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B E Stein

Publications and source records attributed to B E Stein.

At least 19 recordsLinked to original sources

The influence of visual and auditory receptive field organization on multisensory integration in the superior colliculus.

The spatial register of the different receptive fields of multisensory neurons in the superior colliculus (SC) plays a significant role in determining the responses of these neurons to cross-modal stimulus combinations. Spatially coincident visual-auditory stimuli fall within these overlapping receptive fields and generally produce response enhancements that exceed the individual modality-specific responses and can exceed their sum. Yet, in this context, it has not been clear how "spatial coincidence" is operationally defined. Given the large size of SC receptive fields, visual and auditory stimuli could be within their respective receptive fields even when there are substantial spatial disparities between them. Indeed, previous observations have raised the possibility that there may be a second level of determinism in how SC neurons deal with the relative spatial locations of within-field cross-modal stimuli; specifically, that multisensory response enhancements become progressively weaker as the within-field visual and auditory stimuli become increasingly disparate. While the present experiments demonstrated that SC multisensory neurons have heterogeneous receptive fields, and that the greatest number of impulses evoked were by stimuli that fell within the area of cross-modal receptive field overlap, they also indicate that there is no systematic relationship between cross-modal stimulus disparity and the magnitude of multisensory response enhancement. Thus, two within-field cross-modal stimuli produced the same proportionate change (i.e., multisensory response enhancement) when they were widely disparate as they did when they overlapped one another in space. These observations indicate that cross-modal spatial coincidence can be defined operationally by the borders of an SC neuron's receptive fields regardless of the size of those receptive fields and/or the absolute spatial disparity between within-field cross-modal stimuli.

Acoustic Stimulation↗

Corticotectal and corticostriatal projections from the frontal eye fields of the cat: an anatomical examination using WGA-HRP.

Corticofugal projections from the frontal eye fields (FEF) are believed to access the superior colliculus (SC) directly (i.e., monosynaptically) and indirectly (i.e., multisynaptically) through the basal ganglia. The present results suggest that these two pathways are derived from largely segregated populations of corticofugal neurons. Furthermore, while the different subregions of the FEF from which these pathways originate have different termination patterns in the basal ganglia (i.e., striatum, ST), they share a common termination pattern in the SC. Injections of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) into the two major subdivisions of the FEF (presylvian and cruciate sulci) resulted in dense label in both the ST (bilaterally) and the SC (ipsilaterally). Corticostriatal labeling was found in the caudal part of the head of the caudate nucleus (heaviest ipsilaterally), with labeling from cruciate injections located ventromedial to that produced by presylvian injections. Only presylvian injections resulted in labeling in the putamen. Retrograde tracing experiments demonstrated that both presylvian and cruciate corticostriatal projections originated from neurons in lamina III and the upper aspects of lamina V. An additional but small group of presylvian corticostriatal projections was found in lamina VI. Corticotectal terminal labeling was restricted to the deep laminae of the SC and was derived exclusively from lamina V neurons in cortex. They differed from their corticostriatal counterparts in laminar/sub-laminar location and in soma sizes.

Animals↗

Two cortical areas mediate multisensory integration in superior colliculus neurons.

The majority of multisensory neurons in the cat superior colliculus (SC) are able to synthesize cross-modal cues (e.g., visual and auditory) and thereby produce responses greater than those elicited by the most effective single modality stimulus and, sometimes, greater than those predicted by the arithmetic sum of their modality-specific responses. The present study examined the role of corticotectal inputs from two cortical areas, the anterior ectosylvian sulcus (AES) and the rostral aspect of the lateral suprasylvian sulcus (rLS), in producing these response enhancements. This was accomplished by evaluating the multisensory properties of individual SC neurons during reversible deactivation of these cortices individually and in combination using cryogenic deactivation techniques. Cortical deactivation eliminated the characteristic multisensory response enhancement of nearly all SC neurons but generally had little or no effect on a neuron's modality-specific responses. Thus, the responses of SC neurons to combinations of cross-modal stimuli were now no different from those evoked by one or the other of these stimuli individually. Of the two cortical areas, AES had a much greater impact on SC multisensory integrative processes, with nearly half the SC neurons sampled dependent on it alone. In contrast, only a small number of SC neurons depended solely on rLS. However, most SC neurons exhibited dual dependencies, and their multisensory enhancement was mediated by either synergistic or redundant influences from AES and rLS. Corticotectal synergy was evident when deactivating either cortical area compromised the multisensory enhancement of an SC neuron, whereas corticotectal redundancy was evident when deactivation of both cortical areas was required to produce this effect. The results suggest that, although multisensory SC neurons can be created as a consequence of a variety of converging tectopetal afferents that are derived from a host of subcortical and cortical structures, the ability to synthesize cross-modal inputs, and thereby produce an enhanced multisensory response, requires functional inputs from the AES, the rLS, or both.

Acoustic Stimulation↗

Parallel analyses of nociceptive neurones in rat superior colliculus by using c-fos immunohistochemistry and electrophysiology under different conditions of anaesthesia.

Multiple sensory inputs to the superior colliculus (SC) play an important role in guiding head and eye movements toward or away from biologically significant stimuli. Much is now known about the visual, auditory, and somatosensory response properties of SC neurones that mediate these behavioural reactions. Rather less is known about the responses of SC neurones to noxious stimuli, and thus far, most of this information has been obtained in anaesthetised animals. Therefore, the purpose of the present study was to use the c-fos immunohistochemical technique and standard extracellular electrophysiology as parallel measures of nociceptive activity in the SC under different conditions of anaesthesia. In unanaesthetised animals, experimental and control treatments induced a qualitatively similar pattern of Fos-like immunoreactivity (FLI) in the SC, which was quantitatively related to the severity or biologic salience of the treatment; thus, baseline control < control injections of saline < a nonpainful stressor (immobilisation) < noxious injections of formalin. Compared with baseline levels, urethane and avertin anaesthesia induced FLI expression in the SC intermediate layers, although the FLI response to both noxious stimulation and control conditions was differentially suppressed in different layers of the SC by anaesthesia. Parallel electrophysiologic recordings found that anaesthesia was associated with high levels of spontaneous activity in the SC intermediate layers, often in neurones which were also nociceptive. High rates of background spike activity were also induced in the SC intermediate layers by noxious stimulation in chronically recorded awake animals. Although these results point to some differences between the nociceptive responses of SC neurones in anaesthetised and unanaesthetised animals, both data sets support the view that there are different populations of nociceptive neurones in the rodent SC that may be related to different adaptive functions of pain.

Anesthesia↗

Onset of cross-modal synthesis in the neonatal superior colliculus is gated by the development of cortical influences.

Many neurons in the superior colliculus (SC) are able to integrate combinations of visual, auditory, and somatosensory stimuli, thereby markedly affecting the vigor of their responses to external stimuli. However, this capacity for multisensory integration is not inborn. Rather, it appears comparatively late in postnatal development and is not expressed until the SC passes through several distinct developmental stages. As shown here, the final stage in this sequence is one in which a region of association cortex establishes functional control over the SC, thus enabling the multisensory integrative capabilities of its target SC neurons. The first example of this corticotectal input was seen at postnatal day 28. For any individual SC neuron, the onset of corticotectal influences appeared to be abrupt. Because this event occurred at very different times for different SC neurons, a period of 3-4 postnatal months was required before the adult-like condition was achieved. The protracted postnatal period required for the maturation of these corticotectal influences corresponded closely with estimates of the peak period of cortical plasticity, raising the possibility that the genesis of these corticotectal influences, and hence the onset of SC multisensory integration, occurs only after the cortex is capable of exerting experience-dependent control over SC neurons.

Algorithms↗

Neural mechanisms for synthesizing sensory information and producing adaptive behaviors.

The ability to integrate information from different sensory systems is a fundamental characteristic of the brain. Because different bits of information are derived from different sensory channels, their synthesis markedly enhances the detection and identification of external stimuli. The neural substrate for such "multisensory" integration is provided by neurons that receive convergent input from two or more sensory modalities. Many such multisensory neurons are found in the superior colliculus (SC), a midbrain structure that plays a significant role in overt attentive and orientation behaviors. The various principles governing the integration of visual, auditory, and somatosensory inputs in SC neurons have been explored in several species. Thus far, the evidence suggests a remarkable conservation of integrative features during vertebrate evolution. One of the most robust of these principles is based on spatial relationships: a striking enhancement in activity is induced in a multisensory neuron when two different sensory stimuli (e.g., visual and auditory) are in spatial concordance, whereas a profound response depression can be induced when these cues are spatially discordant. The most extensive physiological observations have been made in cat, and in this species the same principles that have been shown to govern multisensory integration at the level of the individual SC neuron have also been shown to govern overt attentive and orientation responses to multisensory stimuli. Most surprising, however, is the critical role played by association (i.e. anterior ectosylvian) cortex in facilitating these midbrain processes. In the absence of the modulating corticotectal influences, multisensory SC neurons in cat are unable to integrate the different sensory cues converging upon them in an adult-like fashion, and are unable to mediate overt multisensory behaviors. This situation appears quite similar to that observed during early postnatal life. When multisensory SC neurons first appear, they are able to respond to multiple sensory inputs but are unable to synthesize these inputs to significantly enhance or degrade their responses. During ontogeny, individual multisensory neurons develop this capacity abruptly, but at very different ages, until the mature population condition is reached after several postnatal months. It appears likely that the abrupt onset of this capacity in any individual SC neuron reflects the maturation of inputs from anterior ectosylvian cortex. Presumably, the functional coupling of cortex with an individual SC neuron is essential to initiate and maintain that neuron's capability for multisensory integration throughout its life.

Adaptation, Physiological↗

Multisensory integration in the superior colliculus of the alert cat.

The modality convergence patterns, sensory response properties, and principles governing multisensory integration in the superior colliculus (SC) of the alert cat were found to have fundamental similarities to those in anesthetized animals. Of particular interest was the observation that, in a manner indistinguishable from the anesthetized animal, combinations of two different sensory stimuli significantly enhanced the responses of SC neurons above those evoked by either unimodal stimulus. These observations are consistent with the speculation that there is a functional link among multisensory integration in individual SC neurons and cross-modality attentive and orientation behaviors.

Acoustic Stimulation↗

Relationship of the meniscofemoral ligaments of the knee to lateral meniscus tears: magnetic resonance imaging evaluation.

This study sought to determine if the presence or absence of meniscofemoral ligaments exerts an influence on the prevalence of tears of the lateral meniscus. We reviewed the sagittal and coronal magnetic resonance images of 173 knees for lateral meniscus tears and the presence of the meniscofemoral ligaments. One or both meniscofemoral ligaments were present in 142 of 173 knees (82%). Fifty-four knees had lateral meniscus tears, 27 of which involved the posterior horn. Thirty-three percent of knees with meniscofemoral ligaments had a lateral meniscus tear, and 23% of knees without meniscofemoral ligaments had a lateral meniscus tear (no significant difference). We found no association between the presence of the meniscofemoral ligaments and tears of the lateral meniscus. Our study questions the importance of preserving or reconstructing these ligaments in instances of meniscal transplantation.

Adolescent↗

Response properties of corticotectal and corticostriatal neurons in the posterior lateral suprasylvian cortex of the cat.

Lateral suprasylvian cortex (LS) is an important source of visual projections to both the striatum and superior colliculus. Although these two LS efferent systems are likely to be involved in different aspects of visual processing, little is known about their functional properties. In the present experiments, 86 neurons in halothane-anesthetized, paralyzed cats were recorded along the posterior aspects of the medial and lateral banks of LS (PMLS and PLLS). Neurons were selected for analysis on the basis of antidromic activation from electrodes chronically implanted in the superior colliculus and caudate nucleus. The segregated nature of corticostriatal and corticotectal neurons was apparent; in no instance could a neuron be antidromically activated from both the superior colliculus and the caudate nucleus. Many common features were revealed between corticotectal and corticostriatal neurons; the majority of neurons in both populations were binocular and contralaterally dominant, showed similar responses to stationary flashed light, and expressed within-field spatial summation and surround inhibition. However, a number of information-processing features distinguished between corticotectal and corticostriatal neurons; the former were generally tuned to lower velocities than were the latter, and, for a given eccentricity in visual space, corticotectal neurons had smaller receptive fields than did corticostriatal neurons. Moreover, most corticotectal neurons displayed a marked preference for movements toward temporal visual space, whereas corticostriatal neurons revealed no specialization for a particular direction of movement. In addition, whereas corticotectal neurons were selective for receding stimuli, corticostriatal neurons were selective for approaching stimuli. The presence of these two corticofugal pathways is discussed in relation to their presumptive functional roles in the facilitation of attentive and orientation behaviors.

Action Potentials↗

Development of multisensory neurons and multisensory integration in cat superior colliculus.

The development of multisensory neurons and multisensory integration was examined in the deep layers of the superior colliculus of kittens ranging in age from 3 to 135 d postnatal (dpn). Despite the high proportion of multisensory neurons in adult animals, no such neurons were found during the first 10 d of postnatal life. Rather, all sensory-responsive neurons were unimodal. The first multisensory neurons (somatosensory-auditory) were found at 12 dpn, and visually responsive multisensory neurons were not found until 20 dpn. Early multisensory neurons responded weakly to sensory stimuli, had long latencies, large receptive fields, and poorly developed response selectivities. Most surprising, however, was their inability to integrate combinations of sensory cues to produce significant response enhancement (or depression), a characteristic feature of the adult. Responses to combinations of sensory cues differed little from responses to their modality-specific components. At 28 dpn an abrupt physiological change was noted. Some multisensory neurons now integrated combinations of cross-modality cues and exhibited significant response enhancements when these cues were spatially coincident and response depressions when the cues were spatially disparate. During the next 2 months the incidence of multisensory neurons, and the proportion of these neurons capable of adult-like multisensory integration, gradually increased. Once multisensory integration appeared in a given neuron, its properties changed little with development. Even the youngest integrating neurons showed superadditive enhancements and spatial characteristics of multisensory integration that were indistinguishable from the adult. Nevertheless, neonatal and adult multisensory neurons differed in the manner in which they integrated temporally asynchronous stimuli, a distribution that may reflect the very different behavioral requirements at different ages. The possible maturational role of corticotectal projections in the abrupt gating of multisensory integration is discussed.

Acoustic Stimulation↗

Visual response properties and visuotopic representation in the newborn monkey superior colliculus.

Visual response properties and visuotopic representation in the newborn monkey superior colliculus. J. Neurophysiol. 78: 2732-2741, 1997. Visually responsive neurons were recorded in the superior colliculus (SC) of the newborn rhesus monkey. The receptive fields of these neurons were larger than those in the adult, but already were organized into a well-ordered map of visual space that was very much like that seen in mature animals. This included a marked expansion of the representation of the central 10 degrees of the visual field and a systematic foveal to peripheral increase in receptive field size. Although newborn SC neurons had longer response latencies than did their adult counterparts, they responded vigorously to visual stimuli and exhibited many visual response properties that are characteristic of the adult. These included surround inhibition, within-field spatial summation, within-field spatial inhibition, binocularity, and an adult-like ocular dominance distribution. As in the adult, SC neurons in the newborn preferred a moving visual stimulus and had adult-like selectivities for stimulus speed. The developmentally advanced state of the functional circuitry of the newborn monkey SC contrasts with the comparative immaturity of neurons in its visual cortex. It also contrasts with observations on the state of maturation of the newborn SC in other developmental models (e.g., cat). The observation that extensive visual experience is not necessary for the development of many adult-like SC response properties in the monkey SC may help explain the substantial visual capabilities shown by primates soon after birth.

Aging↗

Mechanisms of within- and cross-modality suppression in the superior colliculus.

The present studies were initiated to explore the basis for the response suppression that occurs in cat superior colliculus (SC) neurons when two spatially disparate stimuli are presented simultaneously or in close temporal proximity to one another. Of specific interest was examining the possibility that suppressive regions border the receptive fields (RFs) of unimodal and multisensory SC neurons and, when activated, degrade the neuron's responses to excitatory stimuli. Both within- and cross-modality effects were examined. An example of the former is when a response to a visual stimulus within its RF is suppressed by a second visual stimulus outside the RF. An example of the latter is when the response to a visual stimulus within the visual RF is suppressed when a stimulus from a different modality (e. g., auditory) is presented outside its (i.e., auditory) RF. Suppressive regions were found bordering visual, auditory, and somatosensory RFs. Despite significant modality-specific differences in the incidence and effectiveness of these regions, they were generally quite potent regardless of the modality. In the vast majority (85%) of cases, responses to the excitatory stimulus were degraded by >/=50% by simultaneously stimulating the suppressive region. Contrary to expectations and previous speculations, the effects of activating these suppressive regions often were quite specific. Thus powerful within-modality suppression could be demonstrated in many multisensory neurons in which cross-modality suppression could not be generated. However, the converse was not true. If an extra-RF stimulus inhibited center responses to stimuli of a different modality, it also would suppress center responses to stimuli of its own modality. Thus when cross-modality suppression was demonstrated, it was always accompanied by within-modality suppression. These observations suggest that separate mechanisms underlie within- and cross-modality suppression in the SC. Because some modality-specific tectopetal structures contain neurons with suppressive regions bordering their RFs, the within-modality suppression observed in the SC simply may reflect interactions taking place at the level of one input channel. However, the presence of modality-specific suppression at the level of one input channel would have no effect on the excitation initiated via another input channel. Given the modality-specificity of tectopetal inputs, it appears that cross-modality interactions require the convergence of two or more modality-specific inputs onto the same SC neuron and that the expression of these interactions depends on the internal circuitry of the SC. This allows a cross-modality suppressive signal to be nonspecific and to degrade any and all of the neuron's excitatory inputs.

Acoustic Stimulation↗

A simple enzyme histochemical method for the simultaneous demonstration of acetylcholinesterase and monoamine oxidase in fixed-frozen sections.

We describe an enzyme histochemical technique for the simultaneous demonstration of acetylcholinesterase (AChE) and monoamine oxidase (MAO) (Types A, B, or A+B) in fixed-frozen sections. Several regions in the mesencephalon and brainstem were examined for both somatic and neuropil labeling. The results obtained are equivalent or superior to those obtained using previous methods for the individual localization of these enzymes. The simultaneous visualization of AChE and MAO in the same section allows the relationship of the two enzymes to be easily assessed with brightfield microscopy.

Acetylcholinesterase↗

The role of anterior ectosylvian cortex in cross-modality orientation and approach behavior.

Physiological and behavioral studies in cat have shown that corticotectal influences play important roles in the information-processing capabilities of superior colliculus (SC) neurons. While corticotectal inputs from the anterior ectosylvian sulcus (AES) play a comparatively small role in the unimodal responses of SC neurons, they are particularly important in rendering these neurons capable of integrating information from different sensory modalities (e.g., visual and auditory). The present experiments examined the behavioral consequences of depriving SC neurons of AES inputs, and thereby compromising their ability to integrate visual and auditory information. Selective deactivation of a variety of other cortical areas (posterolateral lateral suprasylvian cortex, PLLS; primary auditory cortex, AI; or primary visual cortex, 17/18) served as controls. Cats were trained in a perimetry device to ignore a brief, low-intensity auditory stimulus but to orient toward and approach a nearthreshold visual stimulus (a light-emitting diode, LED) to obtain food. The LED was presented at different eccentricities either alone (unimodal) or combined with the auditory stimulus (multisensory). Subsequent deactivation of the AES, with focal injections of a local anesthetic, had no effect on responses to unimodal cues regardless of their location. However, it profoundly, though reversibly, altered orientation and approach to multisensory stimuli in contralateral space. The characteristic enhancement of these responses observed when an auditory cue was presented in spatial correspondence with the visual stimulus was significantly degraded. Similarly, the inhibitory effect of a spatially disparate auditory cue was significantly ameliorated. The observed effects were specific to AES deactivation, as similar effects were not obtained with deactivation of PLLS, AI or 17/18, or saline injections into the AES. These observations are consistent with postulates that specific cortical-midbrain interactions are essential for the synthesis of multisensory information in the SC, and for the orientation and localization behaviors that depend on this synthesis.

Anesthetics, Local↗

Nociceptive neurones in rat superior colliculus. I. Antidromic activation from the contralateral predorsal bundle.

Accumulating evidence suggests that the rodent superior colliculus (SC) plays as important a role in avoidance and defensive behaviours as it does in orientation and approach. These two complementary behaviours are associated with two anatomically segregated tectofugal output pathways, such that orientation and approach are mediated by the crossed descending projection, whereas avoidance and defence are subserved via the uncrossed projection. Because nociceptive neurones in the SC have been presumed to participate in withdrawal or defensive behaviours, it has been proposed that they have direct access only to the uncrossed efferent pathway. However, in certain behavioural situations, the most adaptive response to injury, or to a painful object in prolonged contact with the skin, is to orient towards the source of discomfort so that the skin can be licked and/or the offending object removed. Presumably then, nociceptive as well as low-threshold neurones would have access to the crossed descending pathway in order to initiate such behaviours. Determining whether or not this is the case was the objective of the present study. Both nociceptive-specific (82%) and wide-dynamic-range (18%) SC neurones were identified using long-duration (up to 6 s), frankly noxious mechanical and thermal stimuli in urethane-anaesthetised Long-Evans hooded rats. The majority (85.7%) of the nociceptive neurones encountered were located within the intermediate layers, which corresponds with the location of the cells-of-origin of the crossed descending projection. Nearly half (44.9%) were activated antidromically from electrical stimulation of the crossed descending pathway at a site in the brainstem below its decussation. The mean conduction velocity of these nociceptive output neurones was 9.02 m/s, which corresponds well to previous estimates of conduction velocity in the crossed tecto-reticulo-spinal tract. These data demonstrate that a significant proportion of nociceptive neurones in the rat SC have axons that project to the contralateral brainstem via the crossed descending projection. Nociceptive neurones could, therefore, effect orientation responses to noxious stimuli via similar output pathways that low-threshold neurones utilize to initiate orientation to innocuous stimuli.

Animals↗

Nociceptive neurones in rat superior colliculus. II. Effects of lesions to the contralateral descending output pathway on nocifensive behaviours.

A wealth of evidence implicates the crossed descending projection from the superior colliculus (SC) in orientation and approach behaviours directed towards novel, non-noxious stimuli. In our preceding paper, we identified a population of nociceptive neurones in the rat SC that have axons that project to the contralateral brainstem via this output pathway. The purpose of the present study was, therefore, to evaluate the prediction that the crossed descending projection of the SC is also involved in the control of orientation and approach movements of the head and mouth made during the localisation of persistent noxious stimuli. An independent-groups design was used to test the effects of interrupting the contralateral descending projection from the SC on the behavioural reactions elicited by noxious mechanical stimuli presented to the tail and hindpaws. In different groups of animals, a microwire knife was used to cut the contralateral descending fibres at two different locations: (1) a sagittal cut at the level of the dorsal tegmental decussation; (2) a bilateral coronal cut of the predorsal bundle at the level of the medial pontine reticular formation. Retrograde anatomical tracing techniques were then used to evaluate the effectiveness of the cuts and to assess possible involvement of non-collicular fibre systems in both lesioned and control animals. Additional behavioural procedures were performed to test for general neurological status and responsiveness of animals to non-noxious stimuli. Anatomical tracing data indicated that the largest population of neurones with fibres severed by both cuts were the cells-of-origin of the contralateral descending projection in the intermediate white layer of the SC. Behavioural results showed that significantly more animals in both lesion groups failed to locate and bite a mechanical clip placed on the tail. Instead of switching to motor behaviours to localise and remove noxious stimuli, they persisted with defensive reactions, which included freezing, vocalisation or forward and backward escape. In contrast, when the clip was placed on the hindpaws, it was successfully localised by most lesioned and control animals; however, lesioned animals had reliably longer latencies and spent less time in close contact with the clip. Consistent with the established role of the contralateral descending projection in non-noxious orientation, lesioned animals also showed orienting deficits to a range of non-noxious sensory stimuli. These data suggest that, under certain behavioural circumstances, nociceptive information from the SC is integral to the elaboration of orienting and approach movements of the head and mouth elicited by persistent noxious stimuli.

Animals↗

Comparisons of cross-modality integration in midbrain and cortex.

Multisensory neurons are abundant in the superior colliculus and anterior ectosylvian cortex of the cat. Despite the fact that these areas receive inputs from different regions, and are likely to be involved in different functional roles, there multisensory neurons have many fundamental similarities. They all have multiple receptive fields, one for each sensory input, and these receptive fields overlap one another. It is this spatial correspondence among receptive fields that determines the manner in which both populations of neurons integrate the inputs they receive from different sensory channels. Several principles of integration characterize both cortical and midbrain multisensory neurons, and these constancies in the fundamentals of cross-modality integration are likely to provide a basis for coherence at different levels of the neuraxis. Yet there are also obvious differences in these populations of multisensory neurons. Cortical receptive fields are significantly larger than those in the midbrain, have a lower incidence of suppressive surrounds, and exhibit less cross-modality inhibitory interactions than in the midbrain. Presumably, these differences reflect a greater emphasis on non-spatial aspects of cross-modality integration in cortex than is required by the orientation and localization functions mediated by the superior colliculus.

Animals↗