Sergey Ya. Reznik

Multigenerational maternal effect on diapause induction in Trichogramma species (Hymenoptera: Trichogrammatidae)

Abstract Maternal photoperiodic response is known to influence the percentage of diapausing prepupae in Trichogramma species. However, the influence of several preceding generations has not yet been studied. We have investigated the stability of photoperiod-induced changes in multiple generations of Trichogramma buesi Voegele and Trichogramma principium Sug. et Sor. Short-day conditions during preimaginal development induced an increase in the percentage of diapausing progeny and grand progeny of both Trichogramma species. A similar trend was also detected in the fourth and fifth generations, but the response was weak although statistically significant. This grand-grandmaternal photoperiodic effect (which has not been demonstrated before for Trichogramma or for any other insect parasitoid) is most probably based on the transgenerational transmission of variations in DNA expression. We conclude that in mass rearing, to facilitate diapause induction before cold storage, it is advisable to rear both maternal and grandmaternal generations under the short-day conditions. In scientific studies, several generations preceding the experiment should be kept under equal conditions to exclude multigenerational maternal effects.

Herbivore Impact Versus Host Size Preference: Endophagous Insects on Heracleum mantegazzianum in Its Native Range

Abstract Classical biological control is a practice to control alien invasive weeds, but many introduced biological control agents exhibit only a weak negative impact on their targets. One reason is that prerelease impact studies in the natural environment are often difficult to carry out. Heracleum mantegazzianum Sommier and Levier (Apiaceae), which is native to the Caucasus, is a perennial noxious weed introduced into Europe and North America. We examined the impact and host size preference of different endophagous insect guilds in the weed’s native range. Instead of the commonly used insect exclosure approach, we estimated plant vigor before and after herbivore attack under natural conditions. Endophagous herbivores were dominated by the weevil species Lixus iridis Olivier, Nastus fausti Reitter, and Otiorhynchus tatarchani Reitter (Coleoptera: Curculionidae), the fly Melanagromyza heracleana Zlobin (Diptera: Agromyzidae), and an unidentified root-boring agromyzid fly species. Most observed insect species exhibited a strong preference for either big or small plants, but none of them caused serious damage within the study period. Occurrence of root-feeding weevils was associated with weak plants, but because of their long larval development, it was not possible to assign this relationship clearly to either feeding damage or host size preference. A comparison with other studies indicated that mature H. mantegazzianum plants are quite tolerant to herbivory. Insects belonging to the feeding guilds studied here will probably not guarantee successful biological control. Further research should focus on earlier stages in the weed’s life cycle.

Maternal thermal and photoperiodic effects on the progeny diapause in Trichogramma telengai Sorokina (Hymenoptera: Trichogrammatidae)

Although maternal photoperiodic and maternal thermal effects on the progeny diapause have been demonstrated in a number of insect species, their interaction has been rarely studied. We investigated this interaction in Trichogramma telengai. In a series of experiments, maternal females were reared at day lengths of 12–18 h and at temperatures of 17, 20, 25 and 30°C. Their progeny developed under day length of 12 h and temperatures of 13, 14 and 15°C. The experiments showed that both short day and low temperature experienced by the maternal generation significantly increased the proportion of diapausing progeny. In particular, the threshold of the maternal photoperiodic response decreased with temperature. Under combinations of photoperiod with daily thermoperiod, the role of the “night” temperature in the induction of diapause in the progeny was much more important than that of the “day” temperature. We conclude that the interaction pattern between the photoperiodic and thermal maternal effects in T. telengai is generally the same as that between the photoperiodic and thermal responses directly influencing diapause induction in other long‐day insects. The threshold temperature of the maternal thermal response of T. telengai was about 25–27°C, while diapause can be induced if larvae develop at temperatures not higher than 15–16°C. This suggests that, at least in the studied Trichogramma species, the maternal thermal effect has no ecological value. In the practice of biocontrol, however, rearing of Trichogramma wasps at high temperature can drastically reduce the proportion of diapausing progeny.

Maternal thermal effect on diapause in Trichogramma species (Hymenoptera: Trichogrammatidae)

Both direct thermal and maternal photoperiodic effects on diapause induction have been thoroughly investigated in many insect species, while maternal thermal effects have been infrequently studied. We studied the effect of temperature during development of maternal generation on the proportion of diapausing progeny in four species of the genus Trichogramma Westw., minute egg parasitoids, widely used for biological control of lepidopteran pests. The maternal generations were reared at day lengths of 12 and 18 h and temperatures of 17, 20, 25 and 30°C, and their progeny developed under day length of 12 h and temperatures of 13 and 14°C. In T. evanescens and T. piceum, the proportion of diapausing progeny decreased with increasing temperature under all tested photoperiods and thermal regimes of progeny development; the high temperature of 30°C totally averted diapause of progeny. In T. buesi and T. principium, low temperatures of 17 and 20°C resulted in relatively high proportion of diapausing progeny only when the maternal generation developed under short‐day conditions. The threshold of the maternal thermal response varied from 17–18 to 22–23°C. Under field conditions, Trichogramma females are exposed to such high temperatures only during summer, when diapause in their progeny is in any case prevented by the maternal photoperiodic response and by the thermal response of the larvae. We conclude that the maternal thermal effect on diapause induction, although to a different extent, is inherent to Trichogramma species but, at least as suggested by laboratory experiments, it does not play any role in the regulation of seasonal development under natural conditions. However, during mass rearing of Trichogramma wasps, it should be taken into account that high temperature, even when combined with short photoperiod, can avert diapause in the next generation.

Costs and benefits of interlarval cannibalism in Harmonia axyridis

Cannibalism, which is rather common in ladybirds, has been usually studied at the individual level: benefits of cannibalism for cannibals were estimated. Our study was conducted at the group level: we evaluated the overall effect of interlarval cannibalism on a group of Harmonia axyridis larvae of the fourth instar deprived of food, including both cannibals and their victims. Experiments showed that the probability of pupation in larvae which were kept individually was significantly higher than in larvae kept in groups of five, other conditions being the same. The proportion of samples in which at least one of five individuals pupated among the larvae kept individually was also higher than among those kept in groups suggesting that the eventual benefit of cannibalism was outweighed by the negative impact of aggressive interlarval interactions. The mean and minimum survival time in samples where none of five larvae pupated were longer when larvae were kept individually than when larvae were kept in groups. However, the maximum survival time (the survival time of the last larva in a sample) increased when larvae were kept in groups, which was the only one benefit of cannibalism found in our study. Under natural conditions, the possible adaptive value of this effect is that in the absence of natural prey, longer larval survival time proportionally increases the possibility of finding a new prey patch thereby ensuring survival of the population.

On the factors inducing the inhibition of diapause in the progeny of diapause females of Trichogramma telengai

The ability to enter diapause in the progeny of Trichogramma Westw. (Hymenoptera: Trichogrammatidae) females that have undergone diapause is markedly reduced or even completely absent. This maternal inhibition of diapause can result from three types of factors: (i) environmental factors that induce maternal diapause; (ii) maternal diapause itself; and (iii) diapause‐terminating environmental factors. The present study aims to determine the factors that prevent diapause in the progeny of a Trichogramma telengai Sor. generation that has undergone diapause. Different tendencies to enter diapause in the maternal generation are induced by different photoperiods (LD 12 : 12 h, 16 : 8 h and 18 : 6 h) during development of the grandmaternal generation and different temperatures (from 10 to 15 °C) during larval development of the maternal generation. To terminate diapause, prepupae of the maternal generation are kept at 4–5 °C in the dark. To estimate the ability to enter diapause, the progeny generation is incubated at 14 °C. The results suggest that the inhibition of the tendency to enter diapause in the progeny of T. telengai females that have undergone diapause is caused by diapause itself rather than by diapause‐inducing or diapause‐terminating environmental factors. This result can be used to clarify the mechanism of the inhibition of diapause in the progeny of females that have undergone diapause.

Diapause induction in Trichogramma telengai: the dynamics of maternal thermosensitivity

Transferring Trichogramma telengai Sorokina (Hymenoptera: Trichogrammatidae) from 17 to 30 °C or vice versa at different stages of pre‐imaginal development shows that the pupa (particularly late pupa) is a sensitive stage of the maternal thermal response. During this period, even a relatively short period (1–2 days) of cold or heat exposure results, correspondingly, in a marked increase or decrease in the percentage of diapausing progeny. Thus, the maternal thermal response has the same features of sensitivity as those of the previously investigated maternal photoperiodic response, suggesting that photoperiodic and thermal responses share the same mechanism. For the practice of rearing and storage Trichogramma as a biocontrol agent, it is important to note that even relatively short‐term environmental heating (i.e. for one to several days) immediately before adult emergence can drastically decrease the percentage of diapausing progeny.