Sonny B. Ramaswamy

Host finding by moths: sensory modalities and behaviours.

Abstract Despite their economic importance, moths have not been the subject of intense studies on host finding as have been their larvae or adults of other groups of insects. This paper details the host-finding behaviour of Heliothis virescens and the kinds of external sensory modalities involved in this process compared with those in other moths. Based on a review of the literature, models for strategies of host finding are discussed, along with the importance of experience and metabolic states on oviposition by moths.

Dynamics of juvenile hormone‐mediated gonadotropism in the lepidoptera

Reproduction in moths and butterflies is a dynamic process that is influenced by various endogenous physiological processes and exogenous factors. The Lepidoptera may be divided into four distinct groups based on their gonadotropic hormones and other reproductive and biological characteristics, regardless of phylogenetic relationships. Some species have integrated their reproduction tightly with the endocrine events of metamorphosis, the waxing or waning ecdysteroid levels. Other species rely instead on juvenile hormone (JH), in part or solely. Species of Lepidoptera that rely on JH as the gonadotropic signal also exhibit polyandry. Several mechanisms have been suggested for the occurrence of polyandry, including availability of male-transferred nutrients (gonadotrophic effect), need for additional sperm, and increased genetic variability. We propose an additional reason for polyandry observed in some lepidopterans. If a female remains a virgin, her endogenous gonadotropic signal diminishes, and eggs that have been produced already may be resorbed to increase longevity. During copulation, the male may trigger a neural/humoral response in the female, thus stimulating release of her endogenous gonadotropic signal, JH, and/or inhibiting degradation of the same, whence she matures new eggs. The mating effect appears to act humorally on the cephalic structures in several species. Whether this change in JH titer is due to an effect on synthesis and release by corpora allata only or occurs in conjunction with inhibition of JH degradation is unknown. Arch. Insect Biochem. Physiol. 35:539–558, 1997.

Hemolymph juvenile hormone titers in pupal and adult stages of southwestern corn borer [Diatraea grandiosella (pyralidae)] and relationship with egg development.

Juvenile hormones I, II and III were monitored in hemolymph of pupal and adult stages of various ages of Diatraea grandiosella females. JH III was the predominant homologue followed by JH II, and JH I was rarely detectable. At day 5 after pupation, no JH was detectable. JH titers increased from 7.5days after pupation to a peak of 24.8ngml(-1) JH II and 26ngml(-1) JH III at adult emergence and then declined to low levels by 24h after emergence. Ovarian development in D. grandiosella parallels changes in hemolymph JH titers, but the role of JH in vitellogenesis is unclear since the time of vitellogenesis initiation has yet to be determined. No apparent vitellogenin deposition was observed in eggs 5days after pupation. Some oocytes were partially vitellogenic by 7.5days after pupation and oocytes continued to grow afterwards, but no oocytes were chorionated during the pupal stage. Chorionated oocytes were observed in 24-h-old female moths. Juvenile hormone is essential for chorion formation in this species, because decapitated pupae treated with 10&mgr;g JH III in corn oil developed chorionated oocytes while decapitated pupae treated with corn oil did not.

Vitellogenin and egg production in the moth, Heliothis virescens

The characteristics of vitellogenin (Vg) and the relationship between Vg production and egg production in the tobacco budworm, Heliothis virescens, were studied. The relationship between Vg production and juvenile hormone (JH) and the impact of mating on Vg and egg production were also investigated. Vg appears in the hemolymph of H. virescens about 6 h after moth eclosion. Vg may be separated into two apoproteins (ApoVg-I and ApoVg-II) by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE). The molecular weights were calculated to be 156,065 ± 800 for ApoVg-I and 39,887 ± 323 for ApoVg-II. SDS-PAGE analysis revealed that the female hemolymph Vg polypeptides appear to be identical to those from eggs but are absent in male hemolymph. Vg concentration was significantly higher in mated females than in virgin females of the same age at 48 h after emergence. Rates of egg production increased as Vg production increased; rates of egg production in mated females were significantly higher than those of virgin females at 48, 72, 96, and 120 h postemergence. Vg production is dependent on JH, because hemolymph from decapitated females lacked Vg while that of decapitated females treated with synthetic JH had Vg at levels comparable to similarly aged, normal H. virescens females. Hemolymph JH titers in mated females were significantly higher compared with those in virgin females at all sampling periods. The high JH level in mated females may explain the high Vg and egg production in mated H. virescens. Arch. Insect Biochem. Physiol. 34:287–300, 1997.

Sensory cues and receptors for oviposition by Heliothis virescens

When given a choice of cotton (host) and ground cherry (non host) in a close range situation, Heliothis virescens (F.), the tobacco budworm, depends primarily on contact chemoreception and mechanoreception, and not olfaction or vision, to discriminate between these two plant species for oviposition. Further, in the presence of a certain level of hairiness (the non‐host plant, ground cherry, has short capitate hairs), the females are capable of discriminating between these two plants based on contact chemosensory cues, if at least one pair of tarsi is intact. The contact chemosensilla on the ovipositor do not seem to be important for this purpose. These results are discussed in relation to the sensory cues and receptors for oviposition in other Lepidoptera, especially moths.

Mating in Heliothis virescens: transfer of juvenile hormone during copulation by male to female and stimulation of biosynthesis of endogenous juvenile hormone.

Studies were undertaken to determine whether adult males of Heliothis virescens transfer juvenile hormone (JH) to females during copulation, and an in vitro radiochemical assay was used to determine whether mating causes an allatotropic effect, i.e., stimulation of JH biosynthesis by corpora allata (CA). In vitro, CA from 3-day-old mated females synthesized and released approximately 2.5 times total JH as that of CA from comparably aged virgin females. Of the homologues, JH II exhibited significant increase in mated females; JH I also increased but not significantly. JH III remained similar to that of virgin females. This is the first demonstration of an allatotropic effect of mating in moths. In contrast to the female, CA of virgin males did not produce any JH, but accessory sex glands (ASG) in 3-day-old males synthesized small amounts of JH. Immediately after adult emergence, male ASG contained approximately 1.5 ng JH I and II, which increased by 12 h after emergence and remained at this high level up to 54 h after emergence. JH III was barely detected in ASG. JH in ASG of mated male immediately after uncoupling was depleted almost completely, and 24 h later recovered to levels comparable to that of 54-h-old virgin male. Virgin female bursa copulatrix did not contain any JH, but mated female bursa, immediately after uncoupling, had JH at levels comparable to that observed in virgin male ASG. By 6 h after uncoupling, JH levels decreased dramatically in mated female bursa. These data suggest the transfer of JH to females by the male.

Neuronal control of post‐coital pheromone production in the moth Heliothis virescens

The mechanism involved in bringing about post-coital suppression of pheromone production, pheromonostasis, was studied in the noctuid moth Heliothis virescens. Mating results in a transient suppression in pheromone production, the signal for which appears to originate in the testes and other components of the males reproductive system. The mating-induced pheromonostasis is due to an ascending signal via the central nervous system that appears to inhibit the release of the pheromonotropin, pheromone biosynthesis activating neuropeptide (PBAN), or other potential pheromonotropic substances, and is not due to a refractoriness in response of the sex pheromone glands to PBAN in the female. A similar mechanism is operative in several species of moths where post-coital pheromonostasis has been observed. Sperm quality is not important for pheromonostasis in H. virescens, because males with apyrene or eupyrene sperm elicit similar pheromonostatic responses. The pheromonostatic activity of the ecdysteroid 20-OH-ecdysone appears to be the result of a direct effect on the sex pheromone glands.

Temporal profiles of juvenile hormone titers and egg production in virgin and mated females of Heliothis virescens (Noctuidae)

Juvenile hormones (JH) I, II, and III were monitored in hemolymph of virgin and mated females of various ages in Heliothis virescens. JH I was the predominant homologue followed by JH II, but JH II was present at a higher level in young virgin females. JH III was detectable only at a low level. In virgin females, hemolymph JH titers were low at emergence (2.2ng/ml-total amount of JH I and JH II), but increased thereafter and reached a maximum at 24h of age (53.5ng/ml). At 30h and 36h of age, JH titers dropped to a low level, but increased again in older virgin females. After mating, JH titers increased significantly. JH titers at 0h after uncoupling (137.4ng/ml) were nearly 3 times as high as those in 24-h-old virgin females. Within 6h after uncoupling, JH titers decreased slightly, but titers increased with age of mated females and reached a level of 320.2ng/ml hemolymph at 72h after uncoupling. The titer of JH I and JH II was correlated highly with total number of eggs produced (r(2)=0.70, P<0.001). Mating stimulated JH production, resulting in an increase in egg production.

Fecundity, longevity and caloric patterns in female Heliothis virescens: Changes with age due to flight and supplemental carbohydrate

Abstract Bomb calorimetry, survivorship and fecundity analysis of female moths indicated that feeding on 5% sucrose reduces the impact of flight activity on longevity and oviposition. Flight has only a slight to moderate effect on longevity or reproductive capacity in fed females; but in unfed females, flight significantly decreased longevity and fecundity. By comparisons to unfed or unflown females, it is concluded that female moths use lipids as the primary flight fuel prior to the initiation of oviposition. Once oviposition begins the female increases its usage of carbohydrates as the flight fuel. Older females which are unable to obtain nectar cannot maintain their flight capacity, resulting in decreased reproductive potential. Therefore, nectar quantity, quality, and likelihood of availability can be important factors determining moth longevity and reproductive and flight capacities in the field.

Spermatophore formation and regulation of egg maturation and oviposition in female Heliothis virescens by the male

This study was undertaken, using various surgical manipulations, to examine the role and functions of accessory sex glands and testes in Heliothis virescens in spermatophore formation and stimulation of egg maturation in mated females. Normal females mated to accessory sex glandectomized (-ASG) males produced fewer eggs and retained most of their eggs in ovarioles compared with females mated to sham-operated (+ASG) or normal males. The ASG are the source of the components for spermatophore formation in H. virescens. Females mated to castrated (-Testes) males showed similar pattern of egg production as did females mated to -ASG males. Females mated to -Testes males and those mated to sham-operated (+Testes) or normal males were found to have spermatophore in the bursa after uncoupling. Normal females mated to allatectomized (-CA) males developed similar numbers of total eggs as did females mated to sham-operated (+CA) males and normal males.