Behavioral switching of biting and of directed head turning in Aplysia: explorations using neural network models.
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Animals↗
Biomedical subjects
S C Rosen
Publications and source records attributed to S C Rosen.
Identification and characterization of cerebral-to-buccal interneurons implicated in the control of motor programs associated with feeding in Aplysia.
We identified candidate neurons in the cerebral ganglion that regulate feeding responses mediated by the buccal ganglion. Backfilling the cerebral-buccal connectives revealed that each cerebral hemi-ganglion contains approximately 20 neurons that project axons to the buccal ganglion. Three M-cluster neurons (CBI-1, CBI-2, CBI-3) and one E-cluster neuron (CBI-4) were identified as cerebral-to-buccal interneurons (CBIs) based on position, morphology, synaptic connections, and ability to drive buccal motor programs (BMPs). CBI-1 responds to touch of the tentacles, lips, and buccal mass. It receives monosynaptic EPSPs from interganglionic, cerebral-to-buccal mechanoafferent (ICBM) neurons and monosynaptically excites buccal cells, some of which are also excited by the ICBMs. Tonic firing of CBI-1 usually evokes a single cycle of BMP activity. CBI-1 phase-shifts the rhythmic BMP driven by firing a dopaminergic neuron in the buccal ganglion. CBI-1 itself exhibits dopamine-like histofluorescence following formaldehyde-glutaraldehyde fixation. CBI-2 is excited by food stimuli applied to the lips. Constant-current intracellular stimulation of CBI-2 produces phasic firing of the cell that reliably evokes a rhythmic BMP that incorporates buccal and cerebral motor neurons, putative pattern-generating and pattern-initiating neurons, and neuromodulatory cells (metacerebral cells). CBI-4 also evokes a rhythmic BMP, but the details of its actions and synaptic effects differ from that of CBI-2. CBI-3 does not evoke a BMP, even though it is excited by food stimuli applied to the lips, and it makes monosynaptic connections (both excitatory and inhibitory) to many follower cells of the other CBIs. Firing of CBI-3 phase-delays the BMP driven by CBI-2. Since its activity is incorporated into BMPs and it provides direct inputs to elements of the feeding circuitry, it may play a role in pattern generation. The distinctive features of the CBIs suggest that the consummatory phase of feeding may be controlled by a population of interneurons that subserve different roles.
Animals↗
Purification and sequencing of neuropeptides contained in neuron R15 of Aplysia californica.
R15 is a large identified neuron present in the abdominal ganglion of the mollusc Aplysia. Previous studies have indicated that this neuron may play a role in water balance and possibly renovascular functions. A peptidic factor contained in the neuron R15 has been shown to increase the water content of Aplysia. To determine the structure of the peptides contained in R15, we purified the extracts of 820 R15 cells by means of two steps of reverse-phase HPLC. The purification yielded a number of peptides, only one of which, R15 alpha 1, resulted in water uptake when injected into animals. Determination of the amino acid content and sequence analysis of the R15 alpha 1 peptide demonstrated that this peptide contains 38 residues, including two cysteines. The peptide failed to react with iodoacetate, indicating that the two cysteines are connected by a disulfide bridge. To confirm the assigned structure, the peptide was synthesized with a disulfide bridge. The chromatographic properties and bioactivity of the synthetic material were identical to those of the native peptide. Several other R15 peptides were inactive in the bioassay for water uptake. The sequence of one of these peptides (R15 beta) was determined, and it was established that the peptide contains 28 residues. Amino acid analysis of three other peaks was performed. One of these peaks contained a peptide (R15 beta f) whose amino acid composition suggests that it is a fragment of the R15 beta peptide. The other two peaks contained peptides with identical amino acid compositions, suggesting that they are variants of a single peptide (R15 gamma). The amino acid sequences of all the peptides identified in neuron R15 correspond to stretches of a polyprotein encoded by a recently sequenced R15 cDNA.
Amino Acid Sequence↗
Selective modulation of spike duration by serotonin and the neuropeptides, FMRFamide, SCPB, buccalin and myomodulin in different classes of mechanoafferent neurons in the cerebral ganglion of Aplysia.
An examination of the cellular properties and synaptic outputs of mechanoafferent neurons found on the ventrocaudal surface of the cerebral ganglion of Aplysia indicated that the cerebral mechanoafferent (CM) neurons are a heterogeneous population of cells. Based on changes in action potential duration in response to bath applications of 5-HT in the presence of TEA, CM neurons could be divided into 2 broad classes: mechanoafferents whose spikes broaden in response to 5-HT (CM-SB neurons) and mechanoafferents whose spikes narrow in response to 5-HT (CM-SN neurons). Morphological and electrophysiological studies of the CM-SN neurons indicated that they were comprised of previously identified interganglionic cerebral-buccal mechanoafferent (ICBM) neurons and a novel set of sensory neurons that send an axon into the LLAB cerebral nerve and have perioral zone receptive fields that are similar to those of ICBM neurons. Changes in spike width due to 5-HT were correlated with changes in synaptic output as indicated by the magnitudes of EPSPs evoked in postsynaptic neurons. Electrical stimulation of cerebral nerves and connectives also produced spike narrowing or broadening, and the sign of the effect was a function of the parameters of stimulation. Both heterosynaptic facilitation and heterosynaptic depression of EPSPs evoked in follower cells could be demonstrated. A variety of putative neuromodulators other than 5-HT were also found to affect the duration of action potentials in both classes of CM neurons. FMRFamide had effects opposite to that of 5-HT. SCPB and a recently characterized Aplysia neuropeptide, buccalin, broadened the spikes of both CM classes. Another neuropeptide, myomodulin, decreased the duration of CM-SB neuron spikes but had no effect on CM-SN spikes. Since the CM neurons appear to mediate a variety of competing behaviors, including feeding, locomotion, and defensive withdrawal, the various neuromodulator actions may contribute to the mechanisms whereby behaviors are selected and modified.
Action Potentials↗
The role of a modulatory neuron in feeding and satiation in Aplysia: effects of lesioning of the serotonergic metacerebral cells.
Food-induced arousal in Aplysia is characterized by a progressive increase in the speed and strength of biting responses elicited by a seaweed stimulus. Data from semi-intact and dissected preparations suggest that the identified, serotonergic, metacerebral cells (MCCs) of the cerebral ganglion contribute to food-induced arousal by enhancing the strength of buccal muscle contractions, and by modulating the output of the central pattern generator for biting movements. In order to test this hypothesis in intact, free-moving animals and to determine if the MCCs play a role in satiation of feeding, the behavior of animals that had their MCCs destroyed by intracellular injection of proteases was compared with that of B Cell-Lesion and Dye injection control animals (Experiment 1) or surgical control animals (Experiment 2). Nonfeeding behaviors such as defensive withdrawal responses, locomotion, and righting reflexes were unaffected by MCC lesioning. Also unaffected by MCC lesioning were appetitive feeding behaviors and the amount of food needed to satiate animals. Significant behavioral deficits in consummatory feeding behaviors, which remained stable for periods exceeding 10 d, were observed in the MCC-lesioned animals but not in controls. Lesioned animals exhibited a slowing of rate of repetitive biting responses by 40% of controls and had reduced magnitudes of repetitive bites, particularly at the end of a testing run of 10 consecutive bites. The deficit in bite magnitude was minimally evident in food-deprived animals (Experiment 1) but became more pronounced as animals were fed to satiation (Experiment 2). MCC-lesioned animals still exhibited a residual build-up of the rate and magnitude of biting responses at the onset of feeding behavior. This suggests that, in addition to the MCCs, there are other sources of modulation that contribute to plasticity of consummatory responses during the food-induced arousal state.
Animals↗
Response properties and synaptic connections of mechanoafferent neurons in cerebral ganglion of Aplysia.
1. The cells of two clusters of small neurons on the ventrocaudal surface of each hemicerebral ganglion of Aplysia were found to exhibit action potentials following tactile stimuli applied to the skin of the head. These neurons appear to be mechanosensory afferents since they possess axons in the nerves innervating the skin and tactile stimulation evokes spikes with no prepotentials, even when the cell bodies are sufficiently hyperpolarized to block some spikes. The mechanosensory afferents may be primary afferents since the sensory response persists after chemical synaptic transmission is blocked by bathing the ganglion and peripheral structures in seawater with a high-Mg2+ and low-Ca2+ content. 2. The mechanosensory afferents are normally silent and are insensitive to photic, thermal, and chemical stimuli. A punctate tactile stimulus applied to a circumscribed region of skin can evoke a burst of spikes. If the stimulus is maintained at a constant forces, the mechanosensory response slowly adapts over a period of seconds. Repeated brief stimuli have little or no effect on spike frequency within a burst. 3. Approximately 81% of the mechanoafferent neurons have a single ipsilateral receptive field. The fields are located on the lips, the anterior tentacles, the dorsal portion of the head, the neck, or the perioral zone. Because many cells have collateral axons in the cerebral connectives, receptive fields elsewhere on the body are a possibility. The highest receptive-field density was associated with the lips. Within each area, receptive fields vary in size and shape. Adjacent fields overlap and larger fields frequently encompass several smaller ones. The features of some fields appear invariant from one animal to the next. A loose form of topographic organization of the mechanoafferent cells was observed. For example, cells located in the medial cluster have lip receptive fields, and most cells in the posterolateral portion of the lateral clusters have tentacle receptive fields. 4. Intracellular stimulation of individual mechanoafferents evokes short and constant-latency EPSPs in putative motor neurons comprising the identified B-cell clusters of the cerebral ganglion. On the basis of several criteria, these EPSPs appear to be several criteria, these EPSPs appear to be chemically mediated and are monosynaptic. 5. Repetitive intracellular stimulation of individual mechanoafferent neurons at low rates results in a gradual decrement in the amplitude of the EPSPs evoked in B cluster neurons. EPSP amplitude can be restored following brief periods of rest, but subsequent stimulation leads to further diminution of the response. 6. A decremented response cannot be restored by strong mechanical stimulation outside the receptive field of the mechanoafferent or by electrical stimulation of the cerebral nerves or connectives...
Animals↗
Steady potential shifts and facilitated learning of delayed response in monkeys.
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Animals↗
Interhemispheric functional differences in prefrontal cortex of monkeys.
Monkeys had nonpolarizable electrodes implanted bilaterally in prefrontal (principal sulcus), precentral, and occipital cortex. They were trained on a spatial delayed-response (DR) task (8-sec intratrial delay), while cortical potentials were recorded. Three groups of monkeys were trained to 90% criterion: (A) 4 monkeys with only the right hand (the left wrist was attached to the testing chair); (B) 2 monkeys with only the left hand; and (C) 2 monkeys with the left and right hands on alternate sessions. Intermanual transfer tests were then given. Averaged steady potential (SP) shifts of several seconds duration were found in prefrontal cortex during cue presentation and the early portion of the intratrial delay and from the precentral area during the choice response. Evaluations of these SP shift magnitudes indicated: (1) Training with only one hand resulted in substantially larger SP shifts in the prefrontal and precentral areas contralateral to the responding hand; (2) alternate hand training resulted in somewhat larger prefrontal SP shifts in the right hemisphere; (3) intermanual transfer had marked effects on the precentral SP shifts, with larger magnitudes in the hemisphere contralateral to the responding hand, but had little effect on the magnitudes of both prefrontal SP shifts. (4) Subsequent training of Group C monkeys with only one hand resulted in greater SP shifts in the prefrontal area contralateral to the responding hand and in decreased SP shifts in the ipsilateral prefrontal area; and (5) additional intermanual transfer tests had no effects on SP shift magnitudes from both prefrontal areas. These findings indicate a dissociation in interhemispheric functions between the precentral and prefrontal cortical areas, with the formal implicated in motor organization for the contralateral limb, and the latter in mediation of mnemonic processes, primarily in one hemisphere. This hemispheric specialization is affected by the hand-training procedure, but other endogenous or experimential factors may be involved.
Animals↗