Flight capability and flight motor pattern in a sedentary South African grasshopper, Phymateus morbillosus– a comparison with migratory species

Abstract Among the Acridoidea, not all species are strong fliers. We have examined the possible causes for loss of flight in a species with a reduced flight system, the South African grasshopper, Phymateus morbillosus (L.). This is a sedentary species that, in the field, displays only marginal flight (in males) or no flight (in females). In a wind tunnel, however, this species can be stimulated to perform flight movements for a short time. In the present study, several morphometric parameters and aspects of the flight motor output have been examined. The data are compared with those of the migratory locust species ( Locusta migratoria L. and Schistocerca gregaria Forsk.). Phymateus morbillosus can exhibit the typical flight motor pattern for short periods of up to 1 min. Morphometric data and wing-beat frequency in P. morbillosus are similar to that of other members of this insect group. However, female specimens of P. morbillosus are too heavy to lift themselves for active flight. We assume that females of this species invest in the augmentation of reproduction rather than investing in the flight system.     

Postembryonic development of centrally generated flight motor patterns in the hawkmoth, Manduca sexta

This study analyses the maturation of centrally generated flight motor patterns during metamorphosis of Manduca sexta. Bath application of the octopamine agonist chlordimeform to the isolated central nervous system of adult moths reliably induces fictive flight patterns in wing depressor and elevator motoneurons. Pattern maturation is investigated by chlordimeform application at different developmental stages. Chlordimeform also induces motor patterns in larval ganglia, which differ from fictive flight, indicating that in larvae and adults, octopamine affects different networks. First changes in motoneuron activity occur at the pupal stage P10. Rhythmic motor output is induced in depressor, but not in elevator motoneurons at P12. Adult-like fictive flight activity in motoneurons is observed at P16 and increases in speed and precision until emergence 2 days later. Pharmacological block of chloride channels with picrotoxin also induces fictive flight in adults, suggesting that the pattern-generating network can be activated by the removal of inhibition, and that proper network function does not rely on GABA_A receptors. Our results suggest that the flight pattern-generating network becomes gradually established between P12 and P16, and is further refined until adulthood. These findings are discussed in the context of known physiological and structural CNS development during Manduca metamorphosis.     

A comparative study of motor patterns during pre-flight warm-up in hawkmoths

Summary Temporal patterns of activation of flight muscles were recorded by means of wires placed extracellularly in thoracic muscles. In the five species of hawkmoths studied, wingstrokes of small amplitude were produced during a preflight warm-up by synchronous contractions of certain groups of muscles which are antagonists in flight. The main depressor muscle, the dorsal longitudinal, was excited in synchrony with some or all of the indirect elevator muscles. Three direct muscles, the subalar, basalar and third axillary muscles, were usually excited out of phase with the dorsal longitudinal muscle. However, details of the motor pattern varied from species to species. During fixed flight phase changes comparable in magnitude to those which occur during the transition from warm-up to flight were observed in Manduca sexta and Smerinthus cerisyi. The results (summarized in Table 2) suggest that a variety of warm-up patterns evolved within the Sphingidae as modifications of a common mechanism generating flight motor patterns.I thank Dr. Harry Lange for assistance in the initial collecting of Manduca sexta and for identifying specimens of this species.     

Octopamine and chlordimeform enhance sensory responsiveness and production of the flight motor pattern in developing and adult moths

Octopamine and an agonist, chlordimeform, increase the responsiveness of adult and pharate adult Manduca sexta to gentle mechanical stimulation of the wing. Higher doses of chlordimeform elicit almost continuous production of the flight motor pattern in both adults and pharate adults, and the effect persists for more than 24 h. The dose of chlordimeform necessary for this effect increases with age. Mechanical stimulation of the wing of pharate adults elicits several cycles of flight motor pattern, but with repeated stimulation the animal habituates. Habituation is slower in chlordimeform-treated animals than in controls. Injection of octopamine (1–8 × 10^?8 mol) or chlordimeform (3 × 10^?9 mol) into the mesothoracic ganglion elicits the flight motor pattern. The excitatory actions of both compounds can be blocked by cyproheptidine. Chlordimeform (5 × 10^?8 mol) in acetone applied to the wing does not cause a noticeably greater increase in the electrical activity of sensory neurons than does acetone applied alone; this result suggests that chlordimeform does not act on these peripheral sites or on axonal membranes in general. We suggest that chlordimeform and octapamine act on the thoracic ganglia to alter the level of excitation on reffectiveness of synaptic transmission among central neurons, including those involved in producing the flight motor pattern.     

Flight motor patterns recorded in surgically isolated sections of the ventral nerve cord ofLocusta migratoria

Summary In the locust,Locusta migratoria, the pairs of connectives between the three thoracic ganglia and in the neck were transected in all possible combinations. Each of these preparations was tested for the production of rhythmic flight motor activity, with sensory input from the wing receptors intact and after deafferentation. The motor activity elicited in these preparations was characterized by intracellular recordings from motoneurons and electromyographic analyses.The motor patterns observed in locusts with either the neck or the pro-mesothoracic connectives severed (Figs. 2, 3, and 4) were very similar to the flight motor pattern produced by animals with intact connectives. The activity recorded in mesothoracic flight motoneurons of locusts with either only the meso-metathoracic connectives cut or both the meso-metathoracic and the neck connectives transected were similar to each other. Rhythmic motor activity could be observed in these preparations only as long as sensory feedback from the wing receptors was intact. These patterns were significantly different from the intact motor pattern (Figs. 5, 6, and 7). Similar results were obtained when the mesothoracic ganglion was isolated from the other two thoracic ganglia, although the oscillations produced under these conditions were weak (Fig. 8 upper). In the isolated metathorax no rhythmic flight motor activity could be recorded (Fig. 8 lower), even when wing afferents were intact.Considering the differences between the motor patterns observed in the various preparations these results suggest that the ganglia of the locust ventral nerve cord do not contain segmental, homologous flight oscillators which are coupled to produce the intact flight rhythm. Instead they support the idea that the functional flight oscillator network is distributed throughout the thoracic ganglia (Robertson and Pearson 1984). The results also provide further evidence that sensory feedback from the wing sense organs is necessary for establishing the correct motor pattern in the intact animal (Wendler 1974, 1983; Pearson 1985; Wolf and Pearson 1987 a).Abbreviations CPG central pattern generator - EMG electromyogram     

Dipteran flight motor pattern: Invariabilities and changes during postlarval development

For Calliphora the wingbeat frequency and the underlying motoneuronal activity were recorded during adult life. Wingbeat frequency increases during the ten days following last molt. The activity of motoneurons serving four selected flight muscles (nonfibrillar and fibrillar ones) also increases with age. The motoneuronal activity of young and old flies was analyzed statistically (serial and cross-correlograms, latency and phase histograms). In addition, several wing manipulations were carried out to evaluate the significance of sensory feedback on pattern generation during maturation. These ontogenetic studies suggest a centrally generated motor pattern that (1) is essentially complete with the molt to adulthood, (2) shows a progressive increase in intrinsic activity, and (3) is modulated by sensory feedback from the wing region by the same amount irrespective of age. Similarities in the postlarval development of the flight pattern of neurogenic and myogenic flyers are discussed.     

Maturation of the flight motor pattern without movement inManduca sexta

The development of the flight motor pattern was studied by recording acutely with fine wire electrodes inserted in the thoracic muscles of pharate moths of known age and by recording chronically for up to 8 days with implanted electrodes. Externally visible morphological characteristics by which the age of a pharateManduca sexta can be established were identified (Table 1). Bouts of activity lasting approximately 30 min to 2 h and alternating with inactive periods of similar duration were recorded as early as the ninth day after pupation and on all successive days until early on the day of eclosion, typically 19 days after pupation (Figs. 1,5). During the 3 days preceding the day of eclosion a rhythmic flight motor pattern was produced (Fig. 2). The rhythmic activity ceased 5^1/2–10^1/2 h before eclosion and only an occasional, large potential change was recorded from the thoracic muscles during this time (Fig. 3). During the 3 days of rhythmic activity the percent-age of time that the animal was active did not change (Fig. 4). The flight motor pattern matured, in that the cycle-time decreased and became less variable (Fig. 6). The approximate flight phase relationship between an elevator muscle and the dorsal longitudinal depressor muscle did not become less variable as the cycle-time improved. The flight motor pattern produced by pharate moths caused neither movement of the scutum nor an increase in thoracic temperature in marked contrast to the consequences of adult motor activity (Fig. 7). Intracellular recording from the dorsal longitudinal muscle of pharate moths 20–30 h before eclosion showed that, after repeated stimulation of the motor nerve at 2/s, only small junctional potentials were elicited (Fig. 8). A burst of 6 stimuli at 50/s elicited 2–5 active membrane responses and a contraction. These observations explain the absence of thoracic movement in immature animals producing the flight motor pattern and the presence of movement in immature animals stimulated to eclose. They also show that the neuromuscular junction matures rapidly during the day before eclosion.     

Reorganization of sensory regulation of locust flight after partial deafferentation

Previous investigations have shown that the flight motor pattern of the mature locust (Locusta migratoria L.) relies heavily on the input of the hindwing tegulae. Removal of the hindwing tegulae results in an immediate change in the motor pattern: the wingbeat frequency (WBF) decreases and the interval between the activity of depressor and elevator muscles (D–E interval) increases. In contrast, removal of the forewing tegulae has little effect on the motor pattern. Here we report adaptive modifications in the flight system that occur after the removal of the hindwing tegulae. Over a period of about 2 weeks following hendwing tegula removal, the flight motor pattern progressively returned towards normal, and in about 80% of the animals recovery of the flight motor pattern was complete. We describe the changes in the activity pattern of flight muscles and in the patterns of depolarizations in flight motoneurons and flight interneurons associated with this recovery. In contrast to the situation in the intact animal, the activity of the forewing tegulae is necessary in recovered animals for the generation of the motor pattern. Removal of the forewing tegulae in recovered animals resulted resulted in similar changes in the flight motor pattern as were observed in intact animals after the removal of the hindwing tegulae. Furthermore, electrical stimulation of forewing tegula afferents in recovered animals produced similar resetting effects on the motor pattern as electrical stimulation of the hindwing tegulae afferents in intact animals. From these observations we conclude that recovery is due to the functional replacement of the removed hindwing tegulae by input from the forewing tegulae.     

A reconsideration of the central pattern generator concept for locust flight

Summary Although it is generally agreed that locusts can generate flight similar rhythmic motor activity in the absence of sensory feedback from the wings, recent studies indicate that functional deafferentation produces significant changes in the flight motor pattern (Hedwig and Pearson 1984). These findings have raised doubts on the adequacy of the central pattern generator concept for the locust flight system (Pearson 1985). In this paper, we re-investigate the effects of deafferentation on the capacity of adult migratory locusts to generate the flight motor pattern. For this purpose, the experimental animals were dissected to various degrees, ranging from head-ventral nerve cord, to isolated pterothoracic nerve cord, and finally single isolated ganglion preparations. Flight motor activity was released by either wind stimulation, the more traditional method, or by applying octopamine (Sombati and Hoyle 1984; Stevenson and Kutsch 1986). In all cases the released motor activity was analysed, giving details of latency, and phase relationships between specific synergistic and antagonistic motor units, and then compared with the flight motor pattern generated by intact tethered locusts.This analysis shows that deafferentation, although reducing the frequency, does not necessarily disrupt the basic flight motor pattern. By using octopamine we could show that even isolated thoracic nerve cord preparations can generate activity, which in all major aspects corresponds to this motor program. This could also be shown for the fully isolated metathoracic ganglion and we provide some evidence that the mesothoracic ganglion may be capable of a similar performance. In addition to releasing flight activity, octopamine was also found to enhance the responsiveness of deafferentated locusts to wind stimulation. This resulted in a considerable elevation of the frequency and prolongation of the flight motor activity to values comparable to the performance of intact tethered locusts.     

Turning manoeuvres in free-flying locusts: two-channel radio-telemetric transmission of muscle activity

A device has been constructed allowing the simultaneous transmission of two separate electrical signals in unrestrained small animals. We employed this device to investigate the motor output in free-flying locusts. The activation pattern of several combinations of different muscles was recorded, including bilateral symmetric muscles and pairs of antagonists. Particular attention was paid to the recruitment of a specific set of flight muscles in both winged segments during rolling manoeuvres. The relationship of the muscle activation with wing movement was analysed in combination with a high-speed video-monitoring. The muscles are activated in advance of the relevant stroke directions, in opposition to previous studies of tethered flying locusts. During turning manoeuvres a statistically significant difference in timing of the bilateral symmetric muscles is not apparent; this contrasts with the distinct difference revealed for the bilateral wing movement. It is discussed that rolling might rely on the fine tuned interaction of several major flight muscles or on the precise activation of a specific wing hinge muscle. Correspondence with investigations of bird flight is discussed.     

Pre-imaginal flight motor pattern in Locusta

In a wind stream, larval stages of Locusta usually show a tonic muscle activity but they can also exhibit a rhythmic motor output. With ageing such a pattern can be released sooner, the trains become longer. The basic rhythm of 10 Hz does not change. The initial co-contraction of specific muscles is substituted later in development by an antagonistic recruitment. This activity resembles the flight motor pattern of young locusts which lack phasic sensory feedback from the wing region. Azadirachtin, an insect growth regulator, has been used to produce a permanent 5th larval instar. However, the extension of the last larval stage does not lead to a further development of the motor pattern to a level comparable to mature animals.     

Demonstration of functional connectivity of the flight motor system in all stages of the locust

Summary The development of the flight motor pattern was studied by recording from the thoracic muscles of locusts of various developmental stages. In response to a short wind stimulus, larval locusts generate unpatterned motor activity, whereas newly moulted adults generate the flight pattern (Fig. 1A). The latter is equivalent to the mature adult flight pattern, although more irregular and of lower frequency. Experiments with highly deafferentated locusts indicate that the switch from the larval tonic to adult phasic flight pattern and subsequent increase in frequency are not dependent on phasic peripheral feedback from moving body structures (Fig. 1B). By using octopamine, flight motor activity could be released without need of the wind stimulus (Fig. 2). This corresponded to the normal wind released flight pattern of intact locusts, although the frequency was lower (Fig. 8). Following octopamine treatment, the response to wind stimulation was enhanced. Wind then released in deafferentated adults long flight sequences of significantly elevated frequency (Fig. 3). Although flight is essentially an adult specific behaviour, octopamine was finally found to release flight motor activity in all larval stages (Fig. 7).We conclude that major steps in the development of the flight motor circuitry are completed by the end of embryogenesis. Thus, in contrast to previous assumptions (cf. Bentley and Hoy 1970; Kutsch 1974a; Altman 1975), postembryonic changes in neither the central, nor peripheral nervous system appear to be of major importance for the ontogeny of the locust flight motor program. Whether developmental changes in the wind sensory system of the head, or levels of neurohormones such as octopamine, are related to the newly acquired responsiveness of freshly moulted adult locusts to the normal flight releasing stimulus is discussed.     

Differential effects of octopamine and tyramine on the central pattern generator for <Emphasis Type="Italic">Manduca</Emphasis> flight

The biogenic amine, octopamine, modulates a variety of aspects of insect motor behavior, including direct action on the flight central pattern generator. A number of recent studies demonstrate that tyramine, the biological precursor of octopamine, also affects invertebrate locomotor behaviors, including insect flight. However, it is not clear whether the central pattern generating networks are directly affected by both amines, octopamine and tyramine. In this study, we tested whether tyramine affected the central pattern generator for flight in the moth, Manduca sexta. Fictive flight was induced in an isolated ventral nerve cord preparation by bath application of the octopamine agonist, chlordimeform, to test potential effects of tyramine on the flight central pattern generator by pharmacological manipulations. The results demonstrate that octopamine but not tyramine is sufficient to induce fictive flight in the isolated ventral nerve cord. During chlordimeform induced fictive flight, bath application of tyramine selectively increases synaptic drive to depressor motoneurons, increases the number of depressor spikes during each cycle and decreases the depressor phase. Conversely, blocking tyramine receptors selectively reduces depressor motoneuron activity, but does not affect cycle by cycle elevator motoneuron spiking. Therefore, octopamine and tyramine exert distinct effects on the flight central pattern generating network.     

Effects of octopamine, dopamine, and serotonin on production of flight motor output by thoracic ganglia of Manduca sexta

Effects of biogenic amines on a centrally generated motor pattern in Manduca sexta were examined by pressure injecting nanomole to micromole amounts of octopamine, dopamine or serotonin into thoracic ganglia. Motor output was recorded extracellularly from a pair of antagonistic flight muscles and their motor neurons. The monoamines were found to alter production of a motor pattern that produces rhythmic wing flapping (10 Hz) and exhibits phase relationships similar to those in the flight pattern of intact moths. In mesothoracic ganglia with sensory nerves intact, octopamine (4 X 10(-9) mol) injected into lateral regions evoked regular firing of a single motor neuron, whereas a higher dose (4 X 10(-8) mol) often elicited the flight motor pattern. In the absence of sensory input, these doses of octopamine had little effect. Low doses (10(-10) mol) greatly enhanced motor responses to electrical stimulation of a wing sensory nerve. Dopamine (2 X 10(-10) mol) injected into the medial region of the mesothoracic ganglion elicited the flight motor pattern in the presence or absence of sensory input. Rhythmic output induced by dopamine (5 X 10(-10) mol) was suppressed by injecting serotonin (5 X 10(-10) mol) into the same region. These findings demonstrate that dopamine, octopamine, and serotonin have different effects on motor output in Manduca and suggest that these amines are involved in initiating, maintaining and terminating flight behavior, respectively. Octopamine may elicit flight production by enhancing the efficacy of sensory transmission thereby increasing excitability or arousal. Dopamine may act on interneurons involved in generating the flight motor pattern.     

Reflex activation of locust flight motoneurones by proprioceptors responsive to muscle contractions

 This report investigates the reflex activation of locust flight motoneurones following their spiking activity. As shown elsewhere, an electrical stimulus applied to a flight muscle produces multiple waves of delayed excitation in wing elevator and depressor motoneurones. Nerve ablation experiments show that this response is initiated by the mechanical movement of the stimulated muscle, and not the antidromic spike evoked in the motoneurone. The delayed excitation still occurs in the absence of inputs from the wing receptor systems, and also when all other sources of afferent feedback are abolished, excepting thoracic nerve 2. Following complete deafferentation, spikes in flight motoneurones had no influence on other flight motoneurones. Numerous afferents in the purely sensory nerve 2 are excited by flight muscle contractions. The responses are consistent for repeated contractions of the same muscle, but differ when other muscles are stimulated. During tethered flight, changes in the activation of single flight muscles are reflected in changes of the nerve 2 discharge pattern. Electrical stimulation of this nerve causes delayed excitation of flight motoneurones, and can initiate flight activity. It is suggested that internal proprioceptors, such as those associated with nerve 2, will contribute to shaping the final motor output for flight behaviour. Accepted: 24 April 1996     

Flight activity of three Spodoptera spp., Spodoptera litura, S. exigua and S. depravata, measured by flight actograph

Flight activities of three Spodoptera species were measured by the aid of flight actograph: S. litura and S. exuiga being regarded as long‐distance migratory insects, and S. depravata being non‐migratory and diapause‐inducible species. In all species tested, flight activities were observed only in scotophase, males showed far higher activities than females, being several times higher at the time of maximum flight activity, which was observed within 2 days after adult eclosion. Total flight activity in males was highest in S. litura, some being flyable even 12 days after eclosion, followed by S. exigua being one‐third compared to the former species, while in S. depravata flight activity was nearly half of that of the second species and most ceased to fly within a week after eclosion. There occurred species‐specific daily rhythms in flight activity during respective scotophase. In S. litura, both females and males exhibited a peak of flight activity shortly after light‐off and exhibited the second flight activity in late scotophase, the females slightly but the males more actively compared to early scotophase. In S. exigua, both sexes did not respond to light‐off, did not show a peak of flight activity in early scotophase, whereas males, but not females prominently increased activity toward the end of scotophase. In S. depravata, both sexes exhibited a peak of flight activity in early scotophase, and the males revived flight activity, being maximum shortly before light‐on, but the females did not show a clear rhythm in flight activity. These features observed in flight activity were discussed in relation with migratory capability.     

EVOLUTION AND COADAPTATION OF THERMOREGULATORY BEHAVIOR AND WING PIGMENTATION PATTERN IN PIERID BUTTERFLIES

This paper addresses the question of how the relationship between morphological structure and functional performance differs in related groups of organisms. I describe the relationship between a suite of phenotypic characters (behavioral posture and the pattern of wing pigmentation) and one function of these characters (thermoregulatory performance) for two groups of butterflies in the family Pieridae, focusing on how behavior and wing pattern interact to affect specific aspects of thermoregulation. Using both natural and experimentally created variation in wing-melanization patterns, I develop and test a series of predictions about the relations among thermoregulatory posture, melanization pattern, body temperature, and flight activity. Results show that increased melanization in different wing regions has positive, negative, or neutral effects in increasing body temperature of Pieris butterflies. The angle of the wings used during basking alters the relative importance of different modes of heat transfer and thereby determines the contribution of different dorsal wing regions to thermoregulation. Experimentally increased dorsal melanization can either increase or decrease the onset of flight activity and can directly alter thermoregulatory posture. For Pieris, dorsal melanization affects basking and flight, while ventral melanization primarily affects overheating. These results are used to generate a functional map relating melanization pattern to thermoregulatory performance in Pieris. Reflectance-basking posture, white background color, and melanization pattern represent coadapted characters in Pieris that interact to determine thermoregulatory performance. The differences in thermoregulatory posture and background color between pierid butterflies in the subfamilies Pierinae and Coliadinae have led to a reorganization and partial reversal of the thermoregulatory effects of melanization pattern. I suggest that this change in the physical mechanism of thermoregulatory adaption in pierids has qualitatively altered the nature of selection on wing-melanization pattern.     

Experiments on the central pattern generator for swimming in amphibian embryos

The central nervous system of paralysed Xenopus laevis embryos can generate a motor output pattern suitable for swimming locomotion. By recording motor root activity in paralysed embryos with transected nervous systems we have shown that: (a) the spinal cord is capable of swimming pattern generation; (b) swimming pattern generator capability in the hindbrain and spinal cord is distributed; (c) caudal hindbrain is necessary for sustained swimming output after discrete stimulation. By recording similarly from embryos whose central nervous system was divided longitudinally into left and right sides, we have shown that: (a) each side can generate rhythmic motor output with cycle periods like those in swimming; (b) during this activity cycle period increases within an episode, and there is the usual rostrocaudal delay found in swimming; (c) this activity is influenced by sensory stimuli in the same way as swimming activity; (d) normal phase coupling of the left and right sides can be established by the ventral commissure in the spinal cord. We conclude that interactions between the antagonistic (left and right) motor systems are not necessary for swimming rhythm generation and present a model for swimming pattern generation where autonomous rhythm generators on each side of the nervous system drive the motoneurons. Alternation is achieved by reciprocal inhibition, and activity is initiated and maintained by tonic excitation from the hindbrain.     

Analysis of Drosophila motor neuron activity patterns with neural analogs

A network of reciprocally inhibitory motorneurons has been previously postulated to account for the firing patterns of motor units during dipteran flight. Possible activity patterns of such a network were analyzed by means of appropriately interconnected neuromimes. This theoretical analysis showed that the proposed network can account for many aspects of dipteran motor unit activity patterns including firing in specific sequences, stability, and resetting of the firing pattern by antidromic spikes. In addition, the analysis showed that one aspect of physiological activity, motor unit phase locking, can not be explained simply on the basis of network properties alone; evidently specific membrane properties of the dipteran motor units also play an essential role in establishing the activity pattern.     

Flight initiations in Drosophila melanogaster are mediated by several distinct motor patterns

We have monitored the patterns of activation of five muscles during flight initiation of Drosophila melanogaster: the tergotrochanteral muscle (a mesothoracic leg extensor), dorsal longitudinal muscles #3, #4 and #6 (wing depressors), and dorsal ventral muscle #Ic (a wing elevator). Stimulation of a pair of large descending interneurons, the giant fibers, activates these muscles in a stereotypic pattern and is thought to evoke escape flight initiation. To investigate the role of the giant fibers in coordinating flight initiation, we have compared the patterns of muscle activation evoked by giant fiber stimulation with those during flight initiations executed voluntarily and evoked by visual and olfactory stimuli. Visually elicited flight initiations exhibit patterns of muscle activation indistinguishable from those evoked by giant fiber stimulation. Olfactory-induced flight initiations exhibit patterns of muscle activation similar to those during voluntary flight initiations. Yet only some benzaldehyde-induced and voluntary flight initiations exhibit patterns of muscle activation similar to those evoked by giant fiber stimulation. These results indicate that visually elicited flight initiations are coordinated by the giant fiber circuit. By contrast, the giant fiber circuit alone cannot account for the patterns of muscle activation observed during the majority of olfactory-induced and voluntary flight initiations.Abbreviations DLM/DLMn dorsal longitudinal muscle/motor neuron - DVM/DVMn dorsal ventral muscle/motor neuron - GF(s) giant fiber interneuron (s) - PSI peripherally synapsing interneuron - TTM/TTMn tergotrochanteral muscle/motor neuron