Migration is a key life history strategy for many animals and requires a suite of behavioural, morphological and physiological adaptations which together form the ‘migratory syndrome’. Genetic variation has been demonstrated for many traits that make up this syndrome, but the underlying genes involved remain elusive. Recent studies investigating migration‐associated genes have focussed on sampling migratory and nonmigratory populations from different geographic locations but have seldom explored phenotypic variation in a migratory trait. Here, we use a novel combination of tethered flight and next‐generation sequencing to determine transcriptomic differences associated with flight activity in a globally invasive moth pest, the cotton bollworm Helicoverpa armigera. By developing a state‐of‐the‐art phenotyping platform, we show that field‐collected H. armigera display continuous variation in flight performance with individuals capable of flying up to 40 km during a single night. Comparative transcriptomics of flight phenotypes drove a gene expression analysis to reveal a suite of expressed candidate genes which are clearly related to physiological adaptations required for long‐distance flight. These include genes important to the mobilization of lipids as flight fuel, the development of flight muscle structure and the regulation of hormones that influence migratory physiology. We conclude that the ability to express this complex set of pathways underlines the remarkable flexibility of facultative insect migrants to respond to deteriorating conditions in the form of migratory flight and, more broadly, the results provide novel insights into the fundamental transcriptional changes required for migration in insects and other taxa.
Overwintering survival of Helicoverpa armigera (Hübner) was studied under field conditions during the winter of 2004-2005 and 2005-2006 to clarify whether a local population of this insect is established in northern Greece. Additionally, the postdiapause eclosion times of the overwintering generation was compared with adult male dynamics using pheromone traps. Our study supplies strong evidence that part, if not all, of the late-season generation of H. armigera overwinters as diapausing pupa in northern Greece, suggesting that a local population exists in this area. Pupae enter diapause during September and October in response to declining temperatures and photoperiod. A limited number of degree-days in autumn prevented part of the population from developing to the diapausing pupal stage. Larvae of H. armigera that were placed in field conditions late in October were not able to pupate and died. The termination of diapause and the eclosion of adults in the following spring were determined by the local soil temperatures. Diapause terminated between mid-April and early May, and adult emergence occurred within a 4- to 6-wk period from late April to early June. Females emerged slightly earlier than the males. The emergence of 10, 25, 50, 75, and 90% of adults required 153, 199, 252, 303, and 347 DD, respectively. Differences in timing and degree of overlap between adult emergence from the experimental population and pheromone trap catches suggests that the spring population of this species also includes immigrants.
Laboratory studies were conducted to assess the effect of temperature on the survival, development, longevity and fecundity of Helicoverpa armigera (Lepidoptera: Noctuidae) at eight different fluctuating temperatures with an amplitude ±9 °C under constant photoperiodic conditions of 16:8 h (L:D). H. armigera achieved complete development from egg to adult emergence between mean 17.5 and 32.5 °C. At mean 35 °C, all newly hatched larvae died and at mean 15 °C entered diapause at pupal stage. The lower developmental thresholds of the immature stages were estimated by a linear model and ranged from 4.63 °C (pupal stage) to 7.69 °C (egg stage). The developmental thresholds estimated by a nonlinear model were slightly higher than those estimated by the linear model. Adult longevity and fecundity were reduced at mean fluctuating temperatures 17.5 and 32.5 °C, but tended to be independent of the pattern of temperature change at moderate temperatures. The maximum reproductive performance, 1130 eggs per female, was observed at mean 25 °C. The intrinsic rates of increase were positive, meaning that H. armigera could be expected to persist or increase in number between mean 17.5 and 32.5 °C, with the maximum value at mean 27.5 °C. H. armigera survives, develops and reproduces within a wide range of fluctuating temperatures, while it completes the above functions with different levels of success at different mean temperatures of diurnal variation. Comparison of our results with similar data from the literature involving constant conditions is discussed. This information will provide a better understanding of H. armigera phenology and population dynamics under natural conditions and is essential to understanding the ecological and evolutionary consequences of climate change on this important species.
Helicoverpa armigera has been controlled effectively with chemical insecticides in the major cotton crop production areas of northern Greece for many years. However, a resurgence of the pest was observed in 2010, which significantly affected crop production. During a 4-year survey (2007-2010), we examined the insecticide resistance status of H. armigera populations from two major and representative cotton production areas in northern Greece against seven insecticides (chlorpyrifos, diazinon, methomyl, alpha-cypermethrin, cypermethrin, gamma-cyhalothrin and endosulfan). Full dose-response bioassays on third instar larvae were performed by topical application. Lethal doses at 50% were estimated by probit analysis and resistance factors (RF) were calculated, compared to a susceptible laboratory reference strain. Resistance levels were relatively moderate until 2009, with resistance ratios below 10-fold for organophosphates and carbamates and up to 16-fold for the pyrethroid alpha-cypermethrin. However, resistance rose to 46- and 81-fold for chlorpyrifos and alpha-cypermethrin, respectively in 2010, when the resurgence of the pest was observed. None of the known pyrethroid resistance mutations were found in the pyrethroid-resistant insects. The possible association between resistance and H. armigera resurgence in Greece is discussed.
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Laboratory studies were conducted to assess the effect of temperature on the survival, development, fecundity, and longevity of Helicoverpa armigera (Hübner) at 11 constant temperatures ranging from 12.5 to 40 degrees C, as well as at five alternating temperature regimes (25-10, 30-15, 32.5-17.5, 35-20, and 35-27.5 degrees C) and under a photoperiod of 16:8 (L:D) h. H. armigera reared at constant temperatures did not develop from egg to adult (emergence) outside the temperature range of 17.5-32.5 degrees C. The alternating conditions expanded this range from 10 to 35 degrees C. The lowest developmental thresholds of the immature stages were estimated by a linear model and ranged from 10.17 (pupal stage) to 11.95 degrees C (egg stage) at constant temperature regimes and from 1.1 to 5.5 degrees C, respectively at alternating temperatures. The values of developmental thresholds estimated using the nonlinear (Lactin-2) model were lower than those estimated by the linear model for constant and alternating temperature regimes except for larval and pupal stages at constant temperatures. Mean adult longevity fluctuated from 34.4 d at 15 degrees C to 7.6 d at 35 degrees C. Females reared under all alternating temperature regimes laid more eggs than females reared at any, except the 25 degrees C, constant temperature treatment. The intrinsic rate of increase was highest at 27.5 degrees C, at both the constant and the corresponding alternating temperature regimes (0.147 and 0.139, respectively). Extreme temperatures had a negative effect on life table parameters.
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