Sound localization deficits during reversible deactivation of primary auditory cortex and/or the dorsal zone

We examined the contributions of primary auditory cortex (A1) and the dorsal zone of auditory cortex (DZ) to sound localization behavior during separate and combined unilateral and bilateral deactivation. From a central visual fixation point, cats learned to make an orienting response (head movement and approach) to a 100-ms broadband noise burst emitted from a central speaker or one of 12 peripheral sites (located in front of the animal, from left 90 degrees to right 90 degrees, at 15 degrees intervals) along the horizontal plane. Following training, each cat was implanted with separate cryoloops over A1 and DZ bilaterally. Unilateral deactivation of A1 or DZ or simultaneous unilateral deactivation of A1 and DZ (A1/DZ) resulted in spatial localization deficits confined to the contralateral hemifield, whereas sound localization to positions in the ipsilateral hemifield remained unaffected. Simultaneous bilateral deactivation of both A1 and DZ resulted in sound localization performance dropping from near-perfect to chance (7.7% correct) across the entire field. Errors made during bilateral deactivation of A1/DZ tended to be confined to the same hemifield as the target. However, unlike the profound sound localization deficit that occurs when A1 and DZ are deactivated together, deactivation of either A1 or DZ alone produced partial and field-specific deficits. For A1, bilateral deactivation resulted in higher error rates (performance dropping to approximately 45%) but relatively small errors (mostly within 30 degrees of the target). In contrast, bilateral deactivation of DZ produced somewhat fewer errors (performance dropping to only approximately 60% correct), but the errors tended to be larger, often into the incorrect hemifield. Therefore individual deactivation of either A1 or DZ produced specific and unique sound localization deficits. The results of the present study reveal that DZ plays a role in sound localization. Along with previous anatomical and physiological data, these behavioral data support the view that A1 and DZ are distinct cortical areas. Finally, the findings that deactivation of either A1 or DZ alone produces partial sound localization deficits, whereas deactivation of either posterior auditory field (PAF) or anterior ectosylvian sulcus (AES) produces profound sound localization deficits, suggests that PAF and AES make more significant contributions to sound localization than either A1 or DZ.     

Cortical control of sound localization in the cat: unilateral cooling deactivation of 19 cerebral areas

We examined the ability of mature cats to accurately orient to, and approach, an acoustic stimulus during unilateral reversible cooling deactivation of primary auditory cortex (AI) or 1 of 18 other cerebral loci. After attending to a central visual stimulus, the cats learned to orient to a 100-ms broad-band, white-noise stimulus emitted from a central speaker or 1 of 12 peripheral sites (at 15 degrees intervals) positioned along the horizontal plane. Twenty-eight cats had two to six cryoloops implanted over multiple cerebral loci. Within auditory cortex, unilateral deactivation of AI, the posterior auditory field (PAF) or the anterior ectosylvian sulcus (AES) resulted in orienting deficits throughout the contralateral field. However, unilateral deactivation of the anterior auditory field, the second auditory cortex, or the ventroposterior auditory field resulted in no deficits on the orienting task. In multisensory cortex, unilateral deactivation of neither ventral or dorsal posterior ectosylvian cortices nor anterior or posterior area 7 resulted in any deficits. No deficits were identified during unilateral cooling of the five visual regions flanking auditory or multisensory cortices: posterior or anterior ii suprasylvian sulcus, posterior suprasylvian sulcus or dorsal or ventral posterior suprasylvian gyrus. In motor cortex, we identified contralateral orienting deficits during unilateral cooling of lateral area 5 (5L) or medial area 6 (6m) but not medial area 5 or lateral area 6. In a control visual-orienting task, areas 5L and 6m also yielded deficits to visual stimuli presented in the contralateral field. Thus the sound-localization deficits identified during unilateral deactivation of area 5L or 6m were not unimodal and are most likely the result of motor rather than perceptual impairments. Overall, three regions in auditory cortex (AI, PAF, AES) are critical for accurate sound localization as assessed by orienting.     

Neuronal activation times to simple, complex, and natural sounds in cat primary and nonprimary auditory cortex

Interactions between living organisms and the environment are commonly regulated by accurate and timely processing of sensory signals. Hence, behavioral response engagement by an organism is typically constrained by the arrival time of sensory information to the brain. While psychophysical response latencies to acoustic information have been investigated, little is known about how variations in neuronal response time relate to sensory signal characteristics. Consequently, the primary objective of the present investigation was to determine the pattern of neuronal activation induced by simple (pure tones), complex (noise bursts and frequency modulated sweeps), and natural (conspecific vocalizations) acoustic signals of different durations in cat auditory cortex. Our analysis revealed three major cortical response characteristics. First, latency measures systematically increase in an antero-dorsal to postero-ventral direction among regions of auditory cortex. Second, complex acoustic stimuli reliably provoke faster neuronal response engagement than simple stimuli. Third, variations in neuronal response time induced by changes in stimulus duration are dependent on acoustic spectral features. Collectively, these results demonstrate that acoustic signals, regardless of complexity, induce a directional pattern of activation in auditory cortex.     

Interhemispheric asymmetry of homotopic transcallosal responses of the auditory cortex in the cat

In acute experiments on 29 d-turbocurarine immobilized cats, with topographical pick-up of potentials from 150–170 symmetrical points of the auditory cortex, the interhemispheric asymmetry of homotopic transcallosal responses were studied. It was established that in the majority of the animals the potentials in the right hemisphere had a greater amplitude and shorter latent periods as compared with the responses of the left hemisphere. A dependence of the properties of the interhemispheric asymmetry on the sex of the animal was noted. In females the asymmetry was characterized by a right hemispheric tendency and by a greater magnitude as compared with males. The maximal magnitude of the asymmetry was detected in the group of pregnant females. The hypothesis is proposed to the effect that the established polarity of transcallosal influenced in the auditory cortex is associated with the dominance of the right hemisphere in the organization of spatial audition.Translated from Fiziologicheskii Zhurnal SSSR, imeni I. M. Sechenova, Vol. 73, No. 7, pp. 851–859, July, 1987.     

Competition and convergence between auditory and cross-modal visual inputs to primary auditory cortical areas

Sensory neocortex is capable of considerable plasticity after sensory deprivation or damage to input pathways, especially early in development. Although plasticity can often be restorative, sometimes novel, ectopic inputs invade the affected cortical area. Invading inputs from other sensory modalities may compromise the original function or even take over, imposing a new function and preventing recovery. Using ferrets whose retinal axons were rerouted into auditory thalamus at birth, we were able to examine the effect of varying the degree of ectopic, cross-modal input on reorganization of developing auditory cortex. In particular, we assayed whether the invading visual inputs and the existing auditory inputs competed for or shared postsynaptic targets and whether the convergence of input modalities would induce multisensory processing. We demonstrate that although the cross-modal inputs create new visual neurons in auditory cortex, some auditory processing remains. The degree of damage to auditory input to the medial geniculate nucleus was directly related to the proportion of visual neurons in auditory cortex, suggesting that the visual and residual auditory inputs compete for cortical territory. Visual neurons were not segregated from auditory neurons but shared target space even on individual target cells, substantially increasing the proportion of multisensory neurons. Thus spatial convergence of visual and auditory input modalities may be sufficient to expand multisensory representations. Together these findings argue that early, patterned visual activity does not drive segregation of visual and auditory afferents and suggest that auditory function might be compromised by converging visual inputs. These results indicate possible ways in which multisensory cortical areas may form during development and evolution. They also suggest that rehabilitative strategies designed to promote recovery of function after sensory deprivation or damage need to take into account that sensory cortex may become substantially more multisensory after alteration of its input during development.     

Auditory corticocortical interconnections in the cat: evidence for parallel and hierarchical arrangement of the auditory cortical areas

Summary The origin and laminar arrangement of the homolateral and callosal projections to the anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) auditory cortical areas were studied in the cat by means of electrophysiological recording and WGA-HRP tracing techniques. The transcallosal projections to AAF, AI, PAF and AII were principally homotypic since the major source of input was their corresponding area in the contralateral cortex. Heterotypic transcallosal projections to AAF and AI were seen, originating from the contralateral AI and AAF, respectively. PAF received heterotypic commissural projections from the opposite ventroposterior auditory cortical field (VPAF). Heterotypic callosal inputs to AII were rare, originating from AAF and AI. The neurons of origin of the transcallosal connections were located mainly in layers II and III (70–92%), and less frequently in deep layers (V and VI, 8–30%). Single unit recordings provided evidence that both homotypic and heterotypic transcallosal projections connect corresponding frequency regions of the two hemispheres. The regional distribution of the anterogradely labeled terminals indicated that the homotypic and heterotypic auditory transcallosal projections are reciprocal. The present data suggest that the transcallosal auditory interconnections are segregated in 3 major parallel components (AAF-AI, PAF-VPAF and AII), maintaining a segregation between parallel functional channels already established for the thalamocortical auditory interconnections. For the intrahemispheric connections, the analysis of the retrograde tracing data revealed that AAF and AI receive projections from the homolateral cortical areas PAF, VPAF and AII, whose neurons of origin were located mainly in their deep (V and VI) cortical layers. The reciprocal interconnections between the homolateral AAF and AI did not show a preferential laminar arrangement since the neurons of origin were distributed almost evenly in both superficial (II and III) and deep (V and VI) cortical layers. On the contrary, PAF received inputs from the homolateral cortical fields AAF, AI, AII and VPAF, originating predominantly from their superficial (II and III) layers. The homolateral projections reaching AII originated mainly from the superficial layers of AAF and AI, but from the deep layers of VPAF and PAF. The laminar distribution of anterogradely labeled terminal fields, when they were dense enough for a confident identification, was systematically related to the laminar arrangement of neurons of origin of the reciprocal projection: a projection originating from deep layers was associated with a reciprocal projection terminating mainly in layer IV, whereas a projection originating from superficial layers was associated with a reciprocal projection terminating predominantly outside layer IV. This laminar distribution indicates that the homolateral auditory cortical interconnections have a feed-forward/feed-back organization, corresponding to a hierarchical arrangement of the auditory cortical areas, according to criteria previously established in the visual system of primates. The principal auditory cortical areas could be ranked into 4 distinct hierarchical levels. The tonotopically organized areas AAF and AI represent the lowest level. The second level corresponds to the non-tonotopically organized area AII. Higher, the tonotopically organized areas VPAF and PAF occupy the third and fourth hierarchical levels, respectively.Abbreviations AAF anterior auditory cortical area - AI primary auditory cortical area - AII secondary auditory cortical area - BF best frequency - C cerebral cortex - CA caudate nucleus - CL claustrum - D dorsal nucleus of the dorsal division of the MGB - ea anterior ectosylvian sulcus - ep posterior ectosylviansulcus - IC internal capsule - LGN lateral geniculate nucleus - LV pars lateralis of the ventral division of the MGB - LVe lateral ventricule - M pars magnocellularis of the medial division of the MGB - MGB medial geniculate body - MGBv ventral division of the MGB - OT optic tract - OV pars ovoidea of the ventral division of the MGB - PAF posterior auditory cortical area - PH parahippocampal cortex - PO lateral part of the posterior group of thalamic nuclei - PU putamen - RE reticular complex of thalamus - rs rhinal sulcus - SG suprageniculate nucleus of the dorsal division of the MGB - ss suprasylvian sulcus - TMB tetrametylbenzidine - VBX ventrobasal complex - VLa ventrolateral complex - VL ventro-lateral nucleus of the ventral division of the MGB - WGA-HRP wheat germ agglutinin conjugated to horse-radish peroxidase - WM white matter - VPAF ventro-posterior auditory cortical area     

Modulation of level response areas and stimulus selectivity of neurons in cat primary auditory cortex

Sounds commonly occur in sequences, such as in speech. It is therefore important to understand how the occurrence of one sound affects the response to a subsequent sound. We approached this question by determining how a conditioning stimulus alters the response areas of single neurons in the primary auditory cortex (AI) of barbiturate-anesthetized cats. The response areas consisted of responses to stimuli that varied in level at the two ears and delivered at the characteristic frequency of each cell. A binaural conditioning stimulus was then presented > or =50 ms before each of the stimuli comprising the level response area. An effective preceding stimulus alters the shape and severely reduces the size and response magnitude of the level response area. This ability of the preceding stimulus depends on its proximity in the level domain to the level response area, not on its absolute level or on the size of the response it evokes. Preceding stimuli evoke a nonlinear inhibition across the level response area that results in an increased selectivity of a cortical neuron for its preferred binaural stimuli. The selectivity of AI neurons during the processing of a stream of acoustic stimuli is likely to be restricted to a portion of their level response areas apparent in the tone-alone condition. Thus rather than being static, level response areas are fluid; they can vary greatly in extent, shape and response magnitude. The dynamic modulation of the level response area and level selectivity of AI neurons might be related to several tasks confronting the central auditory system.     

Anatomical localization of cortical beta rhythms in cat

Beta electrocorticographic rhythms (40 Hz) develop during motionless focused attention in two distinct cortical foci in cats. A cytoarchitectonic study was performed to determine the precise location of these foci. Electrode tips recording beta rhythms were found: (i) in motor areas 4 gamma and 6a beta, in a band extending from the postcruciate cortex to the walls of the presylvian sulcus, crossing the frontal pole (anterior beta focus); (ii) in the posterior parietal associative area 5a, along the divisions of the ansate sulcus, extending to the mesial aspect of the hemispheres (posterior beta focus). The two foci are separated by areas 3, 2 and 1, where beta rhythms were never recorded. The fact that both these areas, containing giant pyramidal cells, develop a specific type of activity during immobility may have a functional meaning: area 5 may be involved in the cat as it is in the monkey in the control of motor behaviour.     

Spectrotemporal receptive fields during spindling and non-spindling epochs in cat primary auditory cortex

It was often thought that synchronized rhythmic epochs of spindle waves disconnect thalamo-cortical system from incoming sensory signals. The present study addresses this issue by simultaneous extracellular action potential and local field potential (LFP) recordings from primary auditory cortex of ketamine-anesthetized cats during spindling activity. We compared cortical spectrotemporal receptive fields (STRF) obtained during spindling and non-spindling epochs. The basic spectro-temporal parameters of "spindling" and "non-spindling" STRFs were similar. However, the peak-firing rate at the best frequency was significantly enhanced during spindling epochs. This enhancement was mainly caused by the increased probability of a stimulus to evoke spikes (effectiveness of stimuli) during spindling as compared with non-spindling epochs. Augmented LFPs associated with effective stimuli and increased single-unit pair correlations during spindling epochs suggested higher synchrony of thalamo-cortical inputs during spindling that resulted in increased effectiveness of stimuli presented during spindling activity. The neuronal firing rate, both stimulus-driven and spontaneous, was higher during spindling as compared with non-spindling epochs. Overall, our results suggests that thalamic cells during spindling respond to incoming stimuli-related inputs and, moreover, cause more powerful stimulus-related or spontaneous activation of the cortex.     

Influence of auditory localization cues on neuronal activity in the auditory thalamus of the cat

1. The response properties of auditory thalamic neurons to the two major localization cues characterizing the azimuth of sound sources in the horizontal plane were investigated in cats. Single-unit responses to auditory stimuli (white noise and tones) presented with interaural phase differences (IPD) or interaural intensity differences (IID) were studied. 2. The proportion of neurons in the medial geniculate body that were sensitive to the localization cues tested was 28% for IPD (n = 253) and 37% for IID (n = 65). Half of the IID-sensitive units were also sensitive to IPD, but when the range of IPDs and IIDs to which each unit responded was converted to the sound-source locations that would generate those ranges they did not always correspond to overlapping azimuth angles. 3. The changes in discharge rate in response to the two localization cues occurred over very broad IPD and IID ranges. If this activity is involved in the representation of acoustic space, then the responses of individual neurons do not provide fine spatial tuning. 4. Contralateral and ipsilateral ear leads were represented in a continuous manner by the maximum discharge rate of IPD-sensitive units. On the other hand, units that were sensitive to IIDs were activated over one of two delimited ranges of IIDs. The first corresponded to IID combinations in which the stimulus was presented at a higher intensity in one ear than in the other (for 15/17 units the contralateral one); these were the lateralized intensity response field units. The second are the centered intensity response field units, whose responses were maximal when the intensity was equal in both ears and decreased when IIDs were introduced.     

Chemoarchitectonic organization of the cat primary auditory cortex

Summary Acetylcholinesterase (AChE) activity, demonstrated histochemically, defines an area of cortex on the middle ectosylvian gyrus that appears to correspond to the cytoarchitectonically defined area 41 and the physiologically defined primary auditory area (AI). In this area there are high levels of AChE in layers III, IV and VI while in the surrounding areas there are comparatively low levels of enzyme in these layers. The monoclonal antibody CAT 301, which was raised against a cell surface proteoglycan, also defines this area. There are high levels of CAT 301 immunoreactivity in cell bodies and the neuropil of layer III and an absence of very large immunoreactive neurons in layer V. Furthermore there are higher levels of the calcium binding protein, parvalbumin and the metabolic enzyme, cytochrome oxidase, in layers III and IV of AI, than in most of the surrounding cortex. By contrast the distribution of the calcium binding protein, calbindin and the distribution of myelinated fibers are similar in area 41 and the surrounding areas.     

Binaural interaction revisited in the cat primary auditory cortex

The binaural interactions of neurons were studied in the primary auditory cortex (AI) of barbiturate-anesthetized cats with a matrix of binaural tonal stimuli varying in both interaural level differences (ILD) and average binaural level (ABL). The purpose of this study was to determine: 1) the distribution of preferred binaural combinations (PBCs) of a large population of neurons and its relationships with binaural interactions and binaural monotonicity; 2) whether monaural responses are predictive of binaural responses; and 3) whether there is a restricted set of representative binaural stimulus configurations that could effectively classify the binaural interactions. Binaural interactions were often diverse in the matrix and dependent on both ABL and ILD. Compared with previous studies, a higher proportion of mixed binaural interaction type and a lower proportion of EO/I type were found. No monaural neurons were found. Binaural responses often differed from monaural responses in the number of spikes and/or the form of the response functions. The PBCs of the majority of EO and PB neurons were in the contralateral field and midline, respectively. However, the PBCs of EE units were evenly distributed across the contralateral and ipsilateral fields. The majority of the nonmonotonic neurons responded most strongly to lower ABLs, whereas the majority of monotonic neurons responded most strongly to higher ABLs. This study demonstrated that in AI a restricted set of binaural stimulus configurations is not sufficient to reveal the binaural responses properties. Also, monaural responses are not predictive of binaural responses.     

Connections between the amygdala and auditory cortical areas in the macaque monkey

Connections between the amygdala and auditory cortical areas TC, and the rostral, intermediate and caudal regions of area TA (TAr, TAi and TAc, respectively) in the macaque monkey (Macaca fuscata and Macaca nemestrina) were investigated following placements of cortical deposits of wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP). Areas TC and TAc received weak projections and these derived only from the lateral basal nucleus. Areas TAi and TAr received projections from the lateral, lateral basal and accessory basal nuclei. In contrast, corticopetal projections to the amygdala originated in areas TAi and TAr, but never in TAc or TC. The projections from areas TAi and TAr terminated only in the lateral nucleus, and in particular at the lateral part of the middle and caudal portions of the amygdala. Thus, the amygdalofugal projections to the auditory cortices are more widespread and more complex than the amygdalopetal projections of the auditory cortices. As judged from experiments in which deposits were made at different sites along the rostrocaudal axis of the auditory cortex, there was a progressive increase seen in density of the amygdala connections with more anteriorly-placed injection sites.     

Several neuronal and axonal types form long intrinsic connections in the cat primary auditory cortical field (AI)

Intrinsic connections in the cat primary auditory field (AI) as revealed by injections Phaseolus vulgaris leucoagglutinin (PHA-L) or biocytin, had an anisotropic and patchy distribution. Neurons, labelled retrogradely with PHA-L were concentrated along a dorsoventral stripe through the injection site and rostral to it; the spread of rostrally located neurons was greater after injections into regions of low rather than high characteristic frequencies. The intensity of retrograde labelling varied from weak and granular to very strong and Golgi-like. Out of 313 Golgi like retrogradely labelled neurons 79.6% were pyramidal, 17.2% multipolar, 2.6% bipolar, and 0.6% bitufted; 13.4% were putatively inhibitory, i.e. aspiny or sparsely spiny multipolar, or bitufted. Individual anterogradely labelled intrinsic axons were reconstructed for distances of 2 to 7 mm. Five main types were distinguished on the basis of the branching pattern and the location of synaptic specialisations. Type 1 axons travelled horizontally within layers II to VI and sent collaterals at regular intervals; boutons were only present in the terminal arborizations of these collaterals. Type 2 axons also travelled horizontally within layers II to VI and had rather short and thin collateral branches; boutons or spine-like protrusions occurred in most parts of the axon. Type 3 axons travelled obliquely through the cortex and formed a single terminal arborization, the only site where boutons were found. Type 4 axons travelled for some distance in layer I; they formed a heterogeneous group as to their collaterals and synaptic specializations. Type 5 axons travelled at the interface between layer VI and the white matter; boutons en passant, spine-like protrusions, and thin short branches with boutons en passant were frequent all along their trajectory. Thus, only some axonal types sustain the patchy pattern of intrinsic connectivity, whereas others are involved in a more diffuse connectivity.Abbreviations AAF anterior auditory cortical field - AI primary auditory cortical field - AII secondary auditory cortical field - ALLS anterolateral lateral suprasylvian visual area - CF characteristic frequency - IFL isofrequency line - INJ injection - LV pars lateralis of the ventral division of the MGB - MGB medial geniculate body - mss middle suprasylvian sulcus - OV pars ovoidea of the ventral division of the MGB - PAF posterior auditory field - PHA-L Phaseolus vulgaris leucoagglutinin - PO posterior nucleus of the thalamus - R rostral - SF suprasylvian fringe - syl sylvian sulcus - TBS TRIS-buffered saline - V ventral - VL ventro-lateral nucleus of the MGB     

Rarity of luxotonic responses in cortical visual areas of the cat

Summary Neuronal responses to continuous, diffuse white light or darkness were studied in cortical visual areas 17, 18, 19 and Clare-Bishop of the unanesthetized cat. In contrast to squirrel monkeys and macaques in which about 40 or 25% of the units in striate cortex are luxotonic (response to continuous light or darkness sustained>2.0 min), all of the visual areas in the cat had fewer than 4.0% of the units exhibiting such luxotonic activity. The functional basis of this difference may be related to differences between the two species in the quantitative balance of antagonistic receptive field properties.This report is dedicated to the guidance and friendship of John R. Bartlett, deceased November 5, 1978 R. Bartlett, deceased November 5, 1978     

Temporal nonlinearity during recovery from sequential inhibition by neurons in the cat primary auditory cortex

Auditory stimuli occur most often in sequences rather than in isolation. It is therefore necessary to understand how responses to sounds occurring in sequences differ from responses to isolated sounds. Cells in primary auditory cortex (AI) respond to a large set of binaural stimuli when presented in isolation. The set of responses to such stimuli presented at one frequency comprises a level response area. A preceding binaural stimulus can reduce the size and magnitude of level response areas of AI cells. The present study focuses on the effects of the time interval between a preceding stimulus and the stimuli of a level response area in pentobarbital-anesthetized cats. After the offset of a preceding stimulus, the ability of AI cells to respond to succeeding stimuli varies dynamically in time. At short interstimulus intervals (ISI), a preceding stimulus can completely inhibit responses to succeeding stimuli. With increasing ISIs, AI cells respond first to binaural stimuli that evoke the largest responses in the control condition, i.e., not preceded by a stimulus. Recovery rate is nonlinear across the level response area; responses to these most-effective stimuli recover to 70% of control on average 187 ms before responses to other stimuli recover to 70% of their control sizes. During the tens to hundreds of milliseconds that a level response area is reduced in size and magnitude, the selectivity of AI cells is increased for stimuli that evoke the largest responses. This increased selectivity results from a temporal nonlinearity in the recovery of the level response area which protects responses to the most effective binaural stimuli. Thus in a sequence of effective stimuli, a given cell will respond selectively to only those stimuli that evoke a strong response when presented alone.     

Effects of auditory cortical lesions on sound localization by the rat

1. Studies with cats, dogs, and monkeys have shown that bilateral ablation of auditory cortex can result in severe deficits in the ability to localize sounds in space. In the present series of studies we sought to extend this observation to include the laboratory rat. 2. Rats were tested in a two-choice sound-localization task, which required a spatial response to a distant goal box. Although the test conditions were very similar to those employed with cat, dog, and monkey, deficits following cortical ablation were minimal. Indeed, following bilateral ablation of both primary and secondary auditory projection areas, rats were still capable of localization at small angles. 3. Several possibilities were considered to explain the apparent species difference in the effect of cortical ablations. It seemed unlikely that the difference was due simply to testing procedures or details of stimulus presentation. Central nervous system factors were discussed, including the possibility that the degree of impairment was related to the extent of cortical development in different species. It was concluded that the effects of auditory cortical ablation are not the same for all mammals and that for some species auditory cortex is not essential for sound localization.     

Representation of cochlea within primary auditory cortex in the cat.

The representation of sound frequency (and of the cochlear partition) within primary auditory cortex has been investigated with use of microelectrode-mapping techniques in a series of 25 anesthetized cats. Among the results were the following: 1) Within vertical penetrations into AI, best frequency and remarkably constant for successively studied neurons across the active middle and deep cortical layers. 2) There is an orderly representation of frequency (and of represented cochlear place) within AI. Frequency is rerepresented across the mediolateral dimension of the field. On an axis perpendicular to this plane of rerepresentation, best-frequency (represented cochlear place) changes as a simple function of cortical location. 3) Any given frequency band (or sector of the cochlear partition) is represented across a belt of cortex of nearly constant width that runs on a nearly straight axis across AI. 4) There is a disproportionately large cortical surface representation of the highest-frequency octaves (basal cochlea) within AI. 5) The primary and secondary field locations were somewhat variable, when referenced to cortical surface landmarks. 6) Data from long penetrations passing down the rostral bank of the posterior ectosylvian sulcus were consistent with the existence of a vertical unit of organization in AI, akin to cortical columns described in primary visual and somatosensory cortex. 7) Responses to tonal stimuli were encountered in fields dorsocaudal, caudal, ventral, and rostral to AI. There is an orderly representation of the cochlea within the field rostal to AI, with a reversal in best frequencies across its border with AI. 8) Physiological definitions of AI boundaries are consistent with their cytoarchitectonic definition. Some of the implications of these findings are discussed.