body ATP-sensitive potassium channels [6]. This triggers AKH release in a calcium-and AMP-activated protein kinase-dependent fashion [13]. Similar to glucagon, AKH acts on energy stores in the fat body by mobilising carbohydrates in form of the haemolymph sugar trehalose (hypertrehalosaemic effect), and by mobilising lipid depots (adipokinetic effect) [8,[14][15][16].OA, an analogue to vertebrate noradrenaline, is considered to act as an insect stress or arousal hormone [17]. OA is produced by octopaminergic neurons (OANs) located mostly along the midline in the brain and ventral nerve cord [18]. Among other functions, OA exerts myotropic effects on skeletal muscles [19] and mediates the adaptation of muscle metabolism to the physiological demands of locomotion [20][21][22].Both AKH and OA are crucial for starvation-induced hyperactivity, as this behaviour is absent in flies with impaired AKH or OA signalling [4,5,[7][8][9]. AKH activates a subset of OANs which are required for starvation-induced hyperactivity [5,9]. These neurons express receptors for AKH and Drosophila insulin-like peptides (DILPs). DILPs act antagonistically to AKH, and suppress starvation-induced hyperactivity via the OANs [9]. Yu and colleagues (2016) showed that the AKH-dependent increase in locomotor activity upon starvation represents a neuromodulatory and behavioural role of AKH and is not an indirect consequence of altered energy metabolism.Remarkably, studies in various insect species suggest that AKH and also OA play a role in the regulation of locomotor activity when the insects have free access to food (reviewed in [23]). For example, the circulating haemolymph titres of AKH correlate positively with daily rhythms in walking activity in the linden bug Pyrrhocoris apterus [24,25]. In Pyrrhocoris [26,27], crickets [28] and the cockroach Periplaneta americana [29,30], injection into the haemolymph or topical application of AKH induces a strong increase in locomotor activity in ad libitum fed animals, independent of the time of day [27,29]. Also in Drosophila, genetic overexpression of AKH enhances locomotor activity when food is freely available [31]. Unlike for starvation-induced hyperactivity, however, it is unclear whether and how AKH:OAN signalling contributes to shape daily activity under ad libitum feeding conditions, or whether increased AKH signalling is a consequence of a locomotor-induced increase in energy demands. Cockroach AKHs are known to increase the spontaneous neuronal activity of octopaminergic dorsal unpaired median neurons in the central nervous system, which express a