Introduction
Sesame (Sesamum indicum L.) is one of the most
imperative oilseed crops in Pakistan that belongs to family Pedaliaceae (Ahmad et al. 2018). Sesamum seeds are considered as the much
primitive oilseed crop dating back to 1600 BC. Now, it is widely grown in the
tropical areas of Asia and Africa (Saydut et al. 2008). It has high nutrition
value as it contains sesamin, sesamolin,
sesaminol and sesamolinol
that maintain the low-density lipoprotein fats (Banerjee and Kole 2009). The consumption of 100 g sesame seed provides
carbohydrates (24.05 g), proteins (18.08 g), fats (50.87 g), total dietary
fibres (5.5 g), calcium (960 mg), iron (19.2 mg), magnesium (362 mg),
phosphorus (659 mg), sodium (12 mg), potassium (582 mg), thiamine (0.240 mg), riboflavin (0.200 mg), pantothenic acid (0.052 mg), vitamin B6 (0.816
mg), vitamin A (3 µg), saturated
fatty acid (1.252 g), monosaturated fatty acids
(3.377 g) and polyunsaturated fatty acids (3.919 g) (Prasad et al.
2012).
There are numerous biotic
constraints in successful cultivation and harvesting of sesame. Among those
production inhibiting biotic restrains, charcoal rot of sesame caused by Macrophomina phaseolina (Tassi) Goid
is the most destructive disease (Ibrahim and Abdel-Azeem 2015; Ouoba
et al. 2019). The losses in yield
under field conditions owing to M. phaseolina are ranging from 22–53% with 37%
disease incidence (Gupta et al. 2018) whereas yield losses of 5–100%
in Egypt has also been observed under favourable environmental conditions (El-Bramawy and Wahid, 2006). The pathogen has anxiously
diminished the yield of sesame approximately 27 million bushels per annum in
USA (Chattopadhyay and Sastry
2002) with the predicted value of $US 146 million (Bashir et al. 2017).
M. phaseolina is a necrotrophic
thermophilic soil-borne plant pathogen having two
asexual subphases i.e.,
saprophytic phases and pathogenic phase. It produces microsclerotia
and mycelia in the saprophytic phase whereas microsclerotia,
mycelia and pycnidia are produced during pathogenic
phase (Kaur et al. 2012). The genus Macrophomina is a
monotypic genus because it comprises only one species namely “phaseolina” (Sutton 1980). The characteristic symptoms
owing to M. phaseolina
on sesame are spindle shaped lesions with brown margins and light grey centres
having dispersed pycnidial bodies. Infected plants
depict wilting, drying, shredding of leaves ultimately the decaying of host
plant after sever infection (Beas et al. 2006). The visual appearance of
M. phaseolina
in petri-plates varies from black to brown or grey but it becomes dark with the
passage of time. Numerous aerial mycelia are produced with sclerotia
imbedded within the hyphae. The hyphae are septate
and black in appearance. Microsclerotia are also in
black colour with different sizes, ranging from 50–150 μm whereas pycnidia are dark to greyish in colour with globose or flattened structure having diameter of 100 to
200 μm
(Yang and Navi 2005; Kaur et al. 2012).
Environmental factors
imperatively influence the genotypic characters of host plant by promoting
virulence factors of pathogens and subsequently reduce the maximum cultivation
of sesame in a certain geographic location. Genotype by environment interaction
(GGE) biplot analysis is an eminent method to
determine yield stability and suitable variety selection consider genotypes as
main effect under multi-geographic locations (Shim et al. 2015).
Similarly, additive main effects and multiplicative interaction (AMMI) biplot analysis is an effective tool which is used to
separate multiplicative components and additive main effects through principal
component analysis. The comparison of multilocation
trials with genotypes as well as main effects and multiplicative interaction is
also helpful to enhance the yield by understanding all yield limiting factors
(Yan and Hunt 1998). The GGE comprises numerous interaction methods of biplot analysis by which genotype by environment (G×E)
interactions can visually be addressed (Yan and Kang 2003; Yan and Tinker
2006). The GGE and AMMI biplots analysis are
extensively used for diversity, stability, adoptability trials, for the source
of resistance and better yield performing cultivars in a set of environmental
interactions (Pourdad and Moghaddam
2013).
Various conventional
(application of plant extracts, use of antagonistic organisms, plant growth
regulators, management through fertilizers, disease management through organic
manures, soil solarization, application of fungicides
etc.) and molecular approaches
(systemic induce resistance, systemic acquired resistance, DNA markers,
proteomics, transcriptomics, comparative genomics,
computational biology etc.) are
available for the management of sesame charcoal rot disease. Among all methods,
few are not practicable and attainable for poor farmers due to higher cost
benefit ratio, less efficiency under diverse climatic conditions whereas others
impose direct or indirect impact on human health such as indiscriminate use of
fungicides for diminishing disease incidence. Thus, it is dire need to explore
such disease inhibiting or management approaches which are economically
adoptable, environmentally friendly and easily available for farmers (Ahmed et al. 2013). Therefore, the ideal and
most appropriate way to combat the disease and hinder the server yield losses
is the use of climate resilient and genetically resistant sesame advanced
genotypes. The source of resistance through screening of existing germplasm against M. phaseolina under natural field conditions is pre-requisite
to accomplish the purpose (Meena et al. 2018). Thus, indispensable
efforts were carried out to assess the genotypes of sesame under natural field
conditions. The research was
conducted to evaluate the existing sesame germplasm/advanced
lines under natural field conditions against charcoal rot disease caused by M. phaseolina (Tassi) Goid
for the source of resistance.
Genotypes possessing resistant genes against virulent pathogen will be
submitted for varietal approval based on data of continuous multilocational
research trials to find the resistant source.
Materials and Methods
Identification, isolation and purification
of M. phaseolina
Sesame plants with distinctive
symptoms of Charcoal rot disease were collected from experimental research filed
and brought in the Oilseeds Plant Pathology Lab. for isolation of virulent
pathogen. Infected roots were washed carefully with running tab water, cut into
pieces of 3–5 mm and surface sterilized with 1% sodium hypochlorite (NaOCl) solution. The roots were air dried for 2–3 min by
keeping on sterilized filter paper in 9 cm petri plate. The infected roots (3–4
pieces) were placed in petri plate containing 20 mL Potato Dextrose Agar (PDA)
medium. Petri plates were incubated at 25°C±2 for 48–72 h (Sarwar
et al. 2005). Maximum fungal growth was observed in plates, a temporary
slide of pathogen was prepared for identification. After identification and
confirmation through literature as well as comparison with the isolated M. phaseolina
under stereomicroscope, the purified fungal colonies were multiplied for
further use in the research (Soesanto et al.
2011).
Preparation of sick field
Highly susceptible single stem
variety of Sesame i.e., TH-6 was sown
in experimental research area of Plant Pathology at Oilseeds Research
Institute, Faisalabad, Pakistan. The suspension of M. phaseolina pure culture possessing
1×106 sclerotia and pycnidia/mL
of H2O was prepared, and concentration was confirmed through
haemocytometer. TH-6 was inoculated through drenching with two successive
irrigation of 7 days interval by placing the bottle having inoculum near inlet
of water. Similarly, 4 mL of sterilized water was poured on 5–8 days old pure
culture of M. phaseolina
in petri plates, shake them gently and 4 plates were transferred in 250 mL
water in beaker. 5 mL pure culture from beaker was also drenched near root zone
of each plant before irrigation to enhance the inoculum density. Disease
symptoms appeared on 30–45 days old plants, the diseased plants were
homogeneously mixed in the soil and the field was irrigated to increase
decomposition of plant debris as well as to enhance pathogen infection density
on plants during research.
Establishment of disease screening
nursery
Ten varieties/advanced lines of
sesame namely TS-5, 10003, 40009, 50009, 50011, 50022, 16001, 40004, Black Till
and TH-6 were collected from Oilseeds Research Institute (ORI), Ayub Agricultural Research Institute (AARI), Faisalabad,
Pakistan. The experiment was sown at six different locations i.e., Faisalabad, Toba Tek Singh, Piplan, Bhakhar, Mandi Baha Uddin and Bahawalpur. Each
row of (3.5 m) 350 cm was prepared and seed was sown through hand drill in
three replications at Experimental Research Area of Plant Pathology, ORI, AARI,
Faisalabad. After 7–10 days, seedlings were observed,
and the experiment was visited daily to visualize the germination rate in the
field. After 15 days of seedling emergence, 30–40 healthy sesame plants were
sustained in each row by maintaining the P×P and R×R distance of 8–10 cm and 45
cm under Randomized Complete Block Design (RCBD). All weak and unwanted plants
were pulled off to diminish the nutritional competition. All the horticultural
practices including hoeing, earthing up, weed
eradication and irrigation were followed to keep the plants in good health.
Data of disease incidence was recorded after 10–15 days interval from all
experimental location at different regions.
Statistical analysis
Experiment was conducted under
randomized complete block design (RCBD). Statistical analysis was performed
using R. Combined analysis of variance was used to assess the level of
significance whereas Genotype × Environment Interaction table was used to
separate means of data regarding disease incidence. Further, Additive Main
Effects and Multiplicative Interaction (AMMI) Biplot
was performed for stability analysis.
Results
Genotypes adaptability/resistance
status through additive main effects and multiplicative interaction (AMMI) analysis
Separate Analysis of Variance for each environment
revealed that the genotypes performed statistically significant response to
control the Disease Incidence (%) of Sesame charcoal rot. The performance of
ten genotypes was different at various locations (Table 1). Combined analysis of variance not only showed that the
performance of the genotypes was not the same (F = 440.053, P < 0.001) but also revealed that
their performance against charcoal rot of sesame varies across the environments
(F = 4.248, P < 0.001) (Table 2).
The significant interaction between genotypes and environments implied that the
performance of genotypes was not stable across environments and all the
genotypes performed differently
across environments. The same phenomena were depicted by genotypes by
environment means and ranks of genotypes means within environments (Table 3 and
4). The genotypes 16001, 10003 and 50009 depicted the highest disease incidence
on sesame at six different locations as compared to remaining genotypes and
showed susceptibility (Disease rating 5). The genotype 16001 expressed the highest disease
incidence in four (Bahawalpur, Bhakkar, Faisalabad
and M.B. Din) out of the six locations and the second highest disease incidence
in the other two locations (Piplan and T.T. Singh)
(rating 5). Similarly, the genotype 10003 showed the highest disease incidence
in two (Piplan and T.T. Singh) out of the six locations and the second highest
disease incidence in three locations (Bhakkar,
Faisalabad and M.B. Din) and the third highest disease incidence in Bahawalpur
(rating 5). Genotype 50009 was the third most sensitive among ten genotypes. It
remained third most diseased in four (Bhakkar,
Faisalabad, Piplan and T.T. Singh) out of six
localities by exhibiting moderately susceptible response (rating 4) while at
Bahawalpur it was found second most diseased genotype after showing susceptible
response (rating 5). The genotypes 40004, TS-5 and 50011 were found to be the
resistant genotypes. The genotype 40004 expressed resistance response at two
locations (Piplan, Bhakkar)
out of six localities and ninth most resistant genotype at Faisalabad and T.T.
Singh (rating 2) as well as it was found seventh and sixth resistant genotype
at M. B. Din and Bahawalpur respectively (rating 3). Genotype TS-5 was found to
be resistant for Bahawalpur, Faisalabad and Piplan
environments while ninth most resistant in two locations (Bhakkar,
M.B. Din) and the most appropriate for T.T. Sing owing to its highest
resistance against charcoal rot disease of sesame (rating 2). Likewise,
genotype 50011 remained highly resistant at Faisalabad and Bahawalpur (rating
2) whereas the eighth at two localities (Bhakkar,
T.T. Singh) and the sixth moderately resistant response at M.B. Din and Piplan (rating 3). The genotype 50022 expressed
moderately resistant response at two locations (Bahawalpur, Bhakkar
(rating 3) while showed resistant response at Faisalabad and T.T. Singh,
(rating 3). It remained ninth resistant genotype at Piplan
and tenth most resistant in the environment of M. B. Din (Disease rating 2). Likewise, the genotype 40009 expressed moderately
resistant response at three (Faisalabad, Piplan, T.T.
Singh) out of six locations while sixth at Bhakkar
and eighth at M.B. Din localities (Disease rating 3). The same genotype
depicted resistant response at Bahawalpur location under disease rating 2. The genotype TH-6 showed moderately susceptible response
at five locations viz., Bahawalpur, Bhakkar,
Faisalabad, Piplan and T.T. Singh whereas it also
expressed moderately susceptible response at third of M.B. Din (Disease rating
4). Black Till genotype depicted resistant response at Piplan
(rating 2) and remained at sixth after showing moderately resistant response
(rating 3) in two locations (Faisalabad, T.T. Singh). It showed moderately
susceptible response at fifth in Bhakkar (rating 4)
as compared to Bahawalpur and M.B. Din localities where it remained at fifth
but exhibited moderately
resistant response (rating 3) (Table 5 and Fig. 1). The Additive Main Effects and
Multiplicative Interaction (AAMI) analysis indicated almost 84% variability or
adaptability in all genotypes at six diverse
Table 5: Modified disease rating scale for charcoal rot of sesame
|
Disease Rating |
Description |
Response |
Symbol |
|
0 |
No symptoms on plants |
Immune |
I |
|
1 |
1% or less plants infected |
Highly Resistant |
HR |
|
2 |
0.01-10% plants infected |
Resistant |
R |
|
3 |
10.01-20% plants infected |
Moderately resistant |
MR |
|
4 |
20.01-50% plants infected |
Moderately Susceptible |
MS |
|
5 |
˃ 50.01% plants
infected |
Susceptible |
S |
%20851-856_files/image002.jpg)
Fig. 1: Charcoal
rot disease incidence on sesame advance genotypes at
six different locations
environments. The main effects i.e.,
genotypes, and the environmental interaction expressed the
suitability/adaptability of genotypes in different environments (Fig. 2).
Charcoal rot caused by M. phaseolina is
a potential threat in sesame growing areas of Pakistan, causing huge losses
with disease incidence ranging from 30–37% in the field under conducive
environmental conditions (El-Bramawy and Wahid, 2006;
Bashir et al. 2017). Shredding of leaves,
wilting, drying and decaying are the characteristic symptoms this disease (Beas
et al. 2006). High temperature, moisture, soil physical and chemical
characteristics, virulent pathogen races, inoculum density and time of
infection influence the development of symptoms (Akhtar et al. 2011). Moreover, resistant
genotypes not only reduce the disease incidence but also tolerate fungicide
toxicity (Banerjee and Kole 2009). The use of molecular techniques for
identification, selection and insertion of resistant genes is costly and
non-durable due to higher genomic evolutionary processes but screening for the
source of resistance through conventional breeding under diverse environmental
conditions is an easily accessible and durable approach to farmers (Meena et al. 2018). That is why, ten sesame
varieties/advanced lines were assessed against Charcoal rot disease in six
different environmental regions and it was observed that resistance is
prevailing in some genotypes (40004, 50011, 50022 and TS-5) but, the resistance
status of genotypes may be decreased if specific genotypes are continuously
used in a geographically specific area as well as due to more virulent
aggressive strains of pathogens. Durable, field or polygenic resistance
diminishes the severity of disease owing to the presence of many genes in host
plant as compared to non-durable or monogenic resistance. However, the multilocational conventional hybridization breeding program
for the source of resistance in host plants is also an imperative possible
solution to reduce the disease (Meena et al. 2018). Resistant source should
be incorporated directly or indirectly through conventional breeding to restore
the genotypes at desirable level. During selection of resistant source, two segregating generations (F3 and F4) were exposed to sesame charcoal
rot pathogen (M. phaseolina) under natural field conditions during two
successive seasons (2004 and 2005). The highly significant
variability regarding disease incidence was observed in five crosses i.e., P1×P2, P1×P4,
P1×P5, P2×P6 and P3×P4. Such crosses in conventional breeding might be
helpful for the selection of stable source of resistance and will aid breeders to achieve better sesame cultivars with charcoal root rot resistance (El-Bramawy and Wahid, 2006). Similarly, thirty-six (36) F1
genotypes were evaluated under artificial screening as well as sick plot
conditions against root rot of sesame and three genotypes namely ORM-7, ORM-14 and ORM-17 were found resistance against
disease (Thiyagu et al. 2007).
%20851-856_files/image004.jpg)
Fig. 2: Stability analysis for additive main effects and
multiplicative interaction
Statistical models like AMMI and
GGE (Gauch et al. 2008) are helpful for
stability analysis of genotypes against aggressive virulent pathogens (Belay et al. 2018). The GGE and AMMI Bi Plot
analysis also play a pivotal role for selection of resistant source in multilocation trails (Yaseen et
al. 2014) as well as to check the variability and stability in sesame
genotypes against charcoal rot disease (Kaur et al. 2012).
Conclusion
It is concluded that the genotypes
40004, TS-5, 50011 and 50022 exhibited resistant response against Charcoal rot
of sesame caused by M. phasiolina at different
localities. The genotype 40004 is much suitable for Bhakkar,
Faisalabad, Piplan and T.T. Singh whereas genotype
TS-5 is better for Bhakkar, Faisalabad, MB Din and
T.T. Singh. Similarly, the genotype 50011 exhibited resistant response against
disease at three location (Bahawalpur, Faisalabad and T.T. Singh) while
genotype 50022 is also good for three localities including MB Din, Piplan and T.T. Singh.
Acknowledgements
The current research is a part of Annual Research Work of Oilseeds Research Institute, Faisalabad. The authors acknowledge
Government of Punjab, Agriculture
Department for funding.
Table 1: Individual
analysis of variance for each environment
|
SOV |
d.f. |
Sum of Squares (SS) |
|||||
|
Genotypes |
9 |
||||||
|
Residuals |
20 |
670 |
263 |
293 |
920 |
||
Table 2: Combined analysis of variance for the assessment of resistant
source
|
SOV |
D.f. |
SS |
MS |
F |
P |
|
Environments |
5 |
336.480 |
67.296 |
6.508 |
0.004 |
|
Replication (Env.) |
12 |
124.080 |
10.340 |
|
|
|
Genotypes |
9 |
103734.959 |
11526.107 |
440.053 |
<0.001 |
|
Gen: Env |
45 |
5006.836 |
111.263 |
4.248 |
<0.001 |
|
PC1 |
13 |
3246.591 |
249.738 |
9.535 |
<0.001 |
|
PC2 |
11 |
946.380 |
86.035 |
3.285 |
<0.001 |
|
Pooled Deviation |
31 |
813.865 |
26.254 |
1.002 |
0.475 |
|
Residuals |
108 |
2828.792 |
26.193 |
|
|
Table 3: Genotypes × environment interaction means
|
Genotypes |
Locations |
|||||
|
Bahawalpur |
Bhakkar |
Faisalabad |
M.B. Din |
Piplan |
T.T. Singh |
|
|
10003 |
64.547 |
67.050 |
66.750 |
72.547 |
77.383 |
71.560 |
|
16001 |
74.423 |
72.037 |
77.200 |
77.277 |
65.550 |
69.063 |
|
40004 |
16.920 |
7.777 |
6.983 |
15.883 |
6.487 |
7.133 |
|
40009 |
7.270 |
17.470 |
16.800 |
13.893 |
16.717 |
19.653 |
|
50009 |
71.407 |
34.760 |
53.247 |
39.713 |
40.953 |
46.983 |
|
50011 |
7.293 |
15.570 |
6.533 |
16.530 |
16.380 |
9.033 |
|
50022 |
16.217 |
16.660 |
8.057 |
7.410 |
7.710 |
9.730 |
|
Black Till |
17.093 |
27.627 |
16.700 |
19.397 |
9.113 |
14.357 |
|
TH-6 |
43.550 |
33.103 |
40.977 |
42.747 |
36.500 |
41.290 |
|
TS-5 |
8.653 |
7.780 |
7.800 |
9.123 |
7.997 |
6.853 |
Table 4: Ranks of sesame genotypes within different environments
|
Ranks |
Locations |
|||||
|
Bahawalpur |
Bhakkar |
Faisalabad |
M.B. Din |
Piplan |
T.T. Singh |
|
|
1 |
16001 |
16001 |
16001 |
16001 |
10003 |
10003 |
|
2 |
50009 |
10003 |
10003 |
10003 |
16001 |
16001 |
|
3 |
10003 |
50009 |
50009 |
TH-6 |
50009 |
50009 |
|
4 |
TH-6 |
TH-6 |
TH-6 |
50009 |
TH-6 |
TH-6 |
|
5 |
Black Till |
Black Till |
40009 |
Black Till |
40009 |
40009 |
|
6 |
40004 |
40009 |
Black Till |
50011 |
50011 |
Black Till |
|
7 |
50022 |
50022 |
50022 |
40004 |
Black Till |
50022 |
|
8 |
TS-5 |
50011 |
TS-5 |
40009 |
TS-5 |
50011 |
|
9 |
50011 |
TS-5 |
40004 |
TS-5 |
50022 |
40004 |
|
10 |
40009 |
40004 |
50011 |
50022 |
40004 |
TS-5 |
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