Population studies of three stored grain pests on two common legumes, the common pea and the Indian pea

Introduction

Legumes (Family: Fabaceae) are a major source of affordable dietary nutrition globally. They also augment soil nitrogen by nitrogen fixation, constituting an integral component of the farming systems. India is the leading producer and consumer of legumes, contributing about 25-28% of the total global production (Naveen et al., 2022) [36] . Common pea (Pisum sativum L.) is among the second most important legume crops in the world. Pea seeds contain high levels of amino acids, proteins, minerals and vitamins (Kalapchieva et al., 2020) [26] . Indian pea (Lathyrus sativus L.) is another crop of immense economic significance as food, feed and fodder in India and many other countries. However, it produces a neurotoxin [β-N-oxalyl-l-α, β-diaminopropionic acid (ODAP)], prolonged consumption of which may lead to the neurological disorder lathyrism in humans and domestic animals (Dixit et al., 2016) [18] . Despite the greater production of pulses, the production needs to be increased to attain the goal of self-sufficiency in India (Soumia et al., 2015) [57] . Moreover, post-harvest losses result in a 10% decline in the stored crops, out of which 26% can be attributed to insect pests (Kalpna et al., 2022) [27] . The cowpea weevil, Callosobruchus maculatus (Fabricius), is regarded as the most important and cosmopolitan storage pest of several species of pulse grains, such as cowpea, chickpea, garden pea and lentils (Osman et al., 2015) [38] . The rice weevil, Sitophilus oryzae L. (Coleoptera: Curculionidae), is a major primary pest of stored rice worldwide. They are capable of infesting corn, wheat, barley, sorghum, oats, rye and dried beans (Zahir et al., 2012) [61] . The red flour beetle, Tribolium castaneum Herbst (Coleoptera: Tenebrionidae), is a prevalent polyphagous secondary pest of stored grains with a worldwide distribution. This species has been recorded on 246 commodities, showing a preference for flours and amylaceous by-products (Duarte et al. 2021) [19] . Insect feeding leads to extensive losses in the quality and quantity of nutritional and aesthetic values of products, weight losses, deterioration of seed viability and secondary infestations (Malaikozhundan and Vinodhini, 2018) [31] .

Insect survival is influenced by the primary and secondary phytochemicals of hosts (Demissie et al., 2015) [16] . The primary metabolites affect the growth and reproduction of herbivorous pests, while the secondary metabolites serve defensive roles against them (Chowdhury et al., 2025) [13] . Stored grains provide shelter, security and abundant food to the pests. Grain transportation and marketing networks then pave the way for the local and global dispersal of these species (Nopsa et al., 2015) [37] . However, farmers use broad-spectrum synthetic pesticides and biorational treatments to manage pest species, without considering the growth rates of pest populations (Carvalho, 2017) [10] . This leads to regulatory complications in the ecosystem (Kang, 2019) [28] . Therefore, it is necessary to identify how the characteristics of the host legumes are linked with their susceptibility (Murad and Batool, 2017) [34] . This is an important aspect of integrated pest management (IPM) that can reduce or eliminate the need for harmful insecticides (Quellhorst et al., 2021) [41] . Thus, the objective of this research was to study and compare the biological characteristics of C. maculatus, S. granarius and T. castaneum when fed on common pea and Indian pea, under the same environmental conditions, and to assess the susceptibility of the pulses to the selected insect pests.

Material and Methods

Collection of the legumes and maintenance of insect cultures Uninfested seeds of common pea (CV-Arka Priya) and Indian pea (CV-BioL-212 (Ratan)) and seeds infested with C. maculatus, S. oryzae and T. castaneum were collected from the FCI godown, Chinsurah, Hooghly (22°54' N, 88°23' E), different commercial stores of Purba Bardhaman (23°13' N, 87°51' E) and from farmers of Indas, Bankura (23.1504° N, 87.6318° E), West Bengal, India. We selected the uninfested seeds after separating the damaged seeds by physical inspection. Then we washed the seeds properly with distilled water, followed by heat drying (45ºC) for 30 mins and UV-treatment (254 nm) for 20 mins. After the treatments, they were refrigerated (0ºC) in airtight containers for further studies. To ensure disinfestation, a control set (with 100 seeds of each legume in separate containers) was also observed during the course of the population studies. The uninfested seeds of the selected legume cultivars, with a moisture content of 8-10%, were divided into two parts. One part of each legume cultivar served as host for each pest on which the population studies were conducted, and the other part was prepared for various phytochemical analyses. The stock cultures of the pests were initiated with adults (F 1 ) from the infested seeds in a growth chamber [30 eggs and 100 seeds in a glass jar (20 cm diameter × 30 cm height)]. The cultures were maintained from F 1 to F 4 generations separately on the selected legumes, and the F 4 generation was used for population studies with three replicates. All the culture sets were maintained under laboratory conditions (27 ± 1°C, 65 ± 5% RH, 12:12 [L: D] photoperiod and 1500 lux light intensity).

Phytochemical estimation of the legumes

Freshly collected uninfested seeds of common pea and Indian pea were initially rinsed with distilled water and dried with a paper towel. They were then dipped in different solvents for the extraction of various primary and secondary metabolites. The concentrations of each metabolite were estimated by using standard procedures: carbohydrates (DuBois et al., 1956) [20] , total proteins (Miller, 1959) [32] , lipids (Folch et al., 1957) [22] , amino acids (Moore and Stein, 1948) [33] , nitrogen (Humphries, 1956) [24] , moisture (Roy and Barik, 2013) [44] , ash (Banerjee and Haque, 1984) [5] , phenols (Bray and Thorpe, 1954) [7] , flavonoids (Zhishen et al., 1999) [64] , tannin (Trease and Evans, 1983) [59] , saponin (Trease and Evans, 1983) [59] , alkaloids (Harborne, 1973) [23] , phytates (Reddy and Love, 1999) [43] and oxalates (Day and Underwood, 1986). Total alkanes (Roy et al., 2013b) [46] , free fatty acids (Roy et al., 2012) [47] , and free and bound amino acids (Roy et al., 2013a) [45] were also quantified. All the phytochemicals have been estimated using various protocols, as in Roy and Barik (2013) [44] . Each test was repeated three times and expressed on a dry or fresh weight basis (Roy, 2022; Chowdhury et al., 2025) [50,13] .

Population growth of the insect pests

Ten pairs of freshly emerged males and females from the stock culture (F 3 ) were placed in an oviposition cage of fine nylon net [(25 × 25 × 25) cm 3 ]. The pests were allowed to oviposit separately on each of the selected legumes. Newly laid eggs (n=30) from F 3 females, belonging to a defined cohort, were collected for the population studies. Therefore, the population studies began with the eggs of 3 rd generation (F 3 ) females, i.e., the F 4 generation. Damaged seeds in the oviposition cage were replaced periodically with fresh ones. The larvae were placed in five glass jars [20 cm diameter × 30 cm height] containing the different seeds, and all the larval instars (I-V) were monitored. The mouths of the containers were covered with cotton cloths. Fresh seeds were again provided periodically. The pupae were then placed in separate glass jars [6 cm diameter × 10 cm height], covered with cotton cloth, until adult emergence. The males and females were differentiated based on their distinctive physical features. Adult females are larger than the adult males, and males have a more slender abdomen than females. Three replicates were maintained on each legume, and the experiments were repeated accordingly. Survival, mortality, fecundity and developmental durations from egg to adult were recorded separately on both hosts. The data on survival, oviposition and developmental durations of all individuals on the selected legumes were subjected to stage-specific, two-sex life tables analyses (Chen et al., 2017; Jaleel et al., 2018) [11,25] . The parameters included probability of survival from birth to age x (l x ), proportion dying at each stage (d x ), mortality (q x ), survival rate (s x ) per day per age class from egg to adult stages (Carey, 1993) [9] . Using these parameters the following statistics like total individuals at age x and beyond k (T x ), average population alive at each stage (L x ), life expectancy (e x ), exponential mortality or killing power (k x ), generation mortality (GM), generation survival (GS), gross reproductive rate (GRR), net reproductive rate (NRR or R 0 ), mean generation time (T c ), doubling time (DT), intrinsic rate of population increase (r m ), finite rate of population increase (λ), weekly multiplication rate (λ 7 ), increase rate per generation (λ Tc ), which were computed by Carey's formulae (Price 1997) [40] . Some other demographic parameters like potential fecundity (P f ), total fertility rate (TFR), mortality coefficient (MC), population growth rate (PGR), effective population (N e ), population momentum factor of increase (PMF), number of individuals after time t (N t ), expected population size in 2nd generation (PF 2 ), Hypothetical females in 2 nd generation (HFF 2 ), general fertility rate (GFR), crude birth rate (CBR), reproductive value (RV), vital index (VI) and trend index (TI) were also analysed using various well-defined formulae (Southwood, 1978; Carey, 2001; Chowdhury et al., 2025) [58,8,13] .

Statistical analysis

Correlation analysis has also been performed to demonstrate how variation in metabolites influences insect population growth (Zar, 1999) [62] . Experimental data on metabolites in the green gram cultivar and population parameters of the pests were homogeneous among treatments, as confirmed by Levene's homogeneity test. The data were normally distributed, as determined by Shapiro-Wilk tests, and were compared using one-way ANOVA (Zar, 1999) [62] . Means associated with all the data were separated using Tukey's (HSD) test when the values were significant (Zar, 1999) [62] . Statistical analyses have been performed using SPSS version 16.0.

Result

Host phytochemicals

The primary and secondary metabolites varied significantly 1). Common pea contained higher levels of the primary and secondary metabolites than the Indian pea. The major primary metabolites (µg/mg dwt), total carbohydrates, proteins and lipids in common pea were 57.589±1.185, 13.789±1.105 and 12.389±1.135, respectively, and in Indian pea they were 50.310±0.788, 6.510±0.708 and 5.110±0.738, respectively. Total amino acids and nitrogen (%) followed a similar order, with their higher concentrations in common pea than in Indian pea (Fig 1(a), Supplementary Table 1). The content of free amino acids was lower than that of alkanes, free fatty acids and bound amino acids in both the legumes, and they followed the same pattern as the major primary metabolites: common pea > Indian pea (Fig 1(b), Supplementary Table 1). The concentrations of phenols, saponins and alkaloids (µg/mg dry weight) were also in the order of common pea (20.689±0.993, 19.829±1.013 and 18.896±0.655) > Indian pea (13.867±0.667, 12.673±1.018 and 11.073±1.068). All the other secondary metabolites followed the same pattern (Fig 1(a), Supplementary Table 1). Ultimately, the ratio of primary to secondary metabolites differed significantly in the selected legumes.

Population growth of the insect pests

On common pea, the longest incubation period (days) was exhibited by T. castaneum (12.279±0.386), followed by C. maculatus (6.375±0.217) and S. granarius (5.375±0.217). The total larval period (days) was also longest in T. castaneum (58. The stage-specific life tables of the pests were examined separately for each pest on both legumes. The values of l x , L x , T x and e x gradually decreased with the developmental stages. The l x values were 0.293±0.025, 0.125±0.013 and 0.221±0.01 for the adults of C. maculatus, S. granarius and T. castaneum, respectively, on common pea; and 0.217±0.026, 0.088±0.015 and 0.259±0.011 on Indian pea. In our study, l x dropped gradually from egg to advanced developmental stages in all pest species on both host legumes. Therefore, they exhibited Type III survivorship curves, like most other insects (Table 1 andTable 2, Fig 3). Similar to the l x , the values of L x and T x also gradually decreased across the developmental stages; e x increased from egg to instar-I, after which it decreased consistently; q x and k x followed a more random pattern. In both hosts, l x , L x , T x , and e x were in the order of T. castaneum > C. maculatus > S. granarius, with exceptions; q x and k x were in the order of S. granarius > C. maculatus > T. castaneum. The values of q x , e x and k x for the adult T. castaneum were 0.884±0.039, 0.558±0.02 and 0.998±0.175, respectively, on common pea; and 0.753±0.031, 0.623±0.016 and 0.615±0.055 on Indian pea. C. maculatus and S. granarius adults had the same values of q x , e x and k x , 1.000±0.000, 0.500±0.000 and 0.201±0.000, respectively, on both common pea and Indian pea. The population parameters presented in Tables 1 (F Comparing the population parameters of each pest on the two hosts, it seems that the common pea was a more suitable host for C. maculatus and S. granarius than the Indian pea. In contrast, the Indian pea was a more suitable host for T. castaneum than the common pea (Supplementary Tables 2, 3 and 4). The primary metabolites (such as carbohydrates, proteins, lipids, alkanes and fatty acids) of the hosts were positively correlated with the population parameters of the pests, except for r m , λ and MC. In contrast, the secondary metabolites (ash (%), phenols, flavonoids, tannins, saponins, alkaloids and phytates) showed negative correlations with GRR, NRR, T c , r m , λ, PGR, PF 2 and HFF 2 of the pests (Supplementary Table 5). The correlation coefficients (r) for individual metabolites and population parameters (i.e., along rows and columns) of the pests varied considerably.

Discussion

Insects seek hosts that can provide optimal nutrition and a suitable mating and oviposition site (Scriber and Slansky, 1981; Anukiruthika et al., 2021) [53,1] . Conversely, the hosts have developed complex networks to regulate a balance between growth and defence (Kessler and Baldwin, 2002)

Acknowledgement

AcknowledgementsWe express our deepest gratitude to the University Grant Commission (UGC: NTA Ref. No. 211610066466), India, for financial assistance. We must acknowledge the departmental staff members who supported us in every way possible during the work. Finally, we thank the farmers who helped us all through. NC, SP and NR designed the entire study, including sample collection, chemical analysis, index calculation, data analysis and drafted the manuscript with the help of institutional support. The author declares that there have been no conflicts of interest.

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