Male-killing virus in a noctuid moth Spodoptera litura.

A female-biased sex ratio is considered advantageous for the cytoplasmic elements that inhabit sexually reproducing organisms. There are numerous examples of bacterial symbionts in the arthropod cytoplasm that bias the host sex ratio toward females through various means, including feminization and male killing. Recently, maternally inherited RNA viruses belonging to the family Partitiviridae were found to cause male killing in moths and flies, but it was unknown whether male-killing viruses were restricted to Partitiviridae or could be found in other taxa. Here, we provide compelling evidence that a maternally inherited RNA virus, Spodoptera litura male-killing virus (SlMKV), selectively kills male embryos of the tobacco caterpillar Spodoptera litura, resulting in all-female broods. SlMKV injected into uninfected S. litura can also be inherited maternally and causes male killing. SlMKV has five genomic segments encoding seven open reading frames, has no homolog of known male-killing genes, and belongs to an unclassified group of arthropod-specific viruses closely related to Tolivirales. When transinfected into larvae, both male and female recipients allow SlMKV to proliferate, but only males die at the pupal stage. The viral RNA levels in embryonic and pupal male killing suggest that the mechanism of male killing involves the constitutive expression of viral products that are specifically lethal to males, rather than the male-specific expression of viral products. Our results, together with recent findings on male-killing partiti-like viruses, suggest that diverse viruses in arthropods tend to acquire male killing independently and that such viruses may be important components of intragenomic conflict in arthropods.

Geographic Variation in Photoperiodic Response for Induction of Pseudopupal Diapause in Epicauta gorhami (Coleoptera: Meloidae).

Geographical variations in the threshold of environmental cues for diapause induction are important in understanding the life history strategy of insects. Larvae of the bean blister beetle, Epicauta gorhami (Coleoptera: Meloidae), feed on grasshopper eggs and undergo hypermetamorphosis. They normally enter diapause as a pseudopupa (fifth instar). However, when the larvae are reared under long-day and high-temperature conditions, they do not enter pseudopupal diapause but pupate directly from the fourth instar. In addition, this insect is known to modify its photoperiodic response for induction of pseudopupal diapause depending on food availability for larvae. In this study, the larval photoperiodic responses for diapause induction were examined for three populations of E. gorhami collected from the northernmost (Morioka, 39.7°N), central (Kazo, 36.1°N), and southernmost (Takanabe, 32.1°N) parts of its distribution range, and the responses were examined under both fully fed and food-deprived (FD) conditions. Diapause incidence decreased in FD larvae for all populations in long-day conditions, and the critical day-lengths for the diapause incidence were calculated as 14.81, 13.97, and 13.99 h in the Morioka, Kazo, and Takanabe populations, respectively. The results indicate the presence of a geographical variation in larval photoperiodic response, in which diapause is induced earlier in higher latitudinal areas. From these results, together with data for developmental threshold temperature and thermal constant, the Morioka and Kazo populations were suggested to have a partial bivoltine life cycle under the conditions of extremely early oviposition season, a sufficient summer temperature, and poor larval food availability.

Adaptive significance of precocious pupation in the bean blister beetle, Epicauta gorhami (Coleoptera: Meloidae), a hypermetamorphic insect.

Larvae of the bean blister beetle, Epicauta gorhami Marseul (Coleoptera: Meloidae), feed on grasshopper eggs in soil and undergo hypermetamorphosis. They normally enter diapause as a pseudopupa at the fifth instar, a form characteristic of hypermetamorphosis for meloid beetles. However, fourth-instar larvae exposed to long days and high temperature avoid pseudopupal diapause and pupate directly from the fourth instar. Fourth-instar larvae also tend to pupate precociously with a smaller body size if they are deprived of food. In these larvae, the critical day-length controlling induction of pseudopupal diapause becomes shorter than that for fully-fed larvae. In this study, we examined how the reaction norm of food-deprived E. gorhami larvae functions in nature by rearing insects from the egg stage outdoors in different seasons with manipulation of the food supply. The results indicated that most fully-fed larvae entered pseudopupal diapause, whereas food-deprived larvae tended to pupate precociously without entering diapause, especially early in the season. The resulting smaller adults reproduced early in the autumn and their progeny attained the pseudopupal stage before winter, indicating that the reaction norm may have an adaptive role in controlling seasonal development in the face of food shortages, producing a bivoltine life cycle.

Effects of temperature and photoperiod on termination of pseudopupal diapause in the bean blister beetle, Epicauta gorhami.

Larvae of the bean blister beetle, Epicauta gorhami (Coleoptera: Meloidae), feed on grasshopper eggs in soil and undergo hypermetamorphosis. This beetle undergoes larval diapause in the fifth instar as a pseudopupa, a form characteristic of hypermetamorphosis in meloid beetles. This pseudopupal diapause is maintained for a long period if the larvae are kept under continuous diapause-inducing short-day conditions. In the present study, the effects of temperature and photoperiod on termination of pseudopupal diapause in E. gorhami were examined using pseudopupae obtained under 12L-12D at 25°C. Diapause was terminated by exposure to low or high temperature, but not by transfer to long-day conditions. The pseudopupal stage comprises diapause and post-diapause phases despite its unchanging appearance. The optimum low temperature for diapause termination was 10-15°C, and a higher or lower temperature was less effective. There was an individual variation in the low temperature requirement for diapause termination and post-diapause development may not occur until this requirement is satisfied. Although under natural conditions pseudopupae may encounter low temperatures that are effective for diapause termination at different ages, our results show that pseudopupae at various ages respond similarly to low temperature exposure. This response may ensure that resumption of development synchronizes in a population.