Pattern motion representation in primary visual cortex is mediated by transcortical feedback.

A highly important question in visual neuroscience is to identify where in the visual system information from different processing channels is integrated to form the complex scenery we perceive. A common view to this question is that information is processed hierarchically because small and selective receptive fields in lower visual areas melt into larger receptive fields in specialized higher visual areas. However, a higher order area in which all incoming signals ultimately converge has not yet been identified. Rather, modulation of subthreshold influences from outside the classical receptive field related to contextual integration occurs already in early visual areas. So far it is unclear how these influences are mediated (Gilbert, 1998; Angelucci and Bullier, 2003; Gilbert and Sigman, 2007). In the present study, we show that feedback connections from a higher motion processing area critically influence the integration of subthreshold global motion cues in early visual areas. Global motion cues are theoretically not discernible for a local motion detector in V1, however, imprints of pattern motion have been observed in this area (Guo et al., 2004; Schmidt et al., 2006). By combining reversible thermal deactivation and optical imaging of intrinsic signals we demonstrate that feedback signals from the posteromedial suprasylvian sulcus are critical for the discrimination between global and local motions already in early visual areas. These results suggest that global features of the visual scenery are fed back to lower visual processing units in order to facilitate the integration of local cues into a global construct.

Low frequency transcranial magnetic stimulation on the posterior parietal cortex induces visuotopically specific neglect-like syndrome.

The visuo-parietal (VP) region of the cerebral cortex is critically involved in the generation of orienting responses towards visual stimuli. In this study we use repetitive transcranial magnetic stimulation (rTMS) to unilaterally and non-invasively deactivate the VP cortex during a simple spatial visual detection task tested in real space. Adult cats were intensively trained over 4 months on a task requiring them to detect and orient to a peripheral punctuate static LED presented at a peripheral location between 0 degrees and 90 degrees , to the right or left of a 0 degrees fixation target. In 16 different interleaved sessions, real or sham low frequency (1 Hz) rTMS was unilaterally applied during 20 min (1,200 pulses) to the VP cortex. The percentage of mistakes detecting and orienting to contralateral visual targets increased significantly during the 15-20 min immediately following real but not sham rTMS. Behavioral deficits were most marked in peripheral eccentricities, whereas more central locations were largely unaffected. Performance returned to baseline (pre-TMS) levels when animals were tested 45 min later and remained in pre-TMS levels 24 h after the end of the stimulation. Our results confirm that the VP cortex of the cat is critical for successful detection and orienting to visual stimuli presented in the corresponding contralateral visual field. In addition, we show that rTMS disrupts a robust behavioral task known to depend on VP cortex and does so for the far periphery of the visual field, but not for more central targets.

Role of the superior colliculus in analyses of space: superficial and intermediate layer contributions to visual orienting, auditory orienting, and visuospatial discriminations during unilateral and bilateral deactivations.

The superior colliculus (SC) has been implicated in spatial analyses of the environment, although few behavioral studies have explicitly tested this role. To test its imputed role in spatial analyses, we used a battery of four spatial tasks combined with unilateral and bilateral cooling deactivation of the upper and intermediate layers of the superior colliculus. We tested the abilities of cats to orient to three different stimuli: (1) moving visual, (2) stationary visual, (3) stationary white-noise aural. Furthermore, we tested the ability of the cats to discriminate the relative spatial position of a landmark. Unilateral cooling deactivation of the superficial layers of the SC induced a profound neglect of both moving and stationary visual stimuli presented in, and landmark objects located within, the contralateral hemifield. However, responses to auditory stimuli were unimpaired. Unilateral cooling deactivation of both the superficial and intermediate layers induced a profound contralateral neglect of the auditory stimulus. Additional and equivalent deactivation of the opposite SC largely restored orienting to either moving visual or auditory stimuli, and restored landmark position reporting to normal levels. However, during bilateral SC deactivation, orienting to the static visual stimulus was abolished throughout the entire visual field. Overall, unilateral SC deactivation results show that the upper and intermediate layers of the SC contribute in different ways to guiding behavioral responses to visual and auditory stimuli cues. Finally, bilateral superior colliculus deactivations reveal that other structures are sufficient to support spatial analyses and guide visual behaviors in the absence of neural operations in the superior colliculus, but only under certain circumstances.

Task-specific reversal of visual hemineglect following bilateral reversible deactivation of posterior parietal cortex: a comparison with deactivation of the superior colliculus.

The purpose of the present study was to compare and contrast behavioral performance on three different tasks of spatial cognition during unilateral and bilateral reversible deactivation of posterior parietal cortex. Specifically, we examined posterior middle suprasylvian (pMS) sulcal cortex in adult cats during temporary and reversible cooling deactivation. In Task 1, the cats oriented to a high-contrast, black visual stimulus moved into the visual field periphery. In Task 2, the cats oriented to a static light-emitting diode (LED). Task 3 examined the cats' ability to determine whether a black-and-white checkered, landmark box was closer to the right or left side of the testing apparatus. Following training on all tasks, cryoloops were implanted bilaterally within the pMS sulcus. Unilateral deactivation of pMS sulcal cortex resulted in virtually no responses to either moved or static stimuli and virtually no responses to landmarks presented in the contralateral hemifield, and a profound contralateral hemifield neglect was induced. Responses to stimuli and landmarks presented in the ipsilateral hemifield were unimpaired. Additive, bilateral cooling of the homotopic region in the contralateral hemisphere, but not an adjacent region, resulted in reversal of the initial hemineglect for the moved stimulus, yet induced a complete failure to orient to peripheral static LED stimuli. Bilateral cooling also reversed the contralateral neglect of the landmark, but then cats could not accurately determine position of the landmark anywhere in the visual field because performance was reduced to chance levels for all landmark loci in both hemifields. In this instance, as the contralateral neglect disappeared during bilateral cooling of pMS cortex, a new spatial discrimination deficit was revealed across the entire visual field. We conclude that pMS cortex contributes in multiple ways to the analyses of space, and that these contributions cannot be safely predicted from analyses of unilateral deactivations or from one task to another. Moreover, it is clear that other structures are capable of guiding orienting to high contrast, moved targets when pMS cortex is eliminated from brain circuitry. However, these same structures are incapable of supporting either orienting to static stimuli or analyses of spatial relations as tested with the landmark task. The impact of reversible deactivation of the superior colliculus on these same tasks is discussed.

Perinatal-lesion-induced reorganization of cerebral functions revealed using reversible cooling deactivation and attentional tasks.

We tested the concept that lesions of primary visual cortical areas 17 and 18 sustained on the day of birth induce a redistribution of cerebral operations underlying the ability to disengage visual attention and then redirect it to a new location. In cats, these operations are normally highly localizable to posterior middle suprasylvian (pMS) cortex. Three stimulation paradigms were used: (i) movement of a high contrast visual stimulus into the visual field; (ii) illumination of a static light-emitting diode (LED) stimulus; and (iii) a control static auditory stimulus. To test for the redistribution of critical neural operations, cryoloops were implanted bilaterally in the pMS sulcus and in contact with ventral posterior suprasylvian (vPS) cortex. Separate and combined deactivations of pMS and vPS cortices in cats with early lesions of primary visual cortex showed that full, unilateral deactivation of pMS cortex only partially impaired the ability to detect and orient to stimuli moved into the contracooled hemifield. Much more complete impairment required the additional deactivation of ipsilateral vPS cortex. Bilateral pMS deactivation alone, or in combination with bilateral vPS deactivation, largely reversed the unilateral contracooled neglect. For the orienting to static, illuminated LED stimuli, unilateral deactivation of pMS cortex was sufficient to fully impair orienting to stimuli presented in the contracooled hemifield. Bilateral pMS deactivation induced an almost complete visual-field-wide neglect of stimuli. On its own, unilateral deactivation of vPS cortex was without effect on either task, although bilateral vPS deactivations introduced inconsistencies into the performance. Termination of cooling reversed all deficits. Finally, neither the initial lesion of areas 17 and 18 nor cooling of either the MS or vPS cortex alone, or in combination, interfered with orienting to sound stimuli. Overall, our results provide evidence that at least one highly localizable visual function of normal cerebral cortex is remapped across the cortical surface following the early lesion of primary visual cortical areas 17 and 18. Moreover, the redistribution has spread the essential neural operations from the visual parietal cortex to a normally functionally distinct type of cortex in the visual temporal system.

Feedback connections act on the early part of the responses in monkey visual cortex.

We previously showed that feedback connections from MT play a role in figure/ground segmentation. Figure/ground coding has been described at the V1 level in the late part of the neuronal responses to visual stimuli, and it has been suggested that these late modulations depend on feedback connections. In the present work we tested whether it actually takes time for this information to be fed back to lower order areas. We analyzed the extracellular responses of 169 V1, V2, and V3 neurons that we recorded in two anesthetized macaque monkeys. MT was inactivated by cooling. We studied the time course of the responses of the neurons that were significantly affected by the inactivation of MT to see whether the effects were delayed relative to the onset of the response. We first measured the time course of the feedback influences from MT on V1, V2, and V3 neurons tested with moving stimuli. For the large majority of the 51 neurons for which the response decreased, the effect was present from the beginning of the response. In the responses averaged after normalization, the decrease of response was significant in the first 10-ms bin of response. A similar result was found for six neurons for which the response significantly increased when MT was inactivated. We then looked at the time course of the responses to flashed stimuli (95 neurons). We observed 15 significant decreases of response and 14 significant increases. In both populations, the effects were significant within the first 10 ms of response. For some neurons with increased responses we even observed a shorter latency when MT was inactivated. We measured the latency of the response to the flashed stimuli. We found that even the earliest responding neurons were affected early by the feedback from MT. This was true for the response to flashed and to moving stimuli. These results show that feedback connections are recruited very early for the treatment of visual information. It further indicates that the presence or absence of feedback effects cannot be deduced from the time course of the response modulations.

Graded sparing of visually-guided orienting following primary visual cortex ablations within the first postnatal month.

We compared the abilities of intact cats and cats that incurred lesions of areas 17 and 18 in adulthood, at one month of age (P28), or on the day of birth (P1), to detect and orient towards visual stimuli either moved into or illuminated in the periphery of the visual field, and to detect and orient towards a stationary, broad-band white-noise auditory stimulus. For all groups of cats, movement of a stimulus into the visual field was a more potent stimulus for evoking visually-guided orienting movements than illumination of a static light-emitting diode (LED). The potency of the auditory stimulus was also extremely high. Proficiency on both visual tasks was graded according to the age at which areas 17 and 18 were ablated in the sequence: adult, P1, P28 and intact in the sequence worst-->best performance. The superior performance of the P1- and P28-groups provided evidence for sparing of visually-guided orienting, but the sparing was incomplete because it did not match performance of intact cats. Lesions of areas 17 and 18 incurred in adulthood had no significant impact on orienting to auditory white-noise stimuli. However, orienting performance to auditory stimuli presented in the peripheral quadrants was slightly superior in the P28 group and reduced in the P1 group. Thus, the visual sparing exhibited by the P1 group may be at the expense of highly proficient orienting to auditory cues. Overall, these results extend our knowledge by showing that in addition to P1-cats, cats that incur lesions of areas 17 and 18 at one month-of-age also exhibit sparing of visually-guided orienting, and that the sparing is not confined to a single stimulation paradigm. Finally, the covariation in the magnitude of pathway modifications with the scale of the orienting proficiency in P1- and P28 cats helps to solidify the linkage between rewired brain pathways and spared visually-guided behaviors.

Translaminar differentiation of visually guided behaviors revealed by restricted cerebral cooling deactivation.

The purpose of the present study was to test the hypothesis that superficial and deep layers within a single cerebral region influence cerebral functions and behaviors in different ways. For this test, we selected posterior middle suprasylvian (pMS) sulcal cortex of the cat, a suspected homolog of the area V5 complex of primates, because the region has been implicated in several visually guided behaviors. Cats were trained on three tasks: (1) discrimination of direction of motion; (2) discrimination of static patterns partially obscured by static or moving masks; and (3) visual detection and orienting. Cooling of cryoloops in contact with pMS sulcal cortex to 8+/-1 degrees C selectively and completely impaired performance on the two motion discrimination tasks (1 and 2), while leaving the detection and orienting task (task 3) unimpaired. Further cooling to 3 degrees C resulted in an additional complete impairment of task 3. The 8 degrees C temperature resulted in silencing of neuronal activity in the supragranular layers (I-III) and the 3 degrees C temperature silenced activity throughout the thickness of pMS sulcal cortex. The variation in behavioral performance with covariation of cryoloop temperature and vertical, but not lateral, spread of deactivation shows that deactivation of superficial cerebral layers alone was sufficient to completely impair performance on the two motion discrimination tasks, whereas additional deactivation of the deep layers was essential to block performance on the detection and orienting task. Thus, these results show a functional bipartite division of labor between upper and lower cortical layers that is supported by efferent connectional anatomy. Similar bipartite division into upper and lower layers may be a general feature of cerebral cortical architecture, signal processing and guidance of behavior.

Gamma surgery for vein of Galen malformations.

OBJECT: The goal of this study was to evaluate the results of gamma surgery in nine patients treated for vein of Galen malformations (VGMs). METHODS: A consecutive series of nine VGMs in eight children aged 4 to 14 years and in one adult were treated with gamma surgery. Six of the patients were male, including the adult, and three were female. Among these patients there were three Yasargil Type I, one Type II, two Type III, and three Type IV malformations. Previous embolization had failed in four cases. Three VGMs were treated with gamma surgery twice. An additional patient with a Type III VGM underwent stereotactic angiography in preparation for gamma surgery but was judged to be suitable for direct embolization. Follow-up angiograms were obtained in eight of the VGMs treated. Four no longer filled; one has probably been obliterated, but this cannot be confirmed because the patient refused to undergo final angiography; one patient has residual fistulas not included in the initial treatment field, which were retreated recently; and two other patients have marked reduction of flow through their VGMs. CONCLUSIONS: Gamma surgery is a viable option in the treatment of VGMs in clinically stable patients. Combined endovascular therapy and gamma surgery is of benefit in complex malformations.

Gamma surgery for hemangiopericytomas.

A retrospective analysis of a consecutive series of 12 patients with 15 intracranial hemangiopericytomas treated at the University of Virginia using Gamma surgery is presented. Clinical and radiographic follow up of 3 to 56 months is available for 10 patients with 12 tumors. There was one tumor present at the time of initial Gamma surgery in each patient. Two new tumors occurred in patients previously treated. Nine of the tumors decreased in volume and three remained stable. Four of the nine tumors that shrank later progressed at an average of 22 months after treatment. Of the tumors that decreased in volume and have not progressed, the response has been for an average of 11 months. The follow-up for two tumors that remained unchanged was 10 and 34 months (average 22 months). A third tumor was unchanged at 42 months but the patient died of new disease adjacent to the treated area in the anterior skull base. There were no complications and the quality of life following the procedure was maintained or improved in every case. Gamma surgery is effective in palliating the patients by decreasing tumor volume and delaying recurrence.

Gamma surgery for intracranial metastases from renal cell carcinoma.

OBJECT: The goal of this study was to evaluate the effectiveness and limitations of gamma surgery (GS) in the treatment of renal cell carcinoma that has metastasized to the brain. METHODS: The authors performed a retrospective analysis of a consecutive series of 21 patients with 37 metastatic brain deposits from renal cell carcinoma who were treated with GS at the University of Virginia from 1990 to 1999. Clinical data were available in all patients. No patient died of progression of intracranial disease or deteriorated neurologically following GS. Eight patients clinically improved. Follow-up imaging studies were available for 23 tumors in 12 patients. Nine patients did not undergo follow-up imaging. One patient lived 17 months and succumbed to systemic disease: no brain imaging was performed in this case. Another patient refused further imaging and lived 7 months. Seven patients lived up to 4 months after the procedure; however, their physicians did not require these patients to undergo follow-up imaging examinations because of their general conditions-all had systemic progression of disease. Of the 23 tumors that were observed posttreatment, one remained unchanged in volume, 16 decreased in volume, and six disappeared. No tumor progressed at any time, and there were no radiation-induced changes on follow-up imaging an average of 21 months after GS (range 3-63 months). CONCLUSIONS: Gamma surgery provides an alternative to surgical resection of metastatic brain deposits from renal cell carcinoma. Neurological side effects were seen in only one case; freedom from progression of disease was achieved in all cases.

On variability in the density of corticocortical and thalamocortical connections.

Variability is an important but neglected aspect of connectional neuroanatomy. The quantitative density of the 'same' corticocortical or thalamocortical connection may vary by over two orders of magnitude between different injections of the same tracer. At present, however, the frequency distribution of connection densities is unknown. Therefore, it is unclear what kind of sampling strategies or statistical methods are appropriate for quantitative studies of connectivity. Nor is it clear if the measured variability represents differences between subjects, or if it is simply a consequence of intra-individual differences resulting from experimental technique and the exact placement of tracers relative to local spatial and laminar variation in connectivity. We used quantitative measurements of the density of a large number of corticocortical and thalamocortical connections from our own laboratories and from the literature. Variability in the density of given corticocortical and thalamocortical connections is high, with the standard deviation of density proportional to the mean. The frequency distribution is close to exponential. Therefore, analysis methods relying on the normal distribution are not appropriate. We provide an appendix that gives simple statistical guidance for samples drawn from exponentially distributed data. For a given corticocortical or thalamocortical connection density, between-individual standard deviation is 0.85 to 1.25 times the within-individual standard deviation. Therefore, much of the variability reported in conventional neuroanatomical studies (with one tracer deposited per animal) is due to within-individual factors. We also find that strong, but not weak, corticocortical connections are substantially more variable than thalamocortical connections. We propose that the near exponential distribution of connection densities is a simple consequence of 'patchy' connectivity. We anticipate that connection data will be well described by the negative binomial, a class of distribution that applies to events occurring in clumped or patchy substrates. Local patchiness may be a feature of all corticocortical connections and could explain why strong corticocortical connections are more variable than strong thalamocortical connections. This idea is supported by the columnar patterns of many corticocortical but few thalamocortical connections in the literature.

Contributions of cat posterior parietal cortex to visuospatial discrimination.

The purpose of the present study was to examine the contributions made by cat posterior parietal cortex to the analyses of the relative position of objects in visual space. Two cats were trained on a landmark task in which they learned to report the position of a landmark object relative to a right or left food-reward chamber. Subsequently, three pairs of cooling loops were implanted bilaterally in contact with visuoparietal cortices forming the crown of the middle suprasylvian gyrus (MSg; architectonic area 7) and the banks of the posterior-middle suprasylvian sulcus (pMS sulcal cortex) and in contact with the ventral-posterior suprasylvian (vPS) region of visuotemporal cortex. Bilateral deactivation of pMS sulcal cortex resulted in a profound impairment for all six tested positions of the landmark, yet bilateral deactivation of neither area 7 nor vPS cortex yielded any deficits. In control tasks (visual orienting and object discrimination), there was no evidence for any degree of attentional blindness or impairment of form discrimination during bilateral deactivation of pMS cortex. Therefore, we conclude that bilateral cooling of pMS cortex, but neither area 7 nor vPS cortex, induces a specific deficit in spatial localization as examined with the landmark task. These observations have significant bearing on our understanding of visuospatial processing in cat, monkey, and human cortices.

The cryoloop: an adaptable reversible cooling deactivation method for behavioral or electrophysiological assessment of neural function.

We describe a very adaptable reversible inactivation technique for the behavioral or electrophysiological analysis of neural circuits. The cryoloop device can be permanently implanted or topically applied in an acute preparation to apply cold to discrete surface regions of the central nervous system (e.g. cerebral cortex or midbrain). The cryoloop consists of a custom shaped, stainless steel, hypodermic tubing and cooling is effected by passing chilled methanol through the lumen of the tubing. Cryoloop temperature is monitored by a microthermocouple attached to the union of the loop, and can be maintained within +/- 1 degrees C of a desired temperature. In chronic preparations, implanted cryoloops have been maintained in cats and monkeys for periods in excess of 2 years. After this period there are no structural, metabolic of functional changes in the deactivated tissue, and full reversibility of cooling-induced effects is maintained. Operation of multiple cryoprobes provides great flexibility of experimental protocols, permits double and triple functional dissociations to be made, and strengthens experimental design considerably.

A method to assess the functional impact of cerebral connections on target populations of neurons.

We describe an innovative and tested approach combining two individually potent techniques to visualize simultaneously the functional impact of multiple projections on target populations of neurons in the brain. The rationale is simple: silence a defined set of efferent projections from one cortical region using cooling deactivation and then measure the impact of the deactivation on activities in multiple target regions using 2-deoxyglucose (2DG). This is a straightforward and sound approach because 2DG uptake by neurons reflects levels of underlying neural activity. All distant modifications evoked by the silencing of the set of efferent projections are examined in anatomical tissue and simultaneously for the multiple target sites to provide a global view of the functional impacts of the set of projections on the targets. With this method, downward adjustments of 2DG uptake levels identify removals of net excitatory signals, whereas upward adjustments identify net removals of suppressive influences. Future possible uses and modifications of the technique, including optical imaging, are discussed. Overall, the technique has the potential to provide fundamental, new measures on cerebral network interactions that both complement and extend current static models of cerebral networks and electrophysiological measures of functional impacts on individual neurons.

System-wide repercussions and adaptive plasticity: the sequelae of immature visual cortex damage.

Damage of primary visual cortex in mature mammals severely disrupts vision by disconnecting much of the cognitive processing machinery of extrastriate cortex from its source of visual signals in the retina. However, equivalent lesions incurred early in postnatal life unmask a substantial latent flexibility of the brain to minimize the disruption by specific and ordered pathway expansions that bypass the lesion. The expansions shape pathways from retina through thalamus to extrastriate cortex and onto the midbrain into new, useful forms that are retained into adult life. These useful modifications support relatively normal signal processing in a variety of structures and the sparing of certain visually guided behaviors, such as aspects of complex-pattern vision and localizing objects introduced into the visual field. Thus, both the brain and the individual are optimized, in the absence of primary visual cortex, by adaptations for useful interactions with the environment. So far, the repercussions of early visual cortex lesions have been most thoroughly documented in cats, although it is likely on the basis of known repercussions and similarity of visual system organization and developmental sequence, that broadly equivalent repercussions and adaptations occur in monkeys and humans following early lesions of primary visual cortex. The knowledge gained has implications for devising therapeutic strategies to attenuate defects in vision induced by cortical lesions.

Neuroplasticity in the cat's visual system. Origin, termination, expansion, and increased coupling of the retino-geniculo-middle suprasylvian visual pathway following early ablations of areas 17 and 18.

We used anterograde and retrograde transsynaptic pathway tracing techniques to reveal the retinal origin and the cortical termination of the expanded retino-geniculo-middle suprasylvian (MS) cortex pathway in adult cats which sustained lesions of areas 17 and 18 on the day of birth (P1) or at 1 month of age (P28). Following anterograde transsynaptic transport of tritiated amino acids from the eye, four major results were obtained: (1) a strong and specific pathway from retina through dorsal lateral geniculate nucleus (dLGN) to the posterior half of MS cortex was identified; this pathway is a substantial expansion of an insignificant pathway present in intact cats; (2) the terminus of the pathway was lower layer III and layer IV; (3) contralateral projections were stronger than ipsilateral projections; (4) projections in P28 cats were stronger than those in P1 cats. Following retrograde transsynaptic transport of WGA-HRP from posterior MS cortex, four additional results were obtained: (1) the pathway was enlarged and visuotopically organized; (2) the pathway arose primarily from alpha- and gamma-retinal ganglion cells; (3) a small number of beta-cells in P1 cats and a modest number in P28 cats also contribute to the pathway; (4) the combined numbers of gamma- and beta-cells relative to alpha-cells was greater in temporal retina than in nasal retina. The combined demonstration of both origin and terminus of the pathway with transsynaptic tracers argued strongly for high levels of coupling between primary and secondary pathway limbs in both P1 and P28 cats. This level of coupling, as well as other features of the pathway, have much in common with the retino-geniculo-17/18 pathway of intact cats. However, the retino-geniculo-MS system in P1 cats transmits primarily Y and W signals, in P28 cats X, Y, and W signals; whereas the retino-geniculo-17/18 pathway transmits primarily X and Y signals. These results have implications for understanding the repercussions of early visual cortex lesions in monkeys and humans.

Cortical feedback improves discrimination between figure and background by V1, V2 and V3 neurons.

A single visual stimulus activates neurons in many different cortical areas. A major challenge in cortical physiology is to understand how the neural activity in these numerous active zones leads to a unified percept of the visual scene. The anatomical basis for these interactions is the dense network of connections that link the visual areas. Within this network, feedforward connections transmit signals from lower-order areas such as V1 or V2 to higher-order areas. In addition, there is a dense web of feedback connections which, despite their anatomical prominence, remain functionally mysterious. Here we show, using reversible inactivation of a higher-order area (monkey area V5/MT), that feedback connections serve to amplify and focus activity of neurons in lower-order areas, and that they are important in the differentiation of figure from ground, particularly in the case of stimuli of low visibility. More specifically, we show that feedback connections facilitate responses to objects moving within the classical receptive field; enhance suppression evoked by background stimuli in the surrounding region; and have the strongest effects for stimuli of low salience.

Dissociation of visual and auditory pattern discrimination functions within the cat's temporal cortex.

In ablation-behavior experiments performed in adult cats, a double dissociation was demonstrated between ventral posterior suprasylvian cortex (vPS) and temporo-insular cortex (TI) lesions on complex visual and auditory tasks. Lesions of the vPS cortex resulted in deficits at visual pattern discrimination, but not at a difficult auditory discrimination. By contrast, TI lesions resulted in profound deficits at discriminating complex sounds, but not at discriminating visual patterns. This pattern of dissociation of deficits in cats parallels the dissociation of deficits after inferior temporal versus superior temporal lesions in monkeys and humans.