Xingfu Jiang

Regulation of Migration in Mythimna separata (Walker) in China: A Review Integrating Environmental, Physiological, Hormonal, Genetic, and Molecular Factors

ABSTRACT Each year the Mythimna separate (Walker), undertakes a seasonal, long-distance, multigeneration roundtrip migration between southern and northern China. Despite its regularity, the decision to migrate is facultative, and is controlled by environmental, physiological, hormonal, genetic, and molecular factors. Migrants take off on days 1 or 2 after eclosion, although the preoviposition period lasts ≈7 d. The trade-offs among the competing physiological demands of migration and reproduction are coordinated in M. separata by the “oogenesis-flight syndrome.” Larvae that experience temperatures above or below certain thresholds accompanied by appropriate humidity, short photoperiod, poor nutrition, and moderate density tend to develop into migrants. However, there is a short window of sensitivity within 24 h after adult eclosion when migrants can be induced to switch to reproductive residents if they encounter extreme environmental factors including starvation, low temperature and long photoperiod. Juvenile hormone (JH) titer is low before migration but high titers are associated with termination of migratory behavior and the switch to reproduction. Early release of JH by the corpora allata in environmentally stressed 1-d old adults, otherwise destined by larval conditions to be migrants, switches them to residents. Offspring inherit parental additive genetic effects governing migratory behavior. However, they also retain flexibility in expression of both flight and reproductive life history traits. The insect neuropeptide, allatotropin, which activates corpora allata to synthesize JH, controls adult flight and reproduction. Future research directions to better understand regulation of migration in this species are discussed.

Beet Webworm (Lepidoptera: Pyralidae) Migration in China: Evidence From Genetic Markers

ABSTRACT Genetic diversity within and among 11 geographic populations of the beet webworm Loxostege sticticalis across five provinces in the northern part of China were evaluated using amplified fragment length polymorphism (AFLP) analysis. Five AFLP primer combinations were used on 88 L. sticticalis samples from different locations, detecting a total of 384 polymorphic and 27 monomorphic fragments. Although extensive genetic diversity occurs among individuals from different geographic populations (P = 93.4%, h = 0.398, I = 0.572), the majority of the genetic diversity is within populations and not between populations (GST = 0.196), which agrees well with the results of analysis of molecular variance (84% of the total genetic variation is within populations), indicating high gene flow (NM = 2.046) among natural populations, which are not genetically differentiated. L. sticticalis in northeastern China, northern China, and northwestern China are part of a single large metapopulation. Cluster analyses based on AFLP data were preformed to graphically show groupings between individuals and between populations. Individuals from the same region were not grouped together very well. Eleven subpopulations were clustered into six broad groups, and there was no significant correlation between geographic distance and genetic dissimilarity (r = 0.1236, P = 0.8512). Principle component analysis also indicated a lack of genetic differentiation between the 11 populations. These results indicated that, although high genetic variability existed among individuals, there was little genetic differentiation among geographic populations, which can be explained by the effects of long distance migration of the beet webworm in China and consequent gene flow.

Expression of pheromone biosynthesis activating neuropeptide and its receptor (PBANR) mRNA in adult female Spodoptera exigua (Lepidoptera: Noctuidae).

The full-length cDNA of pheromone biosynthesis activating neuropeptide receptor (PBANR) was cloned from the beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae); it included an open reading frame of 1,053 bp encoding 350 amino acids. The PBANR of S. exigua (SePBANR) was structurally characteristic of G protein-coupled receptor and its amino acid sequence shared 98% identity with the PBANR of Spodoptera littoralis. Both pheromone biosynthesis activating neuropeptide (PBAN) and PBANR mRNA abundance were measured in the brain-subesophageal ganglion complex, pheromone gland, ventral nerve cord, and ovary of S. exigua female moths by real-time RT-PCR. The abundance of PBAN mRNA in brain-subesophageal ganglion complex and PBANR mRNA in pheromone gland was significantly greater compared to other tissues, suggesting that the ligand-receptor relationship of PBAN and PBANR exists quantitatively in S. exigua. Both PBAN and PBANR expression displayed a remarkable diurnal rhythm, for they were low and stable during the photophase (07:00-21:00) and increased markedly during the scotophase (with a maximum abundance at 23:30) in 3-day-old female moths. The abundance of PBAN and PBANR increased steadily from the 1st day to the 5th day of the adult female life. The pattern of both diurnal and daily expression of PBAN and PBANR mRNA were coincident with enhanced capacity of sex pheromone release and mating of S. exigua moths during the same period. We infer from these results that pheromone biosynthesis and release in S. exigua is regulated by PBAN via up-regulating synthesis.

Effects of larval density on flight potential of the beet webworm, Loxostege sticticalis (Lepidoptera: Pyralidae).

ABSTRACT Tethered-flight techniques were used to investigate the flight potential of 1-d-old adult beet webworm, Loxostege sticticalis L. (Lepidoptera: Pyralidae), reared at densities of 1, 10, 20, 30, and 40 larvae per 650-ml jar. Larval density had a significant effect on the flight potential, including total flight duration, distance, average velocity, and longest flight duration. Adults reared at a density of 10 larvae per jar, and those reared in isolation displayed the greatest and poorest flight capacity, respectively, relative to the other density treatments. Larval density also significantly affected length of the preoviposition period. Females with longer preoviposition period usually showed greater flight potential, and preoviposition period was positively correlated with flight potential. Rody weight, water content, and triglyceride content of the moths were significantly affected by larval density, whereas glycogen content was not. Triglyceride content of adults reared at the different larval densities corresponded to the observed differences in flight potential, whereas water content and body weight were not related to their flight potential. These results suggest that larval density exerts a significant influence on preoviposition period and triglyceride content, which in turn influence flight potential of adult L. sticticalis.

Starvation influences allatotropin gene expression and juvenile hormone titer in the female adult oriental armyworm, Mythimna separata.

The first day of adult life is the sensitive stage for shifting migrants into the resident morphs of the oriental armyworm (OAW), Mythimna separata (Walker). The juvenile hormone (JH) titer, expression of the allatotropin (AT) gene, and their relationship were investigated in adult female migrants starved in the sensitive stage, to understand the underlying mechanism of changing migrants into resident OAWs. Haemolymph JH titers of the starved female adults were mostly elevated earlier than in controls, although not all differences were statistically significant. JH I titers in the starved moths were significantly higher than those in the controls on 1, 2, and 5 days after treatment (DAT), respectively. JH II titers in the starved moths were significantly higher than the controls through the period tested except on 5 DAT. JH II is the most likely regulator in changing migrants into resident morphs. The relative quantities of AT expression in the starved moths were higher through the period tested except on 5 DAT. AT expression and JH titers appear to be positively correlated, especially for those in earlier days of the adult life. We infer that AT is the important regulator of JH levels. A model for the shifting of migrants into resident morphs in the OAW is proposed.

Relative fitness of near isogenic lines for melanic and typical forms of the oriental armyworm, Mythimna separata (Walker).

Abstract Melanism in the oriental armyworm Mythimna separata (Walker) is characterized by a black color in adults instead of the normal body color of off-white. Because inheritance of melanism in this moth follows Mendel’s law, with the melanic allele (t) recessive to the normal allele (T) at a single autosomal locus, a backcrossing procedure was used to develop near isogenic lines (NILs) for melanic (NILs-tt) and typical forms (NILs-TT) of this moth. Melanic males (tt) and typical females (TT) were crossed in the laboratory, their progeny (F1) were interbred, and F2 melanic males were backcrossed to typical female offspring of the parallel line. This cycle was repeated five times to create NILs for melanic and typical forms of this moth. The effects of the melanic allele on relative fitness were evaluated in terms of developmental and reproductive characteristics. Life tables of the NILs were constructed to determine net reproductive rate (R0). Results indicated that NILs-tt possessed apparent developmental and reproductive advantages, including shorter immature development time, higher larval and pupal survival, a shorter adult preoviposition period, and more eggs laid per female compared with the NILs-TT.

Amplified Fragment Length Polymorphism Analysis of Mythimna separata (Lepidoptera: Noctuidae) Geographic and Melanic Laboratory Populations in China

Abstract Genetic diversity within and among three wild-type natural populations and one melanic laboratory population of Mythimna separata (Walker) (Lepidoptera: Noctuidae) were evaluated using amplified fragment length polymorphism (AFLP) analysis. Although extensive genetic diversity occurs among individuals from different geographic populations (P = 54.5%, h = 0.209, I = 0.305), the majority of the genetic diversity is within populations and not between populations (GST = 0.172), indicating high gene flow (NM = 2.403) and suggesting that M. separata in northern China are a part of a single large metapopulation. Genetic diversity in the natural populations was significantly higher than that in the melanic laboratory population (with P = 43.4% versus P = 25.9%, h = 0.173 versus h = 0.086, and I = 0.251 versus I = 0.127), suggesting that the melanic laboratory population is narrowly genetic-based and genetically uniform. Genetic similarities based on AFLP data were calculated, and cluster analysis was preformed to graphically display groupings between individuals and populations. Individuals from the same region were not grouped together in cluster analysis of three natural populations, whereas melanic individuals from laboratory population were grouped together very well. Four subpopulations were clustered into two broad groups. Melanic laboratory population became a single group, which had apparent differentiation from the other group in which three natural subpopulations were included. These results indicated that although high genetic variability existed among the individuals of natural populations, there was little genetic differentiation among three geographic populations that could be explained by the effects of the long distance migration of the oriental armyworm in China enhanced the level of gene flow. Influences of migration on the genetic polymorphism and differentiations that make a significant contribution to evolution in this insect are reviewed.

Determination of sensitive stage for switching migrant oriental armyworms into residents.

Abstract The sensitive stage for switching migrant oriental armyworm, Mythimna separata (Walker), into residents was determined under the stresses of starvation and cold (5°C) in a laboratory population. Presumed migrants developing from gregarious larvae oviposited significantly earlier when adults were starved or shocked by cold only during the first 24 h after emergence. In contrast, preoviposition periods for migrants starved or cooled on days 2–6 of adult life were not significantly affected. The preresponse period of male migrants starved on day 1 of adult life was also significantly shortened, whereas those treated on following days were not significantly influenced. Similarly, total lifetime fecundity of migrants, which were starved or cooled on day 1 after emergence, was significantly increased; however, the same effect was not observed in migrants that were stressed later. Although preoviposition periods of presumed residents, developing from solitary larvae, starved or shocked by cold during the first several days of adulthood were significantly prolonged, this delay was not related to migratory strategy, because lifetime fecundity was not significantly reduced. The results suggest that the first day of adult life in this species is a sensitive stage during which migrants can shift into residents, although residents cannot shift to migrants. Thus, starvation and cold experienced in this stage induces migrants to modify their developmental path into reproduction and residency.

Biocontrol of the oriental armyworm, Mythimna separata, by the tachinid fly Exorista civilis is synergized by Cry1Ab protoxin

Tritrophic interactions between Mythimna separata, its tachinid parasite Exorista civilis and the Cry1Ab were examined. Although 6th instar M. separata mortality increased with increasing Cry1Ab concentration, some tolerance was evident. Likewise, parasitization by E. civilis resulted in only 18% host mortality. However, combination of Cry1Ab and E. civilis parasitization resulted in a significant Cry1Ab dose-dependent increase in mortality over that of either alone, including a 50–56% synergistic increase in efficacy at the two concentrations tested. Pupal weight, adult emergence and lifetime fecundity of M. separata derived from larvae surviving both agents were negatively affected. The ability of E. civilis to parasitize and subsequently develop on the host was not adversely influenced by Cry1Ab. Instead, pupation rate increased significantly among host larvae fed 3.125 μg/g Cry1Ab diet. Overall, our results demonstrate that use of Cry1Ab to control M. separata not only is compatible with the use of the tachinid parasitoid, but that the two methods can act synergistically to manage this destructive pest, provide support for the safety of transgenic Cry1Ab Bt plants in China. This example of two independent pest management strategies acting synergistically against a difficult pest offers a new perspective of broad significance in striving for agricultural sustainability.

METHOPRENE INFLUENCES REPRODUCTION AND FLIGHT CAPACITY IN ADULTS OF THE RICE LEAF ROLLER, Cnaphalocrocis Medinalis (GUENỂE) (LEPIDOPTERA: PYRALIDAE)

Juvenile hormone (JH) influences many aspects of insect biology, including oogenesis-flight syndrome tradeoffs between migration and reproduction. Drawing on studies of many migratory insects, we posed the hypothesis that JH influences migratory capacity and oogenesis in the rice leaf roller, Cnaphalocrocis medinalis. We treated adults moths (days 1, 2 and 3 postemergence) with the JH analog (JHA), methoprene, and then recorded the influences of JHA treatments on reproduction. JHA treatment on day 1 postemergence, but not on the other days, shortened the preoviposition period, although JHA did not influence total fecundity, oviposition period, or longevity. We infer day 1 postemergence is the JH-sensitive stage to influence reproduction. Therefore, we treated moths on day 1 postemergence with JHA and recorded flight capacity, flight muscle mass, and triacylglycerol (TAG) accumulation. JHA treatments did not influence flight speed, but led to reductions in flight durations and flight distances. At day 3 posttreatment (PT), JHA-treated females flew shorter times and less distance than the controls; JHA-treated males, however, only flew shorter times than the controls. JHA treatments led to reductions in flight muscle mass in females at days 2-3 PT and reductions in TAG content in females at day 3 PT, but, these parameters were not influenced by JHA in males. These findings strongly support our hypothesis, from which we infer that JH is a major driver in C. medinalis oogenesis-flight syndrome tradeoffs. Our data also reveal a JH-sensitive stage in adulthood during which JH influences the oocyte-flight syndrome in C. medinalis.