Epigenetic transmission of phase in the desert locust, Schistocerca gregaria: Determining the stage sensitive to crowding for the maternal determination of progeny characteristics

Desert locust female adults respond to crowded conditions by changing progeny characteristics such as egg size, clutch size (no. of eggs per pod), hatchling body size and coloration. This study was conducted to determine the stage sensitive to crowding in this locust. Reproductively active females reared in isolation increased egg size and decreased clutch size and the proportion of green hatchlings after exposure to crowded conditions (in which each female was kept with four male adults). These changes depended not only on the timing of exposure to crowded conditions during the reproductive cycle but also on the length of the exposure. By varying the time and length of the exposure, it was found that crowding had no influence on progeny characteristics during the last two days of egg development at 31 °C and that there was a four-day sensitive stage before this period. The sensitive stage coincided with the time when the affected oocytes were 1.5-4 mm long, while the sensitivity to crowding appeared to be constant over the sensitive stage. The larger the magnitude of the increase in egg size after exposure to crowding, the smaller the proportion of green hatchlings (and the larger the proportion of gregarized dark hatchlings); there was a sigmoidal relationship between the two variables. Based on these results, we propose a model for determining the stage sensitive to crowding in both the female parent and the oocytes.     

Maternal effects on progeny body size and color in the desert locust, Schistocerca gregaria: examination of a current view

Hatchling body color and size of the desert locust, Schistocerca gregaria, are determined by the population density of the mothers during their reproductive period. Smaller green hatchlings are produced by adults at low population density (solitarious conditions) and larger dark hatchlings at high population density (gregarious conditions). One claim states that a pheromonal factor secreted by gregarious mothers into foam plugs of egg pods induces darkening in hatchlings. Previous research suggests that the foam factor can be removed by separating eggs individually within 1h of deposition, causing presumptive gregarious eggs to hatch without darkening. The present study re-examined this claim and possible factors that have been proposed which could account for the difference between our results and those reported earlier. Early separation was performed on eggs with a low mortality rate. The results showed that the egg separation did not increase the incidence of green hatchlings. Once chorionated in the ovary, eggs remained unchanged in size until the second day after oviposition in either isolated or crowded locusts. This and other results suggest that the phase-dependent differences in body size and color of hatchlings are established in the ovary and that modifications by the accessory gland factor either in the oviduct or after deposition are unlikely.     

Phase-specific responses to different qualities of food in the desert locust, Schistocerca gregaria: developmental, morphological and reproductive characteristics

Solitarious female adults are known to produce smaller hatchlings than those produced by gregarious adults of the desert locust, Schistocerca gregaria. This study investigated developmental, morphological and reproductive responses to different qualities of food in hatchlings of different phases. Mortality was higher, the duration of nymphal development longer and adult body weight lighter with a low-quality food than a high-quality food. Gregarious hatchlings showed better survivorship, grew faster and became larger adults than did solitarious ones. The incidence of locusts exhibiting extra molting, which was typically observed in the solitarious phase, was dramatically increased when a low-quality food was given to the solitarious hatchlings. Low-quality food caused locusts to shift morphometric ratios toward the values typical of gregarious forms; smaller F/C (hind femur length/maximum head width) and larger E/F (elytra length/hind femur length). Solitarious hatchlings grown at either high- or low-quality foods and then given high-quality food after adult emergence revealed that food qualities during the nymphal stage influence their progeny quality and quantity via adult body size that influenced reproductive performance. Female adults showed an overshooting response to a shift from low- to high-quality food by increasing egg production that was specific to body size. This study may suggest that gregarious hatchlings are better adapted to adverse food conditions than solitarious counterparts and extra molting is induced even among gregarious hatchlings under poor food conditions.     

Phase-related body-color polyphenism in hatchlings of the desert locust, Schistocerca gregaria: re-examination of the maternal and crowding effects

The mechanism controlling the body color of hatchlings was studied for the desert locust, Schistocerca gregaria. A pheromonal factor secreted by gregarious female adults into the foam plugs of egg pods has been suggested to cause darkening in their progeny. We re-examined the role of this maternal factor by washing or separating eggs at deposition. Eggs produced by crowd-reared female adults were washed with saline or separated individually without being washed immediately after deposition and the body color of the hatchlings from them was compared with that from the eggs unwashed and kept in the egg pod until hatching. Most hatchlings were dark and no significant difference was found in the proportions of dark- and light-colored hatchlings between the treatments and controls. Likewise, eggs separated before the foam plug deposition produced dark-colored hatchlings as in the un-separated controls. These results demonstrated that neither washing nor separation of eggs at deposition affected the hatchling body coloration. The variation in hatchling body color was correlated closely to the body weight at hatching, indicating that hatchling body color had been determined maternally. Green hatchlings reared under crowded conditions remained green until the second stadium at which black patterns were induced. It was concluded that body color at hatching has been determined maternally and crowding during the first nymphal stadium influences nymphal body color but its effect is not manifested until the second stadium. The present study casts doubts on the presence of a recently suggested pheromonal factor on the color of the hatchlings.     

Tactile stimuli perceived by the antennae cause the isolated females to produce gregarious offspring in the desert locust, Schistocerca gregaria

Maternal determination of progeny body size and coloration in the desert locust, Schistocerca gregaria, depends on the crowding conditions experienced during the short sensitive period that occurs two to six days before the deposition of the egg pod. Solitarious (isolated-reared) females produce relatively small eggs that yield solitarious green hatchlings but, females that are exposed to crowded conditions during the sensitive period, produce larger eggs that yield the dark-colored hatchlings characteristic of gregarious forms. The present study aimed to determine the stimuli influencing the maternal determination of progeny characteristics as well as the site at which such stimuli are perceived. By exposing isolated female adults to various combinations of visual, olfactory and tactile stimuli from a crowd of other adults, we found that no crowding effects could be elicited without tactile stimulation. Coating of various body surfaces with nail polish followed by exposure to crowding stimulation suggested that female adults perceive crowding stimuli with their antennae. This finding was supported by another experiment in which the antennae were either removed or covered with wax before the isolated females were exposed to crowded conditions. Neither serotonin nor an antagonist of its receptor affected the density-dependent maternal determination of progeny characteristics when injected into isolated or crowded female adults.     

Sex differentiation studies relating to releaser aggregation pheromones of the desert locust,Schistocerca gregaria

Behavioural responses of nymphs and adults in the gregarious phase of the desert locust,Schistocerca gregaria (Forskal) (Orthoptera: Acrididae) were investigated in a single-chamber bioassay system to a choice of two columns of air, one permeated with airborne volatiles emanating from either sex of nymphs or adults and the other untreated. There was no sexual differentiation in the production of or response to nymphal volatiles. Young adults of either sex did not produce a stimulus with significant activity. Of the older adults, only the males produced the aggregation stimulus to which both sexes were equally responsive. Charcoal-trapped volatiles from the two sexes of nymphs and adults evoked similar aggregation responses. Antennae of the older adults showed significantly higher EAG responses than those of fifth instar nymphs to all four volatile collections, of which volatiles from older adult males were the most stimulatory and evoked the highest EAG amplitudes.     

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.     

Maternal effects on phase characteristics in the desert locust, Schistocerca gregaria: a review of current understanding

Desert locusts demonstrate pronounced density-dependent polyphenism: a complex suite of traits shifts over the lifetime of an individual in response to crowding or isolation. These changes also accumulate across generations through a maternal effect. Female desert locusts alter the developmental trajectory of their offspring in response to their own experience of crowding. The mother possesses a memory of both the recency and extent of crowding and shifts the phase state of her hatchlings accordingly. Extensive experimental work has shown that offspring behaviour is controlled by a low molecular weight, polar compound (or compounds) released from the mother's accessory glands. The chemical identity of this agent is not yet known.     

Is juvenile hormone involved in the maternal regulation of egg size and progeny characteristics in the desert locust?

The role of juvenile hormone (JH) in the maternal regulation of progeny characteristics was examined in the desert locust, Schistocerca gregaria. Female adults of this species are known to produce smaller but more eggs when reared in isolation than do those reared in a group. Eggs laid by isolated females develop green hatchlings typical of solitarious forms, whereas those laid by the latter produce black hatchlings typical of gregarious forms. Topical application of a juvenile hormone analog (JHA), fenoxycarb, or implantation of corpora allata (CA) taken from the migratory locust, Locusta migratoria, caused crowded S. gregaria females to deposit smaller eggs, but did not have a significant effect on the number of eggs per egg pod except at high doses of JHA. The production of smaller eggs by treated and untreated crowded females was closely associated with earlier deposition of the egg pods and shorter oviposition intervals. However, neither JHA application nor CA implantation influenced the progeny characteristics in actively reproducing aged females under crowded conditions, while untreated control females started producing smaller and more eggs upon transfer to isolated conditions. These results may suggest that JH is not directly involved in the maternal regulation of phase-dependent progeny characteristics.     

Physiological factors influencing preference for wet and dry food in Schistocerca gregaria nymphs

The physiological factors responsible for behavioral regulation of water balance by differential feeding in Schistocerca gregaria were investigated. Locusts given the option to regulate their water intake by feeding on high-water content (wet) food or low-water content (dry) food had values for percent body water and hemolymph volumes intermediate between those for locusts given either food exclusively. Locusts given wet food only prior to no-choice tests had relatively small meals and long latencies to feed on wet food and large meals and short latencies to feed on dry food. Increasing volume by adding 75 l normal saline did not alter meal size on either food, while locusts given dry food prior to testing had the opposite behavior. Increasing hemolymph concentration led to smaller meals on both wet and dry food, but diluting the hemolymph by an equal volume of distilled water resulted in larger meals on dry food and smaller meals on wet food. Removing an equal volume of hemolymph had no effect, but combining hemolymph removal with increased osmotic pressure reduced the meal size on dry food and increased meal size on wet food.

---

Résumé L'étude a porté sur les bases physiologiques de la régulation comportementale de l'équilibre hydrique par alimentation différentielle chez S. gregaria. Les teneurs en eau (pourcentage du poids du corps) et les volumes d'hémolymphe des criquets ayant eu la possibilité de moduler leur apport en eau en conssommant des aliments à forte teneur en eau (frais) ou à faible teneur (secs), étaient intermédiaires entre ceux des criquets ayant l'un ou l'autre exclusivement. Les criquets, ayant uniquement des aliments frais avant des expériences sans choix, ont présenté des temps de latence prolongés et des repas relativement brefs sur aliments frais, et des temps de latence brefs et des repas abondants sur aliments secs; les résultats ont été inversés quand l'alimentation préalable était sur nourriture sèche.Ces résultats sont partiellement simulés par des altérations du volume et de la pression osmotique de l'hémolymphe. L'augmentation de la concentration de l'hémolymphe entraîne des repas moins abondants sur aliments frais et secs, vraisemblablement par suite de signaux de satiété. L'augmentation du volume en ajoutant 75 l de solution physiologique n'a pas modifié la taille du repas sur les deux types d'aliments, mais la dilution de l'hémolymphe par un volume égal d'eau distillée a induit des repas plus importants sur aliment sec, et plus petit sur aliment frais. Une augmentation de volume plus importante, de 150 l, réduisait la taille du repas sur aliment frais, mais pas sur aliment sec. Le prélèvement d'environ 75 l d'hémolymphe a été sans effet; mais, combiné avec l'augmentation de la pression osmotique, il a réduit la taille du repas sur aliment sec (comme l'augmentation de la pression osmotique seule), mais a augmenté la taille du repas sur aliment frais. Ce dernier résultat est semblable à ceux obtenus lors des études du déterminisme de l'abreuvement. La discussion porte sur la relation entre le contrôle de l'absorption de l'eau et des aliments.

     

The trans-generational phase accumulation in the desert locust: Morphometric changes and extra molting

To understand the underlying trans-generational phase accumulation, a classical morphometric characteristic, the F/C ratio (F, hind femur length; C, maximum head width), of adult desert locusts (Schistocerca gregaria) was monitored over eight consecutive generations. Adult F/C ratios, which are larger in solitarious locusts than in gregarious ones, were negatively correlated to the darkness of body color at hatching. Two successive generations were required for a complete shift from the gregarious (crowd-reared) to the solitarious (isolated-reared) phase and vice versa in the laboratory. That is (1) female adults needed to be exposed to crowded (or isolated) conditions so that their hatchlings would become large (or small) and dark (or green) in color, and (2) the hatchlings then needed to be exposed to crowded (or isolated) conditions for their entire nymphal stage. Solitarious locusts exhibited extra molting that influenced the F/C ratio in the adult stage, but did not exert significant influences on the trans-generational changes in this trait because the incidence was low. The incidence of extra molting was negatively correlated with nymphal survival rates. The morphometric trans-generational changes may be explained without assuming any accumulating internal factor.     

A review of maternal and embryonic control of phase-dependent progeny characteristics in the desert locust

Hatchlings of the desert locust, Schistocerca gregaria, display phase polyphenism in body coloration and size. This phenomenon has been found to be maternally controlled and two different mechanisms have been proposed to explain the underlying process. One mechanism claims that a water-soluble pheromonal factor secreted by gregarious female adults into the foam plugs of egg pods induces darkening in their progeny. The other mechanism states that hatchling body coloration is pre-determined in the ovaries and that no foam factor is involved in this phenomenon. This mechanism was supported by the observation that hatchling melanization was not prevented by the early washing or separation of eggs, which should have removed the pheromonal factor from the eggs and produced green hatchlings according to the other mechanism. This paper reviews the latest findings related to this phenomenon with special reference to reproductive cycles and genetic differences. The close relationships between egg size and the degree of melanization in hatchlings may provide strong evidence against the possible involvement of the pheromonal factor, because egg size is determined in the ovarioles. Furthermore, the absence of “solitarizing” effects of early washing and separation on hatchling melanization was also confirmed in different genetic strains. A hypothesis proposed by others that such effects occur only in eggs from the first reproductive cycle was tested and rejected. Based on these and other results, a model to explain the mechanisms underlying the maternal control of progeny characteristics and embryonic control of melanization in the hatchling was proposed.     

Vitellogenin Titres in Normal and Accelerated Maturation of Gregarious-Phase Schistocerca gregaria

Isolation of vitellogenin of the Schistocerca gregaria (Forskal) in its gregarious phase was achieved by a combination of gel permeation and anion exchange chromatography. Staining for carbohydrate and lipid moieties showed that the vitellogenin is a glycolipoprotein. The vitellogenin of S. gregaria has a native molecular weight of about 700 kDa. On SDS-PAGE, the protein showed nine apoproteins of about 124, 120, 105, 60, 59, 58, 57, 53 and 34 kD. Determination of the levels of vitellogenin by ELISA in the haemolymph of maturing females showed that those exposed to mature males from 1 to 2 days after ecdysis had increased levels of vitellogenin from day 10 (81.1 ± 4.5). In contrast, females exposed to immature males or kept alone showed an increase (107.3 ± 0.9 and 70.2 ± 2.7) not until day 16 or later, respectively. These results are consistent with the accelerating effect of pheromonal emissions from mature males on the maturation of female S. gregaria.     

Feeding behavior: selection of both wet and dry food for increased growth in Schistocerca gregaria nymphs

An apparent fluctuating preference in acridids for food of different water contents is confirmed for Schistocerca gregaria nymphs. Short- and long-term preference tests reveal that previous diet, through its effect on locust water balance and overall food intake, can influence feeding behavior on high-water content (wet) and low-water content (dry) food. Locusts given only wet or dry food for 3 days prior to testing initially chose the food not received in pretreatment when given a choice of the two. They had decreased latencies to feed, increased meal length and decreased switching between food types as compared with locusts given the option to regulate water intake before testing. It is suggested that the behavior of the single-diet locusts has several related causes, among them, deterrence of the food, disturbed water balance in the locust, and chronic hunger. Long-term preference tests reveal that fifth-instar locusts given a continuous choice between wet and dry food take significantly more of their total dry matter from dry food each day until the last feeding day of the instar (day 9), when they take 97% of their total dry matter from wet food. This behavioral regulation of water and dry matter intake is adaptive: locusts with access to a mixed diet of wet and dry food have increased growth as compared with ones given only wet food. The growth increase is due largely to increased dry matter consumption, with some contribution from increased digestive efficiency and reduction in respiratory rate.

---

Résumé L'étude de larves de dernier stade de Schistocerca gregaria confirme les observations sur les acridiens d'une préférence apparemment labile pour les aliments à différentes teneurs en eau. Des observations sur les choix à court et à long termes rélèvent que le régime alimentaire antérieur, par son action sur le régime hydrique du criquet et sur sa prise totale de nourriture, peut influer sur le comportement alimentaire face à des aliments à forte teneur en eau (humides ou frais) ou à faible teneur (secs). Des criquets, approvisionnés uniquement en aliments secs ou frais pendant les 3 jours précédant l'expérience, choisissent de préférence la catégorie d'aliments non fournie pendant ce prétraitement. Par comparaison avec des criquets ayant eu la possibilité d'ajuster leur apport en eau avant l'expérience, ils présentaient une diminution du temps de latence avant de s'alimenter, une prolongation de la durée des repas et une diminution du nombre de changements de type d'aliments. II est suggéré que le comportement des criquets, à régime alimentaire composé d'un seul type, a plusieurs causes associées, en particulier la répulsion pour le végétal, la perturbation de l'état hydrique du criquet, et la faim chronique. Des tests sur les préférences à long terme montrent que des larves de 5ème stade ayant eu un choix continu entre aliments frais et secs prirent significativement plus de leur matière totale d'aliments secs jusqu'à ce qu'elles parviennent au dernier jour d'alimentation du stade (jour 9), alors elle prélevèrent 97% de leur matière sèche totale d'aliments frais. Cette régulation comportementale de l'absorption d'eau et de matière sèche est adaptative: des criquets ayant accès à une alimentation mixte (nourriture sèche et fraîche) ont unt croissance supérieure à ceux nourris exclusivement d'aliments frais. La croissance supérieure est due pour beaucoup à une consommation accrue de matière sèche, avec une légère contribution de l'amélioration de l'efficacité digestive et de la réduction du taux de respiration.

     

Food mixing strategies in the desert locust: effects of phase, distance between foods, and food nutrient content

Food mixing strategies were compared in the cryptically coloured, relatively sedentary `solitarious' and the highly mobile, conspicuously coloured `gregarious' phases of the desert locust, Schistocerca gregaria. Based on phase related differences in behaviour and nutritional regulatory responses, we predicted that solitarious nymphs, compared to gregarious nymphs, would move less between nutritionally complementary foods, particularly as the distance between the foods increased. We manipulated the nutritional composition [protein (p) and digestible carbohydrate (c) content] of two foods in an experimental arena and varied the distance between the foods using a factorial experimental design. Results indicated that in general, solitarious nymphs showed greater fidelity to individual food dishes than did gregarious insects (i.e., they concentrated their feeding mainly on one dish). However, results also demonstrated that for both phases fidelity to a particular food dish increased as the distance between the dishes increased, and that the number of switches between dishes decreased with increasing distance. In the smallest arenas, though, gregarious nymphs switched more frequently between the two food dishes than solitarious nymphs, even when the two dishes contained the same, near-optimal food (p18:c24). When challenged by having the two dishes either placed furthest apart (2 m) or more divergent in nutritional composition (p29:c13 vs. p7:c35), insects of both phases regulated protein intake more strongly than carbohydrate intake, by eating more from the dish containing higher-protein food.     

Quantitative studies on anti-allatin and lethal effects of a synthetic precocene in different larval instars of the desert locust

The lethal and anti-allatin effects of various doses of 7-ethoxy-precocene II (PIII), topically applied to crowdedSchistocerca gregaria nymphs, 0–27 h after the previous moult, were investigated for the five nymphal instars. The LD (lethal dose) probit lines were parallel for the different instars. The ED (effective anti-allatin dose) probit lines were also parallel. But for each instar the LD and ED probit lines seemed to be convergent toward the higher doses. The LD₅₀ and the ED₅₀ (g PIII/g fresh weight) were not widely separated; the LD₅₀/ED₅₀ ratio was instar dependent and varied between 1.26 (Vth instar) and 5.35 (IIIrd instar). Thus, separation between lethal effect, anti-allatin effect and no effect was rather poor; indeed in many instances all these types of results were obtained in a group of hoppers treated with the same dose in the same instar. A low precocene susceptibility was observed for the Ist instar and for each subsequent instar the susceptibility further decreased; the LD₅₀ and the ED₅₀ were respectively 886 g/g and 328 g/g for the Ist instar, but increased to 14 307 g/g and 11 367 g/g for the Vth instar. The anti-allatin effect of the precocene induced various morphogenetic responses: (1) permanent adultiform in the instar next to that treated; (2) death in the moult from the next to the second consecutive instar; (3) permanent adultiform in the second consecutive instar. The occurrence and/or distribution of these responses were instar dependent and for the first three instars also limitedly dose dependent. The anti-allatin effect of the precocene applied to Vth (last) instar nymphs became overt by sterility and lack of yellow colour in the adult. An overt anti-allatin effect was always coupled with a response dependent increase of the duration of the instar treated and/or of the next instar.

---

Etudes quantitatives des effets létaux ou antiallates d'un précocène de synthèse sur différents stades larvaires du criquet pélerin

Résumé L'étude a porté sur les effets létaux ou antiallates de différentes doses de 7-éthoxy-précocène II (PIII), déposées de 0 à 27 heures après la mue sur la surface dorsale de l'abdomen de larves groupées de chacun des 5 stades larvaires deSchistocerca gregaria. Les courbes des doses létales (LD) ont été parallèles pour les différents stades. Les courbes pour les doses antiallates (ED) étaient elles aussi parallèles. Cependent pour chaque stade, ces deux courbes semblaient converger et même se croiser vers les doses les plus fortes. Les LD₅₀ et ED₅₀ (en g de PIII par gramme de poids frais) n'étaient pas nettement séparées; le rapport LD₅₀/ED₅₀ dépendait du stade et variait entre 1,26 (5è stade) et 5,35 (3è stade). Ainsi la distinction entre effet létal, effet antiallate et absence d'effet était plutôt faible; à la vérité, dans de nombreux cas, tous ces effets ont été obtenus dans un groupe de criquets traités avec la même dose au même stade. Une sensibilité assez faible au précocène a été observée, même au ler stade, et pendant chaque stade suivant la sensibilité ultérieure diminuait; LD₅₀ et ED₅₀ ont été respectivement 886 g/g et 328 g/g pour le ler stade, mais ont augmenté à 14 307 g/g et 11 367 g/g pour le 5è stade. Les effets antiallates ont induit différentes conséquences morphologiques: (1) des adultiformes permanents lors du stade suivant immédiatement le stade traité; (2) mort pendant la mue du ler ou 2nd stade suivant le traitement; (3) des adultiformes permanents dans le 2nd stade après le traitement. L'apparition et/ou la distribution de ces réponses dépendaient du stade, et pour les 3 premiers stades étaient partiellement dépendantes de la dose. L'effet antiallate du précocène au 5è stade (dernier) devenait évident avec la stérilité et la perte de la coloration jaune de l'adulte. Un effet antiallate manifeste était toujours associé à un accroissement d dépendant de la dose de la durée du stade traité et/ou du stade suivant.

     

Effects of hatchling body colour and rearing density on body colouration in last-stadium nymphs of the desert locust, Schistocerca gregaria

Abstract.  The influences of hatchling character and rearing density on body colour at the last-nymphal stadium are investigated for the desert locust, Schistocerca gregaria . Hatchlings are divided into five groups based on the darkness of the body colour and reared either under isolated or crowded conditions. Two types of body colour variation at the last-nymphal stadium are separately analysed (i.e. the background colour and black patterns). Under isolated conditions, the background body colour is either greenish or brownish. Most individuals are greenish and the highest percentage of brownish insects is obtained from hatchlings with the darkest body colour. Under crowded conditions, the background colour is yellow or orange and the percentage of yellowish nymphs tends to decrease when they are darker at hatching. The intensity of black patterns differs depending on the body colour at hatching and subsequent rearing density. Most isolated-reared nymphs exhibit few or no black patterns but nymphs with some black patterns also appear, particularly among those that had been dark at hatching. Under crowded conditions, the black patterns become more intense when they are darker at hatching. Therefore, last-stadium nymphs with typical solitarious or gregarious body colouration appear when they have the phase-specific body colouration at hatching as well. The present results demonstrate that both body colour at hatching and rearing density during nymphal development influence body colouration at the last-nymphal stadium.     

Hormonal control of phase-related changes in the number of antennal sensilla in the desert locust, Schistocerca gregaria: possible involvement of [His7]-corazonin

The effect of [His(7)]-corazonin on the abundance of antennal sensilla in the desert locust, Schistocerca gregaria, was investigated to test the hypothesis that injection of this neuropeptide would mimic a crowding effect. Solitarious locusts (reared in isolation) were injected with [His(7)]-corazonin at the 3rd nymphal instar and the numbers of sensilla on the 2nd, 8th and 14th antennal segments in the adult stage were compared with those for oil-injected solitarious controls or un-injected gregarious locusts (reared in group). The numbers of sensilla on these antennal segments were all reduced significantly after [His(7)]-corazonin injection compared with those for oil-injected controls, but similar to the values for gregarious individuals. Among the four major types of olfactory sensilla, coeloconic, trichoid, basiconic type A and basiconic type B, [His(7)]-corazonin injection influenced the abundance of all but the last type. The effect of [His(7)]-corazonin injection varied with the time of injection; the earlier the injection the larger the effects on the abundance of total antennal sensilla on the 8th segment, although the way in which the injection affected the abundance varied with the sensillum type. A hypothesis explaining how crowding affects the abundance of antennal sensilla and other phase-related characteristics through changes in [His(7)]-corazonin concentrations was proposed.     

Autoradiographic localization of [22,23-(3)H(2)] dihydroazadirachtin binding sites in desert locust testes and effects of Azadirachtin on sperm motility

Localization of [22,23-(3)H(2)] dihydroazadirachtin binding sites in locust (Schistocerca gregaria) testes was investigated by in vitro autoradiography. Preferential binding of the ligand at concentrations of 10(-8) M was located in the testes follicles, localized on the tail portions of developing sperm. This binding was fully displaceable with an excess of unlabelled ligand. The results indicate that Azadirachtin binds preferentially to sites on the organelles associated with maturing locust sperm tails. Azadirachtin, at concentrations of 10(-4)M and above, caused a time-dependent reduction in the motility of eupyrene sperm bundles liberated from the accessory glands of mature male S. gregaria. The effect of azadirachtin on boar sperm motility was also time- and concentratiion-dependent. Forward motility was significantly reduced by concentrations above 10(-5)M at each time interval (P<0.05).     

Irrigation and fertilization effects on seed number,size, germination and seedling growth: implications for desert shrub establishment

Plants with limited resources adjust partitioning among growth, survival, and reproduction. We tested the effects of water and nutrient amendments on seed production, size, and quality in Sarcobatus vermiculatus (greasewood) to assess the magnitude and importance of changes in reproductive partitioning. In addition, we assessed interactions among the environment of seed-producing plants (adult plant scale), seed size, and seedling microenvironment (seedling scale) on successful seedling establishment. Interactions of these factors determine the scale of resource heterogeneity that affects seedling establishment in deserts. Both total number of seeds produced per plant and seed quality (weight and germination) increased significantly in the enriched treatment in a 3-year field experiment. Seedling length 3 days after germination and seed N concentration, other measures of seed quality, were higher for seed from both irrigated and enriched plants than for seed from control plants. Field S. vermiculatus seed production and quality can be substantially increased with irrigation and nutrient enrichment at the adult plant scale and this allows management of seed availability for restoration. However, based on a greenhouse study, seedling environment, not the environment of the seed-producing plant or seed size, was the most important factor affecting seedling germination, survival, and growth. Thus it appears that production of more seed may be more important than improved seed quality, because higher quality seed did not compensate for a low-resource seedling environment. For both natural establishment and restoration this suggests that heterogeneity at the scale of seedling microsites, perhaps combined with fertilization of adult shrubs (or multi-plant patches), would produce the greatest benefit for establishing seedlings in the field.