Ind J Plant Physiol.

DOI 10.1007/s40502-016-0257-9

ORIGINAL ARTICLE

Physiological characterization and grain yield stability analysis

of RILs under different moisture stress conditions in wheat
(Triticum aestivum L.)
Harikrishna1 G. P. Singh1,2 Neelu Jain1 P. K. Singh1 S. V. Sai Prasad1
Divya Ambati1 T. R. Das1 Arun Kumar1 Javiad Akther Bhat1
B. Amasiddha1 Priyanka Vijay1 Nivedita Sinha1 P. C. Mishra3
S. C. Misra4 K. V. Prabhu1

Received: 14 October 2016 / Accepted: 27 October 2016

Indian Society for Plant Physiology 2016

Abstract Drought stress is the major environmental constraint contributing to grain yield instability of wheat. In
the present study, the recombinant inbred line population
derived from DBW43/HI1500 cross was characterized for
various morpho-physiological traits as well as grain yield
stability analysis under moisture stress. The population was
evaluated for grain yield under three different moisture
stress environments viz., restricted irrigation, rainfed and
late sown rainfed during the cropping season of year
20132014. Based on principal component analysis, the
first five components explained over 60.40% of genetic
variation. Grain yield per plot showed significant correlation with biomass and physiological traits viz., NDVI3,
NDVI4, NDVI5, CT1, CT2 and CT3. The combined
analysis of variance on grain yield data showed that mean
squares of environments, genotypes and GEI were highly
significant (p \ 0.01). To determine effects of GEI on
grain yield, data were subjected to AMMI and GGE biplot
analysis, which identified G4, G69, G28, G67, G55 and
G112 as the most stable and high yielding genotypes.
Hence, the physiological traits NDVI and CT can be
effectively used to screen out the line for drought tolerance.

Electronic supplementary material The online version of this

article (doi:10.1007/s40502-016-0257-9) contains supplementary
material, which is available to authorized users.
& K. V. Prabhu
kvinodprabhu@rediffmail.com
1

Indian Agricultural Research Institute, New Delhi, India

Indian Institute of Wheat and Barley Research, Karnal, India

Jawaharlal Nehru Krishi Vishwa Vidyalaya, Jabalpur,

Madhya Prasad, India

Agharkar ResearchInstitute, Pune, Maharastra, India

In addition, the stable wheat genotypes identified could be

used in future wheat breeding programme.
Keywords Physiological traits
Drought
Yield stability

PCA
Ideal genotype

Introduction
Wheat (Triticum aestivum L.) is one of the worlds largest
cereal crops, and most important staple food of about two
billion people (36% of the world population). Globally,
20% of the food calories consumed and 55% of the carbohydrates are provided by wheat (Breiman and Graur
1995). It is the second most important source of food and
income after rice in India. However, this crop is subjected
to number of biotic and abiotic stresses leading to considerable losses in annual wheat production, among which
drought and heat stress are of major significance. In the
world it has been reported that 65 million ha of wheat area
is subjected to drought stress (FAO 2013). Although 80%
of the wheat crop in India is estimated to be cultivated
under irrigated conditions (Reynolds et al. 1999), *66% of
the crop receives only partial (12) irrigation, subjecting
the wheat crop to water stress, which causes reduction in
grain yield (Joshi et al. 2007). So, it is an immediate
requirement to develop wheat genotypes with traits to
withstand serious drought stress as well as produce higher
grain yield under drought stress conditions.
In India, breeding for drought and heat tolerance is an
important abiotic stress aspect. The relationship between
Canopy Temperature (CT) and grain yield under drought
stress conditions is well established (Olivares-Villegas
et al. 2007; Lopes and Reynolds 2010; Lopes et al. 2012).
Cooler canopy temperature (CT) is associated with both

123

Ind J Plant Physiol.

drought and heat tolerance (Pinto et al. 2010). Stay-green

in the post anthesis phase is reported to be associated with
drought tolerance in several crops (Borrel et al. 2000;
Campos et al. 2004) and is utilized for breeding drought
tolerant cultivars in wheat (Christopher et al. 2015).
Canopy temperature (CT) and Normalized Difference
Vegetative Index (NDVI) have been effectively combined
for rapid screening of drought and heat tolerance in wheat
(Reynolds et al. 2007). Better understanding of the genetic
basis of morphological and physiological trait variability
will improve the efficiency of wheat for drought tolerance.
Among the number of methods available for yield stability
analysis the Additive Main effects and the Multiplicative
Interaction (AMMI) model have recently received more significance, as it overcomes the limitations of univariate models
as well as are more informative compared to other methods
(Shukla 1972; Gauch 1988; Zobel et al. 1988). The AMMI
model is comprised of additive main effects of genotype and
environment, and the multiplicative effect of GE interaction
(Zobel et al. 1988). To further graphically analyze GE interaction GGE biplot (genotype main effect plus genotype x
environment interaction) has been proposed and were found
useful for the analysis of GE interactions (Ahmadi et al. 2012;
Butron et al. 2004; Farias et al. 2016; Karimizadeh et al. 2013;
Laffont et al. 2007; Samonte et al. 2005).
By keeping the above into consideration the present
investigation was intended to characterize the recombinant
inbred line (RIL) population derived from parental cross
DBW43/HI1500 for moisture stress using morpho-physiological traits as well as to identify wheat genotypes with
high and stable yield across different moisture stress
conditions.

Materials and methods

Plant material
The plant material in the present study comprised of 157
recombinant inbred lines (RILs), two parents and a local
check variety HD2987 (Supplementary Table S1). The
RILs are derived from DBW43 (BABAX/LR42//
BABAX*2/3/VIVITSI) and HI1500 (selection in HW
2002*2//STREMPALLI/PNC 5) cross. Both parents possess several unique morphological and physiological differences in addition to yield and yield components.
DBW43 is moderately tolerant to drought stress, where as
HI1500 is highly tolerant.
Field experiment and observation recorded
The field trial was evaluated at three different moisture stress
environments viz., restricted irrigation (RI; two irrigation)

123

(E1), rainfed irrigation (RF; one pre sowing irrigation) (E2)

and late sown rainfed (LS-RF; one pre sowing irrigation)
(E3) during Rabi season of 20132014 at the Wheat
Research Farm of Division of Genetics, Indian Agricultural
Research Institute (IARI), New Delhi, India. The experiment
was laid out in alpha-lattice design with two replications in
one and half meter rows two lines each, and phenotyping was
carried out for a number of traits. The standard cultivation
practices prescribed for wheat were followed precisely.
The data were recorded on randomly selected five representative plants in all the genotypes in each replication
for various morpho-physiological traits. The recording of
observation for morphological traits viz., days to heading,
plant height, spike length, grain weight per ten spike (GW/
10SP), biomass, yield per plot; and physiological traits viz.,
canopy temperature (CT) and Normalized Difference
Vegetative Index (NDVI) were carried out following
standard trait dictionary of CIMMYT (Pask et al. 2012).
The NDVI are recorded at different stages of plant growth
viz., NDVI1 (vegetative stage), NDVI2 (booting stage),
NDVI3 (heading stage), NDVI4 (grain filling stage),
NDVI5 (double-dough stage) and NDVI6 (maturity stage).
CT is also recorded at different plant growth stages viz.,
CT1 (vegetative stage), CT2 (heading stage), CT3 (grain
filling stage) and CT4 (double-dough stage). CT was
measured using hand-held infrared meter (Ayeneh et al.
2002). NDVI was measured with the help of hand held
Trimble GreenSeeker. The mean values of the data
obtained at the three different drought stress environments
were used for the various statistical analyses.
Statistical analysis
The data collected were subjected to compute Pearson
coefficients of correlation among the sixteen morphophysiological quantitative traits using the SPSS 16.0 software. Principal component analysis (PCA) was used to
detect underlying sources of morphological variability, and
to investigate patterns of genetic diversity (Mohammadi
and Prasanna 2003). Bray-Curtis distances and unweighted
pair group method with arithmetic mean (UPGMA) was the
clustering method and all these analyses were done using
the PAST software (Hammer et al. 2001).
To determine the effects of GEI on grain yield, the yield
data recorded from all three environments (E1, E2 and E3)
were subjected to AMMI and GGE biplot analysis using
Gen Stat 14th edition (VSN International, Ltd, Hemel
Hempstead, UK) and R software 3.3.0 respectively. Then
GGE biplot were used to graphically show the genotypes
and environments (Yan 2001). Angles between environment vectors were used to judge correlations (similarities/
dissimilarities) between pairs of environments (Yan and
Kang 2003).

Ind J Plant Physiol.

was contributed by plant height, spike length and biomass.

Lastly, the variability of PC5 was mainly contributed by
grain weight per ten spike and NDVI6.

Results and discussion

PCA analysis
Principle component analysis (PCA) is a viable technique
that provides information about traits which the breeder
needs to take into consideration while breeding for specific
purpose (Salimi et al. 2012). In our study, PCA of the
morpho-physiological traits revealed that first five PCs
explained 60.40% percent of variation among 160 wheat
genotypes, and these results were supported by the findings
of Siahbidi et al. (2013) and Mishra et al. (2015), who
studied wheat genotypes of Iran and India, respectively
(Table 1). The first principal component (PC1) accounted
for 20.31% of total variance, and characters that contribute
more positively to this component were NDVI3, NDVI4,
NDVI5 and grain yield per plot. The second principal
component (PC2), accounted for an additional 14.96% of
the total variation and is primarily contributed by NDVI1,
CT3 and CT4. Finally, third, fourth and fifth principal
component (PC3, PC4, and PC5) contributed around 10.22,
7.83 and 7.08%, respectively of the variability present
among the accessions for the traits used in the present
study. The PC3 explained the patterns of variation in DH,
NDVI2, CT1 and CT2, and for PC4 the maximum variation
Table 1 Eigenvectors, eigen values, total and cumulative variability
(%) for 160 genotypes of wheat based on morpho-physiological and
yield traits
Principal component (axes)
Eigen value
Variability (%)
Cumulative (%)
Traits

PC1

PC2

PC3

PC4

PC5

3.25

2.39

1.63

1.25

1.33

20.31
20.31

14.96
35.27

10.22
45.49

7.83
53.32

7.08
60.4

Eigenvectors

DH

0.26

0.08

0.30

0.27

0.21

PHT

0.12

-0.09

-0.05

0.63

0.19

SL

-0.01

0.29

0.07

0.44

0.30

GW

-0.06

0.09

-0.41

-0.10

0.61

NDVI1

0.09

0.39

-0.11

-0.17

-0.02

NDVI2
NDVI3

0.34
0.32

0.05
0.19

0.45
0.23

0.06
-0.07

-0.01
-0.33

NDVI4

0.45

0.04

-0.05

-0.11

0.05

NDVI5

0.41

0.02

-0.13

-0.18

0.24

NDVI6

0.23

-0.16

0.10

-0.28

0.33

CT1

-0.12

-0.40

0.24

-0.16

CT2

-0.25

-0.28

0.13

0.11

CT3

-0.27

0.32

0.24

-0.03

0.01

CT4

0.13
0.075

-0.20

0.49

0.01

0.07

-0.03

Biomass

0.17

-0.28

-0.21

0.34

-0.25

Yld

0.19

0.03

-0.51

0.11

-0.30

Correlation among the traits

Phenotypic correlation coefficients among the sixteen
morpho-physiological traits of the wheat genotypes are
given in Table 2. Significant positive correlations were
observed for traits like biomass, NDVI3, NDVI4, NDVI5
and NDVI6 with yield, similar to that observed by ElHendawy et al. (2015). Canopy temperatures (CT) at both
vegetative and reproductive stages viz., CT1, CT2, CT3
and CT4 were negatively correlated with yield. Biomass
was found to be positively correlated with PH, NDVI2,
NDVI3, NDVI4, NDVI5 and NDVI6, which is also
reported earlier by Cammarano et al. (2011). CT at all the
four stages was found to be positively correlated with each
other and a negative significant correlation was observed
with traits like biomass, NDVI and grain yield per plot,
similar to that observed by Ramya et al. (2016). However,
GW per 10 spikes showed positive and negative correlation
with NDVI and CT, respectively. Similarly, SL showed
negative and positive correlation with CT and NDVI,
respectively. Hence, one of the best strategies to increase
the grain yield in wheat crop under moisture stress is
through the genetic manipulation of physiological traits
viz., NDVI and CT.
AMMI analysis
The combined analysis of variance (ANOVA) for grain
yield at three different moisture stress environments
showed that mean squares of environments, genotypes and
genotype 9 environment interaction (GEI) were highly
significant (p \ 0.01) which accounted for 71.68, 5.40 and
10.29% of model sum of squares (SS), respectively
(Table 3). Similar results were reported by (Verma et al.
2015; Mehari et al. 2015; Kaya et al. 2002). This case,
along with a highly significant GEI, required using of
stability analysis. The environments explained the large
variation for yield and indicate the diverse nature of the
environments, with large differences among environmental
means causing most of the variation in grain yield. The
environment yield means (averaged across genotypes)
varied from 395.1 g/plot under late sown rainfed conditions to 760.4 g/plot under restricted irrigation conditions
(Table 4), whereas genotype yield means (averaged across
environments) varied from 424.8 (G100) to 674.2 g/plot
(G69) (Table 1). In our study, the magnitude of SS due to
GE interaction was 1.91 times greater than that for genotypes, which revealed the significant differences in genotypic response across environments as well as indicates the

123

0.29**

-0.04

-0.24**

Table 3 AMMI analysis of variance for grain yield tested at eight

environments

-0.34**

-0.19*
0.02

-0.29**

0.02

-0.21**
0.09

0.13

0.23**

0.26**
0.16*

0.07

0.08

0.12

0.12

0.17*
0.03

0.06

-0.11

-0.09

0.09

-0.04

-0.04
-0.07

0.24**

0.01

BIOMASS

YLD

Ind J Plant Physiol.

Source

df

MS

Total

959

31,118**

Treatments

479

54,437**

Genotypes

159

10,135**

0.51**

0.10

Block

0.02

-0.26**

0.34**
0.04

-0.17*

-0.26**

-0.23**
-0.34**

-0.38**

0.02
-0.01

-0.13
-0.09

-0.12

0.33**
-0.19*

-0.04

-0.28**
0.18*

-0.05

-0.11
-0.11

-0.11

CT3

CT4

Environments

10.29

IPCA1
IPCA2

160
158

11,235**
9063*

58.52
41.47

0.31**

-0.08

0
477

0
17,902

** 0.01P

existence of mega-environment (Yan and Hunt 2002;

Mohammadi et al. 2009). The AMMI analysis partitioned
the SS of GEI into two interaction principal components
axes (IPCA), and both of these IPCA were significant
(Table 3). Results from AMMI model showed that the
IPCA1 of the interaction captured 58.52% of the interaction sum of squares. Similarly, the second interaction
principal component axis (IPCA2) explained a further
41.47% of the genotype-environment interaction sum of
squares. At the same time, IPCA1 had sum of squares
greater than that of genotypes. The mean squares for the
IPCA1 and IPCA2 were significant at p = 0.01 and
p = 0.05 level, respectively and cumulatively contributed
to 99.99% of the total genotype-environment interaction.
Based on AMMI analysis, the best ten selections in terms
of yield performance under moisture stress in each environment were compared (Table 4). Some of the selections
were consistently performing in more than one environment
under moisture stress. The G4 showed the best and consistent
yield performance as well as ranked within the best ten
selections in two out of three environments viz., E1 and E2.
According to AMMI analysis and mean performance (grain
yield per plot), the G69 were the most stable and high
yielding genotype across all the three moisture stress environments followed by G4, G28, G67, G55 and G112. These
genotypes show high yielding and consistent performance in
all the three moisture stress conditions.

0.17*

-0.38**
-0.13

-0.21**

0.28**

0.29**

0.59**

-0.34**

-0.29**
-0.22**
0.16*
0.28**
0.36**

-0.13
0.24**

0.00
0.33**
0.44**

0.02

-0.07

0.48**

-0.01

-0.25**
0.09

0.11
-0.04

0.01
0.25**

0.21**

0.17*

0.26**

0.07

0.12

-0.29**

-0.08
-0.24**
0.01

0.03

0.01

0.07
0.09
0.13
0.07

-0.12

-0.02
0.05

0.05
0.27**

0.05
0.12

0.29**
0.27**

-0.02
0.10

0.42**

0.35

9659**

GGE biplot analysis

** 0.01P

* 0.05 P

YLD

BIOMASS

CT4

CT3

CT2

NDVI6

CT1

NDVI4

NDVI5

NDVI3

NDVI2

NDVI1

GW

SL

PH

DH

34,280**

* 0.05 P

0.16*

0.02

-0.01
0.01
0.10

-0.08

-0.09

-0.07
0.13

CT2
NDVI1

NDVI2

NDVI3

NDVI4

NDVI5

NDVI6

CT1
GW
SL
PH

5.4
71.68

318

Residuals

DH

Table 2 Pearson correlation coefficients among the morpho-physiological and yield traits

10,696,155**

87.37

Interactions

Error

123

Explained %

The GGE biplot indicates the best performing genotype in

each environment and group of environments (Fig. 1). The
polygon is formed by connecting the scores of the genotypes furthest from the origin (Fig. 1), with all remaining
genotypes within it, and indicates which genotypes won
where based on their association with the site scores (Yan

Ind J Plant Physiol.

Table 4 Mean yield
performance in an environment
(Em) and first ten AMMI
selections per environment

Environment

Em

10

E1 (RI)

760.4

G99

G1

G19

G58

G98

G158

G4

G27

G38

G88

E2 (RF)

565.0

G69

G80

G4

G67

G123

G93

G55

G28

G91

G31

E3 (LS-RF)

395.1

G118

G126

G58

G42

G73

G78

G75

G56

G19

G59

Fig. 1 Polygon view of GGE biplot showing which won where

pattern for genotypes and environments

et al. 2000). The genotype at the vertex of the polygon

performs best in the environment falling within the sectors
(Yan 2002; Yan and Tinker 2006). The biplots revealed the
existence of GE crossover as well as mega-environment for
grain yield. The hexagon has seven genotypes viz., G99,
G136, G71, G34, G69, G4, and G1 at the vertices. The G99
performed best in E1, while G34 being the best in E3. G69
performed best in E2. The biplot is effectively divided into
seven sectors by the equality lines, of which three retained
all the environments. Thus the testing environments may be
partitioned into three mega-environments: one mega-environment comprise of E1with G99 as the winning genotype. Second mega-environment encompassed E3 with
G34 as the winning genotype, while last mega-environment
was represented by E2 with G69 as the winning genotype
(Fig. 1).
The genotype ranking for grain yield is shown on the
graph of so-called ideal genotype (Fig. 2). An ideal
genotype is high performer with high stability across
environments (Yan and Tinker 2006). Such an ideal
genotype is defined by having the greatest vector length of
the high yielding genotypes and with zero GEI, as represented by an arrow pointing to it (Fig. 2). A genotype is
more desirable if it is located closer to the ideal genotype.
Thus, using the ideal genotype as the centre, concentric
circles were drawn to help visualize the distance between
each genotype and the ideal genotype (Yan and Tinker
2006). Hence, in the ranking of genotypes for both mean

Fig. 2 GGE biplot based on genotype-focused scaling for comparison of the genotype with ideal genotype

yield and stability performance across the three environments G4 followed by G28, G67, G112, G55 and G69 were
ranked closest to ideal genotype, indicating them as the
most desirable genotypes out of 160 genotypes studied
(Karimizadeh et al. 2013).
Based on mean performance (grain yield kg/ha), AMMI
and GGE biplot analysis it is evident that genotypes G4,
G69, G28, G67, G55 and G112 were the highly adapted,
most stable and high yielding genotypes in all studied
environments. The use of these wheat genotypes by farmers would result in stable performance under moisture
stress conditions. Hence, these genotypes can also be used
to identify QTLs/genes for above morpho-physiological
traits contributing to drought tolerance as well as can be
used as donors for breeding in drought tolerance.

Conclusion
RIL population used in this study revealed considerable
segregation and variability for morphological and physiological traits, which can lead to their effective and potential
utilization in the breeding for drought stress tolerance. The
positive and negative significant correlation of grain yield
per plot with NDVI and CT, respectively indicated that the
strategy to increase grain yield in wheat crop under moisture stress involves genetic manipulation of physiological

123

Ind J Plant Physiol.

and agronomical traits. On the basis of grain yield stability

RILs G4, G69, G28, G67, G55 and G112 were highly
adapted, most stable and high yielding under all three
moisture stress environments viz., RI, RF and LS-RF and
can be used in convergent wheat breeding programme to
develop drought tolerant varieties.

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