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P Zarzecki

Publications and source records attributed to P Zarzecki.

At least 19 recordsLinked to original sources

Cortical influences on the vestibular nuclei of the cat.

Our goal was to study potential substrates for cortical modulation of vestibular reflexes in the cat. In initial experiments, injections of wheat-germ-agglutinate-horseradish-peroxidase into Deiters' nucleus and the rostral descending nucleus revealed bilateral colonies of retrogradely filled neurons in cortical areas 6, 2, and 3a (about 60 cells per colony). In cats anesthetized with chloralose-urethane, we stimulated areas 2 and 3a with trains of pulses while recording from ipsilateral vestibular-nucleus neurons, which were characterized by their responses to sinusoidal tilts and tested for the presence of antidromic responses to stimulation of the upper cervical cord. A majority of the neurons was affected by cortical stimulation, showing either facilitation, inhibition, or a mixture of the two. Stimulation in area 2 was more effective than stimulation in area 3a. Despite the anatomic presence of direct cortico-vestibular projections, properties of facilitation and inhibition suggest that both were evoked by polysynaptic pathways. Cortical effects were broadly distributed to vestibular neurons without regard to responses of these neurons to sinusoidal tilts. There was no significant difference between effects on lateral and medial vestibulospinal tract neurons, but, as a group, vestibulospinal neurons were much more likely to be affected by cortical stimulation than neurons not antidromically activated from the C2 segment. We conclude that, by their influence on vestibulospinal neurons, neurons in cortical areas 2 and 3a should be able to modulate, in behaving animals, vestibular reflexes acting on the neck and limbs.

Animals↗

Structural studies of the O-specific polysaccharide of Hafnia alvei strain PCM 1206 lipopolysaccharide containing D-allothreonine.

The structure of the O-specific side-chain of the Hafnia alvei strain PCM 1206 lipopolysaccharide has been investigated. Methylation analysis, partial acid hydrolysis, FAB-MS/MS and 1H-NMR and 13C-NMR spectroscopy were the principal methods used. D-Allothreonine (D-aThr), amide-linked to the D-galacturonic acid, was identified as a constituent in the polysaccharide and the following structure of a pentasaccharide repeating unit was established: [structure: see text].

Carbohydrate Sequence↗

Intrinsic discharge patterns and somatosensory inputs for neurons in raccoon primary somatosensory cortex.

1. Discharge patterns of neurons are regulated by synaptic inputs and by intrinsic membrane properties such as their complement of ionic conductances. Discharge patterns evoked by synaptic inputs are often used to identify the source and modality of sensory input. However, the interpretation of these discharge patterns may be complicated if different neurons respond to the same synaptic input with a variety of discharge patterns due to differences in intrinsic membrane properties. The purposes of this study were 1) to investigate intrinsic discharge patterns of neurons in primary somatosensory cortex of raccoon in vivo and 2) to use somatosensory postsynaptic potentials evoked by stimulation of forepaw digits to determine thalamocortical connectivity for the same neurons. 2. Conventional intracellular recordings with sharp electrodes were made from 121 neurons in the cortical representation of glabrous skin of digit four (d4). Intracellular injection of identical current pulses (100-120 ms in duration) elicited various patterns of discharge in different neurons. Neurons were classified on the basis of these intrinsic patterns of discharge, rates of spike adaptation, and characteristics of spike waveforms. Three main groups were identified: regular spiking (RS) neurons, intrinsic bursting (IB) neurons, and fast spiking (FS) neurons. Subclasses were identified for the RS and IB groups. 3. Neurons were tested for somatosensory inputs by stimulating electrically d3, d4, and d5. Excitatory postsynaptic potentials (EPSPs) were elicited in 100% of the neurons by electrical stimulation of d4, the "on-focus" digit. EPSPs were usually followed by inhibitory postsynaptic potentials (IPSPs). Many neurons (41%) responded with EPSP-IPSP sequences after stimulation of d3 or d5, the "off-focus" digits. 4. Latencies of somatosensory EPSPs and IPSPs were used to determine the synaptic order in the cortical circuitry of RS, IB, and FS neurons. EPSPs with monosynaptic thalamocortical latencies were recorded in RS, IB, and FS neurons. 5. We conclude that precise patterns of neural discharge in primary somatosensory cortex cannot be reliable estimates of sensory inputs reaching these neurons because patterns of discharge are so strongly influenced by intrinsic membrane properties. Ionic conductances governing patterns of neuronal discharge seem almost identical in intact cortex of raccoon, rat, and cat, and in slices of rodent cortex, because similar patterns of discharge are found. The consistency of patterns of discharge across species and types of preparation suggests that these intrinsic membrane properties are a general property of cerebral cortical neurons and should be considered when evaluation sensory coding by these neurons.

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Synaptic mechanisms of cortical representational plasticity: somatosensory and corticocortical EPSPs in reorganized raccoon SI cortex.

1. Reorganizations of representational maps have been described for a variety of sensory and motor regions of cerebral neocortex in several species. The purpose of this study was to investigate synaptic mechanisms of the reorganizations of primary somatosensory cortex that follow removal of a digit or the joining of two digits into a syndactyly. We examined neurons in the cortical representation of digit 4 (d4). Intracellular recording was used to compare somatosensory and corticocortical excitatory postsynaptic potentials (EPSPs) in normal raccoons, with EPSPs recorded in two experimental groups of animals surviving for a mean of 22 wk after removal of d4, or union of d4 with digit 3 (d3). 2. In normal animals with d4 intact, EPSPs were evoked from this on-focus digit in 100% of cortical neurons. EPSPs were evoked from d3 and digit 5 (off-focus digits) in only a minority of neurons in normal raccoons. The incidence of somatosensory EPSPs from off-focus digits increased dramatically after removal of d4 or its union with d3. Latencies of EPSPs evoked from off-focus digits decreased after d4 removal, so that they were as short as latencies from d4 in normal animals. In contrast, for the group of animals with d3-d4 syndactyly, latencies of EPSPs from off-focus digits were not shorter than responses from these digits in normal animals. 3. Corticocortical EPSPs were no more common in animals with d4 removed than in intact animals. Furthermore, corticocortical EPSPs after d4 removal did not differ in their latencies, amplitudes, half-widths, or integrated amplitudes. The only detected change was that corticocortical EPSPs had faster rising phases after removal of d4. In contrast, after d3-d4 syndactyly, corticocortical EPSPs were more common than in normal animals. 4. Digit removal and digital syndactyly had distinctive effects on somatosensory and corticocortical EPSPs. These results do not identify unique synaptic mechanisms for cortical representational plasticity, nor do they specify the involved CNS site(s). Several synaptic mechanisms consistent with the results are considered in the DISCUSSION, including synaptic proliferation to form new synaptic connections and enhanced effectiveness of existing corticocortical synapses.

Afferent Pathways↗

Intracortical mechanisms for the recruitment of motor cortex neurons.

Neurons project out of motor cortex to the spinal cord and to other targets. Not all projection neurons recruit in the same way during behavior, but instead recruitment patterns depend on the projection target of the neurons. The problem is to understand how neurons projecting to different targets are recruited selectively. We have investigated possible mechanisms for the recruitment of motor cortex neurons with electrophysiological approaches in anesthetized cats. To determine if neurons projecting out of motor cortex to different targets have selective input connectivity from extrinsic sources we electrically stimulated corticocortical, callosal and thalamocortical pathways. Subthreshold effects of input pathways were detected by monitoring latency variations of antidromic responses. Intracortical connections to identified output neurons were evaluated by cross-correlation and a new variation of the antidromic latency method. Output neurons in different layers along single electrode tracks usually had different inputs from extrinsic sources. Neurons in close proximity were most likely to share the same inputs, especially when they projected axons to the same target. These results support the conclusion that combinations of inputs from extrinsic sources could selectively recruit efferent neurons from separate cortical layers or from within groups of nearby neurons, according to the target of their axonal projections. In contrast, the data on intracortical connectivity suggest that common drive causes a more synchronous activation of nearby cortical neurons. Combining the conclusions on effects of inputs from extrinsic and intracortical sources leads to the speculation that motor cortex neurons that might at one time be recruited selectively by action of extrinsic afferent pathways to cortex could at another time be bound into synchrony by a common drive shared with their neighbours.

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Synaptic potentials evoked by convergent somatosensory and corticocortical inputs in raccoon somatosensory cortex: substrates for plasticity.

1. "Unmasking" of weak synaptic connections has been suggested as a mechanism for the early changes in cortical topographic maps that follow alterations of sensory activity. For such a mechanism to operate, convergent sensory inputs must already exist in the normal cortex. 2. We tested for topographic and cross-modality convergence in primary somatosensory cortex of raccoon. The representation of glabrous skin of forepaw digits was chosen because, even though it is dominated by inputs from the glabrous skin of a single digit, it nevertheless comes to respond to stimulation of other digits when, e.g., a digit is removed. 3. Intracellular recordings were made from 109 neurons in the representation of glabrous skin of digit 4. Neurons were tested for somatosensory inputs with electrical and natural stimulation of digits. 4. Excitatory postsynaptic potentials (EPSPs) were evoked in 100% of the neurons (109/109) by electrical stimulation of glabrous skin of digit 4, and in 79% (31 of 39) by vibrotactile stimulation. 5. Glabrous skin of digit 4 was not the sole source of somatosensory inputs. A minority of neurons generated EPSPs after electrical stimulation of hairy skin of digit 4 (10 of 98 neurons, 10%). Electrical stimulation of digits 3 or 5 evoked EPSPs in 22 of 103 neurons (21%). Natural stimulation (vibrotactile or hair bending) was also effective in most of these latter cases (digit 3, 6/7; digit 5, 9/10). 6. Intracortical microstimulation of the "heterogeneous zone" was used to test for corticocortical connections to neurons in the glabrous zone.(ABSTRACT TRUNCATED AT 250 WORDS)

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The distribution of corticocortical, thalamocortical, and callosal inputs on identified motor cortex output neurons: mechanisms for their selective recruitment.

Motor cortex neurons were identified antidromically in anesthetized cats by their axonal projections to one of six targets: (1) somatosensory cortex, (2) opposite motor cortex, (3) red nucleus, (4) lateral reticular nucleus, (5) spinal cord, and (6) ventrolateral thalamus. Three inputs to motor cortex were tested for their influences on the identified cortical efferent neurons. The tested inputs originated from ipsilateral somatosensory cortex, opposite motor cortex, and ventral thalamus. Subthreshold effects of input pathways were detected by monitoring latency variations of antidromic responses. The three afferent sources, when activated by electrical stimulation, were not equally effective on motor cortex neurons. Ipsilateral corticocortical and thalamocortical excitation were found for the majority of neurons; the influenced proportions ranged from 55% to 100%, according to the target of the output neurons. Effects from the opposite hemisphere were found for only 5% to 30% of the neurons in the same projection classes. Many neurons (36 of 81, or 44%) were excited from more than one source, but few (5 of 37, or 14%) were influenced by all three possible sources of input, even in small regions of cortex innervated by all three of the inputs. Among 19 electrode tracks where all three inputs were present, there were only 2 tracks where all the neurons shared the same combination of inputs. Even for neurons in closest anatomical proximity ("clusters"), it was unusual (only 7 of 25 clusters) for all the neurons to have the same input pattern. Among the seven clusters where all the neurons shared the same input pattern, five of the clusters projected to the same target. These variable combinations of inputs to motor cortex neurons support the conclusion that efferent neurons could be recruited selectively from separate cortical layers or from within clusters of nearby neurons, according to the target of their axonal projection.

Afferent Pathways↗

Multiple inputs to a population of thalamocortical neurons projecting to cat somatosensory cortex.

Ninety thalamocortical (TC) neurons were recorded extracellularly in ventrobasal thalamus of halothane-anesthetized cats. Projections of all of these neurons to specific subdivisions of somatosensory cortex were identified by their antidromic invasion following intracortical microstimulation restricted to these subdivisions. Collision-extinction tests were used to document excitatory inputs to TC neurons from afferent fibers of forelimb nerves stimulated electrically. Thirty-nine TC neurons (43% of sample) were excited from at least one forelimb nerve. Fifteen TC neurons were activated from two or more forelimb nerves. Combinations of effective nerves included ones innervating topographically different regions of forelimb. Neurons projecting to area 1-2 were least likely to be activated from more than one nerve. Seven TC neurons activated by electrical stimulation of nerve trunks were tested also with two distinct forms of mechanical somatic stimuli, i.e., hair bending and vibration, and with intradermal electrical stimuli. These tests revealed convergent inputs from hairy and glabrous skin. We conclude that there is a population of neurons, located in ventrobasal thalamus, which is capable of conveying multiple inputs to each of the subdivisions of primary somatosensory cortex. These neurons could be involved in forming properties of feature-extracting neurons of somatosensory cortex.

Action Potentials↗

Influence of somatosensory cortex on different classes of cat motor cortex output neuron.

1. Multiple output pathways originate from motor cortex. In this study on cats, six classes of corticofugal neurons were identified by antidromic activation. Corticocallosal neurons of layer III were activated antidromically by stimulation of contralateral motor cortex. Layer V neurons were identified by antidromic activation from cerebral peduncle, red nucleus, lateral reticular nucleus of medulla, or spinal cord. Corticothalamic neurons were identified in layer VI. All the identified neurons were tested for input from primary somatosensory cortex. 2. Neurons of all corticofugal groups received excitatory inputs from primary somatosensory cortex. The shortest latency corticocortical effects of 1.2-2.5 ms were found for corticocallosal neurons of layer III, and for layer V neurons which projected axons through the cerebral peduncle, to red nucleus, and to spinal cord. 3. Nearby neurons, projecting to the same of different targets, were affected nonuniformly by corticocortical inputs. This finding supports the conclusion that specificity of afferent connections within cerebral cortex is not determined by anatomic segregation of cell bodies nor by projection target of efferent neurons. 4. These selectively distributed input connectivities suggest that even a small region of motor cortex could send different signals to its diverse targets.

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Convergence of sensory inputs in somatosensory cortex: interactions from separate afferent sources.

Intracellular recording techniques were used to test for cross-modality and topographic convergence among inputs to area 3a of cerebral cortex. Recordings were made within the projection area of group I afferent fibers of the deep radial nerve in barbiturate-anesthetized cats. Epsps were evoked in 90% of neurons (81/90) by electrical stimulation of more than one nerve of the contralateral forelimb. The deep radial nerve evoked the shortest latency epsps within this region of cortex and the only ones likely to be mediated by a monosynaptic thalamocortical pathway. However, the epsps evoked from other forelimb nerves (of deep or cutaneous origin) had mean latencies only a few milliseconds (1.3-3.0 ms) longer. Furthermore, there were a variety of interactions among inputs from separate afferent sources. The observed interactions included spatial facilitation, occlusion and afferent inhibition. The consequence of these interactions was that neuronal responses were shaped by combinations of effects from different topographic regions of the forelimb or of different modalities. The findings are interpreted as indicating a sharing of neurons among pathways to cortical neurons from separate afferent sources. Interactions between ascending pathways by way of such shared neurons may contribute to the modulation or plasticity of somatosensory responsiveness during behavior or after deafferentation.

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Responses of cat motor cortex neurons to cortico-cortical and somatosensory inputs.

Intracellular techniques were used to investigate a cortico-cortical path from sensory cortex to motor cortex of cats. Cortico-cortical epsps were evoked in motor cortex neurons by microstimulation of area 3a. Epsps with latencies between 1.2 and 2.4 ms were identified as monosynaptic. These short latency cortico-cortical effects were recorded in layers II through VI of the motor cortex. Neurons with monosynaptic cortico-cortical epsps also received excitatory inputs from forelimb nerves, usually from both muscle and cutaneous afferent fibers. The epsps evoked from forelimb nerves in motor cortex neurons were preceded by neural activity in somatosensory cortex. Time delays between arrival of inputs in sensory cortex and in motor cortex were compared to the latencies of cortico-cortical epsps in the same motor cortex neurons. It was apparent that the timing was appropriate for the identified cortico-cortical path to have relayed some sensory inputs to motor cortex.

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Interactions among convergent inputs to somatosensory cortex neurons.

Postsynaptic potentials (psps) produced by electrical stimulation of 4 forelimb nerves were recorded intracellularly from neurons in the primary somatosensory cortex of sodium pentobarbital anesthetized cats. Convergent inputs were found from nerves subserving different modalities and different regions of the forelimb. Psps from separate afferent sources usually did not sum linearly but rather interacted with one another. These interactions could have occurred at the cortical level or earlier in the ascending pathways and are interpreted with regards to the control of somatosensory responsiveness by multiple converging inputs.

Animals↗

Convergence of sensory inputs upon projection neurons of somatosensory cortex: vestibular, neck, head, and forelimb inputs.

Cortico-cortical neurons and pyramidal tract (PT) neurons of the cat cerebral cortex were tested for convergent inputs from electrically stimulated vestibular, neck, head and forelimb nerves. Neurons were recorded within forelimb and vestibular projection regions of cortical area 3a. Consideration was given to both suprathreshold and subthreshold inputs. Neither vestibular, neck nor head inputs were detected in the forelimb region of area 3a. In contrast, within the vestibular projection region of area 3a, 43% (6/14) of the cortico-cortical neurons and 63% (24/38) of the PT neurons received excitatory vestibular input. Inputs from the skin of the pinna (greater auricular nerve) were detected only for PT neurons (66%, 25/38). No inputs were detected from afferent nerves supplying the dorsal neck muscles biventer cervicis and complexus. Cortico-cortical and PT neurons receiving vestibular input also received convergent inputs originating from forelimb group I deep and low threshold cutaneous afferent fibers. Further, one half of the PT neurons with vestibular input (12/24) received input from three somatic sources: forelimb group I deep, forelimb low threshold cutaneous and greater auricular (head) nerves. The input connectivities suggest a role for these projection neurons of somatosensory cortex in the coordination of head and forelimb movements. The convergence of vestibular information with somatic input from the forelimb implies that vestibular-influenced neurons of area 3a projecting to the motor cortex or through the pyramidal tract would signal head position or movement with respect to proprioceptive feedback from the limbs.

Afferent Pathways↗

Convergence of sensory inputs upon projection neurons of somatosensory cortex.

Cortico-cortical neurons and pyramidal tract neurons of the cat were tested for convergent inputs from forelimb afferents. Neurons were recorded in cortical areas 1, 2, and 3a. Consideration was given to both suprathreshold and subthreshold inputs evoked by electrical stimulation of forelimb nerves. Individual cortico-cortical neurons and also pyramidal tract neurons were characterized by convergence of multiple somatosensory inputs from different regions of skin, from several muscle groups, and between group I deep afferents and low threshold cutaneous afferents. Certain patterns of afferent input varied with cytoarchitectonic area. There was, however, no difference between area 3a and areas 1-2 in the incidence of cross-modality convergence in the form of input from cutaneous and also deep nerves. Many of the inputs were subthreshold. Arguments are presented that these inputs, though subthreshold, must be considered for a role in cortical information processing. The convergent nature of the sensory inputs is discussed in relation to the proposed specificities of cortical columns. The patterns of afferent inputs reaching cortico-cortical neurons seem to be appropriate for them to have a role in the formation of sensory fields of motor cortex neurons. PT neurons of somatosensory cortex have possible roles as modifiers of ascending sensory systems, however, the convergent input which these PT neurons receive argues against a simple relationship between the modality of peripheral stimuli influencing them and the modality of the ascending tract neurons under their descending control.

Afferent Pathways↗

Peripheral input pathways projecting to the motor cortex in the cat.

The possibility that the motor cortex receives peripheral input directly from the thalamus was examined using the evoked potential method and the following results were obtained. Potentials in the motor cortex evoked by stimulation of superficial radial (SR) or group II deep radial (DR) nerve were neither abolished nor delayed by ablation of the sensory cortex. Potentials in the motor cortex evoked by stimulation of group II DR nerve were most severely reduced by interruption of the spinocervcial tract. Potentials evoked by stimulation of SR nerve were more severely reduced in the sensory cortex than in the motor cortex by section of the dorsal funiculus or cooling of the cuneate nucleus. The size of evoked potentials in the motor cortex increased rapidly when stimulus intensity to DR nerve exceeded the threshold to group II fibers. The results suggest that some inputs from the SR and group II DR nerves reach the motor cortex without a relay through the sensory cortex.

Animals↗

Spinal branching of pyramidal tract neurons in the monkey.

The branching pattern of individual pyramidal tract (PT) neurons of the monkey motor cortex was studied by activating these neurons antidromically from within the cervical motor nuclei and also from other regions of the spinal cord. 1. Fifty-four neurons were activated from motor nuclei in the cervical cord. Twenty-eight of these were activated from one segment and six (11%) were activated from motor nuclei of different segments. The remaining 20 neurons were activated from motor nuclei and also from unspecified region(s) of the gray matter. 2. Another 156 neurons were activated from unspecified regions(s) of cervical gray matter which could have been motor nuclei or outside the nuclei, and 64 of these were activated from more than one segment. 3. The branching patterns of PT neurons sending axons directly to motor nuclei innervating distal forelimb muscles suggested that they branch less than the rest of PT neurons.

Animals↗

Projection of individual pyramidal tract neurons to lumbar motor nuclei of the monkey.

The projection of individual pyramidal tract (PT) neurons from the hindlimb area in the precentral gyrus of the cerebral cortex to the lumbar spinal cord was studied in the monkey by systematically searching for sites within identified regions of the spinal gray from which the PT neurons could be antidromically activated by local stimulation. All investigated neurons belonged to the fast conducting fraction of PT neurons. The following results were obtained. 1. Each PT neuron could be activated from more than one region of the spinal gray matter, including identified spinal motor nuclei and areas dorsomedial to these nuclei, but do not the intermediate nucleus or regions dorsal to it. "Passage areas" and "termination areas" were defined. 2. Half of the PT neurons with termination areas within motor nuclei had these areas in more than one nucleus. There were thus strong suggestions for synaptic contacts of some PT neurons with motoneurons of more than one muscle. 3. Four groups of three or four neurons were recorded simultaneously by the same cortical electrode. Comparisons of passage and termination areas within groups revealed both similarities and differences in projections of neighboring neurons. Every neuron was activated from some region(s) where others of the group were not. Common passage areas, or passage and termination areas, for two or three neurons of a group within at least one motor nucleus were found for all groups. Termination areas in the same motor nucleus have been found for the majority of the neurons of only one group. These common projection areas are compatible with, but not prove, that group of adjacent PT neurons has common target cells in the spinal cord.

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