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International Journal of Agricultural

Science and Research (IJASR)


ISSN(P): 2250-0057; ISSN(E): 2321-0087
Vol. 6, Issue 3, Jun 2016, 415-422
TJPRC Pvt. Ltd.

QUANTITATIVE ANALYSIS OF RESISTANCE TO POWDERY


MILDEW IN ADULT MELON PLANTS
K. T. SHASHIKUMAR1, M. PITCHAIMUTHU2 & R. D. RAWAL3
1

Division of Horticulture, University of Agricultural Sciences, G.K.V.K., Bangalore, Karnataka, India

2,3

Division of Vegetable Crops, Indian Institute of Horticultural Research, Bangalore, Karnataka, India

ABSTRACT
The experimental material consisted of six generations namely P1, P2, F1, F2, BC1P1 and BC2 P2 of four melon
crosses viz., RM 43 x IIHR 121, IIHR 681 x IIHR 121, IIHR 681 x IIHR 122 and Punjab Sunehri x IIHR 122.
Quantitative analysis of resistance to powdery mildew in adult melon plants confirmed the insufficiency of simple
additive-dominance model. Consistently high and negative MPH in crosses RM 43 x IIHR 121 and IIHR 681 x IIHR 121
indicated the importance of dominance effects for the expression of resistance to powdery mildew. Dominance x
dominance (l) gene effect was found to be predominant in the cross Punjab Sunehri x IIHR 122. Dominance (h and/or l)
and additive (d) gene effects were found to be important in the cross RM 43 x IIHR 121. Dominance (h) and additive x
additive (i) gene effects were predominant in crosses IIHR 681 x IIHR 121 and IIHR 681 x IIHR 122. Punjab Sunehri x
largest component of total genetic variance in all the crosses. High braod (H) and narrow (h2) sense heritabilities
indicated that transfer of resistance to recipient parent by donor parent is highly possible.
KEYWORDS: Melon Adult Plants Gene Effects Heritabilities Powdery Mildew Podosphaera Xanthii

Original Article

IIHR 122, IIHR 681 x IIHR 121 and IIHR 681 x IIHR 122 showed duplicate type of epistasis. Additive variance was

Received: Mar 17, 2016; Accepted: May 19, 2016; Published: May 30, 2016; Paper Id.: IJASRJUN2016050

INTRODUCTION
Powdery mildew is one of the most important production constraints in muskmelon throughout India.
Powdery

mildew

caused

(formerly

Sphaerotheca

by

fuliginea)

Podosphaera
and

xanthii

Golovinomyces

(Castagne)
cichoracearum

Braun
(DC)

and
V.P.

Shishkoff
Heluta

(formerly Erysiphe cichoracearum) are the most common and serious in Germany, France, Czech Republic, India,
New Zealand and Montenegro (Lebada and Kristova 2000; Mccreight 2006; Nayar and More 1998; Rankovic
2003). The Podosphaera xanthii has several races, some of which attack all cucurbits while others have a host range
restricted to certain types of cucurbits. Infection by powdery mildew is greatly influenced by the stage of the plant,
humidity and temperature. The fungi can sporulate and cause infection in very dry as well as wet atmosphere
(Singh 1987).
Powdery mildew need to be controlled on both leaf surfaces to avoid premature death of leaves. Systemic
or translaminar fungicides generally have high risk for developing resistance because they have specific modes of
action and powdery mildew fungi have a high potential for resistance development. This is especially true for
predominate powdery mildew fungus Podosphaera xanthii (McGrath 2001). The P. xanthii strains have been
detected with resistance to as many as four classes of fungicides (del Pino 1999, O Brien 1995, Tsay and Tung
1992). The relative ease with which fungicide resistant strains can develop in affected field makes powdery mildew
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K. T. Shashikumar, M. Pitchaimuthu & R. D. Rawal

an important disease. Therefore, resistant varieties are being developed and are becoming an increasingly important
component of management programs.
Genetic mechanism of powdery mildew resistance varies with the genotypes and races of the pathogen.
The inheritance of resistance to powdery mildew in melon has been studied in several breeding populations
(Bohn and Whitaker 1964; Cohen and Cohen 1986; Epinat et al. 1993; Floris and Alvarez 1991; Kenigsbuch and Cohen
1989; Kenigsbuch and Cohen 1992; Perchepied et al. 2005; Sivakami et al. 1979) and somewhat different conclusions
have been reached in each case. Although the adult plant stage is usually the important stage for resistance screening, often
seedlings were assessed for disease reactions in controlled conditions. Gene action studies on resistance to powdery
mildew in adult melon plants have not been reported.
Generation mean analysis has been used to detect the types of gene action involved in several quantitatively
inherited traits including disease resistance (Dias et al. 2004; Shashikumar et al. 2010; St. Amand and Wehner 2001) in
melon. Understanding the genetic architecture of a trait is very useful in plant breeding programme for achieving effective
results. Generation mean analysis is one such approach which provides information about the nature and magnitude of
gene effects involved. The present study included a generation mean analysis of four melon crosses with the following
objectives: (1) to determine the types of gene action involved in the expression of resistance to powdery mildew in adult
melon plants (2) to determine the variance components and heritabilities for resistance to powdery mildew and (3) to
determine the number of effective factors controlling resistance to powdery mildew.

MATERIAL AND METHODS


Plant Materials
Five inbred lines viz. Punjab Sunehri, RM 43, IIHR 681, IIHR 122 and IIHR 121 were used as parental lines to
develop four F1 Hybrids: Punjab Sunehri x IIHR 122, RM 43 x IIHR 121, IIHR 681 x IIHR 121 and IIHR 681 x IIHR 122.
Inbred line RM 43 and IIHR 681 are resistant, IIHR 121 and IIHR 122 are moderately resistant and Punjab Sunehri is a
susceptible variety. The F1s were crossed to their parental lines P1 and P2 to get BC1P1 and BC2P2 generations, respectively.
On the same F1s, F2 seeds were generated by self-pollination. The experimental material comprised of six generations
(P1, P2, F1, F2, BC1P1 and BC2P2) derived from each of the four crosses.
Experimental Plan
Seedlings of six generations of each of the four crosses were raised in 50-unit plastic potting trays inside a
greenhouse for both the greenhouse and field experiment. At 2 3 leaf stage, seedlings were transplanted to the main field
at Vegetable Block, Indian Institute of Horticultural Research (IIHR), Bangalore, India during 2006-07. Seedling spacing
was 3.0 m between beds (centre to centre) and 0.45 m within bed. Field plots with 10 plants each were arranged in three
randomized blocks. The P1, P2 and F1 planted one plot per block, F2 planted in eight plots per block and BC1P1 and BC2P2
each planted in four plots per block in back cross generations were raised in three replicates for the greenhouse screening
experiment. For the greenhouse screening experiment, 10 plants each in P1, P2 and F1, 80 plants in F2 and 40 plants in back
cross generations were raised plastic seedling trays in three replicates.
Inoculation in Field
The pathogen (Podosphaera xanthii) was isolated from a melon growers field near Bangalore, India. Colonies of
Podosphaera xanthii were maintained on cucumber plants in the greenhouse at 23 25 oC temperature and 50 70%
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relative humidity. Inoculum of P. xanthii was prepared by gently washing powdery mildew infected cucumber leaves in
distilled water to release the spores. Inoculation was performed on both abaxial and adaxial leaf surfaces spraying
conidiospores suspension containing 5,000 6,000 sporangia per milliliter using a hand sprayer. Following the above
procedure, second inoculation was performed three days after first inoculation. Arka Jeet, a susceptible variety planted at
regular interval in the experimental plot for uniform spread of disease.
Inoculation in Greenhouse
A piece of powdery mildew infected cucumber leaf of size 1.5 cm x 1.5 cm was stapled to the abaxial surface of
leaves of all the test plants at 2 3 leaf stage. Immediately, stapled plants were transferred to a growth chamber
(50 70 % relative humidity, 23 25 C temperature and 12/12 hr day/night). The next day, seedlings were brought back
to the greenhouse benches. This method of inoculation was found ideal to initiate P. xanthii infection on abaxial surface of
leaves in melon plants under the greenhouse conditions. Three days after the first inoculation, second inoculation was
performed to ensure uniform spread of disease on adaxial surface of leaves by breath blowing over powdery mildew
infected cucumber leaves bearing freshly produced conidia. Immediately, inoculated plants were transferred to a growth
chamber (50 70 % relative humidity, 23 25 C temperature and 12/12 hr day/night). The next day, seedlings were
brought back to the greenhouse benches.
In both the screening experiments, 10 12 days after second inoculation, clear fungal growth was noticed on both
sides of leaves of the inoculated plants. Symptoms first appeared as white or fluffy circular patches or spots on the under
surface of the leaves and later spread to upper surface of leaves. Inoculated plants were maintained on greenhouse benches
for six weeks. A single application of micronutrients @ 0.5 ml/l of water was supplied at 2 3 leaf stage. Seedlings were
hand watered daily. A nutrition solution containing 150 mg N, 150 mg P and 150 mg K per liter of water was supplied
every week.
Disease Assessment
Infection on plants was measured 45 days after inoculation in field experiment and 30 days after inoculation in the
greenhouse experiment. A linear 0 5 scale indicating average rating of all the leaves was used to assess the disease in
both experiments where, 0 = free from fungal infection, 1 = 1 10 %, 2 = 11 25 %, 3 = 26 50 %, 4 = 51 75% and
5 = > 75 % of total leaf area covered with fungal growth.
Percent Disease Index (PDI) was calculated using the formula proposed by Wheeler (1969): PDI = Sum of
numerical values/ (Number of leaves rated x Maximum rating) x 100. Plants were clustered into three groups based on
mean percent disease index: resistant, moderately resistant and susceptible. Mean PDI between 0 15 were grouped as
resistant, mean PDI between 15 30 as moderately resistant and mean PDI above 30 as susceptible.
Statistical Analysis
Estimates of PDI were transformed to arcsine values prior to generation mean analysis. The means and variances
were calculated as suggested by Hayman (1958). The presence of epistasis was detected by using A, B, C and D scaling
tests proposed by Mather (1949) and Hayman and Mather (1955). Just the significance of either one or two scaling test
implies the insufficiency of simple additive-dominance model. The model proposed by Hayman (1958) estimated gene
effects. The gene effects were defined in Haymans notations as [m] = mean of the F2 generation, [d] = additive gene
effect, [h] = dominance gene effect, [i] = additive x additive gene effect, [j] = additive 9 dominance gene effect and
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K. T. Shashikumar, M. Pitchaimuthu & R. D. Rawal

[l] = dominance x dominance gene effect. The type of epistasis was determined only when dominance [h] and dominance x
dominance [l] effects were significant, when these effects had the same sign, the effects were complementary while
different signs indicated duplicate epistasis (Kearsey and Pooni 1996). Generation mean analysis was computed using
statistical program SPAR1 (Indian Agricultural Research Institute, New Delhi, India).
Additive (2A), dominance (2D) genetic variances and narrow sense heritability estimates were calculated by
using formulae proposed by Warner (1952): Narrow sense heritability = 2A 2F2 where, 2A is additive variance. Broad
sense heritability was calculated by method proposed by Weber and Moorthy (1952): H = (2F2 2E) 2F2 in which 2F2 is
phenotypic variance of F2 population and 2E is environmental variance [2E = (2P1 + 2P2 + 2F1) 3]. Estimates of
minimum number of effective factors (n) were calculated according to the formula suggested by Castle (1921) and Wright
(1921). Mid-parent heterosis (MPH) was estimated as the percentage deviation of the F1 from the mid-parent.

RESULTS AND DISCUSSIONS


Generation Means
Means and their standard errors for parental lines, F1, F2, and backcross generations for resistance to powdery
mildew are presented in Table 1. Inbred line RM 43 expressed least mean PDI (< 15) followed by IIHR 681. IIHR 121 and
IIHR 122 expressed moderate resistance with mean PDI between 15 and 30. Mean PDI of Punjab Sunehri was
susceptible with very high man PDI (> 30) in both screening experiments. Parental lines were significantly different for
mean PDI in all the crosses (Table 1). The mean PDI of F1 generation was lower than mid-parent values in crosses RM 43
x IIHR 121, IIHR 681 x IIHR 121 and IIHR 681 x IIHR 122 indicated predominance of dominant factors for resistance to
powdery mildew. The mean PDI of F2 generation was significantly different from the mean PDI of F1 generation in crosses
IIHR 681 x IIHR 121 and IIHR 681 x IIHR 122. In all the crosses, the mean PDI of F2 generation exceeded the mean PDI
of F1 generation. The mean PDI of BC1P1 generation in crosses RM 43 x IIHR 121, IIHR 681 x IIHR 121 and IIHR 681 x
IIHR 122 skewed towards their resistant parents conferred the importance of genetic dominance for resistance to powdery
mildew. Similarly, in Punjab Sunehri x IIHR 122, the mean PDI of BC2P2 generation was skewed towards its resistant,
IIHR 122.
Gene Effects and Heterosis
Negative dominance (h) or dominance x dominance (l) gene effects in all the crosses and in both experiments
tends to increase negative heterosis (Table 2). Oppositely signed dominance (h and l) effects cancelled each other and are
not detected by means resulting low net dominance effects. Epistasis of duplicate type was detected in crosses Punjab
Sunehri x IIHR 122, IIHR 681 x IIHR 121 and IIHR 681 x IIHR 122. Dominance effects may be exploited in crosses RM
43 x IIHR 121 and IIHR 681 x IIHR 121, if hybrid melon is the objective of the breeding program.
The MPH varied from -7.43 to -93.34 percent and -22.91 to -87.55 percent in the greenhouse and field
experiment, respectively (Table 2). Significant and negative MPH indicated dominance and/or epistatic dominance effects
for resistance to powdery mildew in these crosses. Consistently high MPH observed in the crosses RM 43 x IIHR 121 and
IIHR 681 x IIHR 121 indicated strong dominance effects which should be exploited through heterosis breeding. Breeding
methods directed towards recovering hybrid vigour may increase gain from selection.
Variance Components, Heritabilities and Effective Factors
Additive variance was largest component of total genetic variance in all the crosses and in both screening
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Quantitative Analysis of Resistance to Powdery Mildew in Adult Melon Plants

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experiments (Table 3). Estimates of dominance variance were negative in three of the four crosses, whereas estimates of
environmental variance were low in all the crosses. Estimates of broad- (H) and narrow sense (h2) heritability were high
(>0.60) in all crosses except for Punjab Sunehri x IIHR 122 (Table 3). The estimates of broad sense heritability varied from
0.72 to 0.97 and 0.74 to 0.93 in the greenhouse experiment and field experiment, respectively (Table 3). The estimates of
narrow sense heritability varied from 0.72 to 1.73 and 0.05 to 1.83 in the greenhouse experiment and field experiment,
respectively (Table 3). Low dominance and environmental variance resulted in higher broad and narrow sense heritabilities
in all the crosses. These findings are supported by high heritabilities and large additive variance for resistance to powdery
mildew in melon (Sivakami et al. 1979). Selection for resistance to powdery mildew should be effective because of high
narrow sense heritability and predominance of additive variance across screening experiments. High heritabilities indicated
that transfer of resistance to recipient parent by donor parent is highly possible. Recurrent selection would prove useful to
exploit large additive variance. The estimates of minimum number of effective factors (n) varied from 0.21 to 2.55
averaging 1.38 in the greenhouse experiment and 0.35 to 3.91 averaging 2.13 in field experiment (Table 3).

CONCLUSIONS
Generation mean analysis assumes absence of dominance and epistasis and equal genetic effects which will
underestimate the number of effective factors (Mather and Jinks, 1971). The number of effective factors appears to be one
to four, however factors were much lesser than actual number because of presence of significant epistasis and dominance
in all the crosses. Number of effective factors controlling resistance to Podosphaera xanthii can be further defined using
QTL analysis (Perchepied et al. 2005; Longzhou et al. 2008) and molecular marker linked resistance gene
(Teixeira et al. 2008). Genetic dissection of resistance to powdery mildew in melon allows the development of
marker-assisted selection for breeding, the characterization of the genes underlying resistance QTLs of genetic resources
and the isolation of the corresponding genes. Quantitative analysis of resistance to powdery mildew confirmed the
insufficiency of simple additive-dominance model. These results will help to further our understanding of the genetics of
resistance to P. xanthii in Cucumis melo.

ACKNOWLEDGEMENTS
The first author is grateful to the Director, Indian Institute of Horticultural Research, Bangalore for their support
and facilities provided during this Ph.D study. The first author thanks Dr. K.R.M. Swamy, Indian Institute of Horticultural
Research, Bangalore and Mr. Seenappa, University of Agricultural Sciences, Bangalore, for their helpful comments and
suggestions.
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APPENDICES
Table 1: Mean, Standard Error (SE) and Variance () For Resistance to Powdery Mildew In
Four Melon Crosses Evaluated In the Greenhouse and Field Conditions
Punjab Sunehri X IIHR 122
Generation
Mean SE
Greenhouse Experiment
P1
41.68 0.54(27)e1
P2
18.70 0.36(25)a
F1
27.19 0.67(30)c
F2
29.27 0.57(220)c
BC1P1
40.50 0.80(108)d
BC2P2
22.55 0.44(106)b
Field experiment
P1
66.55 0.82(26)d
P2
14.38 0.33(29)a
F1
31.20 0.60(30)b
F2
28.37 0.72(233)b
BC1P1
53.59 0.93(106)c
BC2P2
30.28 0.35(109)b
1

RM 43 X IIHR 121
Mean SE

IIHR 681 X IIHR 121


Mean SE

IIHR 681 X IIHR 122


Mean SE

1.78 0.22(26)b
21.90 0.55(28)f
0.67 0.08(30)a
5.07 0.17(226)d
3.77 0.18(112)c
10.17 0.25(109)e

3.61 0.50(26)a
21.90 0.55(28)d
7.97 0.42(30)b
18.83 0.63(216)e
6.50 0.45(109)b
13.67 0.29(101)c

3.61 0.50(26)a
18.70 0.36(25)e
8.33 0.67(28)c
11.93 0.32(229)d
5.51 0.33(111)b
10.83 0.17(105)cd

1.55 0.22(28)a
21.43 0.67(25)e
1.43 0.09(28)a
7.20 0.21(228)c
2.24 0.25(116)b
10.73 0.23(114)d

3.55 0.42(26)a
21.43 0.67(25)d
5.60 0.33(29)b
8.97 0.42(231)c
4.44 0.29(111)ab
6.38 0.31(116)b

3.55 0.42(26)bc
14.38 0.33(29)f
4.53 0.27(30)c
9.60 0.23(225)d
2.80 0.12(112)a
13.41 0.31(109)e

Means followed by same letter in each column of greenhouse and field experiment are not significantly different

according to pairwise t test comparison at P 0.05.


Number in parentheses indicate the number of plants screened for each generation

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K. T. Shashikumar, M. Pitchaimuthu & R. D. Rawal

Table 2: Estimates of Gene Effects and Mid-Parent Heterosis (MPH) for Resistance to Powdery Mildew of
Four Melon Crosses Evaluated in the Greenhouse and Field Conditions
Greenhouse Experiment
Parameter

Punjab
Sunehri x
IIHR 122

RM 43 x
IIHR
121

IIHR 681
x IIHR
121

[m]
29.27**
5.07**
18.83**
**
**
[d]
17.95
6.40
7.17**
**
**
[h]
6.02
3.57
39.76**
**
**
[i]
9.03
7.60
34.98**
**
**
[j]
29.44
3.66
1.98
[l]
20.36**
10.47**
36.09**
Epistasis
D
C
D
MPH
7.36*
94.34**
37.54**
*
-Significant at P 0.05, **-Significant at P 0.01

Field Experiment
IIHR 681
x IIHR
122

Punjab
Sunehri x
IIHR 122

RM 43 X
IIHR 121

IIHR 681
X IIHR
121

11.93**
5.32**
17.86**
15.04**
2.23
21.33**
D
7.43*

28.37**
23.31**
45.00**
54.26**
2.27
78.68**
D
22.91**

7.20**
8.49**
12.92**
2.86**
1.46*
2.76*
D
87.55**

8.97**
1.94**
21.13**
14.24**
7.01**
28.78**
D
55.16**

IIHR
681 X
IIHR
122
9.60**
10.61**
11.41**
5.98**
4.19**
2.55*
D
49.50**

D = Duplicate epistasis, C = Complementary epistasis


Table 3: Estimate of Additive ( 2a), Dominance ( 2d), Environmental ( 2e) and Genetic ( 2g) Variances,
Broad Sense (H) and Narrow Sense (H2) Heritabilities for Resistance to Powdery Mildew in
Adult Plants of four Muskmelon Crosses Evaluated in the Greenhouse and Field Conditions
Greenhouse Experiment
Field Experiment
Punjab
IIHR 681
Punjab
IIHR 681
Parameters Sunehri
RM 43 x IIHR 681 x
RM 43 x
x IIHR
Sunehri x
x IIHR
x IIHR
IIHR 121
IIHR 121
IIHR 121
122
IIHR 122
121
122
2AP
53.31
23.16
140.89
31.78
106.21
12.64
61.01
2D
9.88
7.83
60.27
13.95
1.53
5.10
26.53
2E
8.28
1.44
5.11
5.62
11.95
2.52
6.26
HQ
0.72
0.91
0.97
0.76
0.90
0.74
0.93
h2
0.74
1.38
1.73
1.36
0.05
1.27
1.83
nR
2.55
2.18
0.21
0.90
3.20
3.91
0.35
P
Variance components calculated as follows: 2A = 22F2 (2B1 + 2B2), 2D = (2B1 + 2B2 2F2 2E) and

IIHR
681 x
IIHR
122
11.72
1.95
2.13
0.82
0.98
1.25

2E = (2P1 + 2P2 + 2F1) 3


Q

Heritabilities calculated as follows: H = (2F2 2E) 2F2, h2 = 2A 2F2

Minimum number of effective factors calculated as follows: n = (P1 P2)2 [8 (22F2 2B1 2B2)]

Impact Factor (JCC): 4.7987

NAAS Rating: 3.53

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