Gregarious Females Research Articles

With or without you: gut microbiota does not predict aggregation behavior in European earwig females

Abstract The reasons why some individuals are solitary, and others gregarious are the subject of ongoing debate as we seek to understand the emergence of sociality. Recent studies suggest that the expression of aggregation behaviors may be linked to the gut microbiota of the host. Here, we tested this hypothesis in females of the European earwig. This insect is ideal for addressing this question, as adults both naturally vary in the degree to which they live in groups and show interindividual variation in their gut microbial communities. We video-tracked 320 field-sampled females to quantify their natural variation in aggregation and then tested whether the most and least gregarious females had different gut microbiota. We also compared the general activity, boldness, body size, and body condition of these females and examined the association between each of these traits and the gut microbiota. Contrary to our predictions, we found no difference in the gut microbiota between the most and least gregarious females. There was also no difference in activity, boldness, and body condition between these two types of females. Independent of aggregation, gut microbiota was overall associated with female body condition, but not with any of our other measurements. Overall, these results demonstrate that a host’s gut microbiota is not necessarily a major driver or a consequence of aggregation behavior in species with inter-individual variation in group living and call for future studies to investigate the determinants and role of gut microbiota in earwigs.

Olfactory Mating Signals in the Migratory Locust Locusta migratoria.

Swarming locusts cause huge plagues across the world threatening food production. Before swarms form, locust populations exhibit a dramatic phase change from a solitary to a gregarious phase. The cause of this phase change is a complicated interplay of conspecific and environmental cues and is, especially for one of the major pests, the migratory locust Locusta migratoria, still not well understood. Here we study the behavior of both solitary and gregarious L. migratoria towards the headspace odors of conspecifics. As we do not find a general attraction of gregarious animals to the headspace of gregarious conspecifics, swarm formation does not seem to be mainly governed by olfactory aggregation cues. When testing for potential mating signals, we observe that the headspace of virgin gregarious females is highly attractive only towards virgin males of the same phase, while mated gregarious males and solitary males, regardless of their mating state, do not become attracted. Interestingly, this phase-specific attraction goes along with the finding, that mating behavior in experiments with inter-phasic pairings is extremely rare. Our data suggest that odor emissions in L. migratoria play a significant role in a mating context.

Bonobo social organization at the seasonal forest-savanna ecotone of the Lomami national park.

Examining the relationship between food and primate social organization helps us understand how the environment shaped hominin social evolution. However, there is debate as to whether the social differences between our two closest relatives, bonobos (Pan paniscus) and chimpanzees (Pan troglodytes), are due to differences in food availability between their respective habitats or to nonenvironmental factors. The most prominent theory is that bonobo communities have more socially cohesive, stable parties, centered on gregarious females because they evolved in food-rich habitat where individuals, especially females, are less burdened by competition with groupmates. However, more research on bonobos in habitats with seasonal variation in food is needed. This study measured food availability and bonobo social organization at Luzaka, a new site in a seasonal forest fragment. Fruit abundance and dispersion were recorded for a year at Luzaka with the same methods used at Wamba, a bonobo site in more seasonally stable habitat and terrestrial herbaceous vegetationdensity was measured. At Luzaka, bonobo parties were also recorded for a year using camera traps. Fruit was more seasonal and dispersed at Luzaka than at Wamba. However, the social organization of Luzaka bonobos resembled social organization of bonobos at less seasonal sites. There were minor effects of fruit clumping on party size without effects on the proportion of females in parties suggesting that at Luzaka, the clumping of fruit slightly affected social cohesiveness but does not disproportionately affect females. Bonobo social cohesiveness and female gregariousness appears consistent and compatible with seasonal habitat.

Aggregation pheromone 4-vinylanisole promotes the synchrony of sexual maturation in female locusts.

Reproductive synchrony generally occurs in various group-living animals. However, the underlying mechanisms remain largely unexplored. The migratory locust, Locusta migratoria, a worldwide agricultural pest species, displays synchronous maturation and oviposition when forms huge swarm. The reproductive synchrony among group members is critical for the maintenance of locust swarms and population density of next generation. Here, we showed that gregarious female locusts displayed more synchronous sexual maturation and oviposition than solitarious females and olfactory deficiency mutants. Only the presence of gregarious male adults can stimulate sexual maturation synchrony of female adults. Of the volatiles emitted abundantly by gregarious male adults, the aggregation pheromone, 4-vinylanisole, was identified to play key role in inducing female sexual maturation synchrony. This maturation-accelerating effect of 4-vinylanisole disappeared in the females of Or35-/- lines, the mutants of 4-vinylanisole receptor. Interestingly, 4-vinylanisole displayed a time window action by which mainly accelerates oocyte maturation of young females aged at middle developmental stages (3-4 days post adult eclosion). We further revealed that juvenile hormone/vitellogenin pathway mediated female sexual maturation triggered by 4-vinylanisole. Our results highlight a 'catch-up' strategy by which gregarious females synchronize their oocyte maturation and oviposition by time-dependent endocrinal response to 4-vinylanisole, and provide insight into reproductive synchrony induced by olfactory signal released by heterosexual conspecifics in a given group.

Re-examination of the maternal control of progeny size and body color in the desert locust Schistocerca gregaria: Differences from previous conclusions

The desert locust shows conspicuous phase polyphenism of various traits in response to crowding conditions. Gregarious females lay larger eggs that produce black hatchlings, whereas solitarious females lay smaller eggs that produce green hatchlings. Previous studies have shown that changes in egg size and hatchling body color occurred easily in the laboratory upon exposure of the female parent to crowding or isolation for as few as 2 days. Based on these observations, these studies concluded that female adults perceive crowding stimuli with their antennae and require light for perception of the stimuli, with contact chemicals present on the integument of sexually mature males that are responsible for the crowding effect. We undertook this study to identify the chemical structures of the reported contact substances, which remain unknown to date. However, we could not reproduce the main results reported in the aforementioned studies and found that egg size and hatchling body color did not alter easily, even after crowding or isolation of the female parent for 2 days or longer. We were not able to observe a change in the progeny crowding characteristics by stimulating the antennae of female adults through either physical contact with male adults or using hexane extracts of their body surfaces. Similarly, the importance of light for gregarization or solitarization was also ruled out. These results indicate that some of the conclusions of previous studies should be reconsidered.

Sexual Behavior of the Desert Locust During Intra- and Inter-Phase Interactions

Mating and reproduction behaviors and strategies are fundamental aspects of an organism’s evolutionary and ecological success. In locusts, intra- as well as inter-phase reproductive interactions among gregarious and solitarious locust populations have a major impact on the locust population dynamics. However, practically all previous work on locust sexual behavior has been limited to the gregarious phase. Here we provide a first detailed description of pre-copulatory behavior of solitarious desert locusts. We compare our findings with those of previous reports of pre-copulatory behavior of gregarious locusts, focusing on the behavioral elements that serve in inter-sex signaling and communication. We also studied inter-phase (mixed pairs) reproductive interactions. Solitarious males were found to invest more in pre-copulatory courtship and signaling compared to their gregarious counterparts; and solitarious females played a more dominant role in the inter-sex communication than gregarious females. The solitarious females were also less likely to demonstrate rejection-related behavioral patterns than gregarious females. As a consequence of the particular characteristic behavior of each phase, the most successful among intra- and inter-phase pairs were gregarious males with solitary females. Least successful were solitary males paired with gregarious females, indicating a strong asymmetry in inter-phase reproductive interactions.

White-nosed coatis in Arizona: tropical carnivores in a temperate environment

Intraspecific variation in ranging and social behavior can be perplexing, but also provides an opportunity to assess which behavioral attributes are labile in the face of geographic variation in resources. White-nosed coatis (Nasua narica) are group-living carnivores of tropical origin that possess an unusual social system. In the resource-rich tropics, larger-bodied males are solitary, whereas females live in groups (“bands”) along with young of both sexes, but leave them to give birth and wean their young. Males often disperse socially, but not spatially. We studied coatis in the Chiricahua National Monument, Arizona, a resource-poor, arid, and highly seasonal landscape. Using live capture and radiotelemetry in conjunction with microsatellite DNA analyses, we found that Chiricahua coatis exhibited the species-typical pattern of solitary males and gregarious females. Young males left their natal bands as yearlings, and some were found as adults within their natal home range. On the other hand, home ranges were expanded greatly. Bands, and some males, focused their movements in areas several km² in extent over periods of weeks or months but shifted those areas markedly among seasons. Some males followed the typical mammalian pattern of natal dispersal beyond their natal area. Rates of movement were higher and female associations appeared to be more flexible than in the tropics. Adult females sometimes ranged singly, not only around the time of parturition, but also when population density was low and occasionally otherwise. In addition, home range overlap was high among both sexes during some seasons, and female bands sometimes fused for prolonged periods. Core patterns of sociality are constant in both resource-rich tropical and resource-poor temperate populations, but coatis appear to make major adjustments in scale of movement and frequency of association in response to resource variation.

MicroRNA-276 promotes egg-hatching synchrony by up-regulating brm in locusts

Developmental synchrony, the basis of uniform swarming, migration, and sexual maturation, is an important strategy for social animals to adapt to variable environments. However, the molecular mechanisms underlying developmental synchrony are largely unexplored. The migratory locust exhibits polyphenism between gregarious and solitarious individuals, with the former displaying more synchronous sexual maturation and migration than the latter. Here, we found that the egg-hatching time of gregarious locusts was more uniform compared with solitarious locusts and that microRNA-276 (miR-276) was expressed significantly higher in both ovaries and eggs of gregarious locusts than in solitarious locusts. Interestingly, inhibiting miR-276 in gregarious females and overexpressing it in solitarious females, respectively, caused more heterochronic and synchronous hatching of progeny eggs. Moreover, miR-276 directly targeted a transcription coactivator gene, brahma (brm), resulting in its up-regulation. Knockdown of brm not only resulted in asynchronous egg hatching in gregarious locusts but also impaired the miR-276-induced synchronous egg hatching in solitarious locusts. Mechanistically, miR-276 mediated brm activation in a manner that depended on the secondary structure of brm, namely, a stem-loop around the binding site of miR-276. Collectively, our results unravel a mechanism by which miR-276 enhances brm expression to promote developmental synchrony and provide insight into regulation of developmental homeostasis and population sustaining that are closely related to biological synchrony.

Biochemical changes in developing embryos of Schistocerca gregaria (Orthoptera: Acrididae) induced by pheromone produced by ovipositing gregarious females

Trans-generational transfer of gregarious-phase traits in the desert locustSchistocerca gregaria(Forskål, 1775) is mediated by primer gregarizing pheromonal signals produced by ovipositing females that experience crowding. We monitored time-course proteomic events in eggs from solitary-reared locusts that had been exposed for 1, 3, 5, 7, 10 and 12 days to different levels of the sand-associated gregarizing signal originating from 0, 3, 5 or 10 ovipositions by crowd-reared females. Evidence for the phase transition was sought by comparing the protein patterns of embryos thus exposed with those from crowd-reared (gregarious) controls; this comparison was continued until the stage of the first instars. Expressed proteins were analysed by two-dimensional protein gel electrophoresis, and patterns from the different treatments within stages were compared by profile matching andχ2analyses. Eggs derived from crowd- and solitary-reared females showed essentially similar protein patterns at early stages of embryogenesis; however, mature stages (particularly, days 10 and 12) and hatchlings demonstrated significantly different patterns. Protein patterns of eggs from solitary-reared females that were incubated in sand contaminated with the pheromonal signal and of the hatchlings that emerged were similar to those derived from gregarious females and dependent on the level of the pheromone to which the embryos had been exposed. The results confirm the gregarizing effect of the signal and constitute a useful basis for unravelling the mechanism of the signalling cascades associated with gene expressions triggered by the pheromone.

Adaptive difference in daily timing of hatch in two locust species, Schistocerca gregaria and Locusta migratoria: The effects of thermocycles and phase polyphenism

This study examined the effects of temperature and phase polyphenism on egg hatching time in the desert locust, Schistocerca gregaria, and the migratory locust, Locusta migratoria. The two species exhibited differences and similarities in hatching behavior when exposed to different temperature conditions. In 12-h thermocycles of various temperatures, the S. gregaria eggs hatched during the cryoperiod (low temperature period), whereas L. migratoria eggs hatched during the thermoperiod (high temperature period). The eggs of both species hatched during the species-specific period of the thermoperiod in response to a temperature difference as small as 1°C. Furthermore, the locusts adjusted hatching time to a new thermal environment that occurred shortly before the expected hatching time. In both species, the hatching of the eggs was synchronized to a specific time of the day, and two hatching peaks separated by approximately 1day were observed at a constant temperature after the eggs were transferred from thermocycles 3days before hatching. Eggs laid by gregarious females hatched earlier than those laid by solitarious females in S. gregaria but this difference was not observed in L. migratoria.

Eggs and hatchlings variations in desert locusts: phase related characteristics and starvation tolerance.

Locusts are grasshopper species that express phase polyphenism: modifying their behavior, morphology, coloration, life history and physiology in response to crowding. Desert locusts, Schistocerca gregaria, epigenetically modify progeny quality and quantity in response to crowding. Gregarious (crowded) females produce larger but fewer progeny than do solitarious (isolated) ones. The variability of progeny quality within single egg pod and the reasons why gregarious progeny have a better survival rate than solitarious ones remains unclear. This study investigated 1) the effects of rearing density on the variation in egg size within single egg pods 2) the starvation tolerance of hatchlings from mothers with different phases and 3) the physiological differences in hatchling energy reserve. Isolated females produced smaller but more eggs than did crowded ones. The variation in egg size within egg pods was greater in the latter than in the former. A negative relationship between egg size and number of eggs per egg pod was observed for both groups. Under starvation conditions, gregarious hatchlings survived significantly longer than solitarious ones. Among the solitarious hatchlings, the survival time was longer with increased hatchling body size. However, small individuals survived as long as large ones among the gregarious hatchlings. The percentage of water content per fresh body weight was almost equal between the two phases, before and after starvation. In contrast, the percentage of lipid content per dry body weight was significantly higher in gregarious hatchlings than in solitarious ones before starvation, but became almost equal after starvation. These results demonstrate that female locusts not only trade-off to modify their progeny size and number, but also vary progenies' energy reserves. We hypothesize that gregarious females enhance their fitness by producing progeny differently adapted to high environmental variability and particularly to starvation conditions.

Parental phase status affects the cold hardiness of progeny eggs in locusts

Summary1. The capacity to adapt to low temperature is critical for the persistence of insect populations in heterogeneous environments. Locusts show remarkable phenotypic plasticity, termed ‘phase change’, in response to local population density.2. In this study, the hypothesis that population density, as a social factor, affects the cold hardiness of progeny is validated in the migratory locust, Locusta migratoria, which shows remarkable density‐dependent phase changes between gregarious and solitarious phases.3. We demonstrated that eggs of gregarious and solitarious migratory locusts significantly differ not only in size and weight, but also in cold hardiness. Eggs of solitarious locusts are more resistant to cold stress compared with those of gregarious locusts, with longer 50% lethal time at different low temperatures and under different acclimation treatments of cooling rates or cold exposure time, lower upper limit of chill injury zone, and lower sum of injurious temperature resulting from temperature–time regression. The lowest cooling rate (0·05 °C min−1) yields the highest survival of cold for eggs.4. A custom‐made microarray covering 9154 unigenes of the migratory locust demonstrated quite different gene expression profiles in the two phases in response to normal or low temperature. Under cold stress, the gregarious‐phase eggs have higher transcriptional levels of heat‐shock proteins, DOPA decarboxylases and tyrosine hydroxylase, whereas the solitarious‐phase eggs exhibit stimulated lipid metabolism and carboxylic acid transport.5. Hybridization between the two phases showed that the cold hardiness of eggs from the hybrid with solitarious females is significantly higher than that of the hybrid with gregarious females, and the cold hardiness of eggs from each reciprocal hybrid is close to their maternal origins. These results indicate that the cold hardiness of progeny is affected by the parental phase status.

Artificial miniaturization causes eggs laid by crowd-reared (gregarious) desert locusts to produce green (solitarious) offspring in the desert locust, Schistocerca gregaria

The mechanism underlying the phase-dependent polyphenism in hatchling body coloration was studied by testing for a possible causal relationship with egg size in the desert locust, Schistocerca gregaria. Crowd-reared (gregarious) females typically produce large, black offspring, whereas females reared in isolation (solitarious) deposit small, green offspring. We first tested for possible genetic differences in the role of egg foam by washing or separating eggs from two strains of locust. No solitarizing effect was found in either of the strains tested, supporting a previous finding, using another laboratory strain, to show that the hatchling body coloration and size are pre-determined in the ovary of the mother and no egg foam factor is involved in the control of the hatchling body coloration. Topical application of fenoxycarb, a juvenile hormone analog (JHA), and implantation of extra corpora allata (CA), taken from Locusta migratoria, caused gregarious female adults of S. gregaria to produce small eggs. Some eggs laid by CA-implanted females produced green hatchlings. All large eggs chosen among those deposited by gregarious females produced black hatchlings. When eggs were either kept on dry filter paper at nearly saturated relative humidity during embryogenesis or pricked with a needle so that some egg yolk was squeezed out, some produced small, green hatchlings. These results suggested that the amount of egg yolk or the availability of yolk material may determine the body coloration of hatchlings.

Morphological and behavioural characteristics of a gynandromorph of the desert locust, Schistocerca gregaria

Abstract Morphological and behavioural characteristics are investigated for a gynandromorph of the desert locust, Schistocerca gregaria, appearing under isolated rearing conditions in the laboratory. It has both male and female external reproductive organs bilaterally. The body size and dimensions are similar to a normal male. Morphometric traits (fore wing length/maximum head width ratio and fore wing length/hind femur length ratio) of the gynandromorph are typical for the values of solitarious locusts. When the gynandromorph is placed into an arena holding ten sexually mature gregarious females, it shows a distinct male behaviour: it jumps on a female and tries to mate with her. When kept together with males, males recognize this gynandromorph as a female because some of them try to mount, although no successful copulation is observed. The results suggest that the gynandromorph might have had a female‐specific pheromone. Dissection reveals that the gynandromorph has no testis but abnormal ovaries containing vitellogenic oocytes. These observations indicate that the gynandromorph obtained has a mixture of male and female morphological characteristics and behaves like a male but is recognized as a female by conspecific males.

The nature of the gregarizing signal responsible for maternal transfer of phase to the offspring in the desert locust Schistocerca gregaria.

We examined aggregative behavior of hatchlings of the desert locust Schistocerca gregaria emerging from solitaria egg pods that had been incubated in sand previously used for consecutive ovipositions by gregarious females. Hatchlings derived from such eggs were significantly more gregarious than those derived from pods laid in clean sand. The gregarizing effect of the sand-associated factor originating from 3, 5, or 10 ovipositions by gregarious females increased in a dose-dependent fashion. Washing the sand with organic solvents following such ovipositions, or flushing it with nitrogen gas, led to substantial loss of its gregarizing effect, showing that the active signal is volatile and of medium polarity. The gregarizing activity of the exposed sand correlated with the presence of C-8 unsaturated ketones, (Z)-6-octen-3-one and (E,E)-3,5-octadiene-2-one and its E,Z isomer, previously shown by us to form part of the releaser pheromone system that mediates group oviposition in S. gregaria. These ketones were present in relatively large amounts in the eggs obtained from egg pods of gregarious females and were also detected in the extracts of accessory glands of gregarious females, a candidate source of the gregarization factor. It is proposed that the pheromone is secreted at the onset of oviposition. This would account for its distribution predominantly within the eggs and surrounding sand at the site of oviposition. The study sheds new light on the pheromonal mechanism associated with transgenerational transmission of gregarious characters in crowded S. gregaria populations and provides yet another case in this insect of dual releaser and primer roles played by the same pheromone blend.

Time-course haemolymph juvenile hormone titres in solitarious and gregarious adults of Schistocerca gregaria, and their relation to pheromone emission, CA volumetric changes and oocyte growth

The haemolymph JH III titres in solitarious and gregarious adult desert locusts, Schistocerca gregaria, were examined in relation to corpus allatum (CA) volumes, aggregation–maturation pheromone production in males and oocyte growth in females. The JH titres of gregarious females were generally higher than those of solitarious females at all ages studied. The titre patterns, however, were similar: relatively high on day 10, dropping to low levels between days 20 and 25, before rising again by day 25. In the solitarious males, the JH titre was very low on day 10 after fledging, but increased gradually and reached a maximal amount on day 30. The JH titre in gregarious males was low on day 10, elevated on day 15 coinciding with the start of the production of the pheromone, and dropped to a relatively low level on day 20 around the time of maximal pheromone production, then rising again by day 25. These results suggest that biosynthesis of the pheromone is associated with a high JH titre peak in the haemolymph. Although a clear relationship was found during the first gonadotropic cycle between JH titres, on one hand, and CA volume and oocyte growth, on the other, in both phases, no such correlation could be discerned in the second cycle.

Changing Oviposition Preferences of the Desert Locust, Schistocerca gregaria, Suggest a Strong Species Predisposition for Gregarization

Field surveys at five sites within desert locust breeding habitats around Port Sudan during three successive seasons indicated that early in the rainy season the incoming solitary females oviposited predominantly in the vicinity of Heliotropium spp. (∼66%) and millet (∼32%) seedlings. Solitary nymphs also preferred to feed on these plants. Follow-up cage experiments were conducted in the field in which solitary and gregarious female locusts were presented with choices of selected desert plants and egg pods. When presented with bulrush millet, Heliotropium spp., Zygophyllum simplex, and untreated moistened sand, solitary females oviposited adjacent to the first two plants (40% and 60%, respectively). However, when offered a choice of either or both of these plants together with egg pods derived from gregarious and/or solitary insects, solitary females showed a significantly higher preference for ovipositing near gregaria egg pods than near the plants, with solitaria egg pods eliciting the least response. In contrast with solitary females and in the absence of gregaria egg pods, gregarious females preferred to oviposit in untreated moist (control) sand (74–77%) away from the plants (6–14%) or solitaria egg pods (∼4%). However, when present, gregaria egg pods elicited significantly more oviposition. These and previous results indicate a hierarchy of phase-dependent oviposition preferences in the desert locust and are interpreted in terms of a strong propensity of the species to exploit opportunities under appropriate conditions to facilitate congregation and the gregarization of the progeny.

Evidence for Additional Pheromonal Components Mediating Oviposition Aggregation in Schistocerca gregaria

Oviposition responses of gregarious adult gravid females of the desert locust Schistocerca gregaria to sand previously used for oviposition by gregarious females (contaminated sand, with eggs and froth removed) and to sterilized sand (noncontaminated sand) were compared in bioassays. Gregarious females preferentially oviposited into contaminated sand. The number of egg pods laid by the females was found to correlate with the frequency of use of the contaminated sand for oviposition. Charcoal-trapped volatiles from the contaminated sand also elicited similar responses from gregarious females. Gas chromatography–electroantennographic detection (GC-EAD) analysis of the trapped volatiles revealed the presence of three electrophysiologically active compounds that were identified by GC-MS as (Z)-6-octen-2-one, (E,E)-3,5-octadien-2-one and its geometric isomer (E,Z)-3,5-octadien-2-one. The relative amounts of the three EAG-active compounds in the volatiles increased with consecutive oviposition by the gregarious females. These results indicate mediation of additional pheromonal components in the oviposition behavior of gregarious desert locust that are distinct from those that were previously identified in the froth volatiles.

Ecdysteroids during ovarian development and embryogenesis in solitary and gregarious schistocerca gregaria

Maternal ecdysteroids identified in the vitellogenic oocytes of Schistocerca gregaria included more than 80% polar conjugates, up to 5% free ecdysteroids, and up to 15% non-hydrolyzable polar metabolites. The representations of ecdysone (E), 20-hydroxyecdysone (20E), and 2-deoxyecdysone (2dE) in the conjugates was about 16:3:1, and in the free ecdysteroids about 3:1:1. The quantity of ecdysteroids in the ovaries before egg-laying reached 2.3 ng 20E equiv. per mg tissue in the solitary, and 8.9 ng/mg in the gregarious females. Newly laid eggs contained 14 ng and 89 ng, respectively, of 20E equiv. per egg. Nearly all egg ecdysteroids were in form of conjugates and their content declined during the first half of embryonic development. The amount of ecdysteroids sharply increased to over 70 ng 20E equiv./egg in the solitary, and to nearly 400 ng/egg in the gregarious phase. In the second half of embryonic development, the representation of conjugates in total ecdysteroids was reduced to 45-55%, whereas that of free E + 20E rose to 30-40%. Free 2dE remained low but, in the gregarious embryos, free 26E increased to 10% of all ecdysteroids. The conjugates of solitary embryos contained nearly exclusively E and 20E (in ratio 2:1), whereas those of the gregarious embryos included E, 20E, 2dE, and 26E (in ratio 12:7:4:1). Towards the end of embryonic development, the amounts of conjugates and of free ecdysteroids decreased, while that of polar metabolites rose. A sharp drop in ecdysteroid content was associated with hatching but the more than five times higher ecdysteroid level in the gregarious than in the solitary phase was maintained in the newly hatched larvae. Arch. Copyright 1999 Wiley-Liss, Inc.

A Gregarizing Factor Present in the Egg Pod Foam of the Desert Locust Schistocerca gregaria

The behavioural phase state and coloration of hatchling Schistocerca gregaria were examined in a series of experiments to determine the means by which phase characteristics are passed between generations. Both crowding of solitary-reared females at the time of oviposition and high egg pod densities promoted behavioural gregarization, although the former appeared to be a rather more potent factor. In contrast, egg pod density alone appeared to promote the development of hatchlings with dark patterns characteristic of the gregarious phase. The phase characteristics of hatchlings were unaffected when sand previously used for oviposition was used to collect further egg pods. Early separation of presumptive gregarious eggs from egg pods laid by crowd reared females led to solitarization of the hatchlings, indicating that a factor, either in or around the eggs, removed by early separation promoted gregarization. Both the eggs and foam plugs of egg pods from crowd reared, gregarious females appeared to be a source of this gregarizing factor. In contrast, there was no evidence for a solitarization factor in egg pods from solitary-reared S. gregaria. Saline extracts of egg pod foam plugs produced an active factor which promoted gregarization both in eggs from solitary-reared females and in eggs from gregarious females which were separated and washed to remove the factor. Solitary eggs were influenced by the gregarizing factor in foam plug extracts for up to 1 day after oviposition. Saline extracts of foam plug retained their activity for up to 1 day. Initial studies on the properties of this factor were made. We conclude that the foam plugs of egg pods from crowd-reared, gregarious locusts contain a small (<3 kDa), hydrophilic gregarizing factor which is produced at the time of oviposition and which predisposes hatchlings to attain characteristics of the gregarious phase.