Visuomotor properties of corticotectal cells in area 17 and posteromedial lateral suprasylvian (PMLS) cortex of the cat.

We studied the visuomotor activity of corticotectal (CT) cells in two visual cortical areas [area 17 and the posteromedial lateral suprasylvian cortex (PMLS)] of the cat. The cats were trained in simple oculomotor tasks, and head position was fixed. Most CT cells in both cortical areas gave a vigorous discharge to a small stimulus used to control gaze when it fell within the retinotopically defined visual field. However, the vigor of the visual response did not predict latency to initiate a saccade, saccade velocity, amplitude, or even if a saccade would be made, minimizing any potential role these cells might have in premotor or attentional processes. Most CT cells in both areas were selective for direction of stimulus motion, and cells in PMLS showed a direction preference favoring motion away from points of central gaze. CT cells did not discharge with eye movements in the dark. During eye movements in the light, many CT cells in area 17 increased their activity. In contrast, cells in PMLS, including CT cells, were generally unresponsive during saccades. Paradoxically, cells in PMLS responded vigorously to stimuli moving at saccadic velocities, indicating that the oculomotor system suppresses visual activity elicited by moving the retina across an illuminated scene. Nearly all CT cells showed oscillatory activity in the frequency range of 20-90 Hz, especially in response to visual stimuli. However, this activity was capricious; strong oscillations in one trial could disappear in the next despite identical stimulus conditions. Although the CT cells in both of these regions share many characteristics, the direction anisotropy and the suppression of activity during eye movements which characterize the neurons in PMLS suggests that these two areas have different roles in facilitating perceptual/motor processes at the level of the superior colliculus.

Burst and tonic response modes in thalamic neurons during sleep and wakefulness.

Thalamic neurons can exhibit two distinct firing modes: tonic and burst. In the lateral geniculate nucleus (LGN), the tonic mode appears as a relatively faithful relay of visual information from retina to cortex. The function of the burst mode is less understood. Its prevalence during slow-wave sleep (SWS) and linkage to synchronous cortical electroencephalogram (EEG) suggest that it has an important role during this form of sleep. Although not nearly as common, bursting can also occur during wakefulness. The goal of this study was to identify conditions that affect burst probability, and to compare burst incidence during sleeping and waking. LGN neurons are extraordinarily heterogenous in the degree to which they burst, during both sleeping and waking. Some LGN neurons never burst under any conditions during wakefulness, and several never burst during slow-wave sleep. During wakefulness, <1% of action potentials were associated with bursting, whereas during sleep this fraction jumps to 18%. Although bursting was most common during slow-wave sleep, more than 50% of the bursting originated from 14% of the LGN cells. Bursting during sleep was largely restricted to episodes lasting 1-5 s, with approximately 47% of these episodes being rhythmic and in the delta frequency range (0.5-4 Hz). In wakefulness, although visual stimulation accounted for the greatest number of bursts, it was still a small fraction of the total response (4%, 742 bursts/17,744 cycles in 93 cells). We identified two variables that appeared to influence burst probability: size of the visual stimuli used to elicit responses and behavioral state. Increased stimulus size increased burst probability. We attribute this to the increased influence large stimuli have on a cell's inhibitory mechanisms. As with sleep, a large fraction of bursting originated from a small number of cells. During visual stimulation, 50% of bursting was generated by 9% of neurons. Increased vigilance was negatively correlated with burst probability. Visual stimuli presented during active fixation (i.e., when the animal must fixate on an overt fixation point) were less likely to produce bursting, than when the same visual stimuli were presented but no fixation point present ("passive" fixation). Such observations suggest that even brief departures from attentive states can hyperpolarize neurons sufficiently to de-inactivate the burst mechanism. Our results provide a new view of the temporal structure of bursting during slow-wave sleep; one that supports episodic rhythmic activity in the intact animal. In addition, because bursting could be tied to specific conditions within wakefulness, we suggest that bursting has a specific function within that state.

Widespread distribution of visual responsiveness in frontal, prefrontal, and prelimbic cortical areas of the cat: an electrophysiologic investigation.

By using multiple-unit recording techniques, we explored the visual responsiveness of regions of cortex in and around the area described by others as the cat's "frontal eye fields" (Schlag J, Schlag-Rey M [1970] Brain Res 22:1-13; Guitton D, Mandl G [1978] Brain Res 149:295-312; Pigarev IN [1984] Neirofiziologiia 16:761-766). Our exploration included most of the cat's motor areas (subdivisions of areas 4 and 6) as well as prefrontal and prelimbic regions. Visual responses were routinely obtained from portions of each of the areas we explored, including prefrontal and prelimbic cortex. The qualitative characteristics of visual responses appeared to vary with cytoarchitectonic area. With few exceptions, receptive fields in these areas were large (most exceeding 2,500 deg2) and included the area centralis. Such large fields and inclusion of central vision at nearly all sites precluded retinotopic organization and prevented delineating distinct visual field representations. The most reliable and robust visual activity was observed on the ventral bank of the cruciate sulcus in area 6aalpha. The regions reported to correspond to the "frontal eye fields" did not exhibit any unique visual properties that distinguished them from surrounding areas. The widespread distribution of visually driven activity we observed is consistent with the known pattern of both cortical and subcortical inputs to this broad region of cortex. The observation of visually responsive activity across broad regions of cortex that is nominally motor is consistent with recent studies involving awake animals.

Thalamic control of cat lateral suprasylvian visual area: relation to patchy association projections from area 18.

In this study, we examined functional contributions of major subdivisions of the lateral geniculate nucleus to the cat's lateral suprasylvian visual area (LS) in relation to the patchy horizontal distributions of association inputs. Multiple-unit activity driven via the contralateral eye was assessed during reversible blockade of the retinotopically corresponding part of layer A, the C layers as a group, or the medial interlaminar nucleus (MIN). Inactivating each of these targets reduced activity at some cortical sites, with inactivation of layer A having, on average, the largest effect. Activity was rarely abolished by inactivation of a single target, indicating that most LS sites receive multiple inputs. Dependence on layer A was strongly correlated with the horizontal distribution of association inputs from area 18. Closely spaced injections of anatomical tracers into extensive regions of area 18 resulted in patches of terminal label in lateral suprasylvian cortex. Activity inside the patches was relatively dependent on layer A, whereas that outside the patches was not. Dependence on the MIN and layer A were negatively correlated, suggesting that inputs dominated by the MIN and layer A were concentrated in independent sets of patches. These results indicate that the anatomically observed patchy projections reflect the functional consequences of geniculate lamination. The A layers are high-acuity relays, whereas the MIN is probably a specialization for dim-light vision (Lee et al., 1984; Lee et al., 1992). We propose that the partial overlap of inputs dominated by the A layers and the MIN allows dynamic shifts in their relative contributions to LS responses, optimizing the balance of high-acuity and high-sensitivity channels over a wide range of illumination conditions.

Thalamic control of cat area-18 supragranular layers: simple cells, complex cells, and cells projecting to the lateral suprasylvian visual area.

The goal of this study was to determine the effects of inactivating layer A or the C layers of the cat lateral geniculate nucleus on the supragranular layers of area 18, including cells antidromically activated from the lateral suprasylvian visual area (LS). Isolated cells were visually driven via the contralateral eye while the retinotopically corresponding regions of layer A or, in some cases, the C layers were reversibly inactivated with injections of cobaltous chloride. Simple cells were frequently encountered and were on average more dependent upon layer A than were complex cells, a result qualitatively similar to that found previously in area 17 (Malpeli, 1983; Malpeli et al., 1986). However, the influence of the C layers on area 18 was much more apparent than for area 17. In area 18, as in area 17, the dependence of simple cells on particular geniculate layers appears to follow the terminal patterns of the major direct geniculate inputs. Those simple cells most dependent on layer A were located in lower layer 3. Simple cells in upper layer 3, like complex cells, showed little dependence on layer A, but were strongly dependent upon the C layers. All cells antidromically activated from LS were simple cells with rapidly conducting axons. They had, on average, the same moderately strong dependence on layer A as the patches of LS receiving area 18 input (Lee et al., 1997), supporting the conclusion that the influence of layer A in these patches is largely transmitted via association inputs from area 18. These results demonstrate that simple cells play a major role in association pathways.

Corticostriatal and corticotectal neurons in area 6 of the cat during fixation and eye movements.

We studied the visuomotor properties of 54 corticostriatal (CS) and 38 corticotectal (CT) neurons in a region of area 6 that largely corresponds to the cat's frontal eye fields in five cats trained to do simple oculomotor tasks. Overall, these cells were similar to the general population of area 6 neurons described in the previous paper (Weyand & Gafka, 1998), with very few showing pre-saccadic activity. Likewise, CS and CT cells were similar to each other, although only CS cells showed activity exclusively related to the delivery of the reward and CT cells were more likely to be active during saccades. Variability in visual response latencies and the observation that some cells showed initial visual suppression suggest CS and CT cells reflect the output of a variety of intracortical circuits. Despite similar response properties and overlapping laminar origin, CS and CT circuits appear largely independent. Among 32 cells that we could electrically activate (either synaptically or antidromically) from the superior colliculus, only two could also be activated from stimulating electrodes in the striatum. Similarly, 23 of 25 cells electrically activated from the striatum could not be activated from the superior colliculus. Although few of these efferent cells exhibited pre-motor activity, many exhibit properties that could contribute to gaze control.

Activity of neurons in area 6 of the cat during fixation and eye movements.

We studied the visuomotor properties of 645 neurons in area 6 of five cats trained in oculomotor tasks. The area we recorded from corresponds well with territories believed to contain the feline homologue of the frontal eye fields observed in primates. Despite an expectation that cells with pre-saccadic activity would be common, only a small fraction (approximately 5%) of the cells displayed activity that could be linked to subsequent saccadic eye movements. These pre-motor cells appeared to be distributed over a broad region of cortex mixed in with other cell types. As in primates, saccade-related activity tended to occur only during "purposeful" saccades. At least 30% (208/645) of the neurons were visual, with many of these cells possessing huge receptive fields that appeared to include the entire contralateral visual field. Visual responsiveness was generally attenuated by fixation during the oculomotor tasks. Although attentional mechanisms may play a role in this attenuation, this cortical area also exhibits powerful lateral interactions in which spatially displaced visual stimuli suppress each other. Most cells, visually responsive or not, were affected by fixation. Nearly equal proportions of cells showed increases or decreases in activity during fixation. For many of the cells affected by fixation, the source of this modulation appears to reflect cognitive, rather than sensory or motor processes. This included cells that showed anticipatory activity, and cells that responded to the reward only when it was presented in the context of the task. Based on the paucity of pre-saccadic neurons, it would be difficult to conclude that this region of cortex in the cat is homologous to the frontal eye fields of the monkey. However, when considered in the context of differences in the oculomotor habits of these two animals, we believe the homology fits. In addition to pre-motor neurons, the properties of several other cell types found in this area could contribute to the control of gaze.

Responses of neurons in primary visual cortex are modulated by eye position.

1. We tested the effects of eye position on the visual excitability of 88 neurons in the primary visual cortex of awake cats trained in oculomotor tasks. For most cells, we examined responses evoked by retinotopically identical stimuli for centered gaze, 8 degrees to the left of center, and 8 degrees to the right of center. 2. An effect of eye position was observed for 40% of the cells. For 13%, responsiveness varied by a factor of 2 or more. Most commonly, response was maximal with gaze shifted to one side, minimal when shifted to the opposite side, and intermediate for centered fixation. The exceptions were four cells for which excitability varied symmetrically with fixations to either side of center. 3. Variability in excitability associated with eye position is a wide-spread phenomenon, having been observed in the lateral geniculate nucleus, V1, and extrastriate cortex. These results are consistent with the belief that such variability is utilized in constructing a head-centered frame of reference from a retinotopic input.

A new method of mounting and directing chronically implanted microdrives.

A new method of mounting a microdrive on the skull and adjusting the trajectories of microelectrodes is described. The key to this system is a swiveling guide tube held in a small, skull-mounted base by a low-melting-point metal alloy. The microelectrode is advanced via a modified, commercially available, miniature microdrive screwed onto the guide tube. To change the trajectory of the electrode, the alloy is melted in place, and the guide tube swiveled to a different angle. The microelectrode or the trajectory of the pass may be changed in a few minutes while easily maintaining aseptic conditions. The entire assembly is small enough so that several can be simultaneously implanted on the skull of a cat. A metal crown is used to fix the head during recording sessions or to hold a fiberglass cap that protects all skull implants between sessions. Since the microdrives need not be removed between recording sessions, an electrode pass may be continued from day to day. Although guide tubes are generally employed only for subcortical structures, this arrangement also works well when recording from cortex in deeper parts of gyri.

Area 18 corticotectal cells: response properties and identification of sustaining geniculate inputs.

1. We examined the response properties and geniculate inputs of 35 antidromically identified corticotectal (CT) cells within area 18 of the paralyzed, anesthetized cat. Twenty-three were either standard complex or hypercomplex, 11 were special complex, and 1 was simple. 2. The response properties of CT cells in area 18 were in general quite similar to those examined in a previous study of area 17 CT cells, including similar proportions of standard and special complex CT cells, virtually identical length-response functions, and similar orientation and direction tuning. 3. Area 18 CT cells are rapidly conducting. They are considerably faster than area 17 CT cells. 4. We investigated the composition of thalamic inputs to CT cells by reversibly inactivating a portion of layer A and/or the C layers of the dorsal lateral geniculate nucleus with injections of cobaltous chloride. Blocking layer A strongly attenuated the visual responsiveness of about half of the cells tested. Blocking the C layers alone generally had only moderate effects, but simultaneous blockade of layer A and the C layers demonstrated a substantial C-layer input to many cells. Unlike area 17 in which there is a strong correlation between CT cell class and dependence on layer A, no single receptive-field parameter nor set of parameters was correlated with dependence on layer A. However, cells least affected by simultaneous blockade of layer A and the C layers were special complex, suggesting that, as in area 17, area 18 special complex CT cells integrate more geniculate inputs than standard complex CT cells. 5. We propose that the similarities of response properties of area 17 and area 18 CT cells results from their participation in similar interlaminar columnar circuits and that differences in the patterns of geniculate control reflect differences in the global patterns of geniculate inputs to these two areas.

Corticogeniculate neurons, corticotectal neurons, and suspected interneurons in visual cortex of awake rabbits: receptive-field properties, axonal properties, and effects of EEG arousal.

The intrinsic stability of the rabbit eye was exploited to enable receptive-field analysis of antidromically identified corticotectal (CT) neurons (n = 101) and corticogeniculate (CG) neurons (n = 124) in visual area I of awake rabbits. Eye position was monitored to within 1/5 degrees. We also studied the receptive-field properties of neurons synaptically activated via electrical stimulation of the dorsal lateral geniculate nucleus (LGNd). Whereas most CT neurons had either complex (59%) or motion/uniform (15%) receptive fields, we also found CT neurons with simple (9%) and concentric (4%) receptive fields. Most complex CT cells were broadly tuned to both stimulus orientation and velocity, but only 41% of these cells were directionally selective. We could elicit no visual responses from 6% of CT cells, and these cells had significantly lower conduction velocities than visually responsive CT cells. The median spontaneous firing rates for all classes of CT neurons were 4-8 spikes/s. CG neurons had primarily simple (60%) and concentric (9%) receptive fields, and none of these cells had complex receptive fields. CG simple cells were more narrowly tuned to both stimulus orientation and velocity than were complex CT cells, and most (85%) were directionally selective. Axonal conduction velocities of CG neurons (mean = 1.2 m/s) were much lower than those of CT neurons (mean = 6.4 m/s), and CG neurons that were visually unresponsive (23%) had lower axonal conduction velocities than did visually responsive CG neurons. Some visually unresponsive CG neurons (14%) responded with saccadic eye movements. The median spontaneous firing rates for all classes of CG neurons were less than 1 spike/s. All neurons synaptically activated via LGNd stimulation at latencies of less than 2.0 ms had receptive fields that were not orientation selective (89% motion/uniform, 11% concentric), whereas most cells with orientation-selective receptive fields had considerably longer synaptic latencies. Most short-latency motion/uniform neurons responded to electrical stimulation of the LGNd (and visual area II) with a high-frequency burst (500-900 Hz) of three or more spikes. Action potentials of these neurons were of short duration, thresholds of synaptic activation were low, and spontaneous firing rates were the highest seen in rabbit visual cortex. These properties are similar to those reported for interneurons in several regions in mammalian central nervous system. Nonvisual sensory stimuli that resulted in electroencephalographic arousal (hippocampal theta activity) had a profound effect on the visual responses of many visual cortical neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Cat area 17. I. Pattern of thalamic control of cortical layers.

Reversible inactivation of individual layers of the cat lateral geniculate and medial interlaminar nuclei was used to investigate the necessary and sufficient inputs for maintaining visually driven activity and receptive field properties in area 17. Neither orientation selectivity nor direction selectivity depends on any individual geniculate layer. We identified two groups of cortical layers on the basis of the pattern of thalamic inputs providing visual driving through the contralateral eye. One group, consisting of layers 4 and 6, has geniculate layer A as its only necessary and sufficient input. The other, consisting of supragranular layers, integrates at least two sufficient thalamic inputs, one of which is layer A. Several major receptive field properties are independently generated in these two groups of layers.

Cat area 17. II. Response properties of infragranular layer neurons in the absence of supragranular layer activity.

Response properties of cells in the infragranular layers of cortical area 17 of the cat were examined in the absence of input from supragranular layers. Supragranular activity was silenced either reversibly by cooling the surface of cortex or permanently by making a cryogenic lesion of the supragranular layers. Visually driven responses of cells throughout the cortical column were recorded with a linear array of electrodes. Most infragranular layer cells continued to be visually responsive in the absence of supragranular layer input. These cells were similar to normal infragranular layer cells on measures of visual responsiveness, orientation selectivity, and direction selectivity. Special complex, but not standard complex, cells were absent in layer 5 when supragranular layers were destroyed. We found no evidence for a selective effect of removal of supragranular activity on the response properties of cells in layer 6. We propose that the intracolumnar projection from the supragranular layers drives the special complex cells of layer 5, but is not necessary for the visual driving of most other infragranular layer cells. This projection does not impose selectivity for stimulus orientation or direction on the remaining active cells of the infragranular layers.

Cat area 17. III. Response properties and orientation anisotropies of corticotectal cells.

The receptive field properties of antidromically identified corticotectal (CT) cells in area 17 were explored in the paralyzed, anesthetized cat. To compare these with another population of infragranular cells, we also examined the receptive field properties of cells in layer 6. Sixty percent of our sample of CT cells showed increased response to increased stimulus length (length summation) and were classified as standard complex cells. The other 40% showed little or no length summation, were generally end stopped, and were classified as special complex cells. Standard and special complex CT cells have complementary orientation anisotropies: the distribution of orientation preferences of standard complex cells is biased toward obliquely oriented stimuli, whereas special complex cells are biased toward horizontally and vertically oriented stimuli. The receptive fields of the cells in our sample were primarily along the horizontal meridian so we cannot determine if these anisotropies are defined relative to the vertical meridian or relative to the meridian passing through the receptive field. The effects of these anisotropies in preferred orientation are minimized by the broad orientation tuning of CT cells. There was no simple relationship between the direction bias of CT cells and the reported direction bias of tectal cells. In contrast to the heterogeneity of corticotectal cells, layer 6 cells uniformly showed strong length summation, tight orientation tuning, and little spontaneous activity.

Cat area 17. IV. Two types of corticotectal cells defined by controlling geniculate inputs.

The dependence of cat area 17 corticotectal (CT) cells on specific subdivisions of the dorsal lateral geniculate (LGN) and medial interlaminar nuclei (MIN) was examined using reversible inactivation techniques. Inactivation of layer C of the LGN or layer 1 of the MIN did not block visual activity of CT cells driven through the contralateral eye. Inactivation of layer A of the LGN revealed two populations of CT cells: one strongly dependent on layer A and one whose visually driven activity survived layer A inactivation. CT cells that responded best to short stimuli (special complex cells) were least dependent on layer A, whereas cells that responded best to long stimuli (standard complex cells) were most dependent on layer A. We propose a model of the intracortical circuitry of these two types of CT cells. Standard complex cells, which are heavily dependent on layer A, receive sustaining visual input through layers 4 and/or 6. Special complex cells, which are not dependent on any single layer of the lateral geniculate nucleus, receive sustaining visual input from supragranular layers.

Thalamic inputs to visual areas 1 and 2 in the rabbit.

Thalamic projections to two cortical representations of the visual field, visual areas 1 and 2 (V1, V2), in the rabbit were studied by using the retrograde transport of horseradish peroxidase (HRP). Physiological guidance was employed to inject small amounts of HRP into topographically defined regions of V1 or V2. Injections restricted to V1 revealed a dense projection from the dorsal lateral geniculate nucleus as well as projections from the pulvinar, the posterior thalamic nucleus, and the ventral lateral nucleus. Injections restricted to V2 revealed projections from the pulvinar, the ventral lateral nucleus, and the posterior thalamic nucleus, but only a slight projection from the dorsal lateral geniculate nucleus. V2, but not V1, receives an input from neurons within the fiber plexus between the dorsal lateral geniculate nucleus and the pulvinar. Finally, the neurons in the lateral geniculate nucleus that project to V2 have larger somata on average than those that project to V1 (means = 18.25 micron vs. 14.04 micron, P less than .001).

Rabbit cingulate cortex: cytoarchitecture, physiological border with visual cortex, and afferent cortical connections of visual, motor, postsubicular, and intracingulate origin.

The connections of cingulate cortex with visual, motor, and parahippocampal cortices in the rabbit brain are evaluated by using a modified Brodmann cytoarchitectural scheme, electrophysiological mapping techniques, and the pathway tracers horseradish peroxidase (HRP) and tritiated amino acids. Rabbit cingulate cortex can be divided into areas 25, 24, and 29. Area 29 is of particular interest because area 29d has a lateral extension with a granular layer IV, area 29b has a caudal extension in which the connections differ from anterior area 29b, and there is a prominent area 29e. Cytoarchitectural delineation of the lateral border of area 29d with area 17 closely approximates the medial edge of the visual field representation in area 17 as determined electrophysiologically. The main interconnections between visual and cingulate cortices occur between cingulate areas 24b and 29d and visual areas 18 and medial parts of area 17. Projections between areas 29d and 18 are organized in a loose topographic fashion with rostral parts of each and caudal parts of each being reciprocally connected. Neurons mainly in superficial layer II-III of areas 17 and 18 project to layer I of area 29d, while the reciprocal projection originates from neurons in layer V of area 29d and project mainly to layer I of areas 17 and 18. The medial portion of motor area 8 projects to areas 18 and 29d and has a smaller projection to area 17. Postsubicular area 48 is reciprocally connected with area 29d, and it also projects to areas 29b and c. The subiculum projects to areas 29a and 29c but only to the anterior two-thirds of area 29b not the posterior one-third. Rostral area 29d receives the most extensive intrinsic cingulate projections including those from all major cytoarchitectural divisions. Interconnections between areas 29d and 29b appear to be topographically organized in the rostrocaudal plane. Area 29c projects more heavily to area 29b than vice versa. Finally area 29d projects mainly to area 24b in anterior cingulate cortex. In conclusion, rostral area 29d has extensive connections with visual areas 17 and 18, motor area 8, and all subdivisions of cingulate cortex. In light of these connections, it may play a pivotal role in associative functions of the rabbit cerebral cortex including visuomotor integration.

Receptive-field and axonal properties of neurons in the dorsal lateral geniculate nucleus of awake unparalyzed rabbits.

The intrinsic stability of the rabbit eye was exploited to enable receptive-field analysis of LGNd neurons and optic tract axons in the awake, unparalyzed state. We found eye position to remain within a range of less than 1.0 degrees for periods of 4-5 min, and in some cases for periods in excess of 10 min. Such stability is comparable to that seen in awake monkeys that have been trained to fixate. Receptive fields of dorsal lateral geniculate nucleus (LGNd) neurons were analyzed, and approximately 84% were concentrically organized. This is a higher value than previously reported in this species. In addition, the receptive-field centers of concentric cells were much smaller than those previously reported (mean diameter = 2.5 degrees). Most remaining neurons in the LGNd were either directionally selective (6.5%) or motion/uniform (6.5%). Concentric cells were classified as sustained or transient based on response duration to standing contrast. In the LGNd the receptive fields of sustained concentric cells were predominantly near the horizontal meridian, within the representation of the visual streak, while the receptive-field positions of transient concentric cells were more prevalent in the upper visual field. In the optic tract the receptive-field positions of both sustained and transient cells were more evenly distributed than was seen in the LGNd. Sustained and transient concentric cells in LGNd showed primarily nonlinear spatial summation. The receptive-field properties of LGNd neurons were related to geniculocortical antidromic latency. Most LGNd neurons of all receptive-field classes projected axons to the visual cortex. Thus, any intrinsic interneurons in the rabbit LGNd may have receptive-field properties similar to those of some principal neurons. There was significant overlap in the distribution of antidromic latencies in neurons of different receptive-field classes. Concentric sustained neurons, however, did conduct somewhat more slowly than did concentric transient neurons. Nonvisual sensory stimuli that resulted in EEG arousal (hippocampal theta activity) had a profound effect on the response duration of most (28/32) sustained concentric neurons. For these cells, the sustained response to standing contrast began to diminish and sometimes disappeared after 2-15 s. However, arousing stimuli that resulted in hippocampal theta activity reestablished the sustained response. Such arousing stimuli usually had little or no effect on the discharge of the cell in the absence of visual stimuli. Arousing stimuli had no effect on optic tract axons with sustained concentric receptive-field properties.(ABSTRACT TRUNCATED AT 400 WORDS)

Cat medial interlaminar nucleus: retinotopy, relation to tapetum and implications for scotopic vision.

The medial interlaminar nucleus (MIN) of the cat was electrophysiologically mapped in sufficient detail to resolve individual laminae and to allow reconstruction of isoazimuth and isoelevation lines in coronal, sagittal, and horizontal planes. The electrophysiologically defined laminar pattern was in agreement with that revealed anatomically in the same animal, as well as with the general pattern revealed by anatomical methods in several unmapped nuclei. The MIN is made up of three distinct layers, each receiving inputs from one hemi-retina but none representing an entire hemifield. We confirm the findings of Guillery et al. (19) that the contralateral hemifield is represented in layers 1 and 2 through the contralateral and ipsilateral eyes, respectively, and that layer 3 represents the ipsilateral hemifield through the contralateral eye. Elevation and absolute value of azimuth are represented continuously through the MIN. When an isoazimuth line crosses the border between layer 3 and either layer 1 or 2, the absolute value of azimuth is maintained but the sign of the azimuth changes. Adjacent points on either side of this border represent mirror symmetrical visual directions on opposite sides of the vertical meridian. This indicates that the distance from the vertical meridian is an independently coded parameter within the geniculate complex. There is virtually no nasotemporal overlap in any layer of the MIN. The function relating magnification (mm3 per steradian) to eccentricity is strikingly similar to the function relating retinal ganglion cell density to eccentricity, suggesting that a constant fraction of retinal ganglion cells project to the MIN at all eccentricities. Most of the volume of each MIN layer is devoted to lower visual fields. Analysis of the geniculate retinotopic maps of Sanderson (46) reveals no equivalent bias toward lower visual fields in the dorsal lateral geniculate nucleus. The MIN represents a region of retina roughly coincident with the tapetum, suggesting a role of the MIN in dim-light vision.

Efferent systems of the rabbit visual cortex: laminar distribution of the cells of origin, axonal conduction velocities, and identification of axonal branches.

Several efferent systems of visual area I in Dutch rabbits were studied with anatomical (horseradish peroxidase) and physiological (antidromic) methods. Anatomical studies provided information regarding the laminar origin of the projections to the contralateral hemisphere, visual area II, the dorsal lateral geniculate nucleus, and the superior colliculus. Physiological studies provided information regarding conduction velocities and multiple destinations of efferent axons. Both the callosal projection and the projection to V-II were shown to originate primarily in layer II-III. However, approximately 10-20% of the callosal projection and 20-40% of the projection to V-II originated in layers IV and V. In contrast, the projection to the dorsal lateral geniculate nucleus originated nearly exclusively in layer VI, while corticotectal neurons occurred primarily in layer V. A significant number of corticotectal neurons were, however, found in layer IV. Thus, the above efferent systems were found to differ in their principal laminar origin and in the diffuseness of that origin. The origins of corticocortical projections were considerably more diffuse than those of corticofugal projections. In addition to differences in laminar origin, efferent systems also differed significantly in the conduction velocities of their axons. The projection to visual area II and to the lateral geniculate nucleus consisted primarily of very slowly conducting axons, while the projection to the superior colliculus was fast conduction. The callosal projection consisted of both slow and fact conduction axons. Finally, the question of branching of V-I efferent axons was addressed. Although the laminar origin of the projections to the contralateral hemisphere and to visual area II overlapped considerably, none of these corticocortical axons could be shown to project to both locations or to a subcortical destination. In contrast, approximately one-third of corticotectal axons were shown to project a collateral into the thalamus. Although the destination of this collateral is unclear, it is medial to the lateral geniculate nucleus and may be the pulvinar.