plant. |
Seed
yield (g) |
SYD |
Weight of seeds produced per
plant |
DFF = Days to 50% flowering; PHT
= Plant height; BRN = Branch number; LLT = Leaf length; LWT = Leaf width; PDB =
Pod bearing; 100SW = hundred seed weight; PDL= Pod length; PDW = Pod width; SNP
= Seed number per pod; PWT = Pod weight; SEP = Seed number per plant; STD =
Stem diameter; SYD = seed yield per plant
diversity index. The biplots
were generated using principal coordinate analysis in SAS version 9.6 (SAS
Institute 2021). A dendrogram was constructed using Genstat 18th edition (VSN International,
Hempstead, UK) (2020).
Results
Genetic variability and
genotype by environment interaction
The univariate analysis of
variance (ANOVA) result depicted that there were highly significant (P ≤ 0.01) genetic variability
observed among the tested genotypes for pod length (PDL) and pod weight (PWT) (P ≤ 0.01) (Table 3). Furthermore,
the ANOVA mean squares showed genotype (G), site (S), and genotype × site
interaction (GEI) effects on quantitative traits is presented in Table 3.
Furthermore, the agronomic characteristics of the pigeonpea genotypes showed
variability in different environments due to the significant genotype ×
environment interaction (Table 3) for plant height (PHT) (P ≤ 0.01), pod bearing (PDB) (P ≤ 0.05) and seed number per pod (SNP) (P ≤ 0.05). There were significant (P ≤ 0.05) differences between sites based on days to
flowering (DFF), plant height, branch number (BRN), stem diameter (STD), pod
bearing, pod length, pod weight and significant differences for seed number per
pod. There was a significant (P ≤ 0.05)
site × genotype interaction effect based on plant height, pod bearing and seed
number per pod (Table 3).
Pearson correlation
analysis
Pearson’s correlations (r) of 14 quantitative traits measured in
the study are shown in Table 4. Days to flowering was significantly and
positively correlated with plant height, branch number, stem diameter, and
hundred seed weight. Similarly, Table 3:
Combined analysis of variance for quantitative traits among the studied
pigeonpea genotypes in terms of means squares
SOV |
d.f |
DFF |
PHT |
BRN |
STD |
LLT |
LWT |
PDB |
100 SW |
PDL |
PDW |
SNP |
PWT |
SEP |
SYD |
Site
(S) |
1 |
85323.9** |
175005.4** |
1141.9** |
11556.7** |
0.2 |
722.6 |
41198.2** |
83.4 |
1957.3** |
10.9 |
138.3* |
142.2** |
1.3 |
45.9 |
Genotype(G) |
18 |
76.2 |
1908.7 |
15.6 |
13.5 |
1.9 |
327.2 |
1417.3 |
163.3 |
507.8** |
3.7 |
57.0 |
46.0** |
8.0 |
20.0 |
S ×
G |
18 |
72.9 |
2867.6** |
16.7 |
14.7 |
2.1 |
325.4 |
2733.8* |
165.9 |
322.2 |
4.0 |
61.2* |
30.8 |
8.9 |
17.9 |
SOV = Sources of variation; d.f.
= degree of freedom; DFF = Days to 50% flowering; PHT = plant height; BRN =
Branch number; LLT = Leaf length; LWT = Leaf width; PDB = Pod bearing; 100SW =
hundred seed weight; PDL= Pod length; PDW = Pod width; SNP = Seed number per
pod; PWT = Pod weight; SEP = Seed number per plant; STD = Stem diameter; SYD =
seed yield per plant; * = significantly different from zero at P ≤ 0.05; ** = significantly
different from zero at P ≤ 0.01
Table 4: Pearson correlations for
quantitative traits among the studied pigeonpea genotypes
Variable |
DFF |
PHT |
BRN |
STD |
LLT |
LWT |
PDB |
100 SW |
PDL |
PDW |
SNP |
PWT |
SEP |
DFF |
1.00 |
|
|
|
|
|
|
|
|
|
|
|
|
PHT |
0.701*** |
1.00 |
|||||||||||
BRN |
0.625*** |
0.751*** |
1.00 |
||||||||||
STD |
-0.900*** |
-0.667*** |
-0.492*** |
1.00 |
|||||||||
LLT |
-0.089 |
0.017 |
0.040 |
0.241 |
1.00 |
||||||||
LWT |
-0.034 |
-0.019 |
0.075 |
0.169 |
0.672*** |
1.00 |
|||||||
PDB |
-0.498*** |
-0.405*** |
-0.341*** |
0.504 |
0.190* |
-0.056 |
1.00 |
||||||
100SW |
0.525*** |
0.431*** |
0.296** |
-0.574** |
-0.053 |
-0.003 |
-0.353*** |
1.00 |
|||||
PDL |
0.183 |
0.046 |
-0.011 |
-0.136 |
0.117 |
0.095 |
-0.010 |
0.159 |
1.00 |
||||
PDW |
0.063 |
0.083 |
0.114 |
-0.024 |
0.003 |
-0.110 |
0.014 |
-0.060 |
0.018 |
1.00 |
|||
SNP |
0.086 |
0.133 |
0.089 |
-0.076 |
-0.085 |
-0.202* |
0.020 |
-0.063 |
0.436*** |
0.135 |
1.00 |
||
PWT |
0.189* |
0.068 |
0.013 |
-0.139 |
0.102 |
0.055 |
-0.006 |
0.135 |
0.986*** |
0.161 |
0.526*** |
1.00 |
|
SEP |
0.183 |
0.107 |
0.064 |
-0.130 |
0.060 |
-0.037 |
0.005 |
0.072 |
0.858*** |
0.453*** |
0.669*** |
0.932*** |
1.00 |
SYD |
0.183 |
0.096 |
0.042 |
-0.136 |
0.065 |
-0.013 |
0.001 |
0.092 |
0.928*** |
0.248** |
0.694*** |
0.974*** |
0.976*** |
DFF = Days to 50% flowering, PHT
= plant height, BRN = Branch number, LLT = Leaf length, LWT = Leaf width, PDB =
Pod bearing, 100 SW = hundred seed weight, PDL = Pod length, PDW = Pod width,
SNP Seed number per pod, PWT = Pod weight, SEP = Seed number per plant, STD =
Stem diameter, SYD = seed weight per plant.
Bold value represent significant
association * = P < 0.05, ** = P < 0.01, *** = P < 0.001
Table 5: Factor loadings of the
most important PCs for agro-morphological traits among the studied pigeonpea
genotypes
Traits |
PC1 |
PC2 |
PC3 |
PC4 |
PC5 |
DFF |
0.57 |
-0.72 |
0.05 |
-0.02 |
0.01 |
PHT |
0.47 |
-0.70 |
0.12 |
0.24 |
-0.25 |
BRN |
0.37 |
-0.64 |
0.19 |
0.37 |
-0.31 |
STD |
-0.52 |
0.74 |
0.12 |
0.13 |
-0.07 |
LLT |
-0.00 |
0.21 |
0.87 |
0.18 |
-0.04 |
LLW |
-0.04 |
0.08 |
0.91 |
-0.02 |
0.07 |
PDB |
-0.27 |
0.58 |
-0.01 |
0.17 |
-0.22 |
HSW |
0.37 |
-0.54 |
0.09 |
-0.35 |
0.32 |
PDL |
0.82 |
0.42 |
0.13 |
-0.30 |
0.03 |
PDW |
0.27 |
0.11 |
-0.15 |
0.78 |
0.53 |
SNP |
0.612 |
0.31 |
-0.26 |
0.15 |
-0.45 |
PWT |
0.87 |
0.44 |
0.07 |
-0.17 |
0.06 |
SEP |
0.88 |
0.44 |
-0.05 |
0.14 |
0.10 |
SYD |
0.89 |
0.45 |
-0.02 |
-0.07 |
-0.02 |
Eigenvalue |
4.62 |
3.50 |
1.78 |
1.163 |
0.82 |
Explained variance (%) |
32.97 |
24.96 |
12.694 |
8.307 |
5.87 |
Cumulative variance (%) |
32.968 |
57.931 |
70.625 |
78.932 |
84.80 |
DFF = Days to 50% flowering, PHT
= plant height, BRN = Branch number, LLT = Leaf length, LWT = Leaf width,
PDB = Pod bearing, 100 SW = hundred seed weight, PDL= Pod length, PDW
= Pod width, SNP = Seed number per pod, PWT = Pod weight, SEP = Seed
number per plant, STD = Stem diameter, SYD =– seed weight per plant
days to flowering was
significantly and positively correlated with pod weight and negatively
correlated with pod bearing. Plant height was highly significant and positively
correlated with branch number per plant, stem diameter, and hundred seed
weight, and negatively associated with pod bearing. Branch number had a
negative and significant association with stem diameter and pod bearing, and a
positive correlation with hundred seed weight. Stem diameter had a significant
and positive correlation with leaf length, whereas pod bearing showed a negative
association with hundred seed weight. Leaf length showed positive and
significant correlations with leaf width and pod bearing. Leaf width had a
negative association with seed number per pod. Pod bearing had a highly
significant negative correlation with hundred seed weight. Pod length showed a
positive association with seed number per pod, pod weight, seed number per
plant, seed yield. Pod width showed a positive and highly significant
correlation with seed number per plant and seed yield. Seed number per pod was
positively correlated with pod weight, seed number per plant, and seed yield.
Pod weight had positive correlations with seed number per plant and seed yield.
Seed number per plant was highly significant and positively correlated with
seed yield (Table 4).
Fig. 1: PC biplot for
quantitative traits among the studied pigeonpea genotypes. PC1=first principal
component; PC2=second principal component
Fig. 2: PCA biplot for
qualitative traits among the studied pigeonpea genotypes.
Principal component
analysis
Five most important PCs were
identified contributing 32.9, 24.9, 12.7, 8.3 and 5.9%, to the total variation
of 84.7%, respectively (Table 5). The first PC had pod length, pod weight, seed
number per plant and seed yield contributing to this variation. In the second
PC, days to flowering, plant height, branch number, stem diameter contributed
the most to variation. Leaf length and leaf width contributed the most
variation in third PC. In the fourth PC, pod width was the most contributors to
variation whereas in the fifth PC, pod width and seed number per pod was the
traits that contributed the most variation.
Principal coordinate
analysis
The principal component (PC)
biplot of the quantitative traits showing grouping of genotypes superimposed
with traits is presented in Fig. 1, PC1 had 31.35% and PC2 had 0.26% variances
with the total contributing variation of 51.61%. Gerrano et al. (2022)
reported that the angles lesser than 45o between the vector lines of
the two respective variables indicate positive and high trait correlation and
revealed the ability to discriminate the test genotypes for breeding. Genotypes
ICEAPO1150-1, ICEAPO1154-2, ilonga14-M2, ICEAPO1172-2-4, ICEAPO1544-2, Mali,
ICEAPO4459 and longa14-M1 were grouped Table
6: Frequency percentages of qualitative traits for medium duration
pigeonpea
Trait |
Score |
Frequency
(%) |
Cumulative
frequency
(%) |
H’ |
Vigour
at 50% flowering |
Low |
5.36 |
5.36 |
0.99 |
Intermediate |
23.21 |
28.57 |
|
|
High |
71.43 |
100 |
|
|
Base
flower colour |
Light yellow |
19.65 |
19.65 |
0.97 |
Yellow |
51.78 |
71.43 |
|
|
Orange-yellow |
28.57 |
100 |
|
|
Second
flower colour |
Red |
71.43 |
71.43 |
0.96 |
Purple |
28.57 |
100 |
|
|
Pattern
of streaks |
Sparse |
35.09 |
35.09 |
0.97 |
Medium amount |
15.79 |
50.88 |
|
|
Dense |
22.81 |
73.68 |
|
|
Uniform coverage of second
color |
26.32 |
100 |
|
|
Flowering
pattern |
Determinate |
100 |
100 |
1.00 |
Stem
Thickness rating |
Thick (>13 mm) |
100 |
100 |
1.00 |
Growth
habit |
Erect and compact |
22.81 |
22.81 |
0.98 |
Semi spreading |
1.75 |
24.56 |
|
|
Spreading |
75.44 |
100 |
|
|
Stem
color |
Green |
63.16 |
63.16 |
0.98 |
Sun Red |
36.84 |
100 |
|
|
Pod
form |
Flat |
3.64 |
3.64 |
0.99 |
Cylindrical |
96.36 |
100 |
|
|
Seed
color pattern |
Plain |
3.57 |
3.57 |
0.99 |
Mottled |
7.14 |
10.71 |
|
|
Speckled |
71.43 |
82.14 |
|
|
Mottled and speckled |
17.86 |
100 |
|
|
Seed
shape |
Oval |
21.43 |
21.43 |
0.98 |
Globular |
64.29 |
85.71 |
|
|
Square |
14.29 |
100 |
|
H’ = Shannon Weaver Index
together based on high SYD, HSW,
SNT, PWT, PDW and PDL. Further, ICEAPO1179 and Tumia were identified as best
genotypes for BRN, PHT and SNP. The genotype ICEAP01147, Kiboko, and ICEAP00850
were associated with the variables LLF, DFF, and LLW, while the genotypes
ICEAP00557, Karatu-1, ICEAP00554 and Komboa revealed less association to the
variables recorded indicating that the genotypes were less responsive to the
variables. Genotype ICEAP00850 was associated to PDB. Genotype ICEAP00540 is peculiar
genotypes that was found far from the rest of genotypes from the scatter biplot
(Fig. 1), which can be considered for further evaluation in the breeding
program. Stem diameter and pod bearing were negatively correlated with plant
height, branch number, seed yield, and 100 seed weight. Seed number per pod,
pod length, pod width, pod weight, seed yield, and seed number per plant were
positively correlated with hundred seed weight, while branch number and plant
height were highly positively correlated. The same traits were also correlated
with stem diameter, pod bearing, leaf width and leaf length.
The biplot for the
qualitative traits, the PC1 showed 40.27% and F2 had 26.41% (Fig. 2). The first
quadrant showed base flower colour and vigour at 50% flowering, which are
positively correlated in this quadrant and are associated with the genotypes
Ilonga 144-M1, ICEAP 00850, and ICEAP 01159, while the second quadrant showed
seed shape that was associated with the genotypes positioned in this quadrant.
The third quadrant had pod form and seed colour pattern that are positively
correlated to each other. The genotypes Kiboko and Mali had similar pod form
and seed colour pattern in this quadrant. The 4th quadrant consists of only
second flower colour. All the genotypes scattered in this quadrant were grouped
together based on this trait (Fig. 2). In Fig. 2, the genotypes that are
circled have similar values for PC1 and PC2 scores, which made them to be
positioned on one dot.
Frequencies of qualitative
traits
The frequencies of eleven
qualitative traits measured are shown in Table 6. Vigorousness at flowering was
high with 71.4% of plants being vigorous and intermediate was 23.2%. The base
flower colour was dominated by yellow flowers followed by orange-yellow. The
second flower colour was predominantly composed of red flowers (71.4%). The
pattern of streaks was dominated by sparse streaks (35.1%), followed by uniform
coverage of second colour and dense streaks. All plants of various genotypes
had 100% stems thicker than 13 mm with green stems dominating (63.2%). The
growth habit of the crop was predominantly composed of spreading types (75.4%)
followed by erect and compact at 22.8%. The genotypes were dominated by
cylindrical pods 96.40 with speckled seed colour pattern at 71.4%
Fig. 3: A dendrogram showing
interrelationships and divergence among nineteen pigeonpea genotypes based on
quantitative traits
followed by mottled and speckled
at 17.9%. The shape of the seed was predominantly globular (64.3%) with oval
shape being 21.4%.
Shannon weaver diversity
Shannon weaver diversity indices
are shown in Table 6. The diversity indices range from 0.96 (second flower
colour) to 1.00 (flowering pattern and stem thickness). All traits showed
significant variation except for flowering pattern and stem thickness.
Hierarchical clustering
A dendrogram was constructed
using hierarchical clustering to present differences and interrelationships
among the studied pigeonpea genotypes (Fig. 3). The dendrogram grouped
genotypes into three clusters and four singletons. The first cluster was
composed of six genotype, Longa14-M1, Mali, ICEAP00557, Ilonga14-M2, ICEAP01159,
and ICEAP00850. The second cluster was composed of four genotypes, ICEAP0050-1,
ICEAP01179, kmboa and ICEAP01147. The genotypes, Kiboko, ICEAP01154-2, Tumia,
ICEAP01172-2-4 and ICEAP01544-2 were grouped in third cluster. Four genotypes
were identified as most divergent and grouped as singletons (ICEAP00554,
ICEAP000979-1, ICEAP00540, and Karatu-1). These genotypes were far and
distantly related with the rest of the test genotypes.
Discussion
The significant difference
observed among the pigeonpea genotypes showed the existence of genetic
variation with respect to the measured morphological characteristics. The
knowledge of morphological variation for a trait and trait correlations are
important components of any breeding objective. There were highly significant
differences for sites based on days to flowering, plant height, branch number,
stem diameter, pod bearing, pod length, pod weight and significant differences
for seed number per pod (Table 3). This indicates that the expression of the significant
traits varied with the growing environmental conditions they were tested in.
Their performance was not stable across sites. The presence of highly
significant differences in genotypes based on pod length and pod weight
highlights the presence of genotypic variation among the genotypes evaluated
based on the two traits which can be exploited for cultivar improvement in
future breeding programmes. The significant differences on genotype x site
interaction could be attributed to the different reactions of the genotypes to
sites or due to differences between the sites. In each environment, phenotypic
manifestation is the result of the action of the genotype under the influence
of the environment. However, when considering a series of growing environments,
in addition to the genetic and environmental effects, an additional effect can
be detected from their interaction (Marais et
al. 2013; Nunes et al. 2014).
Significant genotype × environment interaction on yield and yield components in
this study concur with the results reported previously (Vales et al. 2012; Kimaro 2016; Gerrano et al. 2020).
The positive and
significant correlation observed among the quantitative characters indicated
that direct selection for any of these traits could lead to simultaneous improvement
in the other characters of pigeonpea for increased production and productivity.
Yohane et al. (2020) reported the
existence of positive correlations for most of secondary traits that revealed
multiple trait identification and selection for simultaneous trait improvement,
while the weak correlations among the traits would result in an inefficient
selection or low genetic gains that will take long time to fix the traits of
interest. In this study seed yield was positively correlated with seed number per
pod, seed number per plant and pod weight whereas pod bearing was negatively
associated with hundred seed weight. The positive and significant correlations
in the current study indicated the importance of simultaneous improvement for
the traits of interest in the crop (Sodavadiya et al. 2009; Linge et al.
2010; Prasad et al. 2013; Saroj et al. 2013; Ojwang et al. 2016; Kinhoégbè et al.
2020; Yohane et al. 2020).
The Principal component
analysis over sites revealed five most important PCs with pod length, pod
weight, seed number per plant, seed yield, leaf length, leaf width, days to
flowering, plant height, and stem diameter being the most contributing traits
to the total variation observed. This suggests that these traits are useful for
selection. Other reports have indicated that trait contribution to different
PCs varies with genetic diversity within the tested germplasm and the number of
traits evaluated (Upadhyaya et al.
2007). The biplot also showed the different grouping of pigeonpea genotypes
based on specific traits. These findings suggested that both agro-morphological
traits revealed variability among the tested genotypes but complementary
information for breeding.
The most of pigeonpea
genotypes in the current study showed a tendency to spreading growth habit,
yellow based flower colour, with red second flower colour, sparse pattern of
streaks, green stems, with globular and speckled seed color pattern. Similar
results have been reported for qualitative traits (Rupika and Bapu 2014).
Shannon Weaver indices also confirmed the presence of genetic divergence based
on qualitative traits. Thus, in spite of the influence of prevalent
environmental factors, qualitative variables can be used to characterize
pigeonpea genetic resources.
The pigeonpea genotypes
were clustered into three major groups, indicating that these genotypes in the
three groups are distantly related. The genotypes in the same cluster group are
closely related and they maybe of the same source or origin. Hence, selection
of genotypes within these clusters may not be desirable to get higher yield and
economic benefits (Muniswamy et al.
2014; Rupika and Bapu 2014). Therefore, for the crop hybridization programs, the
choice of suitable diverse parents based on their genetic differentiation would
be more fruitful than the choice based on the geographical distances. ICEAP
00540, ICEP00979-1, Karatu-1 and ICEAP00554 would be the ideal genotypes for
use as parents in any pigeonpea breeding programme for agronomic improvement.
The genotypes clustered in the same group showed their genetic similarities
that might be due to free exchange of similar materials with different names
that may have overlapped in the previous diversity distribution pattern of the
domesticated species (Jaradat and Shahid 2006; Aghaee et al. 2010). Reddy and Jayamani (2019) as well as Niranjana et al. (2014) reported the existence of
genetic diversity in pigeonpea using multivariate analysis. Singh et al. (2014); Qutadah et al. (2019); Kinhoégbè et al. (2020) further reported different
grouping of genotypes for the agronomic traits in the assessment of genetic
variability.
Conclusion
In conclusion, there was
sufficient genetic variability existed among pigeonpea genotypes that would
help the improvement through identification and selection of parental lines for
the traits of interest with greater chances of success in pigeonpea breeding.
The study revealed the presence of genetic diversity among the pigeonpea
genotypes studied based on the analysis of variance and multivariate tools used
for analyses. The results indicated that the higher level of genetic diversity
observed within the acquired genotypes from ICRISAT collection in Kenya and
Tanzania would enable efficient utilisation and pigeonpea improvement in
breeding programs in South Africa and other countries. The variability among
the genotypes also will help to identify and select the potential parents for
hybridization. The selection for single trait and improvement for this trait
would require more breeding work to fix the trait, therefore it is suggested
that selection of genotypes for multiple traits as well as directly correlated
traits would accelerate pigeonpea breeding for improvement of traits of
interest simultaneously. Further characterization this crop using molecular
techniques should be conducted to elucidate the
environmental factor for
utilization in the future breeding programs.
Acknowledgements
The first author would like to
thank the Department of Agriculture, Land Redistribution and Rural Development
for funding. Additionally, the authors would like to thank the technical
assistance and trial management of Paul Rantso, Dinah Scott, Deon Du Toit, and
Theodora Mathobisa.
Author Contributions
Dr.
Maletsema Alina Mofokeng is a
Researcher in Plant Breeding at the Agricultural Research Council-Grain Crops,
Potchefstroom, South Africa. Contribution: Conceptualization, experimental
design, planting, trial monitoring, data recording, analysis and drafting of
the manuscript. Dr. Zaid Bello
is a Researcher in Agronomy department of Agricultural Research Council-Grain
Crops, Potchefstroom, South Africa. Contribution: trial management, and review
the manuscript. Dr Kingstone
Mashingaidze is a Senior Research Manager in the Agricultural Research
Council-Grain Crops, Potchefstroom, South Africa. Contribution: review,
supervision and edition inputs into the manuscript. Dr Abe Shegro Gerrano is a Senior Research Specialist in the
Agricultural Research Council-Vegetables, Industrial and Medicinal Plants,
Pretoria, South Africa. Contribution: Conceptualization, experimental design,
data analysis and review the manuscript.
Conflicts
of Interest
The authors have not declared
any conflict of interests.
Funding
Source
This work was supported by the
Department of Agriculture, Land Redistribution and Rural Development.
In Loving Memory
This article is dedicated to our
colleague Dr. Maletsema Alina Mofokeng who passed away on 21 April 2022.
Data Availability
All new research results were presented in this
article.
Ethics Approval
Not applicable.
References
Adebowale OJ, K Maliki
(2011). Effect of fermentation period on the chemical composition and
functional properties of pigeonpea seed flour. Int Food Res J 18:1329‒1333
Aghaee M, R Mohammadi, S
Nabovati (2010). Agro-morphological characterization of durum wheat
accessions using pattern analysis. Aust J Crop Sci 4:505‒514
Ajmal SU, MN Minhas, A
Hamdani, A Shakir, M Zubair, Z Ahmad (2013). Multivariate analysis of genetic
divergence in wheat (Triticum aestivum)
germplasm. Pak J Bot 45:1643‒1648
Choudhary AK, S Kumar, BS
Patil, JS Bhat, M Sharma, S Kemal, TP Ontagodi, S Datta, P Patil, SK
Chaturvedi, R Sultana, VS Hedge, S Choudhary, PY Kamannavar, AG Vijaykumar (2013).
Narrowing yield gaps through genetic improvement for Fusarium wild resistance in three pulse crops of the semi-arid
tropics. Sabrao J
Breed Genet 45:341‒370
Duran C, N Appleby, D
Edwards, J Batley (2009). Molecular genetic markers: Discovery, applications,
data storage and visualisation. Curr Bioinform 4:16‒27
Faris DG, KB Saxena, S
Mazumdar, U Singh (1987). Vegetable Pigeonpea: A promising crop for India.
ICRISAT, Patancheru, India
Gbaguidi AA, A Dansi, I
Dossou-Aminon, DSJC Gbemavo, A Orobiyi, F Sanoussi, H Yedomonhan (2018).
Agromorphological diversity of local Bambara groundnut (Vigna subterranea (L.) Verdc.) collected in Benin. Genet Res Crop
Evol 65:1159–1171
Gerrano AS, A Moalafi, HA
Seepe, S Amoo, H Shimelis (2022). Nutritional and phytochemical ompositions and
their interrelationship in succulent pods of pigeon pea (Cajanus cajan L.] Millsp.). Heliyon 8:1–7
Gerrano AS, WSJV Rensburg,
I Mathew, AIT Shayanowako, MW Bairu, SL Venter, W Swart, A Mofokeng, JJ Mellem,
M Labuschagne (2020). Genotype and genotype x environment interaction effects
on the grain yield performance of cowpea genotypes in dryland farming system in
South Africa. Euphytica 216:80–90
Gerrano AS, WSJV Rensburg,
PO Adebola (2015). Genetic diversity of amaranthus species in South Africa. S
Afr J Plant Soil 32:39‒46
IBPGR (1994).
International Board for Plant Genetic Resources. Rome, Italy
Immad AS, KV Imran, AM
Shakeel, MS Pukhta, AD Zahoor, L Ajaz (2018). Genetic Diversity by Multivariate
Analysis Using R Software. Intl J Pure Appl Biosci 6:181‒190
Jaradat AA, MA Shahid
(2006). Patterns of phenotypic variation in a germplasm collection of (Carthamus tinctorius L.) from the Middle
East. Genet Res Crop Evol 53:225‒244
Kasirivu J, S Materechera,
M Dire (2011). Composting ruminant animal manure reduces emergence and species
diversity of weed seedlings in a semi-arid environment of South Africa. S
Afr J Plant Soil 28:228‒235
Khan SA, J Iqbal, H
Khurshid, N Saleem, MA Rabbani, M Zia, ZK Shinwari (2014). The extent of
intra-specific genetic divergence in Brassica
napus L. population estimated through various agro-morphological traits. Eur
J Acad Res 2:2255‒2275
Kimaro D (2016). Genetic improvement
of pigeonpea (Cajanus cajan (L.)
Millsp.) for fusarium wilt resistance in Tanzania. Ph.D. Thesis
University of Kwazulu-Natal, Pietermaritzburg, South Africa
Kinhoégbè G, G Djèdatin,
LEY Loko, RI Agbo, RK Saxena, RK Varshney, C Agbangla, A Dansi (2020).
Agro-morphological characterization of pigeonpea (Cajanus cajan L. Millspaugh) landraces grown in Benin: Implications
for breeding and conservation. J Plant Breed Crop Sci 12:34‒49
Kovacic Z (1994). Multivariate
analysis, p:293. Faculty of Economics, University of Belgrade, Serbian
Linge SS, HV Kalpande, SL
Sawargaonlar, BV Hudge, HP Thanki (2010). Study of enetic variability and
correlation in interspecific derivatives of pigeonpea (Cajanus cajan (L.) Millsp.). Electr J Plant Breed 1:929‒935
Lin-Qi X, TT Li, Z Wei, N
Guo, M Luo, W Wang, Y Zu, Y Fu, X Peng (2014). Solvent-free microwave
extraction of essential oil from pigeonpea leaves and evaluation of its antimicrobial activity. Ind Crops
Prod J 58:322‒328
Maesen LGJVD (1990).
Pigeonpea: Origin, history, evolution and taxonomy. In: The pigeonpea,
pp:15–46. Nene L, SD Hall, VK Sheilla (Eds), pp:15‒46.
C.A.B. International, Wallingford, UK
Malik R, H Sharma, I
Sharma, S Kundu, A Verma, S Sheoran S, R Kumar, R Chatrath (2014). Genetic
diversity of agro-morphological characters in Indian wheat varieties using GT
biplot. Aust J Crop Sci 8:1266‒1271
Mallikarjuna N, KB Saxena,
DR Jadhav (2011). Cajanus. In: Wild
Crop Relatives: Genomic and Breeding Resources, Legume Crops and Forages,
pp:21‒33. Kole C (Ed). Springer-Verlag,
Berlin, Heidelberg, Germany
Marais LDD, KM Hernandez, E Juenger (2013). Genotype-by-environment
interaction and plasticity: Exploring genomic responses of plants to the
abiotic environment. Annl Rev Ecol Evol Syst 44:5‒29
Mohammadi SA, BM Prasanna
(2003). Analysis of genetic diversity in crop plants-salient statistical tools
and considerations. Crop Sci 43:1235‒1248
Molope M (1987). Soil aggregate
stability. The contribution of biological and physiological processes. S Afr
J Plant Soil 4:121‒126
Mondal MAA (2003).
Improvement of Potato (Solanum Tuberosum
L.) through hybridization and in vitro
culture technique. Ph.D. Thesis. Rajshahi University, Rajshahi,
Bangladesh
Muniswamy S, R Lokesha, P Dharmaraj, S Yamanura, JR Diwan (2014).
Morphological characterization and assessment of genetic diversity in minicore
collection of igeon pea (Cajanus Cajan
(L.) Millsp). Eur J Pharm Biopharm 5:179‒186
Niranjana KB, PS Dharmaraj, VB Wali (2014). Genetic diversity and
variability studies of advanced breeding lines of pigeonpea (Cajanus cajana. L). Intl J Adv Pharm Biol
Sci 3:404‒409
Nunes HF, FFH Freire, VQ Ribeiro, RLF Gomes (2014). Grain yield
adaptability and stability of blackeyed cowpea genotypes under rainfed
agriculture in Brazil. Afr J Agric Res 9:255‒261
Odeny DA (2007). The
potential of pigeonpea (Cajanus cajan (L.) Millsp.) in Africa. Natural resources
forum. Wiley Online Library
Ojwang JD, RO Nyankanga, OM
Olanya, DO Ukuku, J Imungi (2016). Yield components of vegetable pigeonpea
cultivars. Trop Subtrop Agric Environ 67:1‒12
Peeters J, JA Martinelli
(1989). Hierarchical cluster analysis as a tool to manage variation in germplasm collections. Theor Appl
Genet 78:42‒48
Prasad Y, K Kumar, SB
Mishra (2013). Studies on genetic parameters and interrelationships among yield
and yield contributing traits in pigeonpea [Cajanus
cajan (L.) Millsp.]. Bioscan 8:207‒211
Qutadah SM, S Mehandi, IP
Singh, F Singh (2019). Assessment of Genetic Diversity for Polygenic Traits in
pigeonpea [Cajanus cajan (L.)
Millspaugh]. Intl J Curr Microbiol Appl Sci 8:1581‒1588
Rachovska G, D Dimova, B
Bojinov (2002). Application of cluster analysis and principal component
analysis for evaluation of common winter wheat genotypes. In: Proceedings of
the Scientific Session of Jubilee,
Vol. 3, pp:68‒72. Sadovo, Bulgaria
Reddy DSE, P Jayamani
(2019). Genetic diversity in land races of pigeonpea (Cajanus cajan (L.) Millsp.). Electr J Plant Breed 10:667‒672
Rupika K, KJR Bapu (2014).
Assessment of genetic diversity in pigeonpea germplasm collection using
morphological characters. Eur J Pharm Biopharm 5:781‒785
Saidia PS, F Asch, AA
Kimaro, J Germer, FC Kahimba, F Graef, JM Semoka, CL Rweyemamu (2019). Soil
moisture management and fertilizer micro-dosing on yield and land utilization
efficiency of inter-cropping maize-pigeon-pea in sub humid Tanzania. Agric
Water Manage 223:105712
Saroj SK, MN Singh, R Kumar,
T Singh, MK Singh (2013). Genetic variability, correlation
and path analysis for yield attributes in pigeonpea. Bioscan 8:941‒944
SAS Institute (2021). SAS/STAT
user’s Guide, version 9.2 SAS Institute, Cary, North Carolina, USA
Saxena KB, RV Kumar, R
Sultana (2010). Quality Nutrition through Pigeonpea-Review, Vol.
2, pp:1335‒1344. ICRISAT,
Hyderabad, India
Singh AK, S Swain, RK Gautam,
PK Singh, AK Betal, T Bharathimeena, N Kumar, SD Roy (2014). Agro-morphological
characterization of Bay Islands ppigeonpea (Cajanus
cajan) landraces and advanced lines using under Islands conditions. In: 3rd
International Conference on Agriculture and Horticulture, October 27–29,
2014 Hyderabad International Convention Centre, India
Smith JSC, OS Smith
(1989). The description and assessment of distance between inbred lines of
maize. The utility of morphological, biochemical and genetic descriptors and a
scheme for the testing of distinctiveness between inbred lines. Maydica
34:151‒161
Sodavadiya MS, JJ Pithia,
AG Savaliya, A Pansuriya, VK Korat (2009). Studies on characters association
and path analysis for seed yield and its components in pigeonpea (Cajanus cajan (L.) Millsp.). Leg Res 32:203‒205
Syafii M, I Cartika, D
Ruswandi (2015). Multivariate analysis of genetic diversity among some maize
genotypes under maize-albizia cropping system in Indonesia. Asian J Crop Sci
7:244‒255
Syed MQ, M Suhel, IP
Singh, S Farindra (2019). Assessment of Genetic Diversity for Polygenic Traits
in Pigeonpea [Cajanus cajan (L.)
Millspaugh]. Intl J Curr Microbiol Appl Sci 8:1581‒1588
Upadhyaya HD, KN Reddy,
CLL Gowda, S Singh (2007). Phenotypic diversity in the pigeonpea (Cajanus cajan) core collection. Genet
Res Crop Evol 54:1167‒1184
Vales M, R Srivastava, R
Sultana, S Singh, I Singh, G Singh, S Patil, K Saxena (2012). Breeding for
earliness in pigeonpea: Development of new determinate and non-determinate
lines. Crop Sci 52:2507‒2516
Virk PS, GS Khush, SS
Virmani (2003). Breeding strategies to enhance heterosis in rice. In: Hybrid Rice for Food Security, Poverty
Alleviation and Environmental Protection, pp:21‒29.
Virman SS, CX Mao, B Hardy (Eds). International Rice Research Institute Los Banos,
Philippines
VSN International (2020). Genstat
for Windows, 20th edn. VSN International, Hemel Hempstead, UK
Yohane EN, H Shimelis, M
Laing, I Mathew, A Shayanowako (2020). Phenotypic Divergence analysis in
pigeonpea [Cajanus cajan (L.)
Millspaugh] germplasm accessions. Agronomy 10:1682–1699