Identification of biogenic amines involved in photoperiod-dependent caste-fate determination during the adult stage in a temperate paper wasp.

In the temperate paper wasp Polistes jokahamae, caste is influenced by photoperiod during the adult stage, but the mechanisms underlying the caste-fate determination system have been unclear. We measured the brain levels of monoamines and related substances in females kept isolated for two weeks under different photoperiods. Except for in the first-emerging group, the females developed ovaries under long-day conditions, whereas they stored lipids under short-day conditions. The levels of tyramine in the brain were significantly higher under long-day than under short-day conditions and positively correlated with maximum oocyte lengths. These results suggest that tyramine was produced in response to long daylength during the adult stage and associated with ovarian development, which is the principal characteristic of reproductive workers. There was also a significant positive correlation between dopamine levels in the brain and maximum oocyte length, independent of photoperiod, suggesting that dopamine is involved in reproductive function with tyramine resulting in the induction of reproductive workers. Meanwhile, higher levels of tryptophan in the brain were found in short-day conditions and positively correlated with lipid stores. However, serotonin synthesized from tryptophan and N-acetylserotonin were not associated with lipid stores without photoperiodic responses, suggesting that tryptophan is involved in the physiological changes toward gyne under short daylength, independently of serotonin signaling. In conclusion, tyramine and tryptophan are candidates for mediating photoperiod-dependent caste-fate determination in P. jokahamae: the former is involved in generating the worker caste while the latter is involved in generating the gyne caste.

Large mothers produce progeny with high survival rates during the immature stage and large sizes at adulthood in a parasitoid species.

Parasitoid researchers have generally thought that the body size of the mother parasitoid does not affect the fitness performance of the progeny during the immature stage, as long as the progeny develop in the same environment. We reveal for the first time that this is not true for the parasitoid Echthrodelphax fairchildii (Hymenoptera: Dryinidae), which is parasitic on planthoppers. Large females ensured an increased survival rate for their progeny during the immature stage and a large body size at adult emergence. Maternal body size differentially affected the body sizes and survival rates of male and female progeny. Small females did not produce female progeny, and the survival rate of the female progeny increased more steeply with increasing maternal body size than that of the male progeny. Meanwhile, the body size of male progeny increased more steeply with increasing maternal body size. The influence of maternal body size on progeny survival to adult emergence has never been reported in insects before. In addition, large females were more likely to lay female eggs, suggesting that females control the sex ratio of progeny in response to their own body size.

Caste-fate determination primarily occurs after adult emergence in a primitively eusocial paper wasp: significance of the photoperiod during the adult stage.

Independent-founding paper wasps constitute a major group of primitively eusocial insects, and when caste-fate determination occurs in temperate species of these wasps, particularly regarding whether it occurs before or after emergence, remains unclear. No critical morphological differences occur between potential queens of the next generation (often called gynes) and workers in primitively eusocial insects. The gynes of temperate species are characterized by diapausing, and the nutrients available during the larval stage have often been believed to determine caste fate. Short days usually induce diapause in temperate nonsocial insects, although few investigations of the effects of day length on caste-fate determination in paper wasps have been conducted. By exposing individuals to different combinations of short and long days during the immature and adult stages, we show for the first time that short days during the adult stage (but not during the immature stage) facilitated caste-fate determination toward gynes in a paper wasp. Moreover, the decision to diapause partly depended on changes in the photoperiod during the pupal and adult stages. The size of the adult also affected caste-fate determination, with diapause more likely to occur in large adults, but this size effect did not occur when individuals were exposed to many short days during the pupal stage. In addition, all adults except for a small proportion of smaller individuals prepared for diapause under short days. These findings suggest that the photoperiod is a higher priority cue than adult size.

Self-/conspecific discrimination and superparasitism strategy in the ovicidal parasitoid Echthrodelphax fairchildii (Hymenoptera: Dryinidae).

Superparasitism in solitary parasitoids results in fatal competition between the immature parasitoids, and consequently only one individual can emerge. In the semisolitary ovicidal parasitoid Echthrodelphax fairchildii (Hymenoptera: Dryinidae), 2 adults can emerge under superparasitism with a short interval (<24 h) between the first and second ovipositions. We determined the female parasitoid's behavioral responses under self- and conspecific superparasitism bouts with first-to-second oviposition intervals of ≤2 h. The self- and conspecific superparasitizing frequencies increased up to an oviposition interval of 0.75 h, with the former remaining lower than the latter, particularly for oviposition intervals of ≤0.25 h, suggesting the existence of self-/conspecific discrimination. The superparasitizing frequency plateaued for oviposition intervals of ≥0.75 h, with no difference between self- and conspecific superparasitism. The ovicidal-probing frequency did not differ under self- and conspecific superparasitism, and was usually <20%. The females exhibited no preference for the oviposition side (i.e., ovipositing on the side with or without the first progeny) and almost always laid female eggs for any oviposition interval under self- and conspecific superparasitism. The sex ratio was not affected by the type of superparasitism, oviposition sides, or the occurrence of ovicidal probing. These observed results about the oviposition side, ovicidal probing, and sex ratios differed from the predictions obtained assuming that the females behave optimally. Possible reasons for the discrepancies are discussed: likely candidates include the high cost of selecting oviposition sides and ovicidal probing, and, for the sex ratio, the low frequency of encountering suitable hosts before superparasitism bouts.