The use of hormonally induced mosaics to study alterations in the synaptic connections made by persistent sensory neurons during insect metamorphosis

During the initial phase of metamorphosis in the hawkmoth, Manduca sexta, persistent mechanosensory neurons expand their terminal arborizations within the CNS and evoke a reflex response in the pupa which is different than in the larva. In an effort to determine the contribution of sensory neuron modifications to the difference in reflex responses, manipulations of juvenile hormone and 20-hydroxyecdysone were used to generate mosaic animals in which the sensory neurons were advanced or delayed developmentally with respect to the rest of the animal, including circuit components within the CNS. In the larval stage electrical stimulation of the sensory axons evokes a slow depolarization and a prolonged burst of action potentials in the ipsilateral intersegmental muscle motor neurons. By contrast, in pupal preparations the same motor neurons respond to an identical stimulus with a larger, more rapid depolarization which leads to a relatively brief, high-frequency burst of action potentials. Motor responses on the contralateral side of the body are also altered during pupal development. In mosaic animals where larval-like sensory neurons interact with a pupal CNS, a larval reflex response is generated. In the converse situation, pupal-like sensory neurons interacting with a larval or prepupal CNS evoke a motor response that is typical of larvae or prepupae. We conclude, therefore, that pupal development of the sensory neurons is necessary, but not sufficient, for the production of the pupal reflex.     

Multisegmental motor activity in the segmentally restricted gin trap behavior in Manduca sexta pupae

Stimulation of sensory neurons innervating hairs in the gin traps on the abdomen of Manduca sexta pupae evokes a rapid bending of the abdomen that is restricted to one or more of the three articulating posterior segments. However, electrical stimulation of the gin trap sensory nerve in an isolated abdominal nerve cord evokes characteristic motor neuron activity in every abdominal segment. To determine if the segmentally distributed motor activity also occurred in intact animals and how it contributed to the segmentally restricted reflex movement, mechanical stimulation of the sensory hairs in intact animals was used to evoke reflex responses that were recorded as electromyograms synchronized with video recordings of the behavior. Motor activity was monitored during movements to determine if there was activity in many segments when the movement was restricted to one segment. Coordinated muscle activity was evoked throughout the abdomen in response to stimulation of any of the three gin traps, even when movement was restricted to one segment. Differences in the timing of ipsilateral and contralateral motor activity among segments allowed the closing of gin traps to be segmentally restricted. These findings suggest that the neural circuit underlying the gin trap reflex is distributed throughout the abdominal nerve cord. This network generates a complex, yet coordinated, motor pattern with muscular activity in many abdominal segments that produces a localized bending reflex. Accepted: 10 January 1997     

Segmentally distributed metamorphic changes in neural circuits controlling abdominal bending in the hawkmoth Manduca sexta

During the metamorphosis of Manduca sexta the larval nervous system is reorganized to allow the generation of behaviors that are specific to the pupal and adult stages. In some instances, metamorphic changes in neurons that persist from the larval stage are segment-specific and lead to expression of segment-specific behavior in later stages. At the larval-pupal transition, the larval abdominal bending behavior, which is distributed throughout the abdomen, changes to the pupal gin trap behavior which is restricted to three abdominal segments. This study suggests that the neural circuit that underlies larval bending undergoes segment specific modifications to produce the segmentally restricted gin trap behavior. We show, however, that non-gin trap segments go through a developmental change similar to that seen in gin trap segments. Pupal-specific motor patterns are produced by stimulation of sensory neurons in abdominal segments that do not have gin traps and cannot produce the gin trap behavior. In particular, sensory stimulation in non-gin trap pupal segments evokes a motor response that is faster than the larval response and that displays the triphasic contralateral-ipsilateral-contralateral activity pattern that is typical of the pupal gin trap behavior. Despite the alteration of reflex activity in all segments, developmental changes in sensory neuron morphology are restricted to those segments that form gin traps. In non-gin trap segments, persistent sensory neurons do not expand their terminal arbors, as do sensory neurons in gin trap segments, yet are capable of eliciting gin trap-like motor responses. Accepted: 10 January 1997     

Intersegmental interneurons serving larval and pupal mechanosensory reflexes in the moth Manduca sexta

1. Intersegmental interneurons (INs) that participate in the larval bending reflex and the pupal gin trap closure reflex were identified in the isolated ventral nerve cord of Manduca sexta. INs 305, 504, and 703 show qualitatively different responses in the pupa than in the larva to electrical stimulation of sensory neurons that are retained during the larval-pupal transition to serve both reflexes. Action potentials produced by current injected into the 3 interneurons excite motor neurons that are directly involved in the larval and pupal reflexes. The excitation of the motor neurons is not associated with EPSPs at a fixed latency following action potentials in the interneurons, and thus there do not seem to be direct synaptic connections between the interneurons and the motor neurons. 2. IN 305 (Fig. 2) has a lateral soma, processes in most of the dorsal neuropil ipsilateral to the soma, and a crossing neurite that gives rise to a single contralateral descending axon. IN 305 is excited by stimulation of the sensory nerve ipsilateral to its soma in the larva and the pupa. Stimulation of the sensory nerve contralateral to its soma produces an inhibitory response in the larva, but a mixed excitatory/inhibitory response to the identical stimulus in the pupa. 3. IN 504 (Fig. 3) has a lateral soma, processes throughout most of the neuropil ipsilateral to the soma, and a crossing neurite that bifurcates to give rise to a process extending to the caudal limit of the neuropil and an ascending axon. IN 504 is excited by stimulation of the sensory nerve ipsilateral to its soma in both larvae and pupae, while the response to stimulation of the sensory nerve contralateral to its soma is inhibitory in the larva but mixed (excitatory/inhibitory) in the pupa. 4. IN 703 has a large antero-lateral soma, a neurite that extends across to the contralateral side giving rise to processes located primarily dorsally in both ipsilateral and contralateral neuropils, and two axons that ascend and descend in the connectives contralateral to the soma (Fig. 4). IN 703 responds to stimulation of the sensory nerves on either side of the ganglion, but the form of the response changes during the larval-pupal transition. In the larva, the response consists of very phasic (0-2 spikes) excitation, but in the pupa there is a prolonged excitation that greatly outlasts the stimulus (Fig. 6).(ABSTRACT TRUNCATED AT 400 WORDS)     

Morphology and physiology of a hair plate sensory organ located on the antenna of the rock lobster Palinurus vulgaris

At the level of the J1 joint of each antenna of the rock lobster Palinurus vulgaris a hair plate sensory organ (hp) similar to those described in insects has been observed. The hp is located on the internal side of the S2 segment of the antenna, close to the soft articulating membrane of the J1 joint. It is formed by a triangular surface of cuticle about 3mm2 in area, covered with numerous hairs of different lengths (Figs. 1 and 2). Details of the hp were studied by scanning electron microscopy (Fig. 2). Physiological stimulation of the hp hairs occurs during medial movement of the J1 joint. Under this condition the soft articulating membrane rolls over the hairs and bends them progressively back onto the cuticle. Flexion of all the hairs corresponds to a medial movement of the J1 through 40 degrees. During this type of movement, the number of successively flexed hairs increases linearly (Fig. 3). Electrophysiological recordings of the hp sensory nerve correlated with selective mechanical stimulation of individual hairs demonstrated that each hair is innervated by a single sensory fiber. This sensory neurone responds phasically when the hair is flexed back onto the cuticle (as during an S2 medial movement) and when it returns to its resting position (as during an S2 lateral movement). Most of the sensory neurones are sensitive to the movement velocity of the hairs (Figs. 4 and 5). When the hair is maintained flexed its sensory neurone discharges tonically (Fig. 4). Electrical stimulation of the hp sensory nerve induced reflex actions in the external and internal rotator muscles of the segment S1. These effects were found to selectively activate the tonic motor command of these muscles (Fig. 6).     

Functions of the LE sensory neurons inAplysia

Mechanosensory neurons which innervate the siphon and have their cell bodies in the LE cluster of the abdominal ganglion ofAplysia have revealed many cellular and molecular processes that may play general roles in learning and memory. It was initially suggested that these cells are largely responsible for triggering the gill-withdrawal reflex evoked by weak siphon stimulation, and that most of this effect is mediated by their monosynaptic connections to gill motor neurons. This implied a simple link between plasticity at these synapses and modifications of the reflex during learning. We review more recent studies from several laboratories showing that the LE cells are not activated by very weak tactile stimuli that elicit the gill-withdrawal reflex, and that an unidentified population of siphon sensory neurons has lower mechanosensory thresholds and produces shorter latency responses. Furthermore, the direct connections between LE cells and gill motor neurons make a minor contribution when the reflex is elicited in pinned siphon preparations by light stimuli that weakly activate the LE cells. Because weak mechanical stimulation of the unrestrained siphon causes little or no LE cell activation, it is unlikely that, under natural conditions, sensitization or conditioning of reflex responses elicited by light siphon touch depends upon plasticity of LE cell synapses onto either motor or interneurons. The LE cells appear to function as nociceptors because they are tuned to noxious stimuli and, like mammalian nociceptors, show peripheral sensitization following nociceptive activation. This sensitization and the profound activity-dependent potentiation of LE synapses indicate that LE cell contributions to defensive reflexes should be largest during and after intense activation of the LE cells by noxious stimulation (with the LE cell plasticity contributing to long-lasting memory of peripheral injury). The LE sensory neurons offer special opportunities for direct tests of this and other hypotheses about specific mnemonic functions of fundamental mechanisms of neural plasticity.     

Synaptic responses produced in lobster abdominal postural motor neurons by mechanical stimulation of the swimmeret

1. Intracellular recordings were obtained from the somata of identified abdominal postural motor neurons in lobster to examine their subthreshold and suprathreshold responses to tactile stimulation of the swimmeret. 2. Pressure stimulation of the swimmeret surface evoked abdominal extension by producing tonic spiking in the extensor excitors and the synergistic flexor inhibitor (f5) and hyperpolarizing responses in the extensor inhibitor and antagonistic flexor excitors. These responses often continued for several seconds following the termination of the stimulus. The receptive fields of these motor responses extended over most of the swimmeret surface. 3. More localized tactile stimulation of the swimmeret surface elicited EPSPs in f5 and the extensor excitors, and IPSPs in the flexor excitors. The amplitude of these synaptic potentials decreased as the stimulus intensity was reduced. 4. Stimulation of feathered hair (both sexes) and smooth hair (female only) sensilla produced responses characteristic of extension whereas bristly spines on the male accessory lobe excited only two flexor excitors without affecting any of the other postural motor neurons. 5. Summed synaptic responses recorded from the motor neurons differed in their amplitudes and latencies according to the type of mechanoreceptor stimulated-cuticular receptors, feathered hairs or smooth hairs. Stimulation of the swimmeret cuticle produced the strongest responses (shortest latency, largest amplitude), while feathered hair stimulation initiated the weakest responses (longest latency, smallest amplitude). 6. The relatively long latencies (greater than 35 ms) and the complex form of the EPSPs and IPSPs indicate the involvement of multisynaptic interneuronal pathways in the reflex arcs.     

Responses of efferent octopaminergic thoracic unpaired median neurons in the locust to visual and mechanosensory signals

Insect thoracic ganglia contain efferent octopaminergic unpaired median neurons (UM neurons) located in the midline, projecting bilaterally and modulating neuromuscular transmission, muscle contraction kinetics, sensory sensitivity and muscle metabolism. In locusts, these neurons are located dorsally or ventrally (DUM- or VUM-neurons) and divided into functionally different sub-populations activated during different motor tasks. This study addresses the responsiveness of locust thoracic DUM neurons to various sensory stimuli. Two classes of sense organs, cuticular exteroreceptor mechanosensilla (tactile hairs and campaniform sensilla), and photoreceptors (compound eyes and ocelli) elicited excitatory reflex responses. Chordotonal organ joint receptors caused no responses. The tympanal organ (Müller's organ) elicited weak excitatory responses most likely via generally increased network activity due to increased arousal. Vibratory stimuli to the hind leg subgenual organ never elicited responses. Whereas DUM neurons innervating wing muscles are not very responsive to sensory stimulation, those innervating leg and other muscles are very responsive to stimulation of exteroreceptors and hardly responsive to stimulation of proprioceptors. After cutting both cervical connectives all mechanosensory excitation is lost, even for sensory inputs from the abdomen. This suggests that, in contrast to motor neurons, the sensory inputs to octopaminergic efferent neuromodulatory cells are pre-processed in the suboesophageal ganglion.     

Neural correlates of habituation of the proleg withdrawal reflex in larvae of the hawk moth, Manduca sexta

The larval proleg withdrawal reflex of the hawk moth, Manduca sexta, exhibits robust habituation. This reflex is evoked by deflecting one or more mechanosensory planta hairs on a proleg tip. We examined neural correlates of habituation in an isolated proleg preparation consisting of one proleg and its segmental ganglion. Repeated deflection of a single planta hair caused a significant decrease in the number of action potentials evoked in the proleg motor nerve (which carries the axons of proleg retractor motor neurons). Significant response decrement was seen for interstimulus intervals of 10 s, 60 s and 5 min. Response decrement failed to occur in the absence of repetitive stimulation, the decremented response recovered spontaneously following a rest, and electrical stimulation of a body wall nerve facilitated the decremented response (a neural correlate of dishabituation). Adaptation of sensory neuron responses occurred during repeated hair deflections. However, when adaptation was eliminated by direct electrical stimulation of sensory neurons, the response in the proleg motor nerve still decreased significantly. Muscle recordings indicated that the response of an identified proleg retractor motor neuron decreased significantly during habituation training. Thus, habituation of the proleg withdrawal reflex includes a central component that is apparent at the level of a single motor neuron. Accepted: 20 December 1996     

Electrical activity in the chemoreceptors of the blowfly. I. Responses to chemical and mechanical stimulation

The electrical responses of the neurons associated with the various types of chemosensory hairs of the blowfly, Phormia regina Meigen, following stimulation by chemical and mechanical means have been studied. The singly innervated chemosensory hairs on the ovipositor, maxillary palpi, and antennae respond vigorously to chemical stimulation, but not to mechanical stimulation. The triply innervated chemosensory hairs on the labellum, tarsus, and wing have two neurons which respond only to chemical stimuli. The third neuron responds only to mechanical stimulation. The differential responses of the two chemosensory neurons to various chemical stimuli following the removal of the tip of the hair suggest that the structures responsible for chemoreception are located throughout the distal processes of these neurons. The response of the third neuron to mechanical stimulation is similar to the response recorded from the neuron associated with one type of tactile hair which responds to motion and not to steady deformation. Recordings have been made from the neurons associated with purely tactile hairs using the cut hair as an extension of the micropipette. The mechanosensory neuron of the wing chemosensory hair is capable of responding at the rate of at least 600 impulses per sec. and may serve to indicate changes in air flow over the wing surfaces during flight to enable the fly to correct the wing camber and attack angle.     

Auditory input to motor neurons of the dorsal longitudinal flight muscles in a noctuid moth (Barathra brassicae L.)

Summary Motor neurons innervating the dorsal longitudinal muscles of a noctuid moth receive synaptic input activated by auditory stimuli. Each ear of a noctuid moth contains two auditory neurons that are sensitive to ultrasound (Fig. 1). The ears function as bat detectors. Five pairs of large motor neurons and three pairs of small motor neurons found in the pterothoracic ganglia innervate the dorsal longitudinal (depressor) muscles of the mesothorax (Figs. 2 to 5). In non-flying preparations the motor neurons receive no oscillatory synaptic input. Synaptic input to a cell resulting from ultrasonic stimulation is consistent and can be either depolarizing or hyperpolarizing (Figs. 6 to 9). Quiescent neurons only rarely fire a spike in response to auditory inputs. Motor neurons in flying preparations receive oscillatory synaptic drive from the flight pattern generator and usually fire a spike for each wingbeat cycle (Figs. 10 to 12). Ultrasonic stimulation can provide augmented synaptic drive causing a neuron to fire two spikes per wingbeat cycle thus increasing flight vigor (Fig. 11). The same stimulus presented on another occasion can also inhibit spiking in the same motor neuron, but the rhythmic drive remains (Fig. 12). Thus, when the flight oscillator is running auditory stimuli can modulate neuronal responses in different ways depending on some unknown state of the nervous system. Sound intensity is the only stimulus parameter essential for activating the auditory pathway to these motor neurons. The intensity must be sufficient to excite two or three auditory neurons. The significance of these responses in relation to avoidance behavior to bats is discussed.     

Physiological and pharmacological properties of responses to GABA and ACh by abdominal motor neurons in Manduca sexta

1. GABA, ACh, and other agents were applied by pressure ejection to the neuropil of the third abdominal ganglion in the isolated nerve cord of Manduca sexta. Intersegmental muscle motor neurons with dendritic arborizations in the same hemiganglion were inhibited by GABA (Fig. 2) and excited by ACh (Fig. 5).
2. Picrotoxin was a potent antagonist of GABA (Fig. 4A). Bicuculline reduced GABA responses in some motor neurons (Fig. 4C), but had no effect on many other motor neurons. Curare reduced ACh responses (Fig. 6A). Bicuculline was an effective ACh antagonist in most motor neurons tested (Fig. 6B).
3. Motor neurons with dendrites across the ganglion from the ejection pipette exhibited different responses to GABA and ACh. Contralateral motor neurons often showed smaller, delayed hyperpolarizing GABA responses (Fig. 7). On two occasions, contralateral motor neurons had excitatory responses (Fig. 8). Contralateral motor neurons were hyperpolarized by ACh (Fig. 9). The inhibitory responses had only slightly longer latencies than ipsilateral excitatory ACh responses (Fig. 10A). The contralateral inhibitory ACh responses, but not the ipsilateral excitatory ACh responses, were eliminated by TTX (Fig. 10B).
4. A model, which includes inhibitory interneurons that cross the ganglionic midline to inhibit their contralateral homologs and motor neurons (Fig. 11), is proposed to account for contralateral responses to GABA and ACh and antagonistic patterns of activity of motor neurons during mechanosensory reflex responses.

     

Mechanosensory afferents innervating the swimmerets of the lobster

The mechanosensory innervation of the lobster (Homarus americanus) swimmeret was examined by electrophysiologically recording afferent spike responses initiated by localized mechanical stimulation of the caudal surface of the swimmeret. Two functional groups of subcuticular hypodermal mechanoreceptors innervate the swimmeret. Afferents of one group innervate the small discrete "ridges" of calcified cuticle lining the margins of both swimmeret rami. Putative ridge receptors are bipolar sensory neurons responding phasically to deformation of the ridge cuticle with the number and frequency of impulses produced dependent on stimulus strength and velocity. Afferents of the second group, which innervate substantial areas of hypodermis underlying the soft, flexible cuticular regions of the swimmeret, were designated "wide-field" hypodermal mechanoreceptors. These neurons have multiterminal receptive fields and respond phaso-tonically to cuticular distortion. The response properties of both types of hypodermal mechanoreceptors imply that they are activated during the characteristic beating movements of the swimmerets.     

The sensory basis for the trap-jaw mechanism in the ant Odontomachus bauri

Ants of the ponerine genus Odontomachus have evolved a mechanism that allows them to instantaneously close their long mandibles to catch prey or defend themselves. This trap-jaw action is triggered by contact of trigger hairs with a potential prey item. Two of these long mechanosensory hair sensilla reside proximally on each mandible and are supplied by giant sensory cells.Extracellular recordings demonstrate that the sensory cells respond to tactile stimulation. Their phasic responses encode amplitude and velocity of hair-deflection away from the midline, but not hair position. The discharge of action potentials follows stimulus frequencies of more than 300 Hz. During sinusoidal stimulation, the cells adapt very little, sustain discharge rates of more than 200 Hz for more than 20 s, and reach peak spike rates of about 450 Hz.The afferent axons of these sensory cells give rise to huge axon terminals within the suboesophageal ganglion. One of the afferents has a prominent contralateral branch, the other is confined to ipsilateral neuropil. Anatomical data indicate that the 4 afferents may be coupled and may serve as the substrate for a very fast reflex.Abbreviations HRP horseradish peroxidase - LGS lateral giant sensillum - MGS median giant sensillum - SEM scanning electron microscopy - SOG suboesophageal ganglion     

The cord stretch receptors in the abdominal nerve cord of the crayfish Cherax destructor: physiology and relationships

The physiology and relationships of tonic cord stretch receptor neurons in the crayfish Cherax destructor were examined with intracellular and extracellular recording. Cord stretch evoked slow depolarisations leading to action potentials in tonic cord stretch receptor neurons. Intermittent post-synaptic potentials were also seen in cord stretch receptor neurons but were not the primary cause of the action potentials. Cord stretch still evoked action potentials in cord stretch receptor neurons when all synaptic activity, monitored at another known chemical synapse, was blocked using high [Mg(2+)] and low [Ca(2+)] in the bath. One source of facilitating excitatory post-synaptic potentials in the cord stretch receptor neurons was from mechanosensory hairs on the dorsal abdominal surface. Tonic cord stretch receptor neuron activity was associated with an increase in the activity of the abdominal slow extensor inhibitor motor neuron and at least one abdominal flexor excitor motor neuron in its segment, and reduced activity in the abdominal slow flexor inhibitor motor neuron. Activation of individual cord stretch receptor neurons produced a local resistance reflex. Cord stretch, activating many receptors, produced several other outcomes. One was the "extensor state" described in earlier literature. The tonic cord stretch receptor neurons of Cherax destructor appear to be stretch-sensitive interneurons that receive inputs from other elements of the abdominal control system and mediate polysynaptic reflex activity in postural motor neurons.     

Metamorphic changes in the central projects of hair sensilla inTenebrio molitor L. (Insecta: Coleoptera)

Summary This paper describes the afferent projections of hair sensilla of the pro- and mesothoracic legs and the lateral thoracic sclerites of larval and adultTenebrio molitor and the corresponding set of pupal hair sensilla. The sensory neurons that innervate the hair sensilla of larval or adult insects project somatotopically into the thoracic neuropil. Different types of sensilla on the same region of the body surface project to the same zone of the ipsilateral thoracic ventral neuropil but exhibit different arborization patterns. Although there is a profound reorganization of body surface sensilla, the basic somatotopic layout of the larva is maintained in the adult. The sensory neurons that innervate the pupal hair sensilla possess central projections similar to those of the corresponding adult sensory neurons. The central projections of pupal sensory neurons are somatotopically oriented. Their projection pattern is serially homologous in the thoracic and the abdominal ganglia. The central projection pattern of the described pupal sensory neurons is constant throughout pupation. MAb 22C10 immunoreactivity allows an estimate of the timing of the early differentiation of the imaginal sensory neurons originating during pupation. Ablation experiments indicate that pupal sensory neurons influence the central projection pattern of the differentiating imaginal sensory neurons.     

Differential effects of nitric oxide on the responsiveness of tactile hairs

The responses of tactile hairs located on legs of the desert locust Schistocerca gregaria (Forskål) are modulated by nitric oxide (NO). There are two types of tactile hair on the tibia of the hind leg of the locust which differ in their thresholds for mechanical stimulation, their location on the leg and in the effect of NO on their responses to deflection. The spike response rates of mechanosensory neurons of low-threshold hairs decreased when exposed to elevated NO levels caused by perfusion of the leg with saline containing the NO donor PAPANONOate. In contrast, in high-threshold hairs, which show low responsiveness under control conditions, an increase in spike rates was observed during PAPANONOate application. These opposing effects of NO reduce the differences in the spike responses of the two types of tactile hairs to mechanical stimulation and are likely to have an impact on behaviours elicited by mechanical stimulation of the legs.     

Physiology and pharmacology of acetylcholinergic responses of interneurons in the antennal lobes of the mothManduca sexta

1. Neurons in the antennal lobe (AL) of the moth Manduca sexta respond to the application, via pressure injection into the neuropil, of acetylcholine (ACh). When synaptic transmission is not blocked, both excitatory (Fig. 2) and inhibitory (Fig. 3) responses are seen. 2. Responses to ACh appear to be receptor-mediated, as they are associated with an increase in input conductance (Figs. 2B and 3B) and are dose-dependent (Fig. 2 C). 3. All neurons responsive to ACh are also excited by nicotine. Responses to nicotine are stronger and more prolonged than responses to ACh (Fig. 4C). No responses are observed to the muscarinic agonist, oxotremorine (Fig. 4 B). 4. Curare blocks responses of AL neurons to applied ACh, while atropine and dexetimide are only weakly effective at reducing ACh responses (Figs. 5 and 6). 5. Curare is also more effective than atropine or dexetimide at reducing synaptically-mediated responses of AL neurons (Fig. 7). 6. In one AL neuron, bicuculline methiodide (BMI) blocked the IPSP produced by electrical stimulation of the antennal nerve, but it did not reduce the inhibitory response to application of ACh (Fig. 8).     

Control of respiratory motor pattern by sensory neurons in spinal cord of lamprey

Summary Extracellular stimulation over the dorsal funiculus in the spinal cord of lampreys was found to selectively activate prolonged episodes of fictive arousal respiration (Figs. 1, 3). The induced episodes showed comparable increases in cycle frequency and motoneuron burst duration to the spontaneous arousal pattern observed in isolated brain preparations (Fig. 2). Intracellular stimulation of primary sensory neurons with axons in the dorsal funiculus, called dorsal cells, also elicited the arousal pattern (Fig. 4). Mechanoreceptive dorsal cells respond to cutaneous stimulation. When mechanical stimuli were applied to the skin of intact lampreys (Fig. 6) or to lampreys with ipsilateral vagotomy, arousal respiration was induced (Figs. 7, 8). Bilateral, but not unilateral, trigeminal lesion blocked dorsal cell induction of the arousal response (Fig. 5). Spontaneous arousal respiration was recorded from intact, unrestrained lampreys (Fig. 9). These results suggest that fictive arousal respiration is the in vitro correlate of natural arousal respiration in lampreys, and that one mechanism leading to arousal respiration may be the activity of sensory dorsal cells. A model for respiratory motor pattern switching in lamprey is proposed. The model suggests that the normal and arousal patterns are produced by separately engaging rostral or caudal pattern generators in the medulla, rather than by modifying one pattern generator (Fig. 10).     

Labellar lobe spreading in the blowfly: Regulation by taste and satiety

Summary Lobe spreading behavior was studied by recording electromyograms from the muscles which spread the labellar lobes, the retractors of the furca (RF) inPhormia regina. RF responses and lobe spreading could be elicited by stimulating labellar, but not tarsal, taste hairs with sucrose (Fig. 3). RF activity was important to spread the lobes at the beginning of a meal, but was not necessary for continued feeding (Fig. 4).Temporal summation between sugar receptor spikes was necessary to elicit RF responses. Central response decrement occurs independently for different labellar hairs and may participate in the termination of motor responses.RF responses were more probable and more intense when either the sucrose concentration of the stimulus or the number of hairs stimulated was increased (Fig. 7). Stimulation with NaCl had no effect on the response to simultaneous sucrose stimulation of other hairs (Table 1).Feeding caused decreases in the probability and intensity of motor responses, but did not alter chemosensory responses (Figs. 8 and 9). Section of either the recurrent or median abdominal nerves prevented this postingestional inhibition of lobe spreading (Fig. 9).These results are discussed with regard to the possible role that regulation of lobe spreading may play in the control of food intake.This work was supported by United States Public Health Service Training Grant 5T01 GM 00457-13S2 and by a grant from the National Science Foundation to Dr. Vincent G. Dethier. I wish to thank Dr. Dethier for his support and encouragement.