Yudai Nishide

Behavioral phase shift in nymphs of the desert locust, Schistocerca gregaria: Special attention to attraction/avoidance behaviors and the role of serotonin

Schistocerca gregaria exhibits a phase-specific behavior in response to crowding. Nymphs occurring at low population densities (solitarious phase) tend to avoid one another, whereas those occurring at high population densities (gregarious phase) are attracted to one another. This study examined how this attraction/avoidance behavior changed after isolation or crowding. The behavior of the test nymphs was assessed by determining their positioning with respect to a stimulus cup, which contained 12 gregarious nymphs, placed at one end of an elliptical arena and an empty cup placed at the opposite end. Gregarious (crowd-reared) nymphs were most frequently observed close to the stimulus cup, whereas solitarious (isolated-reared) nymphs tended to avoid it. This tendency was easily changed by exposing the nymphs to isolation or crowding. However, contrary to a previously reported conclusion that behavioral gregarization occurs in 4-8 h of crowding, the nymphs required at least 3 days to achieve a significant change in the attraction/avoidance behavior in either direction, from solitarious to gregarious or the converse. The discrepancies between the present study and previous studies appear to result from the different behaviors observed. The present study focused on the attraction/avoidance behavior that is most likely the most important element in the gregarization and solitarization processes, whereas other studies used a mixture of various activity-related behaviors. No evidence was obtained for a trans-generational accumulation in the attraction/avoidance behaviors. Serotonin, a biogenic amine that is suggested by others to be involved in behavioral phase changes in S. gregaria, had no influence on this behavior or body color when injected into the solitarious nymphs.

Re-examination of the roles of environmental factors in the control of body-color polyphenism in solitarious nymphs of the desert locust Schistocerca gregaria with special reference to substrate color and humidity.

This study re-examines the effects of environmental factors including substrate color, humidity, food quality, light intensity and temperature on the green-brown polyphenism, black patterning and background body color of solitarious (isolated-reared) nymphs of Schistocerca gregaria. All individuals reared in yellow-green or yellow containers became green morphs, whereas those reared in white, ivory-colored, blue, grey, brown, zinc-colored and black containers produced brown morphs in similar proportions. The intensity of black patterns was negatively correlated with the brightness of the substrate color of the containers. Humidity, which previous studies claimed controls green-brown polyphenism in this species, exerted no significant influence on either the green-brown polyphenism or the black patterning. Food quality also had little effect on body color. High temperature tended to inhibit darkening. The background body color on the thorax was greatly influenced by the substrate color of rearing containers and a close correlation was found between these two variables, indicating that, in contrast to what has been suggested by others, this species exhibits homochromy to match the body color to the substrate color of its habitat. Similar responses were observed in another strain, although some quantitative differences occurred between the two strains examined. Based on these results, a new model explaining the control of body-color polyphenism in this locust is proposed and the ecological significance of black patterns in solitarious nymphs is discussed.

Do desert locust hoppers develop gregarious characteristics by watching a video

Various sensory stimuli have been suggested to induce gregarious body coloration in locusts, but most previous studies ignored the importance of substrate color. This study tested the effects of visual, olfactory and tactile stimuli from other locusts on the induction of gregarious body coloration in single (isolated-reared) Schistocerca gregaria nymphs housed in yellow-green cups. Odor from gregarious (crowd-reared) locusts, which is believed to induce black patterns in single locusts, had little effect when applied to visually isolated nymphs at the 2nd stadium onward, and all test nymphs remained green without black patterns at the last stadium, as in controls reared without odor and visual stimuli. Visual stimuli alone induced black patterns when a single solitarious nymph was allowed to see other locusts in another cup. The degree of black patterns increased as the number of locusts shown increased, and some test nymphs developed body coloration typically observed in gregarious forms. A classical morphometric ratio (hind femur length/head width) shifted toward the value typical of gregarious forms when the single nymphs were allowed to see 5 or 10 locusts. Single nymphs also developed black patterns when presented green conspecific nymphs and adults of two hemipteran species kept in another cup. No synergetic effects of visual and odor stimuli were detected. Movies of locusts, crickets and tadpoles were found effective in inducing black patterns in single locusts. Ontogenetic variation in the sensitivity to crowding and experimental methodology might be responsible for some discrepancies in the conclusions among different researchers.

Desert locust, Schistocerca gregaria , eggs hatch in synchrony in a mass but not when separated

This laboratory study examined the stimuli involved in controlling egg hatching in the desert locust Schistocerca gregaria. We found that eggs that were kept in their egg pods hatched in synchrony, whereas those that were kept individually hatched over an extended period of time and took significantly longer to hatch. Synchronous hatching was also observed in pairs of eggs that were kept in contact with each other in the same plastic wells, but not in those that were kept individually in separate wells. This demonstrates that the eggs needed to be placed in contact with each other to achieve synchronous hatching and suggests that a hatching individual stimulates adjacent eggs to hatch via some unknown stimulus. Additional experiments suggested that neither auditory nor pheromonal stimuli were important in this process, but rather some sort of vibratory stimulus. This inference was supported by the results of another experiment, which demonstrated that when isolated eggs were vibrated with a vortex mixer, most of them hatched shortly after the stimulation. Our results suggest the possibility that direct mechanical stimulation caused by the first egg to hatch stimulates adjacent eggs to also hatch, thus producing synchronous hatching in the entire egg pod. However, the exact nature of this stimulus remains to be determined.Significance statementSynchronous hatching is often observed in insects. It helps hatchlings to form aggregations and to reduce the risk of predation and cannibalism. Yet, the mechanism controlling this phenomenon has received little attention except for a few subsocial species. This study demonstrates that eggs kept in their egg pods hatched in synchrony, whereas those kept individually in sand hatched over an extended period of time and took longer to hatch. Synchronous hatching was also observed in pairs of eggs that were kept in contact with each other but not placed some distance apart from each other. We also show that artificial vibration induced hatching.

First Record of the Occurrence and Genetics of a Short-Winged Morph in the Migratory Locust, Locusta migratoria (Orthoptera: Acrididae)

Abstract Short-winged adults appeared in the 2nd generation of inbred colonies of migratory locusts, Locusta migratoria, originating from Tsushima Island, Japan. Of 14 family lines, two produced several short-winged adults in the 2nd generation. These short-winged adults also had shorter hind femora than long-winged adults, indicating they may represent ‘the short-winged morph’ rather than monsters with abnormally short wings. The two wing morphs could be separated by the ratio of forewing length to head width. Crosses between the two wing morphs revealed that the short-winged morph was controlled by a simple recessive Mendelian unit. It is unknown whether the short-winged morph occurs in the field.

Body‐color and behavioral responses by the mid‐instar nymphs of the desert locust, Schistocerca gregaria (Orthoptera: Acrididae) to crowding and visual stimuli

The effects of crowding and isolation on body color and behavior were observed for the mid‐instar nymphs of the desert locust, Schistocerca gregaria. Some of the solitarious (isolation‐reared) nymphs that were crowded for 1 or 4 h during the third instar developed black patterns at the fourth instar, but most individuals remained unaffected. Black patterns appeared in all individuals that were crowded for 1 day or longer, but even after 4 days of crowding the black patterning for some individuals was not as intense as that for the gregarious (crowd‐reared) controls. Isolation of gregarious nymphs caused the black patterns to recede or disappear at the last (fifth) nymphal instar, but it was necessary to isolate the nymphs from the beginning of the first instar to obtain body coloration looking like solitarious nymphs in most individuals. Solitarious nymphs that were allowed to see gregarious nymphs developed different intensities of black patterns depending on the body size and number of nymphs shown. The behavioral phase shift from one phase to another was observed when the nymphs were crowded or isolated for 2 days or longer, as previously reported for the last nymphal instars of the same strain. Behavioral gregarization was induced for isolated nymphs that were allowed to see a group of nymphs through a transparent double wall. These results suggested that body‐color phase shift occurred more rapidly for mid‐instar nymphs than for late instar nymphs but the rate of behavioral phase shift was similar for the two instars.

The mechanism controlling phenotypic plasticity of body color in the desert locust: some recent progress

Schistocerca gregaria exhibits density-dependent body color polyphenism. Nymphs occurring at low population densities show green-brown polyphenism. They show phase polyphenism and develop black patterns at high population densities. Recent studies suggest a third type of polyphensim, that is, homochromy, a response to background color. Laboratory experiments that considered homochromy suggest that humidity is not directly involved in green-brown polyphenism and that odor from other individuals does not induce black patterns. Black patterns can be induced in isolated nymphs by video images of locusts and tadpoles. Juvenile hormone and [His7]-corazonin control body color in locusts. The gene encoding the latter has been identified for S. gregaria and Locusta migratoria, and its key role in controlling black patterning has been demonstrated.

Wing dimorphism in the migratory locust, Locusta migratoria: differentiation of wing morph and phase polyphenism

A short‐winged morph was recently discovered in the migratory locust, Locusta migratoria. It is different from the normal, long‐winged morph not only in forewing length but also in hind femur length, displaying a dimorphism. To understand the significance of this dimorphism, other morphological characters were compared between the two morphs, and the time of differentiation of wing‐pad length was investigated. Wing weights were heavier in the long‐winged morph than in the short‐winged morph. This result showed that the short‐winged morph is not formed by a failure of wing expansion. No obvious morph‐specific differences were observed in wing venation, but wing allometry studies indicated that the distal areas of the fore‐ and hindwings were disproportionally reduced in the short‐winged morph compared to the long‐winged morph. The morphological differentiation of the wing pad between the two morphs was observed at the penultimate nymphal stage. The flight muscle was well developed in the two morphs, and no sign of flight muscle histolysis was detected in either morph after adult emergence. An analysis of adult body dimensions suggested that the density‐dependent phase shifts known for the long‐winged morph of this locust were also exhibited by the short‐winged morph, demonstrating that these shifts are not specific to the migratory long‐winged morph.

Yellowing, morphology and behaviour in sexually mature gynandromorphs of the desert locust Schistocerca gregaria

The morphology, colouration and sexual behaviour of two gynandromorph adults of the desert locust Schistocerca gregaria Forskål are described. In each of these specimens, the left half of the external genitalia has male characteristics and the right half has female characteristics. Yellowing of the epidermis occurs to different degrees in normal sexually mature male and female adult S. gregaria under crowded conditions. Two hypotheses are known to explain this phenomenon. One suggests involvement of a sex‐hormone/receptor complex, whereas the other proposes different sensitivities of the epidermis to the same hormonal environment. To examine which hypothesis is more likely, one gynandromorph adult is kept under crowded conditions and the other is maintained under isolated conditions. In the former, the left half of the body turns bright yellow, as in a sexually mature crowded male, and the right half is brownish with little yellowing, as in a sexually mature crowded female of the same age. The bilaterally divided body colouration may support the second hypothesis. In the second gynandromorph, which is isolated after adult emergence, the body colour remains brownish. Upon sexual maturity, this individual is kept together with normal male or female adults for behavioural observations. It attempts to mount a female but is approached and mounted by males.

The hatching time of Locusta migratoria under outdoor conditions: role of temperature and adaptive significance: Locust hatching under outdoor conditions

The present study investigates the time of hatching of the migratory locust Locusta migratoria using egg pods that are artificially buried in the soil under outdoor conditions. Most eggs hatch in the mid‐morning, with a peak between 11.00 and 12.00 h, and none hatch before 09.00 or after 16.00 h. Furthermore, most egg pods complete hatching within a day, although some take 2 or 3 days, and egg hatching is interrupted by rain. There are no large differences in hatching time from May to September. Laboratory experiments in which the eggs are exposed to temperatures simulating outdoor conditions show that soil temperature is the main factor controlling hatching activity. The increase in temperature in the morning appears to trigger egg hatching, as confirmed by laboratory experiments, which may explain the similar hatching times between seasons. The seasonal patterns of temperature variation and hatching time suggest that the hatching time of L. migratoria eggs may be adjusted to allow the hatchlings to be exposed to high temperatures in the afternoon so that they can harden their bodies quickly.