Egg laying decisions by the bean weevil Callosobruchus maculatus

ABSTRACT.

• 1 Previous work has shown that female Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) can accurately assess the number of eggs on a seed and use this information to produce a near uniform distribution of egg-loads, so minimizing larval competition within hosts.
• 2 The decision to lay on a particular seed may be based on one of two possible rules: (i) a‘relative’rule, which compares the number of eggs on the present seed with that on previously encountered seeds; and (ii) an‘absolute’rule, which takes into account the current egg-load alone.
• 3 This paper describes an experimental study of oviposition behaviour which concludes that an absolute measure of egg-load probably determines egg-laying decisions. The implications of this result are discussed.

     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

The toxicity of Jack bean (Canavalia ensiformis) cotyledon and seed coat proteins to the cowpea weevil (Callosobruchus maculatus)

The seeds of the Jack bean, Canavalia ensiformis (L) DC are known to contain several toxic substances that prevent their utilisation as food for humans and animals. The lectin concanavalin A and the enzyme urease are the best known of these proteins. We have found that many proteins present in the seeds of the Jack bean, like trypsin inhibitors and canatoxin, are detrimental to the development of the bruchid insect Callosobruchus maculatus (F) (Coleoptera: Bruchidae). Among these proteins, canavalin (vicilin, 7S globulin) was found to be expressed in the seed coat. We suggest that seed coat canavalin, in addition to other detrimental proteins expressed in this tissue, may have been of importance in the evolutionary discrimination of the seeds of this legume by non-pest bruchids.     

Vibrations mediate prudent resource exploitation by competing larvae of the bruchid bean weevil Callosobruchus maculatus

Larvae of the bean weevil C. maculatus feeding in a single bean apparently respond to vibrations from each other's chewing: one larva feeds normally, the other is inhibited. If the burrows of the larvae intersect, the inhibited larva dies. If the dominant larva pupates or dies without the burrows intersecting, the inhibited larvae then feeds and matures if enough food remains. Since females add second eggs to the largest beans after most beans carry a single egg, competition is most common in the largest available beans, precisely where inhibited larvae can benefit from avoiding a contest.

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Résumé Lors de la compétition, les larves réagissent l'une par rapport à l'autre, même lorsque leurs galeries sont séparées. L'une est inhibée, tandis que l'autre se comporte comme si elle était seule. Le signal provoquant ces réactions, qui peut traverser du papier métallique, est dû vraisemblablement aux vibrations de mastication des larves; ces dernières perçoivent ces signaux tout au long de leur vie active. La larve la plus âgée ne l'emporte pas obligatoirement, ce qui signifie que le signal indique plus que l'âge.Les conditions expérimentales peuvent inverser les réponses des unes et des autres, bien que la larve la plus âgée puisse souvent mourir de ces conditions anormales de compétition. L'avantage potentiel associé à l'inhibition du futur perdant est mis en évidence en collant ensemble des graines. Quand la larve dominante se nymphose, la larve inhibée recommence à s'alimenter et achève son développement. En ayant eu son alimentation interrompue et en restant dans une galerie superficielle, la larve inhibée évite les affrontements et peut se développer lorsque la larve dominante meurt ou se nymphose, puisque la galerie de la larve inhibée n'avait pas été sectionnée.La stratégie de ponte réduit la fréquence de la compétition en uniformisant la distribution des oeufs; elle augmente les chances de survie des larves vaincues avec l'addition des oeufs sur les graines les plus grosses quand les disponibilités sont limitées. L'étude de l'évolution de ces adaptations a été possible puisque, comme on le sait, C. maculatus est génétiquement polymorphe pour les caractères déterminant le comportement de ponte des femelles et la compétition larvaire.

     

Deterrent activity of plant lectins on cowpea weevil Callosobruchus maculatus (F.) oviposition

A set of 14 plant lectins was screened in a binary choice bioassay for inhibitory activity on cowpea weevil Callosobruchus maculatus (F.) oviposition. Coating of chickpea seeds (Cicer arietinum L.) with a 0.05% (w/v) solution of plant lectins caused a significant reduction in egg laying. Control experiments with heat inactivated lectin and BSA indicated that the observed deterrent effects are specific and require carbohydrate-binding activity. However, no clear correlation could be established between deterrent activity and sugar-binding specificity/molecular structure of the lectins. Increasing the insect density reduced the inhibitory effect of the lectins confirming that female insects are capable of adjusting their oviposition rates as a function of host availability.     

The fecundity and oviposition behavior of a South Indian strain of Callosobruchus maculatus

The fecundity of the South India strain of Callosobruchus maculatus (Fab.) (Bruchidae: Coleoptera) is 73 eggs for females developing in 31 days and increases to 94 eggs for females emerging after 41 days. Fed females lay 45–55 more eggs which is the egg-equivalent of the reserves unfed females had to sacrifice for activity. Competition as a larva did not reduce a female's fecundity. Oviposition is inhibited when females are given 25 or fewer beans for oviposition and it is reversed if females find fresh beans. Oviposition is generally released by beans carrying a below average number of eggs, thus, the eggs are almost uniformly distributed over the beans. Eggs are preferentially placed on the largest egg-free beans as expected if oviposition is released when a beetle goes from a smaller to a larger bean. The cues resulating host preferences, cowpea > mung pigeonpea chickpea, are secondary to the cues producing hyperdispersion and the selection of larger beans. We present a model for the oviposition behavior that accounts for the observed responses to species, size, and egg load of oviposition sites.

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Résumé La fécondité de femelles isolées avec un mâle, sans aliment ni eau, et laissées sans perturbation avec 50 graines au moins sert de fécondité standard. Elle augmente avec le temps de développement de l'uf à l'adulte de 73 ufs (32 j) à 94 ufs (44 j). La fécondité potentielle, obtenue avec des femelles alimentées sur sucre est de 125 ufs. Des femelles maintenues avec moins de 25 graines réduisent leur fécondité.Plus de la moitié des femelles distribuent leurs ufs uniformément jusqu'à une densité de 1,5 uf. L'uniformité peut provenir de l'induction de réactions de ponte par les graines portant moins d'ufs que l'effectif moyen. L'hyperdistribution au dessus des densités de 3 ufs/graine révèle l'aptitude à distinguer les graines avec 3 ufs des graines avec 4 ufs. Il y a moins d'écarts par rapport à l'uniformité (erreurs) avec des graines à surface rugueuse. Des femelles avec 10 graines font moins d'erreurs que des femelles avec 50 ou 100 graines.Les ufs sont ajoutés aux plus grosses graines sans ufs; ainsi, plus il y a de graines qui portent des ufs, plus le poids des graines avec ufs diminue. Un tel schéma est prévisible si la ponte est induite par des graines plus grosses que les graines rencontrées précédemment. Les femelles présentent un ordre de préférence clair quand elles sont en rapport avec leurs hôtes classiques: Vigna unguiculata > V. radiata > Cajanus cajan > Cicer arietinum Les caractères spécifiques de l'hôte, plutôt que la taille ou la texture de la surface servent à la distinction entre les graines d'un même hôte.Ces résultats expérimentaux peuvent être utilisés pour la modélisation du comportement de ponte d'une femelle.

     

Effects of search experience in a resource-heterogeneous environment on the oviposition decisions of the seed beetle, Callosobruchus maculatus (F.)

Abstract.  1. This study investigates how female seed beetles, Callosobruchus maculatus , distribute their eggs on various-sized seeds when the size of seed was varied during the egg-laying period.
2. Beetles were allowed to lay eggs on one of three arrays of 64 adzuki beans ( Vigna angularis ). Each array contained four size classes of seed, ranging from small (5.0–5.5 mm diameter) to large (6.5–7.0 mm), but differed in how they were distributed within the environment. In the most heterogeneous condition (the 64-patch design), the four sizes were interspersed, while in the least heterogeneous condition (the four-patch design) they were grouped into four separate blocks. Thus, a beetle exploring the 64-patch design would frequently encounter all four seed sizes, whereas a beetle exploring the four-patch design would only rarely encounter a change in bean size.
3. Beetles experiencing greater seed size heterogeneity were more likely to lay eggs on larger seeds, whereas those in the blocked condition were more likely to oviposit on small seeds. Beetle responses to seed size heterogeneity suggest that the degree of preference for large seeds depends on a female's recent experience.
4. Female beetles exhibited size discrimination throughout their egg-laying process; however, there was a trade-off between seed size and egg discrimination (i.e. avoiding those seeds already containing developing eggs) in response to the change in fitness gained from either laying on larger seeds or lower egg-load seeds during the egg-laying process.
5. Our model provides the first evidence that evolving seed size discrimination ability is adaptive for the seed beetle with egg-discrimination ability.     

Functional anatomy of the egg pore in Callosobruchus maculatus: a trade-off between gas-exchange and protective functions?

Abstract. The chorion of the egg in Callosobruchus maculatus (Coleoptera: Bruchidae) provides protection but also has to allow exchange of respiratory gases. A single opening in the chorion (the egg pore) allows diffusion of gases, and young larvae die if the egg pore is blocked. Comparison of respiration rates using a Gilson respirometer showed an increase in oxygen uptake from the first to the seventh day after the egg was laid; the increase was from 6.8 to 11.2 pl/egg/day in a Brazil strain and from 8.6 to 23.5 pl/egg/day in a Yemen strain. The Yemen strain had double the metabolic rate of Brazil-strain insects in the larvae, which are exposed to seed toxins, but not in the non-feeding adults. It is suggested that the higher metabolic rate in the Yemen larvae is related to their ability to develop in seeds that are toxic to the Brazil strain. The egg pore also differed in size and shape between the two strains. The egg pore was cylindrical in the Brazil strain, but shorter and funnel-shaped with an increased external diameter in the Yemen strain. Calculation showed that the different shape and size of the Yemen egg pore would allow a 2–3-fold increase in gas conductance compared with the Brazil strain, and this increase was consistent with the enhanced oxygen requirement of the Yemen larvae, which was a little over twice that of the Brazil larvae at 7 days. The anatomy of egg pores in Cmaculatus thus seems to represent a trade-off between two functions, allowing adequate gas exchange while maintaining the protective function of the chorion.     

Realized heritability of behavioral responsiveness to an oviposition deterring pheromone in the azuki bean weevil Callosobruchus chinensis

Artificial selection was conducted to reduce the behavioral responsiveness of female bruchid beetles, Callosobruchus chinensis, to the oviposition deterring pheromone excreted by conspecific females. Significant responses to selection were observed after two generations of selection. Realized heritability was estimated as 0.052 ± 0.017 from cumulative responses regressed on cumulative selection differentials. These results indicate that this pheromone communication system has significant additive genetic variance needed for its evolution.     

Pheromone-like substances affecting host-related behaviour of larvae and adults in the dry bean weevil, Acanthoscelides obtectus

Males and females of the dry bean weevil, Acanthoscelides obtectus deposit chemically unknown substance(s) on beans during various activities (most probably during defecation), under laboratory conditions. The marking results in avoidance of these beans by other females for oviposition. Also, first-instar larvae are influenced to disperse more by the deposited substance(s), so that they are more likely to find unmarked beans to enter. No specific marking behaviour by adults was noted. That the deterring substance(s) might originate from microorganismic activity after deposition has occurred, is improbable. Possible explanations for sources of marking substance(s) and the significance of the corresponding behaviour in this species are discussed.

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Résumé Les mâles et les femelles d'A. obtectus déposent des substance(s) encore non-identifiées sur les grains du haricot au cours de leurs diverses activités. Ce comportement de marquage fait que les autres femelles hésitent à pondre des ufs sur ces grains. De plus, les substance(s) déposée(s) poussent les larves du 1^er stade à se disperser plus vigoureusement, par conséquent, elles sont enclines positivement à entrer dans les grains sans marque. On n'a observé aucun comportement marqueur spécifique chez les adultes. La possibilité d'une origine microbiologique des substance(s) répulsive(s) a été eliminée. Les origines possibles des substance(s) de marquage et la signification de ce comportement sont discutés.

     

Behavioral processes and egg-laying decisions of the bean weevil, Callosobruchus maculatus

Empirical studies of behavioral processes were designed in a seed beetle, Callosobruchus maculatus, to distinguish between two alternative tactical models, namely, comparison tactics and threshold tactics of seed choice, and also to deduce the behavioral rule of its oviposition decisions. Search sequences and the oviposition process of the female bruchid beetle were observed and recorded. Analyzing data of the search sequence showed that the female bruchid beetle repeatedly encountered seeds randomly and tended to oviposit on a newly encountered seed in an ovposition bout. These results contradict the usual comparison models, which predict that the female will return to choose any one of the previously encountered seeds after sampling. In addition, the oviposition decision (rejection or acceptance of an encountered seed to oviposit) was analyzed with a generalized linear modeling (GLIM) technique. Modeling showed that the probability of accepting a seed with different numbers of eggs on it changed during her egg-laying process. This evidence supports the idea that the female is using a threshold tactic and that her acceptance threshold is being adjusted by experience gained during the egg-laying process. The analysis of statistical modeling also showed that both the time since the last oviposition and the number of eggs which had been laid by the female had a significant effect on the probability of accepting seeds with different numbers of eggs. Therefore, a time measuring system and the physiological state variable, e.g., eggload of the female, should be included in the behavioral rule to explore how the female makes her decision in the egg-laying process and to examine the importance of choice behavior as a component of selection and adaptation. Received: May 8, 1998 / Accepted: September 20, 1999     

Egg dumping and life history strategy of Callosobruchus maculatus

Abstract.  Previous studies have demonstrated that female Callosobruchus maculatus adjust oviposition rates to cope with changes in host availability. A female lays fewer eggs when host availability is low, and hence decreases larval competition. However, females will also dump eggs on unsuitable substrates under conditions of host deprivation. Because the female does not feed as an adult, egg dumping possibly wastes energy and may thus be a maladaptive behaviour. In this study, the effect of mating and age on the egg-dumping behaviour and the life history strategy of the female are explored. Under host-deprived conditions, mating is seen to trigger egg-dumping behaviour. Also, females mated at 6 days dump significantly fewer eggs and live longer than females mated at 0 or 3 days. Thus, a trade-off between fecundity and longevity is seen among females subjected to different manipulations. In addition, 6-day-old virgin females contain more mature eggs than females mated at 6 days can produce when deprived of hosts. This finding indicates that the female reallocates internal energy resources by oosorption in a resource-limited environment. To test the maladaptive hypothesis, mated females were deprived of a host for 6 days and then given sufficient hosts each day. The results show that the more eggs dumped by a female in the first 6 days, the more eggs are laid later on beans. Egg dumping is thus not maladaptive.     

Response of Callosobruchus maculatus (F.) to varying temperature and relative humidity under laboratory conditions

The effect of temperature and relative humidity were determined on the development of Callosobruchus maculatus (F.) on stored bean (Vigna unguiculata) seeds exposed to five temperatures (20, 25, 30, 35 and 40 °C) and six relative humidity levels (40, 50, 60, 70, 80 and 90%). Oviposition and total adult progeny responded in a curvilinear pattern to temperature and relative humidity while developmental period presented a linear response. Egg laying (117.33 ± 3.21) and adult emergence (35.00 ± 1.70) were least at 20 °C and 90% R.H. but those showed the optimal values at 30 °C and 70% R.H. The implication of these findings is that the growth attributes of C. maculatus relate with the weather variables studied in a curvilinear manner and thus quadratic equations generated could be used for the prediction of optimum temperature and relative humidity in a given area for the management of C. maculatus in stored beans.     

Intra-specific variation in female remating in Callosobruchus chinensis and C. maculatus

The effects of mating duration on female remating (exp. 1) and under different male densities (exp. 2) were examined in two strains of the adzuki bean beetle, Callosobruchus chinensis and in one strain of the bruchid beetle, C. maculatus. In experiment 1, the frequency of female remating was markedly different between the two strains of C. chinensis. Females of the jC strain, reared long-term in the laboratory, did not remate after being allowed to mate freely (=monogamy), whereas females of the isC strain, recently established from the field, showed high remating frequencies (=polyandry). In both strains, the frequency of female remating increased after the duration of the first mating was deliberately shortened. The relation between mating duration and remating frequency was significantly different, however, between the two strains. In a closely related species, C. maculatus, which manifests polyandry, this relation was more similar to that of the field-derived (=isC) than to that of the laboratory-derived (=jC) strain of C. chinensis. The reasons for the inter-strain variation observed in the remating frequencies of C. chinensis are also discussed. In experiment 2, the mating duration of the three strains was compared under different male densities. Only the lab-derived strain demonstrated a significantly shorter mating duration when one female was placed together with five males than when paired with one male. The shorter mating duration (approximately 26 s) was similar to that of females allowed to remate in the monogamous strain in experiment 1.     

Modelling and analyzing density-dependent population processes: Comparison between wild and laboratory strains of the bean weevil,Callosobruchus chinensis (L.)

Three models were constructed for analyzing the population characteristics ofC. chinensis on stored beans; model A describing the whole reproductive process with a single equation, model B describing the three age-specific processes (oviposition, egg survival and larval survival) with separate equations, and model C which describes all these processes not for the whole habitat but for the individual beans comprizing it. The logit equation was employed here as a common basis to describe the density-response relationship involved. All three models showed very good fit to the experimental data obtained for both laboratory and wild strains of the weevil. The parameter values characterizing the population dynamics were, however, widely different between the two strains; the laboratory one which had been reared for some 500 generations showed significantly higher reproductive capacity, less sensitive and gentler response to crowding in both adult and egg stages, and more uniform egg distribution among individual beans, as compared with the wild strain newly introduced. Sensitivity analyses using these models suggested that these changes in population characteristics have been attained by the process of domestication or adaptation to stable laboratory conditions through a long period of time. This process seemed in effect to have optimized the population's performances in the laboratory environment. Evolutionary significance of such optimization was discussed with reference to the selection pressure which may have acted upon individuals.     

Dynamic egg maturation strategies in an aphid parasitoid

Abstract.  Females of the parasitoid Aphelinus albipodus both lay more eggs on younger stages of their aphid host, Aphis glycines , and contain more mature eggs in their ovaries when held with these younger host stages. This suggests that egg maturation is more rapid in the presence of younger host stages. Three factors explaining the difference in the egg loads of A. albipodus females exposed to various host stages are examined. These are (i) the number of host-feeding meals; (ii) the number of eggs laid; and (iii) the host stage utilized. Together, these factors explain 69% of the variance in A. albipodus egg load. The number of host-feeding meals taken by females is a strong predictor of egg load. More host meals are taken on young hosts, suggesting that host feeding contributes to the trend of faster egg maturation in the presence of younger hosts. Host stage has a strong impact on egg load even when the effect of host feeding is accounted for. There is no evidence for an effect of the oviposition rate on egg load. The results indicate that egg maturation by A. albipodus is dynamic; females mature eggs faster when in the presence of preferred hosts. It is hypothesized that this allows A. albipodus females to more closely match their reproductive effort to reproductive opportunities.     

Systems analysis of density-dependent population processes in the azuki bean weevil,Callosobruchus chinensis

Systems analysis of density-dependent population processes was conducted in an experimental population of the azuki bean weevil,Callosobruchus chinensis. Density-dependent population change was formulated for two separate processes; adult production of eggs and hatching of the eggs outside the beans, and survival of the first instar larvae inside the beans and their emergence as new adults. The formulation of each process was based on a common basic equation modified from the logistic-difference equation. Parametric values were estimated from experimental data onC. chinensis in a laboratory population. Simulations of the model showed good fits not only to observed population changes both outside and inside the beans, but also to the observed reproduction curve reflecting the population change as a whole within one generation in an experimental population. Furthermore, population dynamics from generation to generation were simulated from the model reproduction curve. The simulation showed a slight dumped oscillation and converged to a carrying capacity, which fitted well with the experimental population dynamics.