Wing Depressor Research Articles

Neuromuscular mechanisms of an elaborate wing display in the golden-collared manakin (Manacus vitellinus)

Many species perform elaborate physical displays to court mates and compete with rivals, but the biomechanical mechanisms underlying such behavior are poorly understood. We address this issue by studying the neuromuscular origins of display behavior in a small tropical passerine bird, the golden-collared manakin (Manacus vitellinus). Males of this species court females by dancing around the forest floor and rapidly snapping their wings together above their back. Using radio-telemetry, we collected electromyographic (EMG) recordings from the three main muscles that control avian forelimb movement, and found how these different muscles are activated to generate various aspects of display behavior. The muscle that raises the wing (supracoracoideus, SC) and the primary muscle that retracts the wing (scapulohumeralis caudalis, SH) were activated during the wing-snap, whereas the pectoralis (PEC), the main wing depressor, was not. SC activation began before wing elevation commenced, with further activation occurring gradually. By contrast, SH activation was swift, starting soon after wing elevation and peaking shortly after the snap. The intensity of this SH activation was comparable to that which occurs during flapping, whereas the SC activation was much lower. Thus, light activation of the SC likely helps position the wings above the back, so that quick, robust SH activation can drive these appendages together to generate the firecracker-like snap sonation. This is one of the first looks at the neuromuscular mechanisms that underlie the actuation of a dynamic courtship display, and it demonstrates that even complex, whole-body display movements can be studied with transmitter-aided EMG techniques.

Responses ofDrosophilagiant descending neurons to visual and mechanical stimuli

In Drosophila, the paired giant descending neurons (GDNs), also known as giant fibers, and the paired giant antennal mechanosensory descending neurons (GAMDNs), are supplied by visual and mechanosensory inputs. Both neurons have the largest cell bodies in the brain and both supply slender axons to the neck connective. The GDN axon thereafter widens to become the largest axon in the thoracic ganglia, supplying information to leg extensor and wing depressor muscles. The GAMDN axon remains slender, interacting with other descending neuron axons medially. GDN and GAMDN dendrites are partitioned to receive inputs from antennal mechanosensory afferents and inputs from the optic lobes. Although GDN anatomy has been well studied in Musca domestica, less is known about the Drosophila homolog, including electrophysiological responses to sensory stimuli. Here we provide detailed anatomical comparisons of the GDN and the GAMDN, characterizing their sensory inputs. The GDN showed responses to light-on and light-off stimuli, expanding stimuli that result in luminance decrease, mechanical stimulation of the antennae, and combined mechanical and visual stimulation. We show that ensembles of lobula columnar neurons (type Col A) and mechanosensory antennal afferents are likely responsible for these responses. The reluctance of the GDN to spike in response to stimulation confirms observations of the Musca GDN. That this reluctance may be a unique property of the GDN is suggested by comparisons with the GAMDN, in which action potentials are readily elicited by mechanical and visual stimuli. The results are discussed in the context of descending pathways involved in multimodal integration and escape responses.

Role of wing pronation in evasive steering of locusts

Evasive steering is crucial for flying in a crowded environment such as a locust swarm. We investigated how flying locusts alter wing-flapping symmetry in response to a looming object approaching from the side. Desert locusts (Schistocerca gregaria) were tethered to a rotatable shaft that allowed them to initiate a banked turn. A visual stimulus of an expending disk on one side of the locust was used to evoke steering while recording the change in wingbeat kinematics and electromyography (EMG) of metathoracic wing depressors. Locusts responded to the looming object by rolling to the contralateral direction. During turning, EMG of hindwing depressors showed an omission of one action potential in the subalar depressor (M129) of the hindwing inside the turn. This omission was associated with increased pronation of the same wing, reducing its angle-of-attack during the downstroke. The link between spike-omission in M129 and wing pronation was verified by stimulating the hindwing depressor muscles with an artificial motor pattern that included the misfire of M129. These results suggest that hindwing pronation is instrumental in rotating the body to the side opposite of the approaching threat. Turning away from the threat would be highly adaptive for collision avoidance when flying in dense swarms.

Electrophysiological recordings from the giant fiber pathway of D. melanogaster.

When startled adult D. melanogaster react by jumping into the air and flying away. In many invertebrate species, including D. melanogaster, the "escape" (or "startle") response during the adult stage is mediated by the multi-component neuronal circuit called the Giant Fiber System (GFS). The comparative large size of the neurons, their distinctive morphology and simple connectivity make the GFS an attractive model system for studying neuronal circuitry. The GFS pathway is composed of two bilaterally symmetrical Giant Fiber (GF) interneurons whose axons descend from the brain along the midline into the thoracic ganglion via the cervical connective. In the mesothoracic neuromere (T2) of the ventral ganglia the GFs form electro-chemical synapses with 1) the large medial dendrite of the ipsilateral motorneuron (TTMn) which drives the tergotrochanteral muscle (TTM), the main extensor for the mesothoracic femur/leg, and 2) the contralateral peripherally synapsing interneuron (PSI) which in turn forms chemical (cholinergic) synapses with the motorneurons (DLMns) of the dorsal longitudinal muscles (DLMs), the wing depressors. The neuronal pathway(s) to the dorsovental muscles (DVMs), the wing elevators, has not yet been worked out (the DLMs and DVMs are known jointly as indirect flight muscles - they are not attached directly to the wings, but rather move the wings indirectly by distorting the nearby thoracic cuticle) (King and Wyman, 1980; Allen et al., 2006). The di-synaptic activation of the DLMs (via PSI) causes a small but important delay in the timing of the contraction of these muscles relative to the monosynaptic activation of TTM (~0.5 ms) allowing the TTMs to first extend the femur and propel the fly off the ground. The TTMs simultaneously stretch-activate the DLMs which in turn mutually stretch-activate the DVMs for the duration of the flight. The GF pathway can be activated either indirectly by applying a sensory (e.g."air-puff" or "lights-off") stimulus, or directly by a supra-threshold electrical stimulus to the brain (described here). In both cases, an action potential reaches the TTMs and DLMs solely via the GFs, PSIs, and TTM/DLM motoneurons, although the TTMns and DLMns do have other, as yet unidentified, sensory inputs. Measuring "latency response" (the time between the stimulation and muscle depolarization) and the "following to high frequency stimulation" (the number of successful responses to a certain number of high frequency stimuli) provides a way to reproducibly and quantitatively assess the functional status of the GFS components, including both central synapses (GF-TTMn, GF-PSI, PSI-DLMn) and the chemical (glutamatergic) neuromuscular junctions (TTMn-TTM and DLMn-DLM). It has been used to identify genes involved in central synapse formation and to assess CNS function.

Electrophysiological Recordings from the Giant Fiber Pathway of <em>D. melanogaster</em>

When startled adult D. melanogaster react by jumping into the air and flying away. In many invertebrate species, including D. melanogaster, the "escape" (or "startle") response during the adult stage is mediated by the multi-component neuronal circuit called the Giant Fiber System (GFS). The comparative large size of the neurons, their distinctive morphology and simple connectivity make the GFS an attractive model system for studying neuronal circuitry. The GFS pathway is composed of two bilaterally symmetrical Giant Fiber (GF) interneurons whose axons descend from the brain along the midline into the thoracic ganglion via the cervical connective. In the mesothoracic neuromere (T2) of the ventral ganglia the GFs form electro-chemical synapses with 1) the large medial dendrite of the ipsilateral motorneuron (TTMn) which drives the tergotrochanteral muscle (TTM), the main extensor for the mesothoracic femur/leg, and 2) the contralateral peripherally synapsing interneuron (PSI) which in turn forms chemical (cholinergic) synapses with the motorneurons (DLMns) of the dorsal longitudinal muscles (DLMs), the wing depressors. The neuronal pathway(s) to the dorsovental muscles (DVMs), the wing elevators, has not yet been worked out (the DLMs and DVMs are known jointly as indirect flight muscles - they are not attached directly to the wings, but rather move the wings indirectly by distorting the nearby thoracic cuticle) (King and Wyman, 1980; Allen et al., 2006). The di-synaptic activation of the DLMs (via PSI) causes a small but important delay in the timing of the contraction of these muscles relative to the monosynaptic activation of TTM (~0.5 ms) allowing the TTMs to first extend the femur and propel the fly off the ground. The TTMs simultaneously stretch-activate the DLMs which in turn mutually stretch-activate the DVMs for the duration of the flight. The GF pathway can be activated either indirectly by applying a sensory (e.g."air-puff" or "lights-off") stimulus, or directly by a supra-threshold electrical stimulus to the brain (described here). In both cases, an action potential reaches the TTMs and DLMs solely via the GFs, PSIs, and TTM/DLM motoneurons, although the TTMns and DLMns do have other, as yet unidentified, sensory inputs. Measuring "latency response" (the time between the stimulation and muscle depolarization) and the "following to high frequency stimulation" (the number of successful responses to a certain number of high frequency stimuli) provides a way to reproducibly and quantitatively assess the functional status of the GFS components, including both central synapses (GF-TTMn, GF-PSI, PSI-DLMn) and the chemical (glutamatergic) neuromuscular junctions (TTMn-TTM and DLMn-DLM). It has been used to identify genes involved in central synapse formation and to assess CNS function.

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.

Escape flight initiation in the fly

Visually evoked escape flight initiation in Drosophila, according to the accepted account, involves a rapid extension of the middle legs that propels the fly into the air while the wings are still folded. This description has remained unchallenged and is accounted for in terms of the activation of a simple neural circuit, the Giant fibre (GF) system. The accepted description of escape is however inconsistent with the sequence of events recorded when the GF system is stimulated. Specifically, previous electrophysiological recordings have shown that the wing depressor muscles are activated before the wings are in a position to be depressed because they have not yet been elevated. Here we show that the accepted behavioural description is wrong. Escape flight initiation actually begins with wing elevation. The current model of the GF system is revised to account for the actual sequence of events that occur when a fly escapes.

Intrinsic noise at synapses between a wing hinge stretch receptor and flight motor neurons in the locust.

Variability in postsynaptic potential (PSP) amplitude due to intrinsic noise limits the reliability of communication between neurons. I measured PSP variability at synapses between a forewing stretch receptor and wing depressor motor neurons in locusts, a pathway that is important in the control of flying. The intrinsic noise in the stretch receptor output synapse was measured by subtracting the background noise, originating in other synaptic pathways onto the motor neuron, from the variability in the amplitudes of PSPs evoked by the stretch receptor. Intrinsic synaptic noise caused successive PSPs to vary by 4-10 % in basalar and subalar flight motor neurons. Recordings from pairs of these wing depressor motor neurons showed that the amount of transmitter released varied independently between different output sites from the stretch receptor. Histograms of excitatory postsynaptic potential amplitude were normal distributions that lacked separate peaks. I estimate that quantal amplitude is significantly less than 0.1 mV and that several hundred quanta are released for each presynaptic spike. This accords well with a previous estimate of the number of discrete anatomical synapses and would facilitate modulation of output from the stretch receptor.

Peculiarities of ultrastructural organization of the metathoracic ganglion in the locustLocusta migratoria larva

In electron microscopic study of structural organization of the thoracic ganglion of the locust larva of the 1st age (1–2 days after hatching), the data on the structure of motoneurons of the 1st nerve, basal and motor neuropil of the larva were obtained. The effector elements of the larval locust CNS are formed rather early and have the structural plan similar to that in adult insects. However, in the larval motoneurons innervating the flight muscles (longitudinal dorsal muscles, wing depressors) the clearly seen features of immaturity of these nervous elements are revealed. Study of the larval ganglion neuropil has shown that the basal neuropil is morphologically formed sufficiently completely as early as in larvae of the first days after hatching. There are shown longitudinal contacts between axons of the ventral neuropil zone, the presence of axons forming theen-passant contacts as well as the synapses with a heterogeneous set of vesicles in the presynaptic area. The presence of the great number of granular vesicles in the basal neuropil of the locust larva may indicate an important role of catecholamines in the early development of the nervous system in the locust larva.

GABA immunoreactivity in processes presynaptic to the locust wing stretch receptor neuron

Studies on the locust Locusta migratoria (Leitch et al. [1998] Soc. Neurosci. Abstr. 624.10) suggest that gamma-aminobutyric acid (GABA)ergic interneurons, activated via muscarinic receptors, may be involved in modulation of transmitter release at synapses between the forewing stretch receptor (fSR) and wing depressor motoneurons. To help elucidate the role GABAergic interneurons may play in modulation of transmitter release from fSR terminals, the proportion and distribution of GABA-immunoreactive (GABA-IR) inputs to the fSR were analysed using double-labelling (horseradish peroxidase and GABA immunocytochemistry) and electron microscopy. Forty-three percent of synaptic inputs to the fSR were from GABA-IR profiles, the majority of which were located on lateral branches. The highest proportion (57%) of inputs to the fSR, however, were from non-IR processes containing either clear spherical vesicles or mixed clear and dense-cored vesicles. Outputs from the fSR to GABA-IR profiles were also found, although their number was relatively low (7%). Networks were identified in which both the fSR and its non-IR postsynaptic partner received input from the same GABA-IR neuron. Such an arrangement would allow both pre- and postsynaptic inhibition of fSR afferent outputs, for example at fSR/motoneuron synapses. These observations demonstrate that the fSR does receive presynaptic inputs from GABA-IR neurons, thus providing morphological support for pharmacological and electrophysiological findings that GABAergic neurons are involved in the presynaptic modulation of the fSR. Nevertheless, modulation at this synapse may be more intricate and involve other, as yet unidentified, neurotransmitters released from non GABA-IR presynaptic processes and also muscarinic receptors located on the fSR itself.

Co-ordination of the Flight Motor Pattern with Forewing Stretch Receptor Stimulation in Immature and Mature Adult Locusts

Abstract We investigated the effect of proprioceptive feedback on the flight motor pattern during maturation of the locust flight system by stimulating the forewing stretch receptors (fSRs) in immature and mature adults and recording the frequency of wing depressor muscle activity. Stimulation of both fSRs produced co-ordination with the motor pattern in 4 of 10 immature animals and in 6 of 16 mature animals. The strength of co-ordination was measured as the variability of the phase of hindwing depressor muscle activity within the stimulus period. Co-ordination was stronger and the phase values higher in mature animals presented with a stimulus that mimics fSR activity in response to rhythmic wing elevation. We conclude that maturation of the flight motor is due, in part, to a change in the response of the central circuitry to a change in the timing of proprioceptive input.

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.

Modulation of transmitter release from the terminals of the locust wing stretch receptor neuron by muscarinic antagonists

The forewing stretch receptor (SR) neuron makes monosynaptic connections with wing depressor motoneurons; in this article the pharmacology of its output onto the first basalar motoneuron (BA1) has been investigated. The SR, like other insect afferents that have been studied so far, appears to be cholinergic; transmission was suppressed reversibly by the nicotinic antagonist gallamine (10(-4) M) and irreversibly by alpha-bungarotoxin (10(-6) M). The choline reuptake blocker hemicholinium-3 (10(-4) M) also caused a reversible reduction in the amplitude of SR excitatory postsynaptic potentials (EPSPs) recorded in BA1. The receptor subtype nonselective muscarinic antagonists atropine (10(-4) M), scopolamine (10(-4) M), and quinuclidinyl benzilate (10(-5) M), unlike nicotinic antagonists, caused an augmentation in EPSP amplitude. This effect does not appear to be caused by an increase in sensitivity of the motoneuron to acetylcholine (ACh), since atropine produced a marked reduction rather than an increase in the amplitude of responses to ACh pressure applied to the soma of BA1. Scopolamine only caused a modest reduction in the amplitude of ACh somatic responses. The simplest explanation for these observations is that muscarinic antagonists bring about an increase in EPSP amplitude by blockade of presynaptic autoreceptors that normally down-regulate the release of ACh from SR terminals. The effects of muscarinic receptor subtype-selective antagonists indicate that presynaptic receptors in this preparation may have a pharmacological profile more similar to that of vertebrate M2 receptors than to that of M1 or M3 subtypes. The functional significance of autoreceptors in this preparation are discussed.

The high number of neurons contributes to the robustness of the locust flight-CPG against parameter variation

Real pattern-generating networks often consist of more neurons than necessary for the production of a certain rhythm. We investigated the question of whether these neurons contribute to the robustness of a pattern-generating system of using the central pattern generator (CPG) for flight of the locust, generating the deafferented activity pattern of wing elevator and wing depressor motoneurons, as an example of a rhythm-generating system. The neuronal network was reconstructed, based on the known connectivity of the interneurons in the flight CPG, using a biologically orientated network simulator (BioSim 3.0). This simulator allows a physiologically realistic simulation of particular neurons as well as the synaptic connections between them. The flight CPG consists of at least five cyclic loops. The simulation shows that each of them is in principle able to produce a rhythm comparable to the rhythm produced by the whole network, i.e. the `deafferented' flight pattern of elevator and depressor motoneurons. Varying the parameter `synaptic strength' in each of these loops and in the complete system shows that this parameter can be changed within certain ranges without loosing the ability to produce oscillations. These ranges are much smaller in each of the subloops than in the whole network. This result demonstrates that the robustness of the system is increased by supranumerary neurons and connections. Changing the active properties of the simulated neurons so that they are able to produce plateau potentials has no effect on the robustness of the simulated network.

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.

Gender specific developmental transformation of a cockroach bifunctional muscle

AbstractAdult male and female cockroaches,Periplaneta americana, differ in their capacity for sustained flight. Tethered 20 day adult males flap their wings about 15 min; females 12 seconds. Males are less massive and have greater total wing area than do females, with wing loading only about half that of the female. The basalar muscle, a major wing depressor in flight, is also sexually dimorphic. In males it is 1.5× more massive and has a cross sectional area 1.2× greater than the female homologue. Muscle fibers of the adult male and female basalar muscles are ultrastructurally and biochemically distinct. Female fibers have a myofilament array similar to that of other non‐flight locomotory muscles, with 10–12 actin filaments surrounding each myosin filament. Adult male fibers typically have six actin filaments around each myosin filament. The male fibers also appear to be more oxidative. The volume density of mitochondria is about 3× greater and the specific activity for citrate synthase, an oxidative indicator, is 3.5× greater in male fibers than in female fibers. Male basalar muscle shows an enhanced ability to resist fatigue when its motoneurone is stimulated at flight frequency; homologues from the male terminal nymph and 20 day adult female are both highly fatiguable. The basalar muscle of both male and female terminal nymphs are similar in ultrastructural and biochemical profile and are not greatly different from the adult female. In male roaches there may be important neuronal or hormonal inputs at about the time of the terminal molt to adulthood that direct the transformation of the basalar muscle in time for the assumption of a new behavioral role—flight. © 1994 Wiley‐Liss, Inc.

Comparison of motor patterns in larval and adult stage of a beetle, Zophobas morio

AbstractFor a holometabolous insect, the beetle Zophobas morio, several motor patterns were recorded, using EMGs, for several homologous metathoracic muscles of the larval and adult stage. In the larva, most muscles are employed during walking, while in the adult, only the remotor coxae muscle (M119) will be activated during leg movements. Segmental bendings of the larva are mainly due to contractions of the dorsal longitudinal muscles. For adults, the myogenically driven thoracic flight muscles display a phase preference within the wing‐beat cycle, resulting in not only a functional but also activity‐dependent classification of wing depressor and elevator muscles. Time interval histograms of the excitatory junctional potentials (ejp's) show several peaks separated by about 20 ms, representing the wing‐beat period. Application of chlordimeform (CDM), an agonist of the neuromodulator octopamine, elicits in adults longer “flight” sequences without wing movement. During such sequences the 20 ms modulation of the ejp intervals is missing. This points to wing‐beat dependent afferences involved in patterning of the output during normal flight. Application of CDM in the larva results in a tonic activation, which shows several similarities with the adult ‘flight’ without wing‐beat. This finding suggests that the basic neuronal circuitry underlying the imago's flight is present precociously, before the dramatic reorganization of the animal during metamorphosis. © 1993 wiley‐Liss, Inc.

Wing kinematics during natural leaping in the mantids Mantis religiosa and Iris oratoria

High‐speed flash photography was used to analyse wing movements of Mantis religiosa and Iris oratoria at the moment of take‐off during natural leaping. Wing kinematics are compared with those of the similarly designed locust wing. Iris oratoria showed strong coupling between leg extensor and wing depressor muscle activity immediately prior to take‐off, with a possible enhancement of jump momentum. A ‘clap and peel’ was observed in the hind wings of both species during the first downstroke. Supination in the mantid forewing is accomplished by a backward rotation of the whole of the main wing plate about the claval furrow. Both fore‐ and hind wings show pronounced ventral flexure at the lower point of stroke reversal. Camber was developed in the hind wing during the upstroke as well as the downstroke. Possible roles of the claval furrow and transverse flexion in protecting the forewing base against torsional forces generated at stroke reversal are discussed.

Locomotor activities in the decerebrate bird without phasic afferent input

We examined whether forelimb and hindlimb phasic afferent input is a prerequisite for the production of avian locomotor patterns. We eliminated phasic afferent feedback through paralysis of a decerebrate animal. The term “fictive” has been used to describe the neural activity associated with spontaneous or evoked motor output during neuromuscular paralysis. We observed that a paralysed decerebrate bird is capable of producing similar locomotor activity patterns as an unparalysed preparation, regardless of whether the “fictive” locomotion is generated spontaneously, or in response to focal electrical and/or neurochemical stimulation of discrete brainstem locomotor regions. Not all aspects of “fictive” locomotor patterns were identical to the locomotion elicited prior to paralysis. The stimulus current threshold necessary to evoke hindlimb locomotion increased from 69 ± 22 μ A (mean ± S.D.) prior to paralysis to 185 ± 87μ A for “fictive” stepping. For wing activity, the threshold increased from 84 ± 46 μ A during wing flapping to 228 ± 148μ A for “fictive” flight. In addition, the frequency of “fictive” efferent locomotor activity from the leg nerve (1.04 ± 0.44Hz) decreased relative to the frequency of leg activity prior to paralysis (1.55 ± 0.70Hz). Similarly, the frequency of wing activity decreased from 2.73 ± 0.73Hz before paralysis to 1.8 ± 0.69Hz after paralysis. Finally flexor burst duration remained constant during treadmill and “fictive” walking while the extensor burst duration was markedly increased during “fictive” walking. Thus, the relative contributions of leg flexor activity to the overall step cycle (burst proportion = burst duration/cycle duration) decreased during evoked “fictive” stepping, while the burst proportion of the leg extensor increased. Afferent feedback therefore appears to modulate leg extensor burst duration more than leg flexor duration. For the wings, the burst proportion of the major wing depressors remained constant before and after paralysis.

Influence of input from the Forewing Stretch Receptors on Motoneurones in Flying Locusts

ABSTRACT Previous studies on the forewing stretch receptors (FSRs) of locusts have suggested that feedback from these receptors during flight contributes to the excitation of depressor motoneurones and reduces the duration of depolarizations in elevator motoneurones. We have investigated these proposals by measuring the timing of FSR activity relative to depressor activity and by examining the effects of stimulating the FSRs on the membrane potential oscillations in flight motoneurones. Activity in the FSRs was recorded in tethered intact animals flying in a windstream and in preparations that allowed intracellular recordings from motoneurones during flight activity. The timing of FSR activity was similar in both preparations. In most animals we observed that at normal wingbeat frequencies (about 20 Hz) the activity in the FSRs commenced after the onset of activity in the wing depressor muscles. As wingbeat frequency declined there was a progressive advance of FSR activity relative to depressor activity. Most of the spikes in each burst of FSR activity occurred during the time that the membrane potential in depressor motoneurones was repolarizing. Electrical stimulation of the FSRs timed to follow the onset of depressor activity slowed the rate of repolarization, decreased the peak hyperpolarization and increased the rate of the following depolarization in depressor motoneurones. In elevator motoneurones, the same pattern of FSR stimulation produced an additional excitatory input during the depolarization phase and, at low wingbeat frequencies, reduced the duration of the peak depolarizations. The reduction in the duration of the peak depolarization in elevator motoneurones was not strongly correlated to the reduction in cycle period. We propose that the primary reason why input from the FSRs increases wingbeat frequency is because this input reduces the degree of hyperpolarization in depressor neurones and thus promotes an earlier onset of the next depolarization in these neurones.