Genetic analysis for quality traits in aromatic rice (Oryza sativa L.

) genotypes

Anita C.Solanke., P.B.Patel., Sagar Jadhav and Pooja.k.Patel

Department of Genetics and Plant Breeding, N.M. College of Agriculture,

Navsari Agricultural University,
Navsari - 396 450 (Gujarat)
ABSTRACT
The present investigation in aromatic rice (Oryza sativa L.) was undertaken for studying the
magnitude of gene action in four cross combination for yield and quality traits deploying
generation mean analysis following six parameter model for parents (P 1 and P2), F1, F2, BC1
and BC2 generations during three crop season. The results of the scaling tests revealed that
the additive-dominance model was inadequate for all of the characters evaluated in all of the
four crosses, suggested the existence of epistasis in the inheritance of these characters. On the
basis of six parameters model, main effect viz., mean (m), additive (d) and dominance (h) and
all three digenic interactions viz., additive x additive (i), additive x dominance (j) and
dominance x dominance (l) were significant for grain yield per plant in cross-III (IET-26214
x GAR-1); for protein content in cross-III (IET-26214 x GAR-1) and cross-IV (IET-26215 x
GNR-2); for amylose content in cross-II (IET-24617 x NWGR-9081); for milling in cross-III
(IET-26214 x GAR-1); for hulling in all crosses except cross-I (GR-104 x IET-26215)
indicated the involvement of additive, dominance as well as epistasis interaction for
controlling this trait. The duplicate type epistasis was observed in majority of all traits in all
crosses. The present study demonstrates the importance of additive, dominance and epistatic
gene effects for the inheritance of almost all the yield as well as quality characters studied.

Key words: Aromatic Rice, gene action, scaling test, generation mean, quality traits.

INTRODUCTION

Rice is a premium food grain crop and is often the main source of calories and
principal food of millions of people. As more than 50 per cent of the world population
depends on rice for their staple diet. Nowadays, the quality considerations assume enhanced
importance, especially in the countries which are self-sufficient in their production.
Aromatic rices constitute a small but special group of rices which are considered
best in quality. Among the quality rices, Basmati is the unique aromatic quality rice. It is a
nature’s gift to Indian sub-continent. Super fine varieties of rice with aromatic grains,
superior cooking and eating characteristics get a premium price in the market. Basmati
fetches comparatively a higher price, which is not affordable by rural consumers and those
with low income. Therefore breeding for quality has to be emphasized to bring out fine
grained rice fetching optimum price, which will be easily within the reach to the rural
consumers. Hence the rice grain quality has become a necessity for the acceptance of
varieties to be released.Genetic improvement for rice has thoroughly been studied worldwide.
To achieve genetic improvement of yield and quality traits, it is imperative to have
knowledge about the nature of gene interactions for different characters. The generation mean
analysis has been considered to be one of the best methods for estimating the different
components of genetic variance and presence or absence of epistasis. Therefore, the study of
genetics of yield and quality traits is important to formulate a breeding programme to
improve yield while maintaining the quality of rice. Keeping in view the above mentioned
facts, present investigation was formulated to study to the gene action for yield and quality
traits in aromatic genotypes of rice.

MATERIALS AND METHODS

The material comprising of seven diversified aromatic and non-aromatic elite

lines of rice (GR-104, IET-26215, IET-24617, NWGR-9081, IET-26214, GAR-1 and GNR-
2). The four crosses (GR-104 x IET-26215, IET-24617 x NWGR-9081, IET-26214 x GAR-1
and IET-26215 x GNR-2) obtained by crossing of seven diverse parents during summer 2017
at Main Rice Research Centre, Navsari Agricultural University, Navsari. Backcrossing was
done in kharif-2017 with its respective parents. Selfing of F1s was done in the same season
(kharif-2017) to get F2s. The evaluation trial was conducted in kharif-2018 at Main Rice
Research Centre, Navsari Agricultural University, Navsari. The experimental material
consisting of six generations (P1, P2, F1, F2, BC1 and BC2) of each of the four crosses were
sown during kharif-2018 in compact family block design with three replications. Each
replication was divided in four compact blocks. Each four crosses consisting of six
generations were randomly allotted to each plot within a block. Each plot consisted of one
row (10 plants) of parents and F1s, two rows of the backcrosses and four rows of the F2
generations of each cross. The recommended package of practices was followed to raise a
good crop. Observations were recorded on yield and quality traits viz., Grain yield per plant,
kernel length (mm), kernel breadth (mm), kernel L/B ratio, protein content (%), amylose
content (%), milling (%), hulling (%) and head rice recovery (%). Kernel dimensional
analysis was done with the help small millimeter scale. The simplified calorimetric method
described by Juliano9 was followed for the estimation of amylose content. Generation mean
analysis was conducted using six generations viz. parental (P1 and P2), F1, F2, and
backcrosses (BC1 and BC2) of four selected crosses. Average values were subjected to
scaling test presenting in table no.1
RESULTS AND DISCUSSION

In the present investigation, all the four scaling tests (A, B, C and D) were
highly significant for all the characters under study, indicating inadequacy of additive-
dominance model to explain the inheritance of yield and quality characters. The values for
individual scaling tests and estimates of mean (m), additive gene effect (d ), dominance gene
effect (h) and epistatic interactions viz., additive x additive (i), additive x dominance (j)and
dominance x dominance( l )interactions are presented in tables 1 and 2 respectively.

On the basis of individual scaling test A, B, C and D the additive-dominance

model was found inadequate for description of variation in generation mean for all traits in
four crosses, either the entire four or any three, two or one individual scaling test (out of A, B
C and D) were found significant which indicated the presence of digenic interaction which
implies that the additive - dominance model is inadequate.

When the simple additive-dominance model failed to explain the variation

among the generation means, a six parameter model involving three digenic interaction
parameters proposed by Hayman (1958) was applied.

The result obtained from six parameter model revealed that in addition to the
significance of mean (m), additive (d) and dominance (h) effects and all the three digenic
interactions additive x additive (i), additive x dominance (j) and dominance x dominance (l)
were significant for grain yield per plant in cross-III; for protein content in cross-III and
cross-IV; for amylose content in cross-II; for milling in cross-III; for hulling in all crosses
except cross-I. These results are in agreement with those obtained by Nayak et al. (2007),
Singh et al. (2007), Roy and Senapati (2011), Samak et al. (2011), Kiani et al. (2013),
Montazeri et al. (2014), Shahid et al. (2014), Patel et al. (2015), Rani et al. (2015), Sultana et
al. (2016) and Kumar et al. (2017)

The additive (d) effect found significant and positive in cross-I grain yield per
plant, protein content, milling (%); in case of cross-II for kernel L/B ratio, grain yield per
plant, hulling (%) and head rice recovery (%); in case of cross-III for kernel L/B ratio, grain
yield per plant, protein content, head rice recovery (%); in case of cross-IV for kernel length,
kernel L/B ratio, protein content, hulling (%) and head rice recovery (%).

Similarly, the additive (d) effect found significant and negative in cross-I for
kernel breadth, amylose content; in case of cross-II for kernel length, kernel breadth, amylose
content; in case of cross-III for kernel breadth, amylose content, milling (%) and hulling (%);
in case of cross-IV for kernel breadth and amylose content.

The additive component of variation can be exploited by simple pedigree

selection. Mass selection for several early generation aimed at the improvement of
heterozygous population by modifying the frequencies of desirable genes followed by single
plant selection in the resulting material would be cheapest and quickest procedure. However,
the presence of non-fixable (h, j and l) component together with duplicate type of epistasis
may cause delay in the improvement in this trait through selection in early generations. Under
this situation, progeny could be achieved and the selection is delayed to later generations.
These results are in agreement with those obtained by Patel et al. (2015), Rani et al. (2015),
Sultana et al. (2016) and Kumar et al. (2017) for days to flowering, plant height, productive
tillers per plant and harvest index .

The dominance (h) effects found positive and significant for milling (%), head
rice recovery (%), in case of cross-II for amylose content and milling (%); in case of cross-III
for milling (%); in case of cross-IV for grain yield per plant, kernel length, milling (%) and
head rice recovery (%). These results are in agreement with those obtained by Shahid et al.
(2014) and Patel et al. (2015) for productive tillers per plant, grain yield per plant and kernel
L/B ratio; Rani et al. (2015) and Sultana et al. (2016) for kernel L/B ratio and Kumar et al.
(2017) for kernel L/B ratio.

Significant and negative dominance (h) effect was observed for amylose
content in cross-I; grain yield per plant and hulling (%) in cross-II; grain yield per plant and
protein content in cross-III; protein content, hulling (%) in cross-IV, respectively.

The sign of dominance x dominance (l) effect was positive in case of cross-I
for kernel L/B ratio and protein content; in case of cross-II for grain yield per plant and
hulling (%); in case of cross-III for grain yield per plant and protein content and in case of
cross-IV for kernel L/B ratio, protein content and hulling (%). The sign of dominance x
dominance (l) component was positive in these crosses indicating their enhancing effect in
the expression of that character in all four crosses of rice. Non fixable gene effect were
important in the expression of these traits in these crosses could be exploited by bi-parental
mating of recurrent selection or the use of population improvement concept as an alternative
to conventional method.
 The sign of dominance x dominance (l) effect was negative for kernel breadth,
amylose content and head rice recovery in cross-I; kernel length, kernel breadth, amylose
content and milling (%) in cross-II; kernel breadth, amylose content, milling (%) and hulling
(%) in cross-III; kernel breadth, grain yield per plant, amylose content, amylose content and
milling (%) in cross-IV indicating the reducing effect in the expression of these characters.
The sign of dominance x dominance (l) component was positive in above characters
indicating their enhancing effect in the expression of that characters in all four crosses of rice.

The additive x additive (i) interaction had greater effect as compare to additive
x dominance (j) and dominance x dominance. additive x additive (i)effect found significant
and positive for for kernel length in cross-II, for grain yield per plant in cross-IV, for
amylose content in cross-II, for milling (%) in cross-I, cross-II and cross-III, for hulling (%)
in cross-III. This indicated better response to selection pressure in population for these
characters. In these crosses, improvement could be made by cyclic method of breeding in
which desirable recombinants are selected and intercrossed to pool the favourable genes for
synthesizing the elite population.

In the present study, the significant additive and additive x additive epistasis
was observed in cross-II for kernel length, in cross-II and cross-III for grain yield per plant;
in cross-III and cross-IV for protein content; in cross-II for amylose content, in cross-I and
cross-III for milling (%); in cross-II, cross-III and cross-IV for hulling (%); in cross-II and
cross-IV for head rice recovery (%). These results are in agreement with those obtained by
Shahid et al. (2014), Patel et al. (2015) for grain yield per plant and kernel L/B ratio; Rani et
al. (2015), Sultana et al. (2016) for and kernel L/B ratio and Kumar et al. (2017) for kernel
L/B ratio.

The duplicate epistasis was observed in almost all traits in all crosses, making
it difficult to fix genotypes with increased level of character manifestation because the
positive effect of one parameter would be cancelled out by the negative effect of another
parameter.

CONCLUSIONS
Generation mean analysis was carried out by evaluating six basic populations
(P1, P2, F1, F2, B1 and B2) of four cross combinations viz., (GR-104 x IET-26215, IET-
24617 x NWGR-9081, IET-26214 x GAR-1 and IET-26215 x GNR-2) for grain yield and
quality traits. All the four crosses were subjected to A, B, C and D scaling tests to sort out the
model (interacting crosses) for the characters concerned were further subjected to six
parameter models to estimate the main gene effects; [m], [d] and [h] and their interactions [i],
[j] and [l] involved in the cross for the expression of respective trait under study. Scaling test
(A, B, C and D) was applied to test the inadequacy of additive-dominance model. Significant
deviation of the scale (s) from zero indicates the presence of epistatic interaction in respective
crosses. It is interesting to note that all the four crosses scored significant values for all the six
components of gene effect for grain yield and quality traits. The result obtained from six
parameter model revealed that in addition to the significance of mean (m), additive (d) and
dominance (h) effects and all the three digenic interactions additive x additive (i), additive x
dominance (j) and dominance x dominance (l) were significant for grain yield per plant in
cross-III; for protein content in cross-III and cross-IV; for amylose content in cross-II; for
milling in cross-III; for hulling in all crosses except cross-I. Since, the sign of dominance (h)
and dominance × dominance (l) for almost all traits of these four crosses was opposite in all
crosses, therefore, the nature of epistasis was identified as duplicate in these crosses.
Duplicate epistasis as observed may postponed single plant selection and biparental mating or
diallel selective mating could be followed where in few cycles of crossing of promising
segregants in F2 and onward generations that might help in the incorporation of desirable
genes into a single genetic background. In other words, this type of epistasis tends to cancel
or weaken the effect of each other in hybrid combination and hinders the progress made
under selection and therefore, selection would have to be differed till later generations of
segregation where dominance effects are dissipated. However Biparental mating, recurrent
selection and diallel selective mating system might be profitable in exploiting both additive
and non additive gene action to obtain desirable recombinants.

REFERENCES
Cavalli, L. L. (1952). An analysis of linkage of quantitative inheritance.
In: Quantitative inheritance (Eds. E. C. R. Reeve and C. H.
Wedelington). HMSO, London, pp.135-144.
Hayman, B. I. (1958). The separation of epistatic from additive and
dominance variation in generation means. Heredity, 12: 371-390.

Kumar, P. S.; Saravanan, K. and Sabesan, T. (2017). Generation mean

analysis for yield and grain quality characters in rice (Oryza
sativa L.). Plant Archives, 17(1): 557 -560.
Nayak, A. R.; Chaudhury, D. and Reddy, J. N. (2007). Genetics of yield
and yield components in scented rice. Oryza, 44 (3): 227-
230.

Patel, V. J., Mistry, P. M., Chaudhari, M. H. and Dave, V. D. (2015).

Combining Ability Analysis in Rice ( Oryza sativa
L.). Trends in Biosciences, 8(1): 82-87.

Sultana, R .; Ansari, N. A. ; Ramesha, M . S .; Shankar, A. S. and murali

Krishna, K . (2016) . Generation mean analysis of quantitative traits in restorer lines
of rice (Oryza sativa L.) . Asian journal of bio science, 11(1): 151 - 161.

Hayman, B. I. and Mather, K. (1955). The description of genic interactions in

continuous variation. Biometrics., 11(1): 69-82.

Sabesan, T. (2005). Studies on the genetics of quality traits in rice (Oryza sativa L.).
Ph.D. Thesis, Annamalai University, Annamalainagar, India.
Table 1: Scaling test for yield & quality characters in aromatic rice

G rai n yi eld p er pl an t (g)

S cal e Cross- I Cross- II Cross- III Cross- IV
S cali n g T ests (Math er, 1949)
A 4.52** ± 1.73 -12.17** ± 2.10 6.26** ± 1.83 0.66 ± 1.81
B -11.51** ± 1.97 -12.37** ± 1.86 -10.15** ± 2.22 9.06** ± 1.65
C -7.01* ± 3.47 -9.22* ± 3.68 15.69** ± 3.85 7.36** ± 2.84
D -0.01 ± 1.67 7.66** ± 1.76 9.79** ± 1.88 -1.18* ± 1.45
K ern el l en gth (mm) )
A 0.43** ± 0.16 0.51** ± 0.16 0.62** ± 0.14 0.69** ± 0.14
B -0.51** ± 0.11 0.50** ± 0.11 -0.11 ± 0.22 0.03 ± 0.19
C 0.27 ± 0.25 0.37 ± 0.24 0.22 ± 0.28 0.82** ± 0.25
D 0.17 ± 0.12 -0.32** ± 0.11 -0.15 ± 0.15 0.05 ± 0.14
K ern el b read th (mm)
A 0.24** ± 0.08 0.28** ± 0.08 0.12 ± 0.11 0.24** ± 0.07
B 1.50** ± 0.23 1.49** ± 0.22 1.36** ± 0.23 1.65** ± 0.22
C 1.60** ± 0.28 1.75** ± 0.28 1.55** ± 0.30 1.84** ± 0.27
D -0.07 ± 0.18 -0.01 ± 0.18 0.03 ± 0.18 -0.02 ± 0.17
K ern el L / B rati o
A -0.21 ± 0.18 -0.20 ± 0.17 0.12 ± 0.19 -0.03 ± 0.15
B -1.69** ± 0.29 -1.22** ± 0.32 -1.40** ± 0.28 -1.68** ± 0.27
C -1.64** ± 0.38 -1.57** ± 0.39 -1.19** ± 0.39 -1.59** ± 0.36
D 0.13 ± 0.23 -0.08 ± 0.24 0.05 ± 0.22 0.06 ± 0.22
Protei n con ten t (%)
A -0.23 ± 0.28 1.15** ± 0.31 -0.96* ± 0.38 -0.95** ± 0.31
B -1.35** ± 0.27 0.11 ± 0.46 -5.10** ± 0.32 -6.85** ± 0.31
C -1.41** ± 0.43 1.27* ± 0.60 2.21** ± 0.55 0.90 ± 0.55
D 0.08 ± 0.20 0.00 ± 0.28 4.13** ± 0.29 4.35** ± 0.24
Amyl ose con ten t (%)
A 1.66* ± 0.80 1.35* ± 0.57 0.83 ± 0.67 2.15** ± 0.83
B 3.03** ± 0.61 3.22** ± 0.58 3.20** ± 0.47 2.91** ± 0.60
C 4.01** ± 1.30 -1.20 ± 0.89 3.20** ± 1.05 3.69** ± 1.24
D -0.34 ± 0.58 -2.88** ± 0.47 -0.41 ± 0.52 -0.68 ± 0.52
Mi l l in g (%)
A 3.74 ± 4.01 -0.84 ± 3.38 -3.81 ± 3.01 18.07** ± 3.16
B -13.40** ± 4.11 1.52 ± 3.32 14.05** ± 3.57 5.48 ± 3.68
C -22.86** ± 6.02 -24.80** ± 5.85 -13.44* ± 5.32 18.97** ± 5.10
D -6.60* ± 2.75 -12.74** ± 2.80 -11.84** ± 2.42 -2.29 ± 2.51
Hu l li n g (%)
A 10.65** ± 3.29 11.07** ± 3.11 -4.84** ± 2.98 9.86** ± 3.24
B -1.63 ± 3.31 -15.23** ± 4.34 21.22** ± 3.09 -16.13** ± 4.41
C 16.83** ± 5.46 30.66** ± 5.63 -14.52** ± 4.61 30.20** ± 5.76
D 3.90 ± 2.24 17.41** ± 2.68 -15.45** ± 2.15 18.24** ± 2.71
H ead ri ce recove ry (%)
A 18.89** ± 3.13 18.84** ± 3.84 12.61** ± 3.85 23.03** ± 2.80
 B 5.21 ± 3.67 -17.75** ± 4.76 -6.36 ± 3.86 -3.31 ± 3.40
C 16.97** ± 5.14 16.86** ± 6.36 6.59 ± 5.93 28.93** ± 4.84
D -3.57 ± 2.34 7.88** ± 2.43 0.17 ± 2.85 4.60* ± 2.16

*, ** Significant at 5% and 1% levels, respectively

Cross-I: GR-104 x IET-26215, Cross-II: IET-24617 x NWGR-9081,

Cross-III: IET 26214- x GAR-1, Cross-IV: IET-26215 x GNR-2

Table No.2: Estimation of gene effects for yield & quality characters using six parameter model
in aromatic rice

S i x paramet er mod el (H ayman , 1958)

G rai n yi eld p er pl an t (g)
Parameters Cross- I Cross- II Cross- III Cross- IV
25.05* 0.7
m ± ± ± ±
* 0.67 26.50** 0.69 28.76** 7 25.98** 0.16
1.0
d ± ± ± ±
5.10** 0.98 2.69* 1.09 6.26** 9 -0.64 0.44
- 3.9
h ± ± ± ±
5.90 3.50 -8.83* 3.72 14.37** 3 10.93** 1.13
- - 3.7
i ± ± ± ±
0.02 3.33 15.32** 3.52 19.59** 6 2.37* 1.08
1.2
j ± ± ± ±
8.01** 1.14 0.10 1.13 8.20** 6 -4.20** 0.49
5.8 -
l ± ± ± ±
6.97 5.23 39.85** 5.69 23.48** 1 12.09** 1.98
T yp es of
- Duplicate Duplicate Duplicate
ep i stasi s
K ern el l en gth (mm)
6.46* 0.0 0.0
m ± ± ± ±
* 5 6.34** 4 6.27** 0.06 6.51** 0.05
0.53* 0.0 0.0
d ± ± ± ±
* 7 -0.30** 7 0.09 0.11 0.30** 0.10
0.2 0.2
h ± ± ± ±
0.25 5 0.37 4 0.14 0.32 - 0.36 0.29
0.2 0.2
i ± ± ± ±
-0.34 3 0.65** 3 0.30 0.31 - 0.11 0.28
0.47* 0.0 0.0
j ± ± ± ±
* 9 0.01 8 0.37** 0.12 0.33** 0.11
0.3 0.3
l ± ± ± ±
0.42 7 -1.66** 6 - 0.81 0.50 - 0.61 0.46
T yp es of
Duplicate
ep i stasi s
K ern el b read th (mm)
2.19* 0.0 0.0
m ± ± ± ±
* 7 2.20** 7 2.22** 0.07 2.23** 0.07
-
d 0.74* ± 0.1 ± 0.1 ± - ±
* 2 -0.70** 1 - 0.79** 0.12 0.82** 0.11
h 0.001 ± 0.3 0.02 ± 0.3 - 0.08 ± 0.36 - 0.02 ± 0.35
 6 5
0.3 0.3
i ± ± ± ±
0.14 6 0.02 5 - 0.07 0.36 0.05 0.35
-
j 0.63* ± 0.1 ± 0.1 ± - ±
* 2 -0.60** 1 - 0.62** 0.12 0.71** 0.11
-
l 1.88* ± 0.5 ± 0.5 ± - ±
* 5 -1.78** 3 - 1.41** 0.55 1.94** 0.52
T yp es of
Duplicate Duplicate Duplicate Duplicate
ep i stasi s
K ern el L / B rati o
3.22* 0.0 0.0
m ± ± ± ±
* 8 3.19** 9 3.16** 0.08 3.20** 0.08
0.96* 0.1 0.1
d ± ± ± ±
* 5 0.53** 6 0.89** 0.15 1.01** 0.14
0.4 0.4
h ± ± ± ±
0.28 6 -0.03 9 - 0.16 0.45 - 0.15 0.45
0.4 0.4
i ± ± ± ±
-0.27 5 0.15 9 - 0.10 0.44 - 0.12 0.44
0.74* 0.1 0.1
j ± ± ± ±
* 7 0.51** 7 0.76** 0.16 0.83** 0.15
2.17* 0.7 0.7
l ± ± ± ±
* 2 1.27 7 1.38 0.70 1.83** 0.67
T yp es of
Duplicate Duplicate Duplicate Duplicate
ep i stasi s
Protei n con ten t (%)
5.12* 0.0 0.1
m ± ± ± ±
* 7 7.95** 0 8.06** 0.10 8.14** 0.10
0.85* 0.1 0.2
d ± ± ± ±
* 4 0.14 0 1.97** 0.20 2.49** 0.14
- 0.4 0.6 -
h ± ± ± ±
0.64 3 0.70 0 - 9.48** 0.61 8.11** 0.52
0.4 0.5 -
i ± ± ± ±
-0.17 0 0.00 6 - 8.27** 0.58 8.70** 0.49
0.56* 0.1 0.2
j ± ± ± ±
* 7 0.52* 6 2.07** 0.22 2.95** 0.20
0.7 0.9 16.50*
l ± ± ± ±
1.74* 0 -1.27 9 14.33** 0.98 * 0.79
T yp es of
Duplicate Duplicate Duplicate Duplicate
ep i stasi s
Amyl ose con ten t (%)
23.80 0.2 0.1 23.71*
m ± ± ± ±
** 3 23.08** 7 24.04** 0.21 * 0.20
-
d 0.94* ± 0.3 ± 0.3 ± - ±
* 5 -0.98** 3 - 1.17** 0.30 1.10** 0.34
1.2 0.9
h ± ± ± ±
-2.65* 4 4.06** 9 - 0.49 1.08 - 0.74 1.15
1.1 0.9
i ± ± ± ±
0.69 6 5.77** 5 0.83 1.03 1.37 1.04
0.4 0.3
j ± ± ± ±
-0.69 3 -0.94* 8 - 1.19** 0.37 - 0.38 0.43
-
l 5.38* ± 1.9 - ± 1.5 ± - ±
* 0 10.34** 9 - 4.85** 1.60 6.43** 1.83
T yp es of
- Duplicate Duplicate Duplicate
ep i stasi s
 Mi l l in g (%)
55.94 0.9 1.1 58.12*
m ± ± ± ±
** 9 59.31** 3 58.66** 0.94 * 0.90
1.9 1.6
d ± ± ± ±
4.91* 0 0.50 4 - 5.15** 1.52 - 1.47 1.75
26.46 5.9 5.8 21.51*
h ± ± ± ±
** 5 41.26** 9 28.73** 5.19 * 5.34
13.20 5.5 5.5
i ± ± ± ±
* 1 25.49** 9 23.67** 4.84 4.58 5.02
8.57* 2.7 2.2
j ± ± ± ±
* 9 -1.18 5 - 8.93** 2.24 6.29** 2.34
-
l ± 9.7 - ± 8.8 - ± 28.14* ±
-3.54 1 26.17** 0 33.91** 8.09 * 8.65
T yp es of
- Duplicate Duplicate Duplicate
ep i stasi s
Hu l li n g (%)
m 73.97** ± 0.88 78.37** ± 0.95 65.51** ± 0.17 79.05** ± 0.35
d -0.19 ± 1.39 9.92** ± 1.88 -10.20** ± 0.41 9.67** ± 0.65
h -3.97 ± 4.94 -29.30** ± 5.74 29.54** ± 1.35 -31.99** ± 2.06
i -7.81 ± 4.48 -34.82** ± 5.35 30.90** ± 1.08 -36.46** ± 1.91
j 6.14** ± 2.15 13.15** ± 2.54 -13.04** ± 0.87 13.00** ± 0.87
l -1.21 ± 7.78 38.98** ± 9.40 -47.28** ± 2.41 42.72** ± 3.33
T yp es of
- Duplicate Duplicate Duplicate
ep i stasi s
H ead ri ce recove ry (%)
m 57.17** ± 0.86 59.00** ± 0.88 52.38** ± 1.03 56.54** ± 0.79
d -1.52 ± 1.59 8.59** ± 1.69 7.47** ± 1.98 6.36** ± 1.46
h 24.96** ± 5.05 3.02 ± 5.55 13.80* ± 6.09 2.11 ± 4.68
i 7.14 ± 4.68 -15.76** ± 4.87 -0.34 ± 5.71 -9.21* ± 4.31
j 6.84** ± 2.32 18.30** ± 2.96 9.49** ± 2.53 13.17** ± 2.06
-
l 31.24**
±
8.19 14.66
±
9.28 -5.90
±
9.89 -10.52
±
7.59
T yp es of
Duplicate - - -
ep i stasi s

*, ** Significant at 5% and 1% levels, respectively

Cross-I: GR-104 x IET-26215, Cross-II: IET-24617 x NWGR-9081,

Cross-III: IET 26214- x GAR-1, Cross-IV: IET-26215 x GNR-2