Effect of phase status on responses to AKHI in the desert locust,Schistocerca gregaria gregaria

Hyperlipaemic response to adipokinetic hormone (AKH I) was demonstrated in both solitary and gregarious phases of the desert locust, Schistocerca gregaria gregaria. Time-course studies showed that the gregarious locusts had a faster response to the hormone than their solitary counterparts. At peak response time (90 min), the gregarious locusts were more sensitive to AKH I doses below 2 pmol while the solitary locusts had a higher response above this dose. Upon injection of the hormone, lipoprotein conversion occurred, resulting in the formation of the low density lipoprotein (LDLp). The LDLp formed in the gregarious locusts was much larger than that of the solitary locusts. The fat body lipid reserve (expressed as % fat body dry weight) was significantly (P < 0.01) higher in the gregarious (79.02 ± 2.77%) than in the solitary locusts (65.23 ± 2.55%). Triacylglycerol was the major lipid class representing 83.9 and 73.9% of the total lipids in gregarious and solitary locusts, respectively. The higher fat body lipid reserves and efficient LDLp formation in response to AKH in gregarious locusts compared to solitary locusts suggests a physiological adaptation for prolonged flights. © 1996 Wiley-Liss, Inc.     

Composition and emission dynamics of migratory locust volatiles in response to changes in developmental stages and population density

Chemical communication plays an important role in density‐dependent phase change in locusts. However, the volatile components and emission patterns of the migratory locust, Locusta migratoria, are largely unknown. In this study, we identified the chemical compositions and emission dynamics of locust volatiles from the body and feces and associated them with developmental stages, sexes and phase changes. The migratory locust shares a number of volatile components with the desert locust (Schistocerca gregaria), but the emission dynamics of the two locust species are significantly different. The body odors of the gregarious nymphs in the migratory locust consisted of phenylacetonitrile (PAN), benzaldehyde, guaiacol, phenol, aliphatic acids and 2,3‐butanediol, and PAN was the dominant volatile. Volatiles from the fecal pellets of the nymphs primarily consist of guaiacol and phenol. Principal component analysis (PCA) showed significant differences in the volatile profiles between gregarious and solitary locusts. PAN and 4‐vinylanisole concentrations were significantly higher in gregarious individuals than in solitary locusts. Gregarious mature males released significantly higher amounts of PAN and 4‐vinylanisole during adulthood than mature females and immature adults of both sexes. Furthermore, PAN and 4‐vinylanisole were completely lost in gregarious nymphs during the solitarization process, but were obtained by solitary nymphs during gregarization. The amounts of benzaldehyde, guaiacol and phenol only unidirectionally decreased from solitary to crowded treatment. Aliphatic aldehydes (C7 to C10), which were previously reported as locust volatiles, are now identified as environmental contaminants. Therefore, our results illustrate the precise odor profiles of migratory locusts during developmental stages, sexes and phase change. However, the function and role of PAN and other aromatic compounds during phase transition need further investigation.     

Neural correlates to flight-related density-dependent phase characteristics in locusts

Locust phase polymorphism is an extreme example of behavioral plasticity; in response to changes in population density, locusts dramatically alter their behavior. These changes in behavior facilitate the appearance of various morphological and physiological phase characteristics. One of the principal behavioral changes is the more intense flight behavior and improved flight performance of gregarious locusts compared to solitary ones. Surprisingly, the neurophysiological basis of the behavioral phase characteristics has received little attention. Here we present density-dependent differences in flight-related sensory and central neural elements in the desert locust. Using techniques already established for gregarious locusts, we compared the response of locusts of both phases to controlled wind stimuli. Gregarious locusts demonstrated a lower threshold for wind-induced flight initiation. Wind-induced spiking activity in the locust tritocerebral commissure giants (TCG, a pair of identified interneurons that relay input from head hair receptors to thoracic motor centers) was found to be weaker in solitary locusts compared to gregarious ones. The solitary locusts' TCG also demonstrated much stronger spike frequency adaptation in response to wind stimuli. Although the number of forehead wind sensitive hairs was found to be larger in solitary locusts, the stimuli conveyed to their flight motor centers were weaker. The tritocerebral commissure dwarf (TCD) is an inhibitory flight-related interneuron in the locust that responds to light stimuli. An increase in TCD spontaneous activity in dark conditions was significantly stronger in gregarious locusts than in solitary ones. Thus, phase-dependent differences in the activity of flight-related interneurons reflect behavioral phase characteristics.     

Neural correlates to flight‐related density‐dependent phase characteristics in locusts

Locust phase polymorphism is an extreme example of behavioral plasticity; in response to changes in population density, locusts dramatically alter their behavior. These changes in behavior facilitate the appearance of various morphological and physiological phase characteristics. One of the principal behavioral changes is the more intense flight behavior and improved flight performance of gregarious locusts compared to solitary ones. Surprisingly, the neurophysiological basis of the behavioral phase characteristics has received little attention. Here we present density‐dependent differences in flight‐related sensory and central neural elements in the desert locust. Using techniques already established for gregarious locusts, we compared the response of locusts of both phases to controlled wind stimuli. Gregarious locusts demonstrated a lower threshold for wind‐induced flight initiation. Wind‐induced spiking activity in the locust tritocerebral commissure giants (TCG, a pair of identified interneurons that relay input from head hair receptors to thoracic motor centers) was found to be weaker in solitary locusts compared to gregarious ones. The solitary locusts' TCG also demonstrated much stronger spike frequency adaptation in response to wind stimuli. Although the number of forehead wind sensitive hairs was found to be larger in solitary locusts, the stimuli conveyed to their flight motor centers were weaker. The tritocerebral commissure dwarf (TCD) is an inhibitory flight‐related interneuron in the locust that responds to light stimuli. An increase in TCD spontaneous activity in dark conditions was significantly stronger in gregarious locusts than in solitary ones. Thus, phase‐dependent differences in the activity of flight‐related interneurons reflect behavioral phase characteristics. © 2003 Wiley Periodicals, Inc. J Neurobiol 57: 152–162, 2003     

Adipokinetic Hormone and Flight Fuel Related Characteristics of Density-Dependent Locust Phase Polymorphism: A Review

Recent findings on differences between the gregarious and solitary phases of locusts are reviewed in relation to flight fuel utilization, adipokinetic responses, and adipokinetic hormones. Laboratory results obtained with Locusta migratoria migratorioides show that the amount of lipid reserves, resting levels of haemolymph lipids, and hyperlipaemic responses to flight and to injection of corpus cardiacum extract or of synthetic adipokinetic hormones, are higher in crowded than in isolated locusts. No major phase-dependent differences seem to exist in flight-related carbohydrate metabolism. The adipokinetic hormone content of the corpora cardiaca is higher in younger isolated locusts than in crowded ones. Adipokinetic hormone precursor-related peptide content of the corpora cardiaca is also higher in isolated than in crowded locusts. Crowded locusts have higher lipid reserves and higher hyperlipaemic responses to flight than isolated locusts also in Schistocerca gregaria and, following injection of synthetic adipokinetic hormone, the formation of low density lipophorin is higher in crowded than in isolated locusts of this species. The laboratory results obtained with isolated and crowded locusts are extrapolated to understand the ecophysiology of the migrations of solitary and gregarious field populations of L.m. migratorioides according to available information on the differences in the migration of the two phases. It is inferred that in this species solitary locusts have a rather coarse adipokinetic strategy focused on a single prereproductive long-distance migratory flight, whereas gregarious locusts possess a fine adipokinetic balance for reiterative, sometimes unpredictably long-distance, migrations in the prereproductive, as well as reproductive, periods. The differences between the adipokinetic strategies of solitary and gregarious S. gregaria seem to be less dramatic, nevertheless, they indicate a better adaptation of the gregarious phase to prolonged flights.     

A miniaturized assay to quantify effects of chemicals or physical stimuli upon locust activity

Solitary and gregarious locusts differ in many traits, such as body color, morphometrics and behavior. With respect to behavior, solitary animals shun each other, while gregarious animals seek each other＇s company, hence their crowding behavior. General activity, depending on the temperature, occurs throughout the day but is much lower in solitary locusts. Solitary locusts occasionally fly by night while gregarious locusts fly regularly during daytime （swarming）. In search of new assays to identify substances that control or modify aspects of （phase） behavior, we designed a simple activity assay, meant to complement existing behavioral measurement tools. The general activity is reflected in the number of wall hits, that is, the number of contacts between the locust and the vertical walls of a small arena. Using this single parameter we were able to quantify differences in total activity of both nymphs and adults of isolation-reared （solitary）, regrouped- and crowdreared （gregarious） locusts under different conditions. Furthermore, we demonstrate that there are inter- and intra-phase dependent differences in activities of 5th instar nymphs afar injections of the three different adipokinetic hormones.     

Effect of Paranosema locustae (Microsporidia) on the behavioural phases of Locusta migratoria (Orthoptera: Acrididae) in the laboratory

The ability of parasites to modify the behaviour of their hosts is a wide spread phenomenon, but the effects of microsporidian parasites on locust behaviour remain unexplored. Here the frequencies of directional changes (ND) and jumping (NJ) per minute of gregarious locusts infected with 2000 spores of the microsporidian parasite Paranosema locustae were significantly different from those of untreated locusts 10 and 16 days after infection, being similar to values for solitary nymphs. In contrast, the behaviour of locusts inoculated with the lower doses of 200 spores/locust was sometimes like that of solitary nymphs. At other times, behaviour was intermediate between solitary and gregarious, i.e. transitional. The rearing density did not affect the turning and jumping behaviour of infected locusts, and their behaviours were similar to those of solitary locusts at 10–16 days after infection. Our study demonstrates that infection with P. locustae may lead gregarious locusts to change some of their behaviour to that typical of solitary locusts.     

Neurophysiological studies of flight-related density-dependent phase characteristics in locusts

Locusts show an extreme example of density-dependent phase polymorphism, demonstrating within the species differences in morphology as well as biology, dependent on the population density. Behavior is the primary density-dependent change which facilitates the appearance of various morphological and physiological phase characteristics. We have studied density dependent differences in flight related sensory and central neural elements in the desert locust Schistocerca gregaria. Wind generated high frequency spiking activity in the tritocerebral commissure giant (TCG, an identified interneuron that relay inputs from head hair receptors to thoracic motor centers) that was much less intense in solitary locusts, compared to gregarious ones. In addition the solitary locusts' TCG demonstrated much stronger adaptation of its response. In cases when flight was initiated high frequency TCG activity was independent of the locust phase. The tritocerebral commissure dwarf (TCD) is a GABAergic flight related interneuron that is sensitive to ambient illumination intensity. An increase in the TCD spontaneous activity under dark vs. light conditions was significantly higher in gregarious locusts then in solitary ones, implying a flight-related inhibitory mechanism that is far more active in gregarious locusts under dark conditions. Thus, density-dependent phase differences in interneuron activity pattern and properties well reflect and may be at least partially responsible to behavioral flight-related characteristics.     

Are recession populations of the desert locust (Schistocerca gregaria) remnants of past swarms?

The desert locust (Schistocerca gregaria) undergoes crowding-induced phase transformation from solitary to gregarious, which involves changes in behaviour, colour, development, morphometry, fecundity and endocrine physiology. During recession, solitary locusts persist in the central, drier part of the species' range in small pocket populations that are prone to extinction. During the intermittent upsurges and the subsequent plagues, gregarious swarms attain huge population size and invade a vast area causing major damage to agriculture. A highly variable nuclear DNA marker, a noncoding 3' end fragment of an antennapedia-class homeobox gene, was screened in locust samples from Eritrea. Despite the homogenizing potential of plague swarms, the last of which was in 1986-89 and originated in this region, the population genetic structure of solitary phase locusts along the Red Sea coast of Eritrea revealed significant divergence. The pattern of divergence indicated that the invasion of the western and northern plains in the summer of 1995 may not, as reported then, have originated in eastern Chad or western Sudan. A number of interrelated hypotheses have been presented to explain the observed genetic heterogeneity between the sampled populations. We conclude, with caution due to the limited sample sizes, that: (i) geographical isolation between breeding sites during plagues and recession; (ii) the marked differences in the flight behaviour of plague swarms and recession populations; (iii) possible failure of gregarious locusts to solitarize and re-establish in recession areas; and (iv) the effect of repeated extinction and recolonization in the meta-population contribute to the maintenance of the genetic structure of recession populations. Potentially productive future research has been identified.     

Adult female desert locusts require contact chemicals and light for progeny gregarization

Crowding causes many organisms to express phenotypic plasticity in various traits. Phase polyphenism in desert locusts represents one extreme example in which a solitary form (solitarious phase) turns into a gregarious form (gregarious phase) in response to crowding. Conspicuous differences in body size and colour occur even in hatchlings. The phase‐specific differences in hatchling characteristics are caused by the tactile stimuli perceived by the antennae of their mother. However, the nature of the tactile stimuli and the mechanism by which the perceived stimuli are processed as a gregarizing signal remain unknown. To explore this problem, the antennae of solitarious adult females of the desert locust Schistocerca gregaria are touched with the bodies of conspecific locusts at different physiological stages and those of other species. The results suggest that a cuticular chemical factor at a specific developmental stage of conspecific locusts causes the solitarious females to produce large eggs that give rise to black hatchlings characteristic of gregarious forms (progeny gregarization), and that this or a similar compound occurs in other acridids, crickets and cockroaches but not in beetles. The involvement of a chemical substance is also supported by hexane extracts of cuticular surfaces of locusts that induce the same effects. Interestingly, crowding induces such gregarizing effects only when the female receives the appropriate stimulus in the presence of light. Solitarious female S. gregaria with their head capsule coated with phosphorescent paint exhibit progeny gregarization in response to crowding and light pulses in darkness, whereas those treated in the same way without light pulses fail to do so.     

蝗虫多型现象的神经内分泌调控

蝗虫有两种型,即散居型和群居型。蝗灾通常由群居型蝗虫所引发。多年来人们试图找到控制蝗虫由散居型向群居型转变的关键因子,以期控制蝗虫危害。该文主要从神经内分泌的角度概述了蝗虫多型性的生理机制,重点介绍了保幼激素、蜕皮激素和脑神经肽［His7］-corazonin在蝗虫多型性中的主要作用和机制。     

Number and size of intracisternal granules in the adipokinetic hormone-secreting cells of locusts: a phase-dependent characteristic

Abstract.The intracisternal (= ergastoplasmic) granules in the adipokinetic hormone-secreting cells of the glandular lobes of the corpora cardiaca in Locusta migratoria migratorioides represent accumulations of adipokinetic prohormones within cisternae of the rough endoplasmic reticulum. Solitary locusts have more and larger intracisternal granules than gregarious locusts. This coincides with the general locomotor activity and thus the energy metabolism in solitary locusts being quite different from that of gregarious locusts, which apparently has consequences for the amounts of adipokinetic hormones synthesized and/or released and, consequently, for the storage of these hormones in the intracisternal granules. These granules apparently function as supplementary stores of secretory material.