Long-range neuronal circuits underlying the interaction between sensory and motor cortex

In the rodent vibrissal system, active sensation and sensorimotor integration are mediated in part by connections between barrel cortex and vibrissal motor cortex. Little is known about how these structures interact at the level of neurons. We used Channelrhodopsin-2 (ChR2) expression, combined with anterograde and retrograde labeling, to map connections between barrel cortex and pyramidal neurons in mouse motor cortex. Barrel cortex axons preferentially targeted upper layer (L2/3, L5A) neurons in motor cortex; input to neurons projecting back to barrel cortex was particularly strong. Barrel cortex input to deeper layers (L5B, L6) of motor cortex, including neurons projecting to the brainstem, was weak, despite pronounced geometric overlap of dendrites with axons from barrel cortex. Neurons in different layers received barrel cortex input within stereotyped dendritic domains. The cortico-cortical neurons in superficial layers of motor cortex thus couple motor and sensory signals and might mediate sensorimotor integration and motor learning.     

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.     

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.     

Interdigitated paralemniscal and lemniscal pathways in the mouse barrel cortex

Primary sensory cortical areas receive information through multiple thalamic channels. In the rodent whisker system, lemniscal and paralemniscal thalamocortical projections, from the ventral posteromedial nucleus (VPM) and posterior medial nucleus (POm) respectively, carry distinct types of sensory information to cortex. Little is known about how these separate streams of activity are parsed and integrated within the neocortical microcircuit. We used quantitative laser scanning photostimulation to probe the organization of functional thalamocortical and ascending intracortical projections in the mouse barrel cortex. To map the thalamocortical projections, we recorded from neocortical excitatory neurons while stimulating VPM or POm. Neurons in layers (L)4, L5, and L6A received dense input from thalamus (L4, L5B, and L6A from VPM; and L5A from POm), whereas L2/3 neurons rarely received thalamic input. We further mapped the lemniscal and paralemniscal circuits from L4 and L5A to L2/3. Lemniscal L4 neurons targeted L3 within a column. Paralemniscal L5A neurons targeted a superficial band (thickness, 60 μm) of neurons immediately below L1, defining a functionally distinct L2 in the mouse barrel cortex. L2 neurons received input from lemniscal L3 cells and paralemniscal L5A cells spread over multiple columns. Our data indicate that lemniscal and paralemniscal information is segregated into interdigitated cortical layers.     

Axonal conduction properties of antidromically identified neurons in rat barrel cortex

Physiological studies of the rodent somatosensory cortex have consistently described considerable heterogeneity in receptive field properties of neurons outside of layer IV, particularly those in layers V and VI. One such approach for distinguishing among different local circuits in these layers may be to identify the projection target of neurons whose axon collaterals contribute to the local network. In vivo, this can be accomplished using antidromic stimulation methods. Using this approach, the axonal conduction properties of cortical efferent neurons are described. Four projection sites were activated using electrical stimulation: (1) vibrissal motor cortex, (2) ventrobasal thalamus (VB), (3) posteromedial thalamic nucleus (POm), and (4) cerebral peduncle. Extracellular recordings were obtained from a total of 169 units in 21 animals. Results demonstrate a close correspondence between the laminar location of the antidromically identified neurons and their anatomically known layer of origin. Axonal properties were most distinct for corticofugal axons projecting through the crus cerebri. Corticothalamic axons projecting to either VB or POm were more similar to each other in terms of laminar location and conduction properties, but could be distinguished using focal electrical stimulation. It is concluded that, once stimulation parameters are adjusted for the small volume of the rat brain, the use of antidromic techniques may be an effective strategy to differentiate among projection neurons comprising different local circuits in supra- and infragranular circuits.     

Projections from areas 5 and 7 to subdivisions of the cat sensorimotor cortex

Projections from the parietal cortex (areas 5 and 7) to subdivisions of the sensori-motor cortical region were investigated in cats using axonal degeneration techniques. Differences between the density of distribution of association fibers proceeding from these areas were found within the parietal and sensorimotor cortex. Area 5 projects mainly to the posterolateral portion of the cruciate sulcus (areas 4fu and 4) and to fields 4y, 4sfu, 6iffu, 6aa, and 6ab to a lesser extent. Area 7 is connected mainly to the medial portion of the lower lip of the cruciate sulcus (areas 6iffu, 6aa, and 6ab). Somewhat fewer fibers proceed to areas 4fu and 4. Fewer projections proceed from the parietal cortex to the somatosensory than to the motor region. Only a few single fibers connect the primary somatosensory region (fields 2, 3a, and 3b) with area 5, while area 7 does not project into this area. Neither field 5 nor 7 projects to the secondary somatosensory cortical area.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 20, No. 3, pp. 319–326, May–June, 1988.     

Hierarchy within human SI: supporting data from cytochrome oxidase,acetylcholinesterase and NADPH-diaphorase staining patterns

The human primary somatosensory cortex consists of four cytoarchitectonic subdivisions (3a, 3b, 1 and 2) that are likely to contain distinct somatosensory representations. The intraareal organization of these areas as well as that of the primary motor cortex (area 4) has been analyzed using histochemical stains of cytochrome oxidase, acetylcholinesterase and NADPH-diaphorase activity in normal human brains. Cytochrome oxidase activity was revealed in individual cortical neurons and neuropil. Areas 4, 3a and 3b were on average darker than areas 1 and 2. The laminar distribution of cytochrome oxidase activity varied in different areas. A prominent dark band was present in layers IV and lower III in areas 3a and 3b and in layer III in areas 1, 2 and 4. Acetylcholinesterase staining revealed fibers and pyramidal cells in layers III and V; stained layer III pyramids were rare in areas 3a and 3b and numerous in areas 1, 2 and 4. NADPH-diaphorase positive elements included Golgi-like stained non-pyramidal neurons and Nissl-like stained pyramidal neurons; the former were found, in small numbers, in layer II of areas 4, 3a, 3b and 1, and the latter in layers III and V of areas 4 and 3a and in layer V of areas 1 and 2. The dark cytochrome oxidase staining of layer IV and the paucity of acetylcholinesterase positive pyramids in areas 3a and 3b resemble the pattern found in primary visual and auditory areas, whereas the dark cytochrome oxidase staining in layer III and abundance of acetylcholinesterase positive pyramids in areas 1 and 2 that of association areas. These results suggest that the four areas included in human SI constitute hierarchical stages of cortical processing, with 3a and 3b corresponding to primary and 1 and 2 to secondary areas.     

Hierarchy within human SI: supporting data from cytochrome oxidase, acetylcholinesterase and NADPH-diaphorase staining patterns

The human primary somatosensory cortex consists of four cytoarchitectonic subdivisions (3a, 3b, 1 and 2) that are likely to contain distinct somatosensory representations. The intraareal organization of these areas as well as that of the primary motor cortex (area 4) has been analyzed using histochemical stains of cytochrome oxidase, acetylcholinesterase and NADPH-diaphorase activity in normal human brains. Cytochrome oxidase activity was revealed in individual cortical neurons and neuropil. Areas 4, 3a and 3b were on average darker than areas 1 and 2. The laminar distribution of cytochrome oxidase activity varied in different areas. A prominent dark band was present in layers IV and lower III in areas 3a and 3b and in layer III in areas 1, 2 and 4. Acetylcholinesterase staining revealed fibers and pyramidal cells in layers III and V; stained layer III pyramids were rare in areas 3a and 3b and numerous in areas 1, 2 and 4. NADPH-diaphorase positive elements included Golgi-like stained non-pyramidal neurons and Nissl-like stained pyramidal neurons; the former were found, in small numbers, in layer II of areas 4, 3a, 3b and 1, and the latter in layers III and V of areas 4 and 3a and in layer V of areas 1 and 2. The dark cytochrome oxidase staining of layer IV and the paucity of acetylcholinesterase positive pyramids in areas 3a and 3b resemble the pattern found in primary visual and auditory areas, whereas the dark cytochrome oxidase staining in layer III and abundance of acetylcholinesterase positive pyramids in areas 1 and 2 that of association areas. These results suggest that the four areas included in human SI constitute hierarchical stages of cortical processing, with 3a and 3b corresponding to primary and 1 and 2 to secondary areas.     

Axonal conduction properties of antidromically identified neurons in rat barrel cortex

Physiological studies of the rodent somatosensory cortex have consistently described considerable heterogeneity in receptive field properties of neurons outside of layer IV, particularly those in layers V and VI. One such approach for distinguishing among different local circuits in these layers may be to identify the projection target of neurons whose axon collaterals contribute to the local network. In vivo, this can be accomplished using antidromic stimulation methods. Using this approach, the axonal conduction properties of cortical efferent neurons are described. Four projection sites were activated using electrical stimulation: (1) vibrissal motor cortex, (2) ventrobasal thalamus (VB), (3) posteromedial thalamic nucleus (POm), and (4) cerebral peduncle. Extracellular recordings were obtained from a total of 169 units in 21 animals. Results demonstrate a close correspondence between the laminar location of the antidromically identified neurons and their anatomically known layer of origin. Axonal properties were most distinct for corticofugal axons projecting through the crus cerebri. Corticothalamic axons projecting to either VB or POm were more similar to each other in terms of laminar location and conduction properties, but could be distinguished using focal electrical stimulation. It is concluded that, once stimulation parameters are adjusted for the small volume of the rat brain, the use of antidromic techniques may be an effective strategy to differentiate among projection neurons comprising different local circuits in supra- and infragranular circuits.     

Areal distributions of cortical neurons projecting to different levels of the caudal brain stem and spinal cord in rats

Distributions of corticospinal and corticobulbar neurons were revealed by tetramethylbenzidine (TMB) processing after injections of wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) into the cervical or lumbar enlargements of the spinal cord, or medullary or pontine levels of the brain stem. Sections reacted for cytochrome oxidase (CO) allowed patterns of labeled neurons to be related to the details of the body surface map in the first somatosensory cortical area (SI). The results indicate that a number of cortical areas project to these subcortical levels: (1) Projection neurons in granular SI formed a clear somatotopic pattern. The hindpaw region projected to the lumbar enlargement, the forepaw region to the cervical enlargement, the whisker pad field to the lower medulla, and the more rostral face region to more rostral brain stem levels. (2) Each zone of labeled neurons in SI extended into adjacent dysgranular somatosensory cortex, forming a second somatotopic pattern of projection neurons. (3) A somatotopic pattern of projection neurons in primary motor cortex (MI) paralleled SI in mediolateral sequence corresponding to the hindlimb, forelimb, and face. (4) A weak somatotopic pattern of projection neurons was suggested in medial agranular cortex (Agm), indicating a premotor field with a rostromedial-to-caudolateral representation of hindlimb, forelimb, and face. (5) A somatotopic pattern of projection neurons representing the foot to face in a mediolateral sequence was observed in medial parietal cortex (PM) located between SI and area 17. (6) In the second somatosensory cortical area (SII), neurons projecting to the brain stem were immediately adjacent caudolaterally to the barrel field of SI, whereas neurons projecting to the upper spinal cord were more lateral. No projection neurons in this region were labeled by the injections in the lower spinal cord. (7) Other foci of projection neurons for the face and forelimb were located rostral to SII, providing evidence for a parietal ventral area (PV) in perirhinal cortex (PR) lateral to SI, and in cortex between SII and PM. None of these regions, which may be higher-order somatosensory areas, contained labeled neurons after injections in the lower spinal cord. Thus, more cortical fields directly influence brain stem and spinal cord levels related to sensory and motor functions of the face and forepaw than the hindlimb. The termination patterns of corticospinal and corticobulbar projections were studied in other rats with injections of WGA:HRP in SI.(ABSTRACT TRUNCATED AT 400 WORDS)     

Symmetric sensorimotor somatotopy

Background

Functional imaging has recently been used to investigate detailed somatosensory organization in human cortex. Such studies frequently assume that human cortical areas are only identifiable insofar as they resemble those measured invasively in monkeys. This is true despite the electrophysiological basis of the latter recordings, which are typically extracellular recordings of action potentials from a restricted sample of cells.

Methodology/Principal Findings

Using high-resolution functional magnetic resonance imaging in human subjects, we found a widely distributed cortical response in both primary somatosensory and motor cortex upon pneumatic stimulation of the hairless surface of the thumb, index and ring fingers. Though not organized in a discrete somatotopic fashion, the population activity in response to thumb and index finger stimulation indicated a disproportionate response to fingertip stimulation, and one that was modulated by stimulation direction. Furthermore, the activation was structured with a line of symmetry through the central sulcus reflecting inputs both to primary somatosensory cortex and, precentrally, to primary motor cortex.

Conclusions/Significance

In considering functional activation that is not somatotopically or anatomically restricted as in monkey electrophysiology studies, our methodology reveals finger-related activation that is not organized in a simple somatotopic manner but is nevertheless as structured as it is widespread. Our findings suggest a striking functional mirroring in cortical areas conventionally ascribed either an input or an output somatotopic function.     

Areal Distributions of Cortical Neurons Projecting to Different Levels of the Caudal Brain Stem and Spinal Cord in Rats

Distributions of corticospinal and corticobulbar neurons were revealed by tetramethylbenzidine (TMB) processing after injections of wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) into the cervical or lumbar enlargements of the spinal cord, or medullary or pontine levels of the brain stem. Sections reacted for cytochrome oxidase (CO) allowed patterns of labeled neurons to be related to the details of the body surface map in the first somatosensory cortical area (SI). The results indicate that a number of cortical areas project to these subcortical levels: (1) Projection neurons in granular SI formed a clear somatotopic pattern. The hindpaw region projected to the lumbar enlargement, the forepaw region to the cervical enlargement, the whisker pad field to the lower medulla, and the more rostral face region to more rostral brain stem levels. (2) Each zone of labeled neurons in SI extended into adjacent dysgranular somatosensory cortex, forming a second somatotopic pattern of projection neurons. (3) A somatotopic pattern of projection neurons in primary motor cortex (MI) paralleled SI in mediolateral sequence corresponding to the hindlimb, forelimb, and face. (4) A weak somatotopic pattern of projection neurons was suggested in medial agranular cortex (Ag_m), indicating a premotor field with a rostromedial-to-caudolateral representation of hindlimb, forelimb, and face. (5) A somatotopic pattern of projection neurons representing the foot to face in a mediolateral sequence was observed in medial parietal cortex (PM) located between SI and area 17. (6) In the second somatosensory cortical area (SII), neurons projecting to the brain stem were immediately adjacent caudolaterally to the barrel field of SI, whereas neurons projecting to the upper spinal cord were more lateral. No projection neurons in this region were labeled by the injections in the lower spinal cord. (7) Other foci of projection neurons for the face and forelimb were located rostral to SII, providing evidence for a parietal ventral area (PV) in perirhinal cortex (PR) lateral to SI, and in cortex between SII and PM. None of these regions, which may be higher-order somatosensory areas, contained labeled neurons after injections in the lower spinal cord. Thus, more cortical fields directly influence brain stem and spinal cord levels related to sensory and motor functions of the face and forepaw than the hindlimb.

The termination patterns of corticospinal and corticobulbar projections were studied in other rats with injections of WGA:HRP in SI. Injections in lateral SI representing the face produced dense terminal label in the contralateral trigeminal complex. Injections in cortex devoted to the forelimb and forepaw labeled the contralateral cuneate nucleus and parts of the dorsal horn of the spinal cord. The cortical injections also demonstrated interconnections of parts of SI with some of the other regions of cortex with projections to the spinal cord, and provided further evidence for the existence of PV in rats.     

Somatosensory cortical unit responses to stimulation of the vibrissae in cats

Characteristics of extra- and intracellular responses of 57 neurons in the vibrissal projection zone of the first somatosensory area of the cat cortex were investigated. The intensity of both excitatory and inhibitory unit responses was found to diminish during successive stimulation of different parts of the receptive fields in the direction from center toward periphery. Usually, when central parts of receptive fields were stimulated, inhibition in the unit responses was postexcitatory, whereas when peripheral parts were stimulated inhibition could precede excitation. The possibility of an increase in the role of interaction between excitatory and inhibitory processes arising in neurons in response to vibrissal stimulation with an increase in the distance from center to periphery of receptive fields of single cortical cells is discussed. Neurons found during one insertion of the microelectrode were seen to have common center for their receptive fields, but the diameters of the receptive fields of individual neurons could differ significantly. Moreover, during such vertical insertions responses of neurons with primary inhibition to the stimuli presented were recorded.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 2, pp. 124–130, March–April, 1980.     

Responses of neurons in cat primary somatosensory cortex to repetitive stimulation of vibrissae

Extra- and intracellular responses of neurons in the primary somatosensory cortex to repetitive mechanical stimulation of the vibrissae at different frequencies were studied in unanesthetized curarized adult cats. Unlike responses to electrical stimulation of the combined afferent input (the infraorbital nerve) spike discharges of neurons in response to vibrissal stimulation can reproduce rather higher frequencies of stimulation and their initial character changes more often in the course of the repetitive series. Most cortical neurons were characterized by limitation of the area of their peripheral receptive fields with an increase in the frequency of adequate repetitive stimulation. A group of cortical neurons was distinguished by its ability to respond to high-frequency stimulation and to generate burst discharges. Comparison of the frequency characteristics of spike responses of these cells and of inhibitory synaptic action in other cortical neurons led to the conclusion that this group of cells thus distinguished may be inhibitory cortical neurons. The role of interaction between excitatory and inhibitory processes arising in cortical neurons during repetitive stimulation of different areas of their receptive fields is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 2, pp. 164–171, March–April, 1982.     

Spatial principle of organization of specific afferent projections and generation of evoked potentials in the somatosensory cortex

Here we investigate the functional organization of structures involved in sensory analysis in a restricted region of a cortical projection area. We have shown that stimulation of somatosensory areas I and II (SI and SII) may block an afferent volley at the level of the thalamic relay nucleus, and that SII may be selectively blocked by stimulation of SI. Also definite somatosensory connections have been demonstrated between SII, SI, and the motor cortex. We suggest that common mechanisms underlie the generation of focal reactions in projection areas of the cortex induced by stimulation of various structures. The properties of two groups of neurones from area SII are described: those having a short latency and receiving direct projections from the thalamic relay nucleus, and those of long latent period with a well-marked convergence, and reacting to stimulation of various afferent pathways. It is suggested that each path to a local point of a cortical projection areas terminates with its relay element. The signal is then directed to a common intracortical system of neurones where signals from various sources occurs (afferent, interhemispherical, subcortico-cortical, and intracortical) converge and interact. All groups of neurones are involved in the formation of the common components of evoked potentials.Presented to the All-Union Symposium: "Electrical responses of the cerebral cortex to afferent stimuli," Kiev, October, 1969.Institute of Normal and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 155–165, March–April, 1970.     

Motor cortex gates vibrissal responses in a thalamocortical projection pathway

Higher-order thalamic nuclei receive input from both the cerebral cortex and prethalamic sensory pathways. However, at rest these nuclei appear silent due to inhibitory input from extrathalamic regions, and it has therefore remained unclear how sensory gating of these nuclei takes place. In the rodent, the ventral division of the zona incerta (ZIv) serves as a relay station within the paralemniscal thalamocortical projection pathway for whisker-driven motor activity. Most, perhaps all, ZIv neurons are GABAergic, and recent studies have shown that these cells participate in a feedforward inhibitory circuit that blocks sensory transmission in the thalamus. The present study provides evidence that the stimulation of the vibrissa motor cortex suppresses vibrissal responses in ZIv via an intra-incertal GABAergic circuit. These results provide support for the proposal that sensory transmission operates via a top-down disinhibitory mechanism that is contingent on motor activity.     

Influence of acute cerebellar lesions on somatosensory evoked potentials (SEPs) in cats.

We studied the effect of acute unilateral cerebellar lesions on the cerebello-thalamo-cortical projection in cats. The lesions were classified into two groups according to their extent. In group I the lesion only covered the cerebellar cortex, while in group II both the cerebellar cortex and deep cerebellar nuclei were removed. Early (short-latency) and late (long-latency) waves, evoked by an electrical stimulation of a forelimb, were collected contralateral to the stimulated leg hemisphere. Pre- and postsurgery recordings from primary and non-primary (motor and parietal) cortices were compared. Cerebellar impairment had a strong influence on discharges of all the considered cortical areas. Early non-primary and primary responses increased in group I and remained unchanged in group II. Late somatosensory evoked potentials components were suppressed in both groups. An inhibitory influence of the cerebellar cortex on the thalamo-cortical projection was confirmed. Changes within the primary sensory cortex may suggest an engagement of that area in the compensation process of cerebellar dysfunction shortly after cerebellar lesion. An alteration in the unaffected hemisphere activation indicate that the spino-cerebellar and cerebello-cortical inputs, responsible for somatosensory evoked potentials generation, are regulated through contralateral and ipsilateral pathways. These pathways are unmasked by cerebellar lesion.     

Organization of the Connections between the Parietal Associative Zone and the Primary Sensory Zones in the Cat Neocortex

In acute experiments on cats, we studied the impulse activity of 262 neurons of the parietal associative zone (PAZ, field 5). Among them, 129 cells [100 silent units and 29 units generating background activity (BA)] were identified as output neurons, while 133 cells with the BA were interneurons of the intrinsic cortical neuronal circuits. Electrical stimulation of the primary visual, auditory, or somatosensory cortices evoked no impulse responses in silent output PAZ neurons, while output neurons with the BA and interneurons (more than 65 and 80% of the cell units, respectively) generated clear responses (more frequently, phasic). Stimulation of the auditory and visual cortices exerted mostly inhibitory effects, while stimulation of the somatosensory cortex provided mostly excitatory influences. The ratios of neurons generating primary excitatory and inhibitory responses to stimulation of the visual, auditory, and somatic cortices were 0.3:1, 0.6:1, and 3.2:1, respectively. More than 95% of the field-5 neurons were influenced from the primary sensory zones via di- and/or polysynaptic pathways. Monosynaptic excitatory inputs from the visual cortex were identified for 3.8% of interneurons and 6.9% of output PAZ neurons; for the auditory cortical inputs, the respective figures were 1.7 and 3.5%. Monosynaptic connections with the somatic cortex were found only for 4% of the interneurons under study. It has been concluded that interaction of heteromodal signals coming to the PAZ via the corticopetal and associative inputs occurs on neurons of all the cortical layers.     

Gain modulation from background synaptic input

Gain modulation is a prominent feature of neuronal activity recorded in behaving animals, but the mechanism by which it occurs is unknown. By introducing a barrage of excitatory and inhibitory synaptic conductances that mimics conditions encountered in vivo into pyramidal neurons in slices of rat somatosensory cortex, we show that the gain of a neuronal response to excitatory drive can be modulated by varying the level of "background" synaptic input. Simultaneously increasing both excitatory and inhibitory background firing rates in a balanced manner results in a divisive gain modulation of the neuronal response without appreciable signal-independent increases in firing rate or spike-train variability. These results suggest that, within active cortical circuits, the overall level of synaptic input to a neuron acts as a gain control signal that modulates responsiveness to excitatory drive.