SUMMARY: Desert locusts (Schistocerca gregaria) can undergo a profound transformation between solitarious and gregarious forms, which involves widespread changes in behaviour, physiology and morphology. This phase change is triggered by the presence or absence of other locusts and occurs over a timescale ranging from hours, for some behaviours to change, to generations,for full morphological transformation. The neuro-hormonal mechanisms that drive and accompany phase change in either direction remain unknown. We have used high-performance liquid chromatography (HPLC) to compare amounts of 13 different potential neurotransmitters and/or neuromodulators in the central nervous systems of final instar locust nymphs undergoing phase transition and between long-term solitarious and gregarious adults. Long-term gregarious and solitarious locust nymphs differed in 11 of the 13 substances analysed: eight increased in both the brain and thoracic nerve cord (including glutamate,GABA, dopamine and serotonin), whereas three decreased (acetylcholine,tyramine and citrulline). Adult locusts of both extreme phases were similarly different. Isolating larval gregarious locusts led to rapid changes in seven chemicals equal to or even exceeding the differences seen between long-term solitarious and gregarious animals. Crowding larval solitarious locusts led to rapid changes in six chemicals towards gregarious values within the first 4 h(by which time gregarious behaviours are already being expressed), before returning to nearer long-term solitarious values 24 h later. Serotonin in the thoracic ganglia, however, did not follow this trend, but showed a ninefold increase after a 4 h period of crowding. After crowding solitarious nymphs for a whole larval stadium, the amounts of all chemicals, except octopamine, were similar to those of long-term gregarious locusts. Our data show that changes in levels of neuroactive substances are widespread in the central nervous system and reflect the time course of behavioural and physiological phase change.

In social Hymenoptera with no morphological caste, a dominant female becomes an egg layer, whereas subordinates become sterile helpers. The physiological mechanism that links dominance rank and fecundity is an essential part of the emergence of sterile females, which reflects the primitive phase of eusociality. Recent studies suggest that brain biogenic amines are correlated with the ranks in dominance hierarchy. However, the actual causality between aminergic systems and phenotype (i.e. fecundity and aggressiveness) is largely unknown due to the pleiotropic functions of amines (e.g. age-dependent polyethism) and the scarcity of manipulation experiments. To clarify the causality among dominance ranks, amine levels and phenotypes, we examined the dynamics of the aminergic system during the ontogeny of dominance hierarchy in the queenless ant Diacamma sp., which undergoes rapid physiological differentiation based on dominance interactions. Brain dopamine levels differed between dominants and subordinates at day 7 after eclosion, although they did not differ at day 1, reflecting fecundity but not aggressiveness. Topical applications of dopamine to the subordinate workers induced oocyte growth but did not induce aggressiveness, suggesting the gonadotropic effect of dopamine. Additionally, dopamine receptor transcripts (dopr1 and dopr2) were elevated in the gaster fat body of dominant females, suggesting that the fat body is a potential target of neurohormonal dopamine. Based on this evidence, we suggest that brain dopamine levels are elevated in dominants as a result of hierarchy formation, and differences in dopamine levels cause the reproductive differentiation, probably via stimulation of the fat body.