Zhudong Liu

Life Table Studies of the Cotton Bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), on Different Host Plants

Abstract The development, body weight, survivorship, and reproduction of the cotton bollworm, Helicoverpa armigera (Hübner), were evaluated in the laboratory at 27°C and photoperiod L:D = 14:10 on six host plants: cotton (Gossypium hirsutum L.), corn (Zea mays L.), tomato (Lycopersicon esculentum Mill), hot pepper (Capsicun frutescens L.), tobacco (Nicotiana tobacum L.), and common bean (Phaseolus vulgaris L.). Cotton bollworm larvae successfully survived on all six host plants, although mortality was very high on hot pepper and tomato. The developmental time of immature stages ranged from 26.6 d on corn to 35.1 d on tomato. Body weight of young fourth instar larvae ranged from 22.8 mg on corn to 5.9 mg on tobacco, while body weight of young last instar larvae and pupae ranged from 176.7 mg on cotton to 132.5 mg on tomato, and 285.2 mg on corn to 167.1 mg on tomato, respectively. Immature survival from egg to pupa varied from 33.1% on cotton to 1.7% on hot pepper. The average number of eggs oviposited by adults reared on cotton, corn, common bean, tomato, hot pepper, and tobacco were 708.4, 784.8, 778.1, 559.1, 562.5, and 314.3, respectively. The net reproductive rate of this species, measured from egg to egg, varied from 117.6 on cotton to 5.1 on hot pepper. We conclude that H. armigera can complete their life cycle on all six host plants, although tomato and hot pepper were relatively unsuitable.

Pupal diapause of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) mediated by larval host plants: pupal weight is important.

Facultative diapause, a strategy that allows insects to initiate additional generations when conditions are favorable or to enter diapause when they are not, has a profound effect on the ecology and evolution of species. Most previous studies have concentrated on the role of photoperiod and temperature in inducing facultative diapause in insects. In contrast, here we studied pupal diapause mediated by larval host plants in the cotton bollworm Helicoverpa armigera, and confirmed that pupal weight is a critical factor. Two groups of third instar H. armigera larvae, kept at 25 °C with L:D=8:16 and 20 °C with photoperiod of L:D=8:16, respectively, were fed on six host plants and on artificial diet (as a control) to determine how larval host plants affect diapause incidence and related traits (such as pupal weight and developmental duration). The data showed larval host plants affected diapause incidence significantly and the effects could be masked by low temperature. Further analysis showed that pupal size, not the length of the sensitive stage, affected the decision to enter diapause. In a further experiment, third-instar to final-stage larvae deprived of artificial diet for 2 days demonstrated a direct relationship between pupal weight and diapause incidence. These results suggest that larval host plants, by affecting pupal size, may influence diapause occurrence in H. armigera. This has important adaptive significance for both over-wintering survival and the possibility for completing an additional generation.

Effects of transgenic Bt cotton on overwintering characteristics and survival of Helicoverpa armigera.

The effects of transgenic Bt cotton on the overwintering generation of the cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), are unknown. We hypothesized that a Bt cotton diet may adversely affect fitness of this generation and examined fresh weight, lipids, glycogens, low-molecular-weight sugars and SCPs (supercooling points) of pupae, as well as survival of larvae, diapausing pupae and adult emergence in comparison with controls. Field and laboratory experiments showed that larvae fed on Bt cotton had a decreased pupation rate, and fewer entered diapause and emerged as adults compared with larvae fed non-Bt cotton. Furthermore, larvae fed Bt cotton had reduced pupal weight, glycogen content and trehalose levels both in diapausing and in non-diapausing pupae, and only diapausing pupae had an increased SCP compared to controls. The SCPs of diapausing pupae reared on Bt cotton were significantly higher than those reared on non-Bt cotton. The trehalose levels of diapausing pupae reared on Bt cotton were significantly lower than those of larvae reared on non-Bt cotton. Thus, these results suggest that a Bt cotton diet weakens the preparedness of cotton bollworm for overwintering and reduces survival of the overwintering generation, which will in turn reduce the density of the first generation in the following year. Effects of transgenic Bt cotton on the overwintering generation of cotton bollworm appear to have significantly contributed to the suppression of cotton bollworm observed throughout northern China in the past decade.

Host-size mediated trade-off in a parasitoid Sclerodermus harmandi.

Optimality models of host-parasitoid relationships have traditionally assumed that host quality increases as a function of host size at parasitism. However, trade-offs may play a crucial role in species evolution and should be found in host-parasitoid interactions where the host quality may differ between different sizes. Here, we investigated the effects of host size differences of Monochamus alternatus larva on foraging decisions, parasitism and related fitness in a gregarious ectoparasitoid, Sclerodermus harmandi. Two-choice and non-choice experiments were conducted with M. alternatus larvae to evaluate preference and performance of S. harmandi, respectively. Results from two-choice test showed that adult females prefer to attack large larvae rather than small larvae. In no-choice tests, adult females needed more time to paralyze large larvae than small larvae before laying eggs on the body surface of M. alternatus larvae and had lower survival and parasitism rate on those large larvae. Foraging decisions of S. harmandi led to the selection of the most profitable host size for parasitoid development, which showed more offspring gained on large M. alternatus larvae than on small larvae and got higher body weight on those large hosts. This study demonstrates the existence of trade-off occurring during host-parasitoids interactions according to host size related quality.

The Pheromone Frontalin and its Dual Function in the Invasive Bark Beetle Dendroctonus valens

The red turpentine beetle, Dendroctonus valens LeConte, is one of the most destructive invasive forest pests in China, having killed more than 6 million pines since its first outbreak in 1999. Little is known about D. valens pheromone biology and no aggregation pheromone has yet been identified. Analysis by gas chromatograph/mass spectrometer of volatiles collected from live beetles in China showed that female beetles produce frontalin and males do not. Olfactory assays in the laboratory showed that males were attracted to frontalin at a wide range of concentrations, whereas females were attracted to it at a narrow range of concentrations. In field trials, 3-carene, a monoterpene kairomone from a pine tree selected to host the beetles attracted both sexes, and when frontalin was added, the total number of beetles captured increased by almost 200%. However, increasing concentrations of frontalin significantly decreased the percentage of female beetles trapped. These results suggest a new role of frontalin as an aggregation pheromone in addition to a female-produced sex pheromone, which was previously shown in a North American population. The dual functions of the pheromone frontalin produced by D. valens females, as well as its ecological significance for overcoming host resistance, are discussed.

The effects of α-pinene on the feeding performance and pheromone production of Dendroctonus valens

Herbivorous insects exploit multiple plant cues to detect and orient toward suitable hosts and, accordingly, hosts have evolved complex constitutive and inducible defenses in response. In China, the red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae), an invasive bark beetle from North America, attacks mainly Pinus tabuliformis Carrière (Pinaceae), which contains many monoterpenes. In this study, we explored how the monoterpene α‐pinene affects the feeding performance and pheromone production of D. valens. First, the composition and quantities of monoterpenes of both P. tabuliformis healthy trees and fresh stumps were determined and the infestation of D. valens in healthy trees and fresh stumps was investigated, linking the amount of monoterpenes and D. valens infestation. Gas chromatography–mass spectrometry (GC‐MS) analysis showed that P. tabuliformis mainly contained α‐pinene, with concentrations of 0.1 and 0.5 mg g−1 in healthy pine phloem and stump phloem, respectively. Second, the monoterpenes influence on feeding performance was tested using phloem media with α‐pinene concentrations ranging from 0 to 30 mg g−1. The results showed that the percentages of beetles boring and the gallery lengths of both adult females and larvae were negatively correlated with the α‐pinene concentration although body weight changes did not correlate with α‐pinene concentration. Finally, pheromone analysis showed that the production of all pheromones increased with increasing α‐pinene concentrations. This study showed the dual effects of α‐pinene on D. valens: α‐pinene inhibited the bark beetles feeding activities and in turn the bark beetle made use of it to produce pheromones. Our study indicated the importance of promptly removing fresh stumps in the field for the management of the bark beetle.

Host adaptation of a gregarious parasitoid Sclerodermus harmandi in artificial rearing

Adaptations of a parasitoid, Sclerodermus harmandi (Buysson 1903) (Hymenoptera: Bethylidae) to a target host, Monochamus alternatus Hope (Coleoptera: Cerambycidae), were investigated in laboratory colonies. We quantified the physiological and behavioral flexibility of the parasitoid reared on M. alternatus compared to the parasitoid reared on a common host, Saperda populnea (Coleoptera: Cerambycidae). Adult performance (fecundity and pre-oviposition period) and offspring physiological responses (mortality and sex ratio) of S. harmandi reared on M. alternatus improved over multiple generations. S. harmandi females of the 12th generation on the target host showed better performance, i.e., shorter time for searching and handling, higher oviposition preference, and, subsequently, lower offspring mortality and proportion of male emergence on the host than those from the base stock. Our work demonstrated that these traits in S. harmandi were influenced by the integration of cultured generations and host quality, which implies that the potential effectiveness of S. harmandi as a biological control agent can be further improved.

Olfactory cues in host and host-plant recognition of a polyphagous ectoparasitoid Scleroderma guani

Olfactory bioassays were performed to investigate the specific odors utilized as host location cues by the beetle parasitoid, Scleroderma guani (Hymenoptera: Bethylidae), a primary biological control agent against Monochamus alternates (Coleoptera: Cerambycidae), the most important vector beetle of the pinewood nematode, Bursaphelenchus xylophilus (Nematoda: Aphelenchoididae), the causal agent for pine wilt disease. Female parasitoids were tested with volatiles and extracts derived from their host beetle. Behavioral assays (Y-olfactometer bioassay and circle arena) demonstrated the response of female parasitoids to odors from host plants damaged by beetle larvae and from their excreta. When available contact cues were additionally provided, the parasitoids showed particularly strong preferences for samples of fresh brown frass of larval beetles. To confirm the electroantennograms (EAG) activity of identified compounds, analyses were repeated with a synthetic blend composed predominantly of compounds in the crude extracts that had revealed apparent electrophysiological activity. Antennal responses to four monoterpenes and three oxygenated monoterpenes were among the strongest, which indicated their potential for use in development of semichemial-based management of the beetle.

Summer diapause induced by high temperatures in the oriental tobacco budworm: ecological adaptation to hot summers.

Summer diapause in Helicoverpa assulta (Hübner), which prolongs the pupal stage, particularly in males, is induced by high temperatures. In the laboratory, 3rd-, 4th-, 6th-instar and prepupal larvae were exposed to high temperatures – 33 and 35 °C with a photoperiod of LD16:8 – until pupation to induce summer diapause. The results showed that the incidence of summer diapause was influenced by temperature, stage exposed, and sex. The higher the temperature, the more often summer diapause was attained. Sixth-instar and prepupal larvae were the sensitive stages for summer diapause induction. H. assulta summer-diapausing pupae needed diapause development to resume development when temperatures became favorable. Furthermore, both body mass and energy storage capacity (lipid and glycogen) were significantly affected by diapause rather than sex, and were significantly higher in summer-diapausing pupae than in non-diapausing pupae. In addition, the body mass loss and respiration rate showed that the rate of metabolism in the summer-diapausing pupae was consistently lower than in non-diapausing pupae, which were significantly affected by diapause and pupal age. We conclude that summer diapause in H. assulta is a true diapause, and H. assulta has evolved mechanisms to accumulate energy storage and to lower its metabolism to adapt to hot summers.

Sound-Triggered Production of Antiaggregation Pheromone Limits Overcrowding of Dendroctonus valens Attacking Pine Trees

For insects that aggregate on host plants, both attraction and antiaggregation among conspecifics can be important mechanisms for overcoming host resistance and avoiding overcrowding, respectively. These mechanisms can involve multiple sensory modalities, such as sound and pheromones. We explored how acoustic and chemical signals are integrated by the bark beetle Dendroctonus valens to limit aggregation in China. In its native North American range, this insect conducts nonlethal attacks on weakened trees at very low densities, but in its introduced zone in China, it uses mixtures of host tree compounds and the pheromone component frontalin to mass attack healthy trees. We found that exo-brevicomin was produced by both female and male D. valens, and that this pheromone functioned as an antiaggregating signal. Moreover, beetles feeding in pairs or in masses were more likely than were beetles feeding alone to produce exo-brevicomin, suggesting a potential role of sound by neighboring beetles in stimulating exo-brevicomin production. Sound playback showed that an agreement sound was produced by both sexes when exposed to the aggregation pheromone frontalin and attracts males, and an aggressive sound was produced only by males behaving territorially. These signals triggered the release of exo-brevicomin by both females and males, indicating an interplay of chemical and sonic communication. This study demonstrates that the bark beetle D. valens uses sounds to regulate the production of an antiaggregation pheromone, which may provide new approaches to pest management of this invasive species.