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Physico-chemical characteristics of rice (Oryza Sativa L.) grain imparting resistance and their association with development of rice weevil, Sitophilus oryzae (L.) (Coleoptera:Curculionidae)

  • G. Basana GowdaEmail author
  • Naveenkumar B. Patil
  • Totan Adak
  • Guru Pirasanna Pandi
  • Nabaneeta Basak
  • Kingshuk Dhali
  • M. Annamalai
  • G. Prasanthi
  • S. D. Mohapatra
  • Mayabini Jena
  • Somnath Pokhare
  • P. C. Rath
Original Article
  • 145 Downloads

Abstract

Association between rice grain characteristics and Sitophilus oryzae development were deciphered through laboratory experiments. Among the different physical characteristics evaluated, significant and negative correlation of 100 seed weight was observed with weevil emergence (r = − 0.53) and susceptibility index (r = − 0.51). Hence, 100 seed weight was found out to be important characteristics of grain. Similarly, hardness of the grain delayed weevil emergence (r = − 0.41) and reduced the grain weight loss (r = − 0.245). Biochemical characteristic of grain, like protein was found positively associated with weevil emergence (r = 0.741) and negatively associated with median developmental period (r = − 0.537). Amylose being an important component of grain had negative relation with weevil emergence (r = − 0.007); median development period (r = − 0.093). Among different varieties tested, Cross-12 was found to be moderately resistant compared to others. It was the only variety having long broad grain and recorded highest 100 seed weight indicating size of the substrate could hinder the development of weevils. The aim of the current study is to decipher the association of development of S. oryzae and physico-chemical characteristics of grain governing it.

Keywords

Coleoptera Oryza sativa Grain hardness Susceptibility index Seed weight 

Introduction

Cereal grains are generally being stored for longer periods (usually more than a year) before being processed or transported or consumed. All through this period, grains become susceptible to insect pests, which cause quantitative as well as qualitative losses (Nagpal and Kumar 2012; HLPE 2014). As a result, a large portion of this food never reaches final consumers. Losses of cereal grains by insect pests in tropical agriculture conditions can exceed 30% due to lack of better storage facility and the high humidity of the tropics (Ramputha et al. 1999). In India, the storage capacity available with the Food Corporation of India (FCI) is around 28 million tons in comparison to the present requirement of approximately 60–70 million tons. Around 3 million tons of food grain is stored temporarily under cover and plinth (CAP) storage for short duration in procurement areas and rest of the produce is not safely stored leading to loss of 3.9–6.0% and demanding higher protection of grain (Dhingra 2016). Among many cereals, rice (Oryza sativa L.) is the major source of calories for more than one-third population of the world, particularly in Asia. During storage, among various insects damaging to rice grains, the rice weevil, Sitophilus oryzae (L.) is the mainly damaging insect pest (Nwaubani et al. 2014). It is considered as ubiquitous, phytosanitary and quarantine pest of stored rice. The weevil has a cosmopolitan distribution, occurring all over the warm and tropical parts of the world (Hong et al. 2018). The damage in stored rice is caused by both grubs and adults. Damage usually occurs from decrease in the loss in nutritive value, grain weight of the produce and secondary contamination by fungi and mites, lastly losing the commercial value of produce (Souza et al. 2012). Preventing this avoidable loss is most important in order to feed every mouth. The chief management tactic to control insect pests during storage is by phosphine and methyl bromide (chemical fumigants) (Hossain et al. 2014; Ribeiro et al. 2003). However, methyl bromide has been kept under restricted use in India since 2015 (http://cibrc.nic.in/). Development of resistance in rice weevil to chemicals hampers the management of pest, and substitute methods of management are necessary (Ribeiro et al. 2013; Lee et al. 2001).

Resistant crop varieties have the potential to reduce losses during storage due to stored grain insect pests under subsistence farming situations mainly in developing countries (Ahmad and Jaiswal 2018; Kalsa et al. 2019). Grain resistance against stored grain insect pests has rarely been considered as a selection criterion in breeding programs during green revolution and later. In India, rice varieties were occasionally ascertained for their resistance to insect pests of storage. Most of the present day high yielding varieties have higher susceptibility to these pests (Kossou et al. 1993). Resistance to storage insect pests can be attributed to the physico-chemical characteristics of the wheat grain (Saad et al. 2018; Tripathi et al. 2017). Resistance to insects could be manifested as antibiosis, wherein few properties of the grain affect the insect pest development. On the other hand, the grains may have antixenosis, thereby affecting the insect behavior, as a result lesser oviposition and feeding (Lara 1991; Smith 2005). Prospective use of varieties resistant to S. oryzae and other stored grain insect pests were reported earlier (Bottega et al. 2012; Sousa et al. 2010). It is well known that single characteristic is not enough to adequately predict resistance of grain. However, there was no detailed study to determine the resistance which combines grain characteristics and development of S. oryzae. The aim of the current study is to decipher the association of development of S. oryzae and physico-chemical characteristics of grain governing it.

Materials and methods

The experiment was conducted at Indian Council of Agricultural Research (ICAR)- National Rice Research Institute (NRRI), Cuttack, Odisha (20°27′14.0″N 85°56′06.0″E). A total of 15 diverse rice varieties and landraces (Table 1) were selected based on their popularity and grain type. One kilogram of each variety (harvested freshly) were taken, thoroughly cleaned and disinfested by storing at 3 °C for 2 weeks to avoid all non-targets. The grains were further stored under laboratory conditions for 2 weeks to acclimatize. Grain moisture was adjusted to 12%. Healthy rice grains were hulled and milled in grain mill (Satake Corporation, Japan). Sound and healthy grains were selected for conducting experiment without any prior treatment.
Table 1

Detailed pedigree information of rice varieties

Sl. no.

Varieties

Grain classification

Pedigree information

V1

Dhusara

Medium slender (MS)

Landrace

V2

Chinikamini

Short broad (SB)

Landrace

V3

Kalajeera

Short broad (SB)

Landrace

V4

Geetanjali

Long slender (LS)

Mutant selection from Basmati 370

V5

Satyabhama

Medium slender (MS)

IR-31283-350-3-2-1/IR-41054-102-2-3-2

V6

Naveen

Medium broad (MB)

Sattari/Jaya

V7

Swarna Sub 1

Short broad (SB)

Swarna 3/IR4980-7-1-2-3

V8

Ketakijoha

Short slender (SS)

Savithri  /Badshabhog

V9

TN 1

Short broad (SB)

Dwarf chow-wu-gen/Tsai- yuan- chunj

V10

Cross 12

Long broad (LB)

IR-8-288-3/CR 1014

V11

Padmini

Short slender (SS)

Mutant selection from CR-1014

V12

Pyari

Medium slender (MS)

EPL R1 5/IR-12979-24-1 (Brown)

V13

Sarala

Short broad (SB)

CR-151-79/CR 1014

V14

Gayatri

Short broad (SB)

Pankaj/Jagannath

V15

Satabdi

Short slender (SS)

CR-10-114/CR 10115

MS medium slender, SB short broad, LS long slender, MS medium slender, MB medium broad, SS short slender, LB long broad

Test insect and rearing

The test insect species (S. oryzae) were collected from rice godown of ICAR-NRRI, Cuttack, Odisha, India and insect culture was maintained in the grain entomology laboratory of the institute. Throughout the study, culture was maintained at 27 ± 2 °C and 65 ± 5% RH. The weevils were preconditioned for three generations on a susceptible rice variety, TN-1, which was used for rearing laboratory culture. The culture was thoroughly maintained so that same aged weevils were obtained for the experiment.

About 100 g rice grains of respective rice genotypes were taken in a 250 cm3 plastic jar and covered with muslin cloth to avoid escape of the weevils and for proper ventilation and no choice method was followed (Stevens and Mills (1973). Initially, 20 pairs of weevils were placed in a plastic jar containing rice grains. The jars were kept for 7 days to allow mating and oviposition, after which weevils were removed. The remaining content of each jar (rice grains and freshly laid eggs) was kept for further multiplication. The subsequent progenies of weevils were used for conducting experiment. Grains and insects were removed and sieved periodically. Removed insects were again placed in the same containers along with fresh grains. Adult weevils were identified sexually based on the rostrum type by observing under a trinocular microscope at 40 X magnifications (Nikon SMZ-745T) (Reddy 1951).

Characterization of physical parameters of grain

Thirty grains from each variety were taken for length and breadth measurement using digital vernier calipers and expressed in millimeter (mm). Further, length breadth ratio (LBR) of the grain was calculated. Likewise, hundred seed weight (HSW) was also recorded for each variety. Grain hardness was measured by pressure exertion method using a texture analyzer (TA.XT plus, Stable Microsystems, UK; capacity 50 kg; speed range: 0.1–40 mm/s). On each individual rice grain pressure was exerted till grain cracked and reading was recorded at this point (Kaur 2017).

Characterization of bio-chemical properties of grain

Soluble protein was estimated by standard and universally acceptable method (Lowry et al. 1951). Similarly amylose content of rice was determined by the modified method of Juliano (1971). Amylose content was calculated according to the following formula; % Amylose = Absorbance × 75 (factor).

Characterization of developmental parameters of weevil

Adult emergence (AE)

In no-choice tests, 300 kernels of each variety were divided in three replicates of 100 kernels each and their initial weight was measured, they were kept in Petri dishes and closed with lid. Treatments were arranged in Completely Randomized Design (CRD). Six pairs of freshly emerged weevils of the same age were placed on test variety of rice and left for 1 week. After 1 week, the weevils were removed from the grain and Petri dishes were kept undisturbed and monitored regularly until the emergence of first adult from the grains. Inspection of the progenies was made every day and they were counted and removed during each observation. The observations were continued until no adults emerged for ten consecutive days (Nwana and Akibo-Betts 1982).

Median development period (MDP)

It is the time taken for 50% of adults to emerge. It was calculated using the formula (Howe 1971):
$$MDP = \frac{D1A1 + D2A2 + D3A3 + \ldots \ldots \ldots \ldots \ldots \ldots \ldots \ldots .DnAn}{Total \,number\, of\, adults\, emerged},$$
where D1—day at which the adults started emerging (First day) (Siwale et al. 2009), A1—number of adults emerged on D1st day.

Weight loss (WL)

The count and weigh method was followed to determine grain weight loss (Gwinner and Harnish (1996)). When weevil stopped their emergence, numbers as well as weight of undamaged and damaged grains were considered for each replication, weight loss percentage was calculated by:
$$WL\,(\% ) = \frac{{\left( {Wu \times Nd} \right) - \left( {Wd \times Nu} \right)}}{{Wu \times \left( {Nd + Nu} \right)}} \times 100$$

Wu—wt. of undamaged grains, Nu—number of undamaged grains, Wd—wt. of damaged grain, and Nd—number of damaged grains.

Susceptibility index (SI)

The index of susceptibility is an important characteristic that indirectly determines the development of weevil. It was calculated using the method of Dobie (1974). This involves the number of F1 progeny and the length of median developmental time.

For each variety, the SI, was determined as below:
$$SI\, = \,\left( {Log_{e} F} \right)/D\, \times \,100$$

F—total number of F1 adults, D—median development period = ∑ xy/∑ x, x—number of adult weevils emerged, y—number of days of the infestation until adult emergence.

The susceptibility index was used to classify the varieties, where 0–3 (resistant), 4–7 (moderately resistant), 8–10 (susceptible) and > 11 (highly susceptible).

Growth index (GI)

GI, a key factor to determine host suitability determining was calculated as adult emergence (%)/mean development period (days) (Tripathi et al. 2012; Soumia et al. 2015). The growth index of S. oryzae in each replication was worked out by the following formula given by Naveena et al. (2011):
$${\text{Growth index = }}\frac{\text{Percent adult emergence}}{\text{Average developmental period}}$$

Data analysis

The data on hundred seed weight, length breadth ratio, median development period, adult emergence, weight loss, index of susceptibility, growth index, grain hardness, protein and amylose were subjected to one-way ANOVA in randomized unblock design. Further, data were also subjected to multivariate analysis using statistical package SAS 9.1 (http://stat.iasri.res.in/sscnarsportal) and relation among the parameters was noted by the values of co-efficient of regression. For further grouping the physico-chemical and developmental parameters and response of varieties based on their affinities principal component analysis (PCA) was done.

Results

Characterization of physical parameters of grain

Length breadth ratio (LBR)

LBR of 15 rice varieties had ranged from 1.84 to 4.50 mm (Table 2). Significant variation (P < 0.0001) with respect to length and breadth of varieties was recorded. Based on LBR, selected varieties were falling in the category of short slender (Ketakijoha, Padmini and Satabdi), short broad (Chinikamini, Kalajeera, Swarna Sub1, TN1, Sarala and Gayatri), medium slender (Dhusara, Satyabhama and Pyari), long slender (Geetanjali) and long broad (Cross 12). The grain LBR was positively correlated with 100 seed weight (0.301*) and grain hardness (r = 0.529**) (Table 4).
Table 2

Physico-chemical characterization of the different rice varieties

Sl. no

Variety

LBR

HSW (g)

GH (N)

Protein (mg/g)

Amylose (%)

1.

Dhusara (MS)

2.52

0.93

166.15

103.04

21.9

2.

Chinikamini (SB)

1.84

1.01

168.9

97.81

22.35

3.

Kalajeera (SB)

2.19

1.17

139.39

101.05

20.77

4.

Geetanjali (LS)

4.50

1.92

253.71

90.59

23.4

5.

Satyabhama (MS)

3.06

1.8

159.82

69.94

17.62

6.

Naveen (MB)

2.72

1.38

190.09

101.29

24.75

7.

Swarna sub 1 (SB)

2.62

1.51

239.99

71.43

26.17

8.

Ketakijoha (SS)

3.59

1.47

205.26

79.64

22.05

9.

TN 1 (SB)

2.28

1.63

194.82

72.42

25.2

10.

Cross 12 (LB)

2.61

2.00

259.16

55.75

25.27

11.

Padmini (SS)

3.21

1.07

201.10

103.53

24.45

12.

Pyari (MS)

2.78

1.54

166.15

80.39

26.77

13.

Sarala (SB)

3.16

1.48

210.46

78.4

25.87

14.

Gayatri (SB)

2.12

1.81

167.07

80.39

27

15.

Satabdi (SS)

3.85

1.46

220.29

78.89

26.1

F test

**

**

**

**

**

CV (%)

9.37

1.92

0.12

2.01

2.35

LBR length breadth ratio, HSW hundred seed weight, GH grain hardness

**Significant at 0.01 level

Seed weight

The size of the grain is an important factor in determining resistance. The 100 seed weight of test varieties ranged from 0.93 to 2.00 g indicating significant varietal divergence (P < 0.0001) (Table 2). Seed weight was found to have significant strong negative relation with weevil emergence (r = − 0.53**), susceptibility index (r = − 0.51**), growth index (r = − 0.52**) and protein (r = − 0.70) and but significant positive relation with median development period (r = 0.32*), length breadth ratio (r = 0.30*) and grain hardness (r = 0.493**) (Table 4).

Grain hardness

Important factors associated with grain resistance are morphological characteristics, which include hardness of the grains (Marsaro Júnior et al. 2005; Fontes et al. 2003; McGaughey et al. 1990). Hardness of tested varieties in the present study ranged from 139.39 to 259.16 N and were significantly different (P < 0.0001) (Table 2). Grain hardness had significant was positive correlation with length breadth ratio (r = 0.527**), 100 seed weight (r = 0.493**), amylose (r = 0.403**). It had a negative correlation with weevil emergence (r = − 407**), protein content (r = − 0.460**), susceptibility index (r = − 0.337), weight loss (r = − 0.245) and growth index (− r = 0.330*) and (Table 4).

Characterization of bio-chemical properties of grain

Protein Total soluble protein of rice kernel ranged from 55.75 mg/g (Cross 12 variety) to 103.53 mg/g (Padmini variety). All the varieties analyzed for protein were significantly different (Table 2). Protein was significantly and positively related with weevil emergence (r = 0.740**); index of susceptibility (r = 0.717*); growth index (r = 0.741**). But it had a strong significant negative relation with median development period (r = − 0.537**); 100 seed weight (r = − 0.770**) and amylose (r = − 0.221) (Table 4).

Amylose Amylose content of rice kernels varied from 17.62 to 27.00% and was significantly different among tested varieties (P < 0.0001). It exhibited positive correlation with grain hardness (r = 0.403**) and hundred seed weight (r = 0.206). Whereas it has negative correlation with other characteristics like protein (r = − 0.221); seed loss (r = − 0.053); weevil emergence (r = − 0.007); median development period (r = − 0.093) and length breadth ratio (r = − 0.005).

Characterization of developmental parameters of weevil

Adult emergence

Considerable variation was observed with respect to weevil emergence. This difference in the susceptibility of rice varieties is due to differences in the particular variety to resist weevil attack. From infested grains, total number of emerged weevils had ranged from 7.33 to 75.67 (Table 3). There positively significant correlation was noticed between weevil emergence and growth index (r = 0.972**), protein (r = 0.734**) and seed loss (r = 0.434**). Median development period (r = − 0.667**), length breadth ratio (r = − 0.057), hundred seed weight (r = − 0.534**) and grain hardness (r = − 0.407**) had significant and negative correlation (Table 4).
Table 3

Developmental and susceptibility parameters of rice weevil on different rice varieties

Sl. no.

Variety

AE

MDP (days)

SI

WL (%)

GI

1.

Dhusara

74.33

34.00

12.67

9.50

2.19

2.

Chinikamini

61.00

36.67

11.21

5.58

1.66

3.

Kalajeera

50.33

42.00

9.33

13.03

1.20

4.

Geetanjali

58.67

35.00

11.63

9.30

1.68

5.

Satyabhama

41.33

39.67

9.38

10.24

1.04

6.

Naveen

67.33

35.00

12.03

11.51

1.92

7.

Swarna sub 1

50.00

38.00

10.29

6.43

1.32

8.

Ketakijoha

49.33

41.33

9.43

5.56

1.19

9.

TN 1

47.67

36.33

10.64

7.81

1.31

10.

Cross 12

7.33

43.67

4.56

0.59

0.17

11.

Padmini

75.67

32.00

13.52

20.64

2.36

12.

Pyari

52.00

38.67

10.22

7.63

1.34

13.

Sarala

40.33

42.00

8.80

5.96

0.96

14.

Gayatri

72.67

36.33

11.80

18.10

2.00

15.

Satabdi

49.33

39.33

9.91

10.50

1.25

F test

**

**

**

**

**

CV (%)

15.04

5.79

7.75

1.43

16.22

AE adult emergence, MDP median developmental period, SI susceptibility index, WL weight loss, GI growth index

**Significant at 0.01 level

Table 4

Correlation co-efficients of physico-chemical characteristics of grain and insect development parameters in rice due to S. oryzae infestation

 

AE

MDP

LBR

HSW

SI

GI

Amylose

GH

Protein

WL

AE

1.00000

− 0.667**

− 0.057

− 0.534**

0.916**

0.972**

− 0.008

− 0.408**

0.739**

0.434**

MDP

 

1.00000

− 0.032

0.326

− 0.856**

− 0.804**

− 0.094

0.100

− 0.537**

− 0.185

LBR

  

1.00000

0.301*

0.02

− 0.043

− 0.006

0.530**

− 0.07

0.172

HSW

   

1.00000

− 0.510**

− 0.524**

0.207

0.494**

− 0.770**

− 0.153

SI

    

1.00000

0.952**

0.004

− 0.338

0.717**

0.482**

GI

     

1.00000

0.021

− 0.331*

0.741**

0.328*

Amylose

      

1.00000

0.403**

− 0.221

− 0.053

GH

       

1.00000

− 0.460**

− 0.245

Protein

        

1.00000

0.201

WL

         

1.00000

AE adult emergence, MDP median developmental period, LBR length breadth ratio, HSW hundred seed weight, SI susceptibility index, GI growth index, GH grain hardness, WL Weight loss

*Significant at 0.05 level, **Significant at 0.01 level

Median development period (MDP)

The MDP of S. oryzae had ranged from 32.00 to 44.3 days (Table 3). The negative association was observed among MDP and weevil emergence (r = − 0.667**), index of susceptibility (r = − 0.856**), growth index (r = − 0.804**), protein (r = − 0.537**) and seed loss (r = − 0.186). A positive correlation was found with hundred seed weight (r = 0.326) and grain hardness (r = 0.099) (Table 4).

Weight loss

The highest per cent of weight loss was found in genotype Padmini (20.64), while Cross 12 (0.59) reported to have minimum (Table 3). Weight loss was having significant positive relation with the weevil emergence (r = 0.434**). With an increase in susceptibility index, there was an increasing and highly significant percentage of grain weight loss (r = 0.482**). Negative correlation was observed with median development period (r = − 0.185) and grain hardness (r = − 0.245) (Table 4).

Susceptibility index (SI)

The index of susceptibility ranged from 4.56 for variety Cross 12 and 13.52 for variety Padmini (Table 3). Out of the 15 rice varieties, only Cross-12 was found to be moderately resistant as per Dobie (1974) rating scale. SI was significantly and negatively correlated with median developmental time (r = − 0.856**). Same is the relation with seed weight also (r = − 0.510**). However factors such as weevil emergence (r = 0.916**), growth index (r = 0.952**), protein (r = 0.717**) and seed loss (r = 0.482**) showed a positive significant relationship with the susceptibility index (Table 4). In the present findings, most significant positive correlation was found with adult emergence.

Growth index (GI)

Differences among the varieties as per their susceptibility/resistance to S. oryzae on the basis of GI were observed. It ranged from 0.17 for variety Cross 12–2.36 for Padmini. Growth index was positively and significantly correlated with weevil emergence (r = 0.972**), index of susceptibility (r = 0.952**), protein (r = 0.717**) and seed loss (r = 0.482**). But it had a negative correlation with median development period (r = − 0.804**) and grain hardness (r = − 0.330**) and seed weight (r = − 0.524**). It is explicitly pointed that larger and softer grains favour S. oryzae development.

Principal component analysis (PCA) was performed to know the relatedness among different characteristics (physical, biochemical and susceptibility parameters) used in the present study. On the other hand, taking 15 varieties into consideration, PCA was performed in order to assess the relation among different varieties. Two principal components (PCs) were extracted for the physico-chemical characteristics from scree plot with eigen value ≥ 1.0. The diversity of different physico-chemical and developmental parameters is presented in Fig. 1. PC1 depicted variation of 49.57%, whereas PC2 depicted variation of 17.88% (Fig. 1). Component loadings of different factors governing resistance for S. oryzae in rice are represented in Table 5. Most of the parameters namely, AE, MDP, HSW, SI, Protein, GI and WL have higher coefficient values in PC1, whereas, only three components namely, amylose, HI and LBR were represented by PC2. Two main groups, comprising different parameters, were identified based on positional proximity in the 2-D biplot. Parameters such as weevil emergence, index of susceptibility and growth index were located close to one another, whereas, length breadth ratio, amylose, grain hardness and hundred seed weight occupied a different location on the 2-D plot. Apart from these groups, median development period was quite distant from the remaining indicating a different trend. Pearson correlation matrix (Table 4) also supported the above statement in terms of degree of relationship among them.
Fig. 1

2-D plot of principal component analysis based on physico-chemical and developmental parameters for resistance against S. oryzae in rice. AE adult emergence, MDP median developmental period, LBR length breadth ratio, HSW hundred seed weight SI susceptibility index, GI growth index, GH grain hardness, WL weight loss

Table 5

Component loadings of physico-chemical and developmental parameters for resistance against S. oryzae

Sl. no.

Parameter

Principal components

PC1

PC2

1.

AE

0.421

0.105

2.

MDP

− 0.341

− 0.288

3.

HSW

− 0.315

0.306

4.

SI

0.427

0.193

5.

GI

0.425

0.159

6.

Amylose

− 0.054

0.411

7.

GH

− 0.233

0.533

8.

Protein

0.384

− 0.118

9.

WL

0.194

0.138

10.

LBR

− 0.059

0.513

AE adult emergence, MDP median developmental period, LBR length breadth ratio, HSW hundred seed weight, SI susceptibility index, GI growth index, GH grain hardness, WL weight loss

Similarly, two principal components (PCs) were extracted for response of different varieties with respect to different parameters from scree plot with eigen value ≥ 1.0, PC1 explained 91.86% of the variation while PC2 explained 5.06% of variation for varieties (Fig. 2). Component loadings of performance of different varieties based on different parameters are represented in Table 6. Plot for the varieties showed discrete pattern which could be understood by their diverse relation among them. Two distinct main groups comprising of different varieties are being observed here. All the varieties except Cross 12 were made one group. Variety Cross 12 showed distinct trend of relation among varieties. The results of PCA were broadly congruent with the data observed among varieties with respect to different parameters (table). For example, Cross 12 had highest 100 seed weight (2 g); highest grain hardness (259.16 N); lowest protein content (55.75 mg/g), lowest adult emergence (7.33 weevil), highest median development period (43.67 days) and lowest seed weight loss (0.59%).
Fig. 2

2-D plot of principal component analysis based on varietal resistance against S. oryzae. v1-Dhusara; v2-Chinikamini; v3-Kalajeera; v4-Geetanjali; v5-Satyabhama; v6-Naveen; v7-Swarna Sub 1; v8-Ketakijoha; v9-TN 1; v10-Cross 12; v11-Padmini; v12-Pyari; v13-Sarala; v14-Gayatri; v15-Satabdi

Table 6

Component loadings of rice varietal resistance against S. oryzae

Sl. no.

Variety

Principal components

PC1

PC2

1.

Dhusara

0.260

0.067

2.

Chinikamini

0.267

− 0.064

3.

Kalajeera

0.263

0.186

4.

Geetanjali

0.266

− 0.073

5.

Satyabhama

0.264

− 0.110

6.

Naveen

0.265

0.044

7.

Swarna sub 1

0.263

− 0.143

8.

Ketakijoha

0.263

− 0.163

9.

TN 1

0.267

− 0.097

10.

Cross 12

0.157

0.916

11.

Padmini

0.258

0.086

12.

Pyari

0.265

− 0.071

13.

Sarala

0.262

− 0.089

14.

Gayatri

0.263

− 0.146

15.

Satabdi

0.269

0.031

Discussion

In present study it was observed that grain characteristics have association to their susceptibility to weevil infestation. Our results are in conformity with the fact that preference of S. oryzae for large kernels was not constant but was density dependent (Ewer 1945). Similar findings that weevil emergence was negatively correlated with kernel size in wheat and sorghum were also reported by Stejskal and Kucerova (1996) and Prasad et al. (2015), affirming the present results. However, present findings contradict some of the previous studies (Segrove 1951; Pederson 1979; Stejskal and Kucerova 1996), indicating that Sitophilus spp. prefers larger kernels; there is a possibility that these studies might have typically looked at the distribution of eggs on grains when exposed to multiple females. Weevil emergence from the 15 varieties in the present findings support the study of Gudrups et al. (2001) who found size of a grain is crucial factor in governing resistance to Sitophilus zeamais in maize, where larger or bigger grains have better resistance than small grains. The grain size can affect fecundity and survival of the insect pest (Antunes et al. 2016). Larger grains are expected to have greater larval survival and more offspring than in smaller seeds (Segrove 1951; Haine 1991; Campbell 2002). Female assess the quality of rice grain and vary the number of eggs laid. Present results were in line with the results of Prasad et al. (2015), who found a negative relationship between seed weight and Sitophilus emergence from sorghum grains.

Greater kernel hardness was found associated with lower insect fitness (Morallo-Rejesus et al. 1982; Throne et al. 2000). The resistance of rice varieties has been related to mechanical structures of the grains (Lale and Yusuf 2001; Ashamo 2001; Lale and Kartay 2006). Garcia-Lara et al. (2004) found that S. zeamais resistance is controlled by kernel hardness. For S. oryzae, grain hardness was the major resistance parameter (Bamaiyi et al. 2007), corroborating our study where susceptibility to the rice weevil attack (in terms of weevil emergence) and grain hardness had a negative relationship. A very recent report suggested that wheat kernel hardness was found to be positively contributed to their resistance against weevils (Saad et al. 2018). Similar observations were also reported by previous workers (Soujanya et al. 2017; Krishna and Lakshmi 2008; Garcia-Lara et al. 2004). Studies of Morallo-Rejesus et al. (1982) and Antunes et al. (2016) showed susceptibility index and F1 progeny of S. zeamais adults were correlated negatively with grain hardness like in our study. Present study did not agree with study of Gudrups et al. (2001), where in no significant association was found between kernel hardness and weevil emergence in maize also showing differences in grain hardness is attributed to size of the grain.

Grain damage by storage insects may be influenced by the chemical composition (of grains) (Piesik and Wenda-Piesik 2015; Wenda-Piesik et al. 2016). The protein is an important factor in determining the susceptibility level of the grain. It is clear from the present study that kernel with higher protein are more preferred for rice weevil hence showing susceptibility. Results of the present study are in perfect conformity with the findings of Sahoo and Sahoo (2016), who reported that population emergence and the percent grain damage by S. oryzae is significantly and positively correlated with the kernel protein with correlation coefficients of 0.827 and 0.878, respectively. Also our results are in close agreement with the fact that varieties with more protein were highly susceptible in case of Sitotroga cerealella (Murad and Batool 2017) and Sitophilus oryzae (Soujanya et al. 2017). In contrast to the current study, Siwale et al. (2009) reported that resistance to maize weevil in grain is imparted by higher protein. Amylose is also an important component of rice starch governing resistance of rice grain. Present results were in agreement with Peters et al. (1972) who reported higher amylose content in resistant varieties of corn than susceptible ones against weevils and beetles. Amylose (one of the two components of starch) was found negatively related with weevil emergence. Comparable results were reported on the starch as susceptibility parameter to insect pests of storage (Osipitan and Odebiyi 2007; Chijindu and Boateng 2008).

In general rice is more susceptible to weevil attack than paddy; among rice varieties, polished rice had a higher susceptibility (Antunes et al. 2016; Seshagiri Rao 1953). In the current study, weevil emergence from grains correlated positively with grain weight loss. It is in agreement with Singh et al. (1984), who reported numbers of emerging adults govern the extent of damage, and therefore, grains permitting faster and higher levels of adult emergence were damaged significantly. In addition, Mohammad et al. (1988) stated that, grain weight loss is crucial factor which depends on adults emerged (number) and damaged grains. Grain hardness is another important physical characteristic of rice grain governing the resistance. Our results showed that adult emergence was significantly and negatively correlated with hardness affirming the findings of Tripathi et al. (2017) who stated significant negative relation between seed hardness and weevils' development, meaning harder the seeds, more the resistance to S. oryzae.

It can be concluded from the present findings that resistant varieties prolong the developmental period (negative correlation between SI and MDP) and greater mortality of the larvae developing inside the kernel. Present results agree with Dobie (1984) who reported that resistant maize cultivars extend the developmental period of S. zeamis. This means that weevil population will remain low resulting in less damage to rice varieties. Prolonged development periods will lead to reduction in number of generations of insect. Varieties with higher susceptibility index show decreased development of weevils (Abebe et al. 2009) as confirmed in present study as well. Present results are in corroboration with Horber (1988), who reported that index of susceptibility is based on the fact that the higher the F1 progeny, lesser the development period, and the more susceptible the grains would be. Abraham (1991) also reported that the level of damage during storage primarily depends on number of adults emerging during each generation and the duration of each generation; hence grains allowing rapid and increased levels of adult emergence will be heavily damaged. It can be stated that because of availability of more space and food in larger grains, the development of weevil is better compared to smaller grains, (Tripathi et al. 2017; Hossain et al. 2014; Keba and Sori 2013; Derera et al. 2014).

The results presented here indicate that variable response of S. oryzae to rice varieties was observed. This was probably due to the extensive variability in the physical and biochemical characteristics of the grain. Since resistance was correlated to bio-physical grain traits like kernel hardness, seed weight, length, breadth and biochemical parameters viz., amylose, protein, these chacracteristics need to be considered while designing breeding strategies to develop rice varieties for weevil resistance. This study also suggests that single parameter is not sufficient to predict adequately the resistance against S. oryzae in rice. To date, breeding programmes conducted in India have concentrated more on field insect pest than storage pests, and very little attempt has been made on developing S. oryzae resistant lines. The outcome of the present study could be used as source for breeding programs to develop resistant cultivars against this notorius pest and offers sustainable, cost effective, and eco-friendly solution for management of S. oryzae.

Notes

Acknowledgements

Authors gratefully acknowledge Director, ICAR-National Rice Research Institute, Cuttack, India for constant support in formulating the project, funding as well as providing all the facilities (Grant no. Institute project 3.1). Authors duly acknowledge Dr. H. N. Subudhi, Principal Scientist, ICAR-NRRI for supplying seeds of rice varieties used in the study. We thank Dr. N. N. Jambhulkar, Scientist, ICAR-NRRI for his guidance in statistical analysis and Dr. Sutapa Sarkar for sharing pedigree information. The authors declare that they have no conflict of interest.

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Copyright information

© Society for Environmental Sustainability 2019

Authors and Affiliations

  • G. Basana Gowda
    • 1
    Email author
  • Naveenkumar B. Patil
    • 1
  • Totan Adak
    • 1
  • Guru Pirasanna Pandi
    • 1
  • Nabaneeta Basak
    • 2
  • Kingshuk Dhali
    • 3
  • M. Annamalai
    • 1
  • G. Prasanthi
    • 1
  • S. D. Mohapatra
    • 1
  • Mayabini Jena
    • 1
  • Somnath Pokhare
    • 1
  • P. C. Rath
    • 1
  1. 1.Crop Protection DivisionICAR-National Rice Research InstituteCuttackIndia
  2. 2.Crop Physiology and Bio-chemistry DivisionICAR-National Rice Research InstituteCuttackIndia
  3. 3.Faculty of Agricultural EngineeringBidhan Chandra Krishi ViswavidyalayaMohanpurIndia

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