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H M Pinsker

Publications and source records attributed to H M Pinsker.

At least 19 recordsLinked to original sources

Egg laying in Aplysia. I. Behavioral patterns and muscle activity of freely behaving animals after selectively elicited bag cell discharges.

Aplysia egg laying is a complex sequence of head and neck movements initiated by the release of ovulatory and neuroactive hormones from the neurosecretory bag cells. This behavioral pattern is difficult to study in reduced preparations, because they do not show ovulation or egg laying behaviors. This paper describes the use of chronically implanted electrodes to elicit normal neurosecretory activity and provides an analysis of egg laying behaviors and the underlying muscle activity in intact, freely behaving A. californica and A. brasiliana. 1. Bag cell discharges elicited with a fine wire electrode implanted in the connective tissue sheath above the cell bodies were typically without noxious behavioral side effects. 2. Following selectively elicited bag cell discharges, egg laying consisted of four rhythmic head and neck movements that were separated functionally into appetitive behaviors ('waves' and 'undulations') used to explore and prepare the substrate and consummatory behaviors ('weaves' and 'tamps') used to distribute and attach the egg string. The amount of time an animal performed consummatory behaviors was positively related to the amount of eggs deposited. By contrast, the appetitive phase of egg laying was independent of the size of the egg mass. 3. The individual behaviors and their temporal sequence were similar following selectively elicited bag cell discharges, spontaneous discharges of animals with implanted electrodes and during normal egg laying of unoperated animals. 4. Three longitudinal muscle systems occurred within the head and neck. Following a selectively elicited bag cell discharge, spatially and temporally coordinated patterns of EJP bursts of different durations were recorded chronically from each muscle group. These EJP patterns were characteristic for specific head and neck movements used in appetitive and consummatory egg laying behaviors.

Animals↗

Egg laying in Aplysia. II. Organization of central and peripheral pathways for initiating neurosecretory activity and behavioral patterns.

1. Central pathways for bag cell activation were identified by examining the frequency of spontaneous egg laying episodes in animals with central connective lesions. Bilateral lesions of the cerebropleural (but not the cerebropedal) connectives abolished spontaneous egg laying. In contrast, bilateral lesions of all cerebral ganglion peripheral nerves did not abolish spontaneous egg laying, suggesting that sensory input to the cerebral ganglion is not necessary for activating the bag cells. 2. Backfilling either pleuroabdominal connective labelled cell bodies in the cerebral ganglia (via the ipsilateral cerebropleural connective) that could project to the bag cells. Focal extracellular stimulation of these stained clusters activated the bag cells in isolated brains. 3. Central pathways for initiating egg laying behaviors were identified by selectively eliciting bag cell discharges in animals with central connective lesions. Bilateral lesions of the cerebropedal (but not the cerebropleural) connectives completely abolished elicited egg laying behaviors. 4. Pathways for motor output during rhythmic head and neck movements were identified by eliciting bag cell discharges in animals with peripheral nerve lesions. Bilateral lesions of the four tegumentary nerves in combination with the anterior pedal nerve completely abolished elicited egg laying behaviors, indicating that these nerves are necessary for normal motor output. A normal pattern of egg laying behaviors occurred when the four tegumentary and the anterior pedal nerves were left intact and all other pedal ganglion nerves were lesioned bilaterally, indicating that these nerves are also sufficient for normal motor output.(ABSTRACT TRUNCATED AT 250 WORDS)

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Swimming in Aplysia brasiliana: behavioral and cellular effects of serotonin.

1. Aplysia brasiliana is a marine mollusk that swims by repeated metachronal flapping movements of its bilateral fleshy parapodia. Animals with bilateral cerebropedal connective (CPC) lesions do not swim when suspended above the substrate, although tonic CPC stimulation can elicit normal parapodial flapping. Although the parapodial opener-phase (POP) cells, a previously identified group of neurons, fire synchronous bursts of efferent spikes in-phase with parapodial opening movements in both intact animals and dissected preparations, they are not likely to be primary parapodial motoneurons. These cells receive one or more large, apparently monosynaptic excitatory postsynaptic potentials (EPSPs) during CPC stimulation that are effective in producing the swimming motor program (SMP). 2. In suspended CPC-lesioned animals, injections of serotonin (5-HT) that produce an average hemolymph concentration of 10(-5) M induced full-amplitude parapodial flapping. Selected episodes of flapping were similar in frequency to normal suspended swimming. 3. In suspended CPC-lesioned animals, 5-HT injections elicited an apparently normal swimming motor program that was associated with synchronous bursts of large-amplitude efferent spikes in the parapodial nerves. In many semi-intact preparations, exposing the circumoesophageal ganglia to 5-HT elicited a similar rhythmic motor program, but usually at a lower frequency than during normal swimming or during tonic CPC stimulation. 4. In isolated-ganglion preparations, bath application of 5-HT produced immediate depolarization and tonic firing of individual POP neurons, followed by smooth and regular bursting in the apparent absence of synaptic input. In such preparations, the motor program elicited by bath-applied 5-HT differed from the one elicited by tonic CPC stimulation in that the 5-HT-elicited rhythmic bursting usually was not synchronous in different POP neurons. Tonic CPC stimulation during bath applications of 5-HT produced immediate synchronization of bursts among the POP neurons. 5. Hyperpolarization (or depolarization) of a POP neuron during bath application of 5-HT increased (or decreased) the burst period, but membrane polarization did not change the burst period elicited during tonic CPC stimulation.

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Swimming in Aplysia brasiliana: identification of parapodial opener-phase and closer-phase neurons.

1. In freely behaving Aplysia brasiliana, spontaneous swimming in the laboratory occurred primarily in the dark hours of the day-night cycle. Suspending an intact animal above the substrate elicited continuous parapodial flapping with the same frequency and amplitude as spontaneous swimming. Parapodial flapping with decreased frequency and amplitude could still be elicited by suspending minimally dissected, but not more radically dissected, preparations. 2. In otherwise intact animals, severing the cerebropedal connective (CPC) bilaterally abolished suspended parapodial flapping, but normal flapping was elicited by tonic stimulation of the distal CPC. In minimally dissected preparations, tonic CPC stimulation elicited parapodial flapping, but with reduced frequency and amplitude. 3. During normal parapodial flapping, chronically implanted electrodes on parapodial nerves recorded the swimming motor program (SMP). The whole-nerve SMP consisted of rhythmic bursts of large-amplitude efferent units in phase with parapodial opening, with no observable activity during parapodial closing. By contrast, simultaneous electromyogram (EMG) recordings from antagonistic parapodial muscles showed antiphasic bursts of activity during opening and closing. The SMP was inhibited by touching food to the animals' lips. 4. Parapodial nerve backfills, using nickel chloride, labeled several cell clusters in the ipsilateral pedal ganglion. Two of these clusters were located caudally: one tightly clustered medial group had large cell bodies, and another, more distributed, lateral group had small cell bodies. The two clusters were identified in semi-intact preparations and isolated brains, using tonic CPC stimulation to elicit a fictive SMP recorded in parapodial nerves, and intracellular electrodes to characterize and stain individual cells. 5. The large parapodial opener-phase (POP) neurons were normally silent. At the onset of CPC stimulation, POP neurons depolarized and fired tonically, and then burst rhythmically in phase with each other, and one for one with large-amplitude axon spikes observed extracellularly in parapodial nerves during the fictive SMP. Intracellular firing of POP cells, singly or in pairs, never produced observable papapodial movements or one-for-one responses in parapodial muscles. Lucifer yellow-filled POP neurons showed a process (with a pronounced rostral loop) that gave off many short, fine neurites in the pedal neuropile before branching into two or three axons projecting into different parapodial nerves. 6. The smaller parapodial closer-phase (PCP) neurons normally discharged tonically at low frequencies.(ABSTRACT TRUNCATED AT 400 WORDS)

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Chromatophore motor fields in the squid, Lolliguncula brevis.

Chromatophore motoneurones in Lolliguncula brevis are known to originate in the suboesophageal lobes of the brain and to project directly to the mantle and fin through bilateral stellate ganglia and fin nerves. The chromatophore motor fields of stellar and fin nerves were investigated by stimulation of the cut end of individual nerves in a semi-intact preparation. This elicited expansion of yellow and brown chromatophores in distinct motor fields. Brown chromatophores extended over the entire mantle, whereas yellow chromatophores were limited to the dorsal and lateral mantle areas. Combined nerve stimulation and lesions demonstrated substantial overlap between adjacent chromatophore motor fields and innervation of individual chromatophores by different stellar nerves.

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Localization and stimulation of chromatophore motoneurones in the brain of the squid, Lolliguncula brevis.

The relatively simple chromatophore system of the squid, Lolliguncula brevis, was studied with combined behavioural, morphological and electrophysiological methods in order to understand how the chromatophore patterns in the skin are organized at the level of the posterior chromatophore lobes (PCL). There are nine simple chromatic components of patterning in L. brevis. Retrograde transport of horseradish-peroxidase from chromatophores in the mantle skin established that the chromatophore motoneurones are located in the PCL. Focal threshold stimulation of the PCL in perfused, semi-intact preparations showed that the motor fields of individual chromatophore motoneurones are compact, including 2-60 chromatophores, generally of the same colour. Adjacent motoneurones in the lobe do not necessarily have adjacent motor fields in the skin.

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Spontaneous and elicited bag cell discharges in gonadectomized Aplysia.

The neuroendocrine bag cells of the hermaphroditic marine gastropod, Aplysia, secrete peptide hormones that induce release of ripe eggs from the ovotestis. The egg string is subsequently deposited on the substrate by means of a complex sequence of rhythmic head and neck movements. Gonadectomy (removal of the ovotestis) was performed in two closely related species of Aplysia to prevent completely the synthesis, build-up and release of eggs. Chronically implanted electrodes were used either to monitor spontaneous bag cell discharges (A. brasiliana) or to selectively elicit bag cell discharges (A. californica) in gonadectomized and mock-operated animals. Gonadectomized animals showed the normal occurrence of spontaneous bag cell discharges in the complete absence of eggs, indicating that feedback from ripe eggs in the ovotestis is not necessary for normal activation of the bag cells. However, gonadectomized animals showed a significant decrease in specific head and neck movements following elicited bag cell discharges. This finding indicates that, once the bag cells fire and the eggs are released, input from the eggs is necessary for normal expression of the behaviour associated with egg deposition.

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Temperature dependence of egg laying in Aplysia brasiliana and A. californica.

The temperature dependence of egg laying was examined in winter-caught Aplysia. Cold-water Aplysia californica and warm-water A. brasiliana were individually housed in the same large aquarium for 16 days at 15 degrees C, and then for 16 days at 20 degrees C. Initially, the majority of the A. californica were not reproductively mature (as determined by injections of atrial gland extracts) whereas all of the A. brasiliana were reproductively mature. When the temperature was increased from 15 to 20 degrees C, both species showed a marked increase in the frequency of egg laying. At both temperatures, A. brasiliana laid eggs more frequently but produced smaller egg masses than A. californica. We conclude that increased egg laying in A. californica was attributable both to facilitation of oogenesis in previously reproductively immature animals and to increased activity of the bag cells which release an egg-laying hormone. Increased egg laying in A. brasiliana was attributable primarily to increased bag cell activity.

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Neuroethological studies of reflex plasticity in intact Aplysia.

Multi-unit recording of siphon nerve activity in intact Aplysia california with chronically implanted cuff electrodes provided a monitor of activity in a central pattern generator, the Interneuron II (Int II) network, which produces large siphon and gill contractions both spontaneously and after tactile stimulation of the siphon. The phase-response curve of the Int II oscillator for single stimuli at different phases of the cycle showed a "refractory" period early in the cycle after which most stimuli phase advanced the oscillator and caused a short-latency Int II burst and a large contraction. The amplitude of gill withdrawal and the duration of siphon withdrawal in response to different stimulus intensities depended on whether an Int II burst was triggered. Activation of the Int II oscillator transformed the reflex from one that was graded smoothly with stimulus intensity to one in which nearly maximal responses were elicited even by weak stimuli. Entrainment and habituation training both involved monotonous repetition of a stimulus at specific intervals. With repeated siphon stimuli, nearly maximal reflex responses were maintained in intact animals as long as the Int II oscillator was entrained, whereas habituation was associated primarily with failure to entrain the oscillator. Long-term sensitization of the reflex was characterized by large and prolonged withdrawal responses. Sensitized animals showed significantly more triggered Int II bursts than did controls. In addition, digital spike-train analysis indicated that individual siphon motoneurons showed significantly increased background activity which often persisted for several minutes.

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Selective recording and stimulation of individual identified neurons in freely behaving Aplysia.

A neuroethological technique is described for selective recording and stimulation of an individual neuron in freely behaving Aplysia by means of a fine wire glued into the connective tissue sheath above the identified cell body. A whole-nerve cuff electrode simultaneously monitored functionally related multiunit axon activity. For biophysical analysis the soma was impaled with a microelectrode when the ganglion was subsequently exposed. The technique is illustrated for several identified neurons involved in different behaviors.

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Phase plane description of endogenous neuronal oscillators in Aplysia.

Phase plane techniques are used to describe graphically the limit cycle behavior of identified endogenous neuronal oscillators in the isolated abdominal ganglion of Aplysia. Intracellularly recorded membrane potential from a bursting neuron and its first derivative with respect to time are used as coordinates (state variables) in phase space. The derivative is either measured electronically or calculated digitally. Each trajectory in phase space represents the entire output of the bursting neuron, i.e., both the rapid action potentials and slow pacemaker potentials. Phase plane portraits are presented for the free run limit cycle before and after a change in a system parameter (applied transmembrane current) and also for phase resetting produced by direct synaptic inhibition from an identified interneuron. The complex topology of the trajectory suggests that the bursting oscillator is a higher order system. Therefore, the second time derivative is used as another state variable. This type of phase plot can help to relate biophysical and mathematical analyses.

Abdomen↗

Neuronal correlates of siphon withdrawal in freely behaving Aplysia.

1. Central neuronal mechanisms of siphon withdrawal in Aplysia were studied for the first time in intact, freely behaving animals by means of population recordings from implanted whole-nerve cuff electrodes. Intracellular follow-up studies were then conducted when the same animal was reduced to a semi-intact preparation. 2. Background spontaneous activity in the siphon nerve consisted of low-frequency firing of a population of efferent units containing identified siphon motoneurons. 3. Spontaneous patterned bursts of efferent activity occurred irregularly and were associated with all-or-nothing contractions of the parapodia, gill, and siphon. Spontaneous bursts were due to centrally generated activity in the interneuron II (INT II) network, an oscillatory network with endogenous pacemaker properties. 4. In intact animals, even weak tactile stimuli to the siphon typically triggered an INTII burst shortly after the stimulus-locked efferent activity. Thus, the stimulus can phase-advance the INT II oscillator. In semi-intact preparations, short-latency INT II bursts were triggered less less frequently and required more intense stimuli. 5. With weak to moderate-intensity stimuli in intact animals, the presence of short-latency triggered INT II bursts largely determined the duration of the siphon component and amplitude of the gill component of the withdrawal reflex. 6. When stimuli were repeated over a range of interstimulus intervals (from 60 to 1 min), the likelihood of triggering a short-latency INT II burst die not change systematically. Thus, the ability of the siphon stimulus to stably entrain the all-or-none INT II component over a wide range of intervals will interact behaviorally with the decrement of the monosynaptic component of the reflex with repetition.

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In vivo responses of paired giant mechanoreceptor neurons in Aplysia abdominal ganglion.

Two neurons with cell bodies symmetrically located in the abdominal ganglion and giant axons in the left (L1) and right (R1) pleurovisceral connectives of Aplysia californica were examined in vivo and in vitro. Direct stimulation of R1 and L1 in the intact animal does not elicit any observable behavior, suggesting that they are neither motoneurons nor command neurons. These cells respond in vivo to sudden onset mechanical stimulation of widespread regions of the body. R1 and L1 spikes are initiated in at least three different loci: (1) the peripheral axon in the foot, (2) the neuropil of the pleural and/or pedal ganglion, and (3) the neuropil of the abdominal ganglion. Furthermore, R1 and L1 probably have two different mechanisms for spike initiation: (1) sensory (foot), and (2) synaptic (abdominal and/or head ganglia). The different loci for spike initiation account for the bidirectional conduction of R1 and L1 spikes. As sensory (mechanoreceptor) neurons, R1 and L1 have peripheral axons in the ipsilateral posterior pedal nerve, show low threshold responses to stimulation of the ipsilateral posterior foot, they are rapidly adapting their responses do not decrease with repetition, and they are not blocked by high Mg++/low Ca++ solutions. As synaptically-driven neurons, R1 and L1 have widespread bilateral responsiveness, their responses decrease with repetition and their inputs are blocked with high Mg++/low Ca++ solutions. These neurons integrate sensory and synaptic inputs and conduct bidirectionally, however, their output connections must be specified before their behavioral function can be understood.

Action Potentials↗

Short-term modulation of endogenous bursting rhythms by monosynaptic inhibition in Aplysia neurons: effects of contingent stimulation.

A presynaptic neuron fires a high-frequency train of spikes that produces long-lasting synaptic inhibition that modulates the bursting rhythm in a small population of endogenous bursting neurons in the left upper quadrant of the isolated abdominal ganglion of Aplysia. Single inputs decrease or increase the duration of the burst cycle as a function of the precise phase of the input (the phase response curve). Two phases of the burst cycle were used to analyze the effects of repeated contingent (phase-locked) stimulation. One contingency involved synaptic input early in the burst cycle that inhibited spikes and decreased the duration, whereas the other contingency involved input late in the cycle that increased the duration. Under both contingencies of stimulation, buildup and short-term persistence were found, however these cumulative effects were not dependent upon the phase of the burst cycle. The locus of the short-term plasticity that underlies the buildup and persistence is in the pacemaker properties of the postsynaptic cell rather than in the synapse. The plastic change appears to involve a nonspecific postinhibitory rebound that follows a single input and builds up with repetition. These results support the suggestion that endogenous rhythms of pacemaker cells can undergo plastic changes and can therefore serve as a means of short-term information storage in the nervous system. However, this neuronal circuit does not have the specificity required to mediate operant conditioning.

Action Potentials↗

Aplysia bursting neurons as endogenous oscillators. I. Phase-response curves for pulsed inhibitory synaptic input.

1. The left upper quadrant bursting neurons in the abdominal ganglion of Aplysia are isochronous, nonlinear oscillators. Transmembrane current and temperature are parameters of the bursting oscillator. 2. The phase-response curve (PRC) for pulsed inhibitory synaptic input from an interneuron describes the phase shift produced by synaptic input at different phases of the burst cycle. 3. The characteristic shape of the PRC consists of two linear functions that intersect at the point in the cycle where the burst of spikes ends. Whether the net effect of the synaptic input at a given phase is phase advance or phase delay depends on 1) the number of spikes inhibited, and 2) the duration of the inhibition relative to the duration of the free-run period. 4. The shape of the PRC remains constant when a stepwise change in a parameter is introduced, when the duration of the synaptic input is increased, when the fast component of the IPSP is blocked, and when a long hyperpolarizing pulse is used to mimic the slow IPSP. 5. The shape of the PRC is changed when short hyperpolarizing pulses or antidromic action potentials are used and when only the pacemaker oscillation is present in the bursting neuron. 6. Therefore, the synaptic modulation of the bursting rhythm is determined by the voltage change produced by the IPSP and its inhibition of spikes in the bursting neuron.

Action Potentials↗