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I Kupfermann

Publications and source records attributed to I Kupfermann.

At least 55 records · Page 3Linked to original sources

Effects of cerebral neuron C-PR on body postural muscles associated with a food-induced arousal state in Aplysia.

1. Firing of cerebral neuron, C-PR, produced complex bilateral movements of various regions of the body of the marine mollusc Aplysia californica. The movements were similar to those seen when the animal assumes the head-up feeding posture during food-induced arousal. Muscles of the neck largely contracted in transverse and longitudinal directions, and large transverse movements were also induced in the middle part of the foot. On the other hand, firing of C-PR appeared to relax the anterior part of the foot in transverse and longitudinal directions. 2. We identified pedal-ganglion motor neurons that innervate various regions of the animal, and explored the synaptic connections of C-PR with these neurons. Firing of C-PR produced synaptic potentials bilaterally in most of the identified motor neurons. 3. Motor neurons for the neck were largely excited by C-PR firing. C-PR firing also excited the motor neurons that produce transverse movements of the middle part of the foot. On the other hand, C-PR inhibited the spontaneous spike activity of the motor neurons for the anterior part of the foot. 4. One neck motor neuron was found to receive a monosynaptic excitatory postsynaptic potential (EPSP) from C-PR, but the postsynaptic potentials (PSPs) induced by C-PR in the other identified motor neurons were mediated polysynaptically. 5. We also found that the C-PR can modulate movements evoked by firing of the motor neurons for the ipsilateral neck and anterior foot. C-PR enhanced both transverse and longitudinal contractions of the neck. 6. For the anterior foot region, C-PR had different modulatory effects on the longitudinal and the transverse contractions. C-PR largely enhanced or initially depressed and then enhanced longitudinal contractions, whereas C-PR depressed transverse contractions. 7. The overall results support the hypothesis that C-PR is involved in controlling the head-up posture when the animal is aroused by food.

Animals↗

The buccalin-related neuropeptides: isolation and characterization of an Aplysia cDNA clone encoding a family of peptide cotransmitters.

The buccalin-related peptides, buccalin A and buccalin B, are members of a family of cotransmitters that modulate neuromuscular transmission in Aplysia. In this study, a third buccalin-related peptide, buccalin C, was purified from neuronal elements in the accessory radula closer, a muscle involved in the animal's feeding behavior. Oligonucleotide probes based upon the amino acid sequence of buccalin C were used to isolate cDNA clones that encode a buccalin precursor polypeptide. The buccalin precursor contains 19 distinct buccalin-related peptides, several of which are present in multiple copies. The buccalin gene appears to be present in a single copy, with one allele containing a small insert. Expression of this gene occurs in a tissue-specific manner and mRNA transcripts are abundant within neurons in the Aplysia CNS. This large family of neuropeptides may exert extraordinarily complex modulatory actions at synapses where they serve as cotransmitters.

Amino Acid Sequence↗

The myomodulin-related neuropeptides: characterization of a gene encoding a family of peptide cotransmitters in Aplysia.

The myomodulin-related peptides comprise a family of cotransmitters that modulate neuromuscular signaling in the feeding system of Aplysia. In this study, cDNA clones encoding a myomodulin precursor polypeptide were isolated and characterized. This precursor contains seven different myomodulin-related peptides, one of which, myomodulin A, is present in 10 contiguous copies. The sequence of a myomodulin genomic clone indicates that all of these myomodulin-related peptides are encoded on a single exon. The myomodulin gene is expressed in a tissue-specific manner and myomodulin mRNA is localized to specific neurons in the Aplysia CNS. The presence of multiple related neuropeptides can greatly increase the range and precision of signaling at synapses where they act as modulator cotransmitters.

Amino Acid Sequence↗

Distribution of buccalin-like immunoreactivity in the central nervous system and peripheral tissues of Aplysia californica.

The neuropeptide buccalin A was originally purified and sequenced from a nerve-muscle system used in feeding-related behaviors of Aplysia californica in which it has been proposed that it acts as a modulatory cotransmitter. The distribution of buccalin-like immunoreactivity in the central ganglia and in peripheral tissues of Aplysia californica was examined by whole mount immunohistochemical techniques. Immunoreactive material was located in specific cell bodies and clusters of neurons in each of the ganglia. Immunoreactive fibers were present in each of the connectives between ganglia, in tracts coursing through the ganglia, and in the majority of the peripheral nerves. Most fibers were smooth in contour, but some had regularly spaced swellings. Varicosities containing immunoreactive material were located on specific neuronal somata and on certain tissues associated with the feeding, circulatory, digestive, and reproductive systems. The specific and widespread distribution of buccalin-like immunoreactivity supports the hypothesis that members of the buccalin peptide family act as neuromodulators or neurotransmitters in a variety of central and peripheral circuits in Aplysia.

Animals↗

Peptidergic co-transmission in Aplysia: functional implications for rhythmic behaviors.

Despite their ubiquitous presence in the central and peripheral nervous systems, the behavioral functions of peptide co-transmitters remain to be elucidated. The marine mollusc Aplysia, whose simple nervous system facilitates the study of the neural basis of behavior, was used to investigate the role of peptidergic co-transmission in feeding behavior. Several novel modulatory neuropeptides were purified, and localized to identified cholinergic motorneurons. Physiological and biochemical studies demonstrated that these peptides are released when the motorneurons fire at frequencies that occur during normal behavior, and that the peptides modify the relationship between muscle contraction amplitude and relaxation rate so as to maintain optimal motor output when the intensity and frequency of feeding behavior change.

Amino Acid Sequence↗

Orientation of Aplysia californica to distant food sources.

The behavior of the marine mollusc Aplysia was examined under different experimental conditions designed to determine the food searching strategy of the animals. In a small, open field tank with still water, the animals took an average of 42 min to find a piece of seaweed, even though the stimulus was never located more than 30 cm away from the animal. Observations of the animals indicated that their search was not directed, without a clear tendency towards the food, and during the course of a search, they often crawled through most of the area of the tank. The search time, the distance travelled, and the strategy of the search of the animals was similar for different types of seaweed. If animals were aroused into activity by the presence of seaweed extract, the time for them to contact a piece of odorless glass fiber paper in the open field was not significantly different than that for a piece of seaweed. The probability at which the animals contacted the seaweed, as a function of the distance travelled, resembled the detection probability determined according to a theory of random search. We thus propose that the aroused animals move in a random pattern until they are very close to the food. This strategy can be advantageous in still water since chemicals do not provide distinct gradients that can serve as cues for chemotactic orientation from distances greater than a few centimeters from the source. In a Y-maze in still water, Aplysia did not perform above chance in selecting the arm that contained the seaweed.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Localization of myomodulin-like immunoreactivity in the central nervous system and peripheral tissues of Aplysia californica.

The distribution of myomodulin-like peptides in the nervous system of Aplysia californica was examined by using immunocytochemical techniques. Neurons and cell clusters containing immunoreactive material were located in each of the major central ganglia. Myomodulin-like immunoreactivity was also present in fibers in each of the connectives between the ganglia and in peripheral nerves. Varicosities containing immunoreactive material were located on specific regions of peripheral tissues associated with the feeding, digestive, cardiovascular, and reproductive systems. Double-labeling experiments were used to demonstrate myomodulin-like immunoreactivity in two identified neurons, the motor neuron B16 in the buccal ganglion and the widely acting interneuron L10 in the abdominal ganglion. Structures in the eye and cerebral ganglion that may correspond to the optic circadian pacemaker system were also stained. The central and peripheral distribution of myomodulin-like immunoreactivity indicates that this family of neuropeptides is present in specific efferent, afferent, and interneuronal elements that participate in a diversity of neural circuits in Aplysia.

Amino Acid Sequence↗

Activity of identified cerebral neuron correlates with food-induced arousal in Aplysia.

Firing of the cerebral-pedal regulator neuron, C-PR, evokes a constellation of responses which are characteristic of the food-induced arousal state that occurs following exposure of Aplysia to seaweed. To provide further evidence that C-PR plays a role in generating the food-induced arousal state, extracellular recordings from the cerebral-pedal connective, which contains the axon of C-PR, were obtained in freely moving animals. The C-PR spike in the connective recorded in vivo was then identified by comparing the wave form to the obtained by firing C-PR in an in vitro preparation. We report here that C-PR activity is evoked by food stimulation, and increased firing of the C-PR is closely correlated with appetitive head lifting, the first manifestation of the food arousal state.

Animals↗

Egg laying hormone inhibits a neuron (C-PR) involved in multiple manifestations of food-induced arousal in Aplysia.

Egg laying behavior is known to suppress feeding in Aplysia, but both behaviors have common responses involving head movements and posture. Egg laying hormone (ELH) applied in vitro to the isolated nervous system of Aplysia reduces the spontaneous and the evoked activity of the C-PR, a neuron implicated in postural responses during feeding. The inhibitory effect of ELH on the C-PR appears to be mediated by interneurons primarily located in the pedal/pleural ganglia, which contain all the known direct follower cells of the C-PR. Our results do not support the idea that postural responses during feeding and egg laying are mediated by the activation of a common arousal element, the C-PR. In fact, the C-PR seems to be a specific element for the food-arousal state, and the inhibition of the C-PR by ELH may contribute to the suppression of appetitive feeding responses during egg laying.

Action Potentials↗

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↗

Differential firing patterns of the peptide-containing cholinergic motor neurons B15 and B16 during feeding behavior in Aplysia.

During egestive responses neuron B16 fires at 20 Hz, while neuron B15 is not active. During ingestive responses B16 fires for 0.5-1.0 s at 15-20 Hz, then B15 and B16 fire together, with B15 firing at 7.5-12 Hz. The duration of activity during ingestive responses depends on consumption of food: when food is not consumed, bursts are shorter (e.g. 2 vs 4 s). This study establishes a basis for investigating the role of peripheral neuromodulation under physiologically relevant conditions.

Acetylcholine↗

Buccalin is present in the cholinergic motor neuron B16 of Aplysia and it depresses accessory radula closer muscle contractions evoked by stimulation of B16.

The accessory radula closer (ARC) buccal muscle motor neuron B16 is buccalin-immunoreactive and it synthesizes a peptide with chromatographic properties identical to those of buccalin. Terminal varicosities in the ARC muscle are buccalin-immunoreactive, and buccalin reduces muscle contractions evoked by stimulation of neuron B16, probably by a presynaptic action.

Animals↗

An identified neuron (CPR) evokes neuronal responses reflecting food arousal in Aplysia.

Feeding behavior of Aplysia is associated with an arousal state characterized by a constellation of maintained behaviors and by a potentiation or depression of responses to specific stimuli. A neuron (the cerebral-pedal regulator or CPR) that has widespread actions on various systems connected with feeding has been identified. CPR excites neurons that modulate or drive (i) body posture, (ii) biting, and (iii) cardiovascular behaviors. CPR also inhibits neurons concerned with defensive responses. Food stimuli, which elicit food arousal in the animal, produce prolonged excitation of the CPR. The results suggest that the CPR may evoke a central motive state representing the neuronal correlate of feeding motivation.

Animals↗

Multiple roles of a histaminergic afferent neuron in the feeding behavior of Aplysia.

The cellular and circuit properties of individual identified neurons in invertebrates can be readily studied; hence it is possible to determine how the complex properties of nerve cells function in the generation of behavior. Recent studies of the cellular basis of feeding behavior in the marine mollusc Aplysia have focused on a neuron, C2, that has a variety of complex properties that determine the behavioral functions of the neuron. C2 conveys mechanosensory information from the mouth of the animal. It receives a complex pattern of inputs during feeding behavior, and generates diverse outputs that may shape behavior. It can act to filter out slow or sporadic sensory inputs, and its own outputs can be 'gated' by synaptic input. C2 uses histamine as its transmitter, and some of its synaptic outputs are modulatory and contribute to the expression of an arousal state induced by food. Other outputs shape feeding behavior directly by affecting motor neurons, as well as presynaptically inhibiting the outputs of feeding motor programs. Thus, the complex properties of this neuron may contribute to the flexibility and adaptability of feeding in Aplysia. Studies of C2 have expanded our concepts of the properties of sensory neurons.

Action Potentials↗

Release of peptide cotransmitters from a cholinergic motor neuron under physiological conditions.

In previous studies, we demonstrated that B15, one of the two cholinergic motor neurons of the accessory radula closer muscle of Aplysia, synthesizes two peptides, small cardioactive peptides A and B (SCPA and SCPB), that, when exogenously applied, increase the size and relaxation rate of muscle contractions elicited by motor neuron stimulation. In the present experiments, we obtained evidence that the SCPs are released under physiological conditions. Specifically, we characterized firing patterns of motor neuron B15 during normal behavior, simulated them in vitro, and demonstrated that this type of neuronal activity produces decreases in SCP levels in neuronal processes and terminals. We also obtained evidence that suggests that enough SCP is released under physiological conditions to modulate neuromuscular activity in the accessory radula closer. We demonstrated that physiological activity of neuron B15 produces significant increases in muscle cAMP levels. Furthermore, increases in the size and relaxation rate of muscle contractions can be produced by changes in stimulation parameters that are also likely to maximize effects of released endogenous SCPA and SCPB.

Animals↗

Appetitive feeding behavior of Aplysia: behavioral and neural analysis of directed head turning.

The appetitive phase of feeding behavior in Aplysia consists of a behavioral sequence in which the quiescent animal starts to locomote and then assumes a characteristic feeding posture. In this position, head-turning responses can be elicited by a localized food stimulus (seaweed) delivered to the lips or tentacles. In response to brief (open loop) stimulation with seaweed, the animal turns toward the stimulus but greatly overshoots the target. However, the angular velocity and the final turning angle are a function of the eccentricity of the stimulus, progressively increasing with greater eccentricities. In a food-aroused animal, a brief tactile stimulus evokes turning and biting responses similar to those triggered by seaweed, which provides both tactile and chemical stimulation. Upon repeated tactile stimulation, however, the response magnitude decrements rapidly, whereas the magnitude remains high when turning responses are repeatedly elicited by food stimuli. A purely chemical stimulus sometimes can elicit a turning response, but chemical stimuli alone are much less efficacious than tactile stimuli alone. When the stimulus is maintained in a stationary position (closed loop), the animal turns until its mouth is oriented over the food. A turning response to a lateral stimulus can be reduced by an immediately following medial stimulus. To explain the above findings, we propose a form of response substitution, in which the response to the first, lateral stimulus is substituted by a weaker response to a more medial stimulus. No turning response is evoked when the animal is stimulated while performing spontaneous or evoked bites, though biting per se does not interrupt ongoing turning movements. In animals with lesions of the cerebral-buccal connectives, a food stimulus on the mouth is also followed by a reduction of the capacity of stimuli to elicit turning responses. In these lesioned animals, the food stimulus appears to elicit a bite command, though the biting behavior itself does not occur. Thus, it appears that the bite-related gating of stimuli is of cerebral origin, rather than due to the generation of the buccal motor program. The force necessary to power the turning movements was calculated from the trajectories of the movements. The results indicate that a power phase during the first half of the duration of the total movement is sufficient to generate a turn. The power phase can be followed by a brief gliding phase, and finally the movement appears to be actively terminated.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗