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C H Page

Publications and source records attributed to C H Page.

At least 19 recordsLinked to original sources

Motoneuron reinnervation of phasic uropod muscles in crayfish.

Motoneuron reinnervation of the lateral abductor and adductor muscles in the exopodite of the crayfish uropod was obtained by cross-tying the cut proximal ends of one set of uropod nerve roots to the cut distal ends of their contralateral homologues. In normal (nonsurgically treated) animals the abductor was innervated by two excitatory motoneurons and an inhibitor while two excitors innervated the adductor. For each muscle one of the excitors produced large excitatory junction potentials (EJPs) while the other evoked small EJPs. When stimulated repetitively only the smaller EJP in the adductor generated a facilitating response. Within less than 10 weeks postsurgery the muscles were each reinnervated by two excitatory motoneurons. While the abductor motoneurons generated synaptic potentials with similar amplitudes and time courses to those of normal animals, they differed from those in normal animals in that they facilitated when stimulated repetitively. In contrast to the large and small EJPs evoked in the normal animal, the two motoneurons that reinnervated the adductor muscle elicited similar amplitude EJPs, neither of which facilitated in response to repetitive stimulation.

Animals↗

Mechanosensory afferents innervating the swimmerets of the lobster. I. Afferents activated by cuticular deformation.

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.

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Mechanosensory afferents innervating the swimmerets of the lobster. II. Afferents activated by hair deflection.

Feathered hair sensilla fringe both rami of the lobster (Homarus americanus) swimmeret. The sensory response to hair displacement was characterized by recording afferent impulses extracellularly from the swimmeret sensory nerve while deflecting sensilla with a rigidly-coupled probe or controlled water movements. Two populations of hairs were observed: "distal" hairs localized to the distal 1/3 of each ramus and "proximal" hairs near its base. Distal hairs are not innervated by a mechanosensory neuron but instead act as levers producing strain within adjacent cuticle capable of activating a nearby hypodermal mechanoreceptor. Hair deflections of 25 degrees or more are required to evoke an afferent response and this response is dependent on hair deflection direction. The frequency and duration of the afferent discharge evoked are determined by the velocity of hair displacement. Each proximal hair is innervated by a single mechanosensory neuron responding phasically to hair deflections as small as 0.2 degrees in amplitude. Deflection at frequencies up to 5 Hz elicits a single action potential for each hair movement; at higher frequencies many deflections fail to evoke an afferent response. These sensilla, which are mechanically coupled, may be activated by the turbulent flow of water produced by the swimmerets during their characteristic beating movements.

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Abdominal postural motor responses initiated by the muscle receptor organ in lobster depend upon centrally generated motor activity.

1. Stretch stimulation of the abdominal muscle receptor organ of the lobster Homarus americanus initiated spike discharge of its tonic sensory neuron (SR1). This sensory response evoked a series of tonic postural reflex responses in the motor neurons that innervate the superficial extensor and flexor muscles of the abdominal postural system. The type of motor response depended on whether a flexion or extension pattern of spontaneous activity was being generated by the postural efferents. Spontaneous shifts between these centrally generated motor activities completely changed the SR1-evoked reflex responses. 2. During spontaneous centrally initiated flexion activity, tonic SR1 neuron discharge elicited an assistance response that included excitation of a medium-sized flexor excitor (f3) and the peripheral extensor inhibitor (e5), and inhibition of at least one extensor excitor. Neither the other flexor excitors nor the peripheral flexor inhibitor (f5) were affected by SR1 excitation. 3. During spontaneous centrally initiated extension activity, SR1 activity elicited a response that included excitation of the extensor excitors and the flexor peripheral inhibitor (f5) only, f3 and e5 spontaneous activities were unchanged. This response was a resistance reflex, since SR1 discharge normally resulted from an imposed abdominal flexion. 4. The SR1-initiated control of postural motor activity in lobster differs from previously published results in the crayfish Procambarus clarkii.

Abdomen↗

Sensory inhibition of flexion producing interneurons in lobster abdomen.

Mechanosensory stimulation of an abdominal swimmeret initiates a fictive extension which includes flexion inhibition. The role of flexion producing interneurons (FPIs) in this motor program was examined by recording from a pair of FPIs which excite flexor motor neuron f3. The afferent-FPI-f3 pathway includes at least 5 levels of neural activity. Since swimmeret stimulation hyperpolarizes the FPIs, and mechanosensory afferents are not known to be inhibitory, sensory interneurons must connect the afferent with the FPIs. The generalized receptive fields and long latencies of the FPI response support polysynaptic afferent-FPI connections. The poor correlation between f3FPI spikes and f3 EPSPs they evoke, and the 15 ms delay for initiation of these EPSPs, suggest that additional premotor interneurons are interposed between f3FPI and f3. Since restricted stimulation of swimmeret sensilla generates IPSPs in f3 without affecting f3FPI activity, f3FPI and f3 must be inhibited by different interneurons. Sensory evoked FPI inhibition contributes to the flexion inhibition component of the swimmeret evoked responses since hyperpolarization of f3FPI to block f3FPI spiking during spontaneous flexion activity decreases ongoing f3 spike discharge. Coupling between f3FPI and f3 activities during spontaneously initiated postural flexions supports this conclusion.

Abdomen↗

GABA mediated inhibition of abdominal postural flexion in lobster.

Swimmeret mechanostimulation initiates an abdominal extension program which includes flexion inhibition. Agonists and antagonists were used to examine the GABAergic nature of inhibitory responses recorded intracellularly from a flexion producing interneuron (FPI 303) and flexor motor neuron (f3) pair, and extracellularly from the other flexor efferents. The GABA antagonist picrotoxin (PTX) enhanced spontaneous flexion. As PTX levels increased, the swimmeret evoked response shifted from inhibition of flexion (less than 10 microM), to inhibition followed by excitation (10-30 microM), to flexion excitation (greater than or equal to 50 microM). The irreversibility of PTX effects, and the absence of bicuculline or baclofen induced changes in flexion activity, suggests that the receptors differ from mammalian GABA receptors. Both GABA and its agonist muscimol suppressed flexion activity and reduced intracellular potential amplitudes. Proof that PTX acts by binding the GABA receptor was obtained by observing that the addition of GABA or muscimol to preparations pretreated with PTX did not affect either spontaneous or swimmeret evoked activities, or intracellular potential amplitudes. These results imply involvement of GABAergic interneurons in the abdominal motor programs which inhibit flexion.

Abdomen↗

Intersegmental modulation of abdominal postural responses initiated by mechanostimulation of the swimmeret in lobster.

In a multiganglionic preparation of the lobster abdominal nerve cord, composed of the first through fifth ganglia (A1-A5) and attached second swimmeret, tactile stimulation of the cuticular surface of the swimmeret initiates a postural motor program in A2 for abdominal extension, whereas deflection of feathered hair sensilla that fringe the swimmeret rami does not affect postural motor activity recorded from A2 (Kotak and Page, 1986a). This report demonstrates that partial isolation of A2 from adjacent abdominal ganglia by sectioning the A1-A2 or the A2-A3 connectives both increases the strength of the extension response evoked by cuticular stimulation and disinhibits a postural flexion inhibition response initiated by feathered hair stimulation. Complete isolation of A2, by cutting the A1-A2 and the A2-A3 connectives, further increases the strength of these postural responses. Intersegmental inhibition of these responses originates in the ganglia adjacent to A2, since mechanoresponsiveness of A2 is not affected by resection of a more distant connective (A3-A4). These results provide evidence for the presence in adjacent abdominal ganglia of intersegmental interneurons that regulate the access of swimmeret sensory activity to the postural motor neurons in A2.

Abdomen↗

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.

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The motor program and role of sensory feedback in claw extension of the crayfish.

The motor program and role of the meropodite sensory organs (the chordotonal organs MC-1, MC-2 and the myochordotonal organ MCO) in extension of the merocarpodite (M-C) joint of the claw was examined during an antennal-evoked defense response. Recordings from the meropodite extensor muscle indicated that the tonic extensor motoneuron was primarily responsible for M-C joint extension. The phasic extensor motoneuron was co-activated with the tonic motoneuron in only 48% of the responses. Tonic motoneuron discharge was elevated in those responses in which the phasic motoneuron was active. Inactivation of individual meropodite sensory organs did not alter the tonic motoneuron response. Simultaneous inactivation of all 3 organs reduced tonic motoneuron discharge by 47%. Phasic motoneuron discharge was reduced following inactivation of the individual MC-1, MC-2 and MCO organs as well as in the sham operated group. Following inactivation of all 3 organs phasic motoneuron activity ceased. These results suggest that feedback from the M-C sensory organs during an active extension response is positive and redundant.

Astacoidea↗

Sexually dimorphic mechanosensitive swimmeret sensilla affect abdominal posture in the lobster.

The sensilla on the male and female second swimmerets are sexually dimorphic. Female swimmerets contain many long "smooth hairs" (long simple setae) on the coxa and rami. The endopodite of the male swimmeret has an accessory lobe covered with short "bristly spines" (serrate setae). In both sexes the swimmeret rami are lined by "feathered hairs" (plumose setae). The influence of mechanosensory stimulation of these sensilla upon abdominal tonic motor activity was analyzed in an in vitro swimmeret-nerve cord preparation. Movement of several clusters of smooth hairs produced an abdominal extension program by exciting the flexor inhibitor f5, inhibiting the flexor excitors, and activating several extensors. Stimulation of the male bristly spines excited the medium-sized flexor excitors f3 and f4. In both sexes the feathered hairs did not generate any response to mechanical stimulation. We infer that in nongravid females the smooth hairs are involved in receiving mechanosensitive cues to support abdominal extension. Bristly spines may contribute to postural adjustments that assist mating. The long latencies of these responses and their propagation to adjacent ganglia suggest that they are mediated by postural interneurons rather than by direct afferent terminations on postural motoneurons.

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Tactile stimulation of the swimmeret alters motor programs for abdominal posture in the lobster Homarus americanus.

The influence of mechanosensory stimulation of a second segment swimmeret upon the abdominal postural program was examined in an isolated abdominal nerve cord-swimmeret preparation. The swimmeret was stimulated in several different ways to assess the extent of influence exerted on abdominal positioning. Localized tactile stimulation of the swimmeret surface with a mechanical probe usually generated flexion inhibition where the flexor inhibitor (f5) was activated while the small and medium flexor excitors were inhibited. Flexion inhibition was much stronger in females than males. In 50% of the animals a weak flexion excitation was seen. After 3-6 hours the response of one-third of these preparations changed to flexion inhibition. Strong manual stimulation of the swimmeret surface inhibited all of the flexor excitors (f1, f2, f3, f4, and f6) while exciting the inhibitor f5 and increasing extensor activity. Similar extension responses were observed in both sexes. Repeated tactile stimulation of the swimmeret surface elicited a response similar to that evoked during manual stimulation. The strongest extension response was produced at 2 Hz which falls within the normal range of swimmeret beating in intact lobsters. Similar extension responses were also obtained during spontaneous swimmeret beating and rhythmic manual movement of the swimmeret.(ABSTRACT TRUNCATED AT 250 WORDS)

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Postural interneurons in the abdominal nervous system of lobster. I. Organization, morphologies and motor programs for flexion, extension and inhibition.

Using intracellular recording and dye-filling techniques, a survey of postural interneurons was undertaken by impaling their somata in the 2nd abdominal ganglion of lobster. During the course of study approximately fourty different intersegmental interneurons in this ganglion were sampled. Of these, 8 evoked unique, patterned responses in the postural (superficial) motoneurons; each could be identified morphologically. Five of the 8 interneurons had caudally directed axons; 4 of these projected beyond the 4th abdominal ganglion. The remainder projected rostrally, beyond the 1st abdominal ganglion. The postural interneurons were classified according to the motor program they elicited. Five were flexion producing interneurons (FPIs), one was extension producing (EPI), and two generated only inhibitory motor outputs. All motor responses were bilateral and occurred in several segments, including A2. Two neurons, FPIs 201 and 301, produced the full motor reciprocity that typically is observed when flexion command fibers are stimulated. However, three of the FPIs and the single EPI did not express complete reciprocity in synergistic and antagonistic motoneurons. The results indicate that some interneurons displaying all of the properties of command neurons are located entirely within the abdominal nervous system. The overall organization of posture-evoking interneurons appears to be similar to that found in crayfish, suggesting an even more fundamental homology in the neuronal connectivities of these two species than has been established previously.

Abdomen↗

Postural interneurons in the abdominal nervous system of lobster. II. Evidence for neurons having both command and driver roles.

The nature of the synaptic relationship between 7 identified postural interneurons and 5 pairs of superficial motoneurons was examined by obtaining dual intracellular recordings from interneuron-motoneuron pairs in the lobster 2nd abdominal ganglion. For six different interneuron-motoneuron pairs EPSPs recorded from motoneurons occurred with a short (1 to 3 ms) fixed latency following each presynaptic spike recorded from the interneuron. This suggests that there is a monosynaptic relationship between these interneurons and motoneurons. Monosynaptic pathways accounted for 27% of all excitatory connections. Preliminary evidence indicates that the monosynaptic potentials are mediated by an excitatory chemical synapse since: all IPSPs occurred with latencies greater than 5 ms, there was no evidence for electrical coupling, and one of the interneurons produced facilitating PSPs. A majority of all monosynaptic connections were made by two of the flexion producing interneurons (FPIs), 201 and 301. The synaptic outputs of these FPIs were similar in that both made monosynaptic connections with a different bilaterally homologous pair of motoneurons. Both also produced larger EPSPs and more vigorous spiking in contralateral members of the bilateral motoneuron pairs. A previous study demonstrated that interneurons 201 and 301 are the only postural interneurons yet identified that express motor programs indistinguishable from command neurons. Taken together, these results suggest that certain intersegmental interneurons share properties with command neurons and driver neurons, and that there may not be a sharp morphological or functional distinction between these two cell types.

Abdomen↗

Postural interneurons in the abdominal nervous system of lobster. III. Pathways mediating intersegmental spread of excitation.

The multisegmented abdomen of crayfish and lobster assumes a variety of postures as components of different behavioral acts. Experimentally these postures can be maintained by activating any of a number of premotor positioning interneurons. The pathways by which the motor output in two or more segments is coordinated were here investigated for a small group of identified postural interneurons whose somata lie in the 2nd abdominal ganglion (A2). Stimulation of all postural interneurons examined evokes a motor output in other abdominal ganglia through which the axon of the neuron passes as well as in the ganglion of origin (ganglion containing the neuron's cell body). The spread of motor excitation away from the originating ganglion occurs via two general pathways. In the first pathway connections to postural motoneurons are made directly by processes of the postural interneuron which pass into ganglia distal to the originating ganglion. Examples of this are shown for two flexion producing interneurons (FPIs) 201 and 301. Each of these FPIs makes monosynaptic connections with motoneurons in A2 and with a homologous set of motoneurons in A3. All postural interneurons fired a set of corollary discharge interneurons (CDIs) whose activities were recorded from the abdominal connectives. Two FPIs, 202 and 301, and a third interneuron, 503, produced motor outputs in ganglia to which they did not project. The motor specificity established in A2 by stimulation of FPIs 202 and 301 (whose axons pass caudally) was preserved in more rostral ganglia, such as A1. Therefore, different sets of CDIs can be specifically recruited to spread the same motor program that is initiated in the originating ganglion to ganglia that do not receive projections from the stimulated postural interneuron. CDIs, in addition, have the capacity to elicit motor programs in distal ganglia that are markedly different from that expressed in the ganglion of origin. For example, although 503 produced an inhibitory output in the abdominal ganglia that it innervated (A1 and A2), a flexion response was generated by it in more caudal ganglia. The caudal flexion response was mediated in part through a monosynaptic activation of FPI 201 and through other unidentified CDIs. Thus, the interneuronal circuitry for postural control is composed of numerous components, some of which have regional control over different portions of the abdominal nerve cord. Depending upon the required movement, select components are coactivated, either serially or in parallel, to effect a variety of spatially distinct positions.

Abdomen↗

Sexual dimorphism of pleopod motor neurons in lobster.

Sexual dimorphism of the efferent neurons that innervate the pleopods in the first abdominal segment of the lobster was examined with nickel/cobalt backfilling techniques. An average of 28.8 somata (maximum of 31) were stained in the female while the mean number of male efferent somata was 25.8 (maximum of 28). These differences are statistically significant. All except one of these somata are situated close to the base of the ipsilateral first root. They are organized into large anterior (13-21 somata) and small posterior (8-13) clusters. One soma is located on the contralateral side of the ganglion.

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Role of the hemigiant neurons in the crayfish defense response.

The medial and lateral hemigiant neurons of the crayfish were isolated in the circumesophageal connective and electrically stimulated to examine their ability to elicit a defense response. In contrast to previous reports, neither a defense response nor any other thoracic or abdominal movements were observed following stimulation of either hemigiant. Small fibers that lie adjacent to the hemigiants in the circumesophageal connective evoked defense responses when they were electrically stimulated. Two types of defense responses could be evoked by stimulation of small fibers.

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Differential backfilling of interneuron populations based upon axon projections in a lobster abdominal ganglion.

Intersegmental interneurons in the lobster second abdominal ganglion were differentially stained by simultaneously backfilling their axons in the anterior and posterior hemiconnectives with nickel and cobalt ions. When precipitated with rubeanic acid the nickel and cobalt formed different colors in the cell bodies depending upon the location of the axon(s) in the connectives. In the most completely stained preparations 55 ascending, 56 descending, and 25 bidirectional interneurons were observed. Soma diameters ranged from 15-120 micron. One of the somata was located in the connective. The differential staining technique has advantages over conventional backfilling techniques for examining the morphological relationships of populations of neurons having different axon projections.

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Anatomical organization of neurons descending from the supraesophageal ganglion of the lobster.

A population of somata in the supraesophageal ganglion of the lobster was filled by retrograde diffusion of cobalt through the circumesophageal connectives. These somata are located predominantly in the dorsal anterior median region of the protocerebrum. They are organized into a contralateral and two ipsilateral clusters. These 3 clusters include approximately one-half of the neurons in the dorsal anterior median region.

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