Foveal tracking cells in the superior temporal sulcus of the macaque monkey

Summary Visual responses were recorded from neurons in the superior temporal sulcus (STS) of awake, behaving cynomolgus monkeys trained to fixate a small spot of light. Visual receptive fields, directionality, and responses during visual tracking were examined quantitatively for 50 cells in the foveal portion of the middle temporal (MT) visual area and surrounding cortex. Directionality indices and preferred directions for tracked and nontracked stimuli were compared. Eighteen cells (18/50 = 36%) were found to respond preferentially during tracking (tracking cells), 7 within MT, 9 in area FST on the floor of the STS, and 2 in unidentified areas. Three distinctly different tracking response profiles (VTS, VTO, and T) were observed. VTS and VTO cells had foveal receptive fields and gave directionally selective visual responses. VTS cells (3 in foveal MT, 6 in FST, 1 in an unidentified area) had a preferred visual direction that coincided with the preferred tracking direction, and began responding 50–100 ms before the onset of tracking. VTO cells (4 in foveal MT, 0 in FST, 1 in an unidentified area) had a preferred visual direction opposite to the preferred tracking direction, and began responding 0–100 ms after the onset of tracking. T cells (0 in MT, 3 in FST) had no visual responses and began responding simultaneously with the onset of tracking. It is suggested that this region of the brain could be the primary location for converting direction-specific visual responses into signals specifying at least the direction of an intended pursuit movement.     

Representation of the fovea in the superior temporal sulcus of the macaque monkey

Summary The response properties of 633 neurons from striate and prestriate cortex were recorded in 3 hemispheres of two awake cynomolgus monkeys while they fixated or tracked a small spot of light. Of 254 penetrations located at 1 mm intervals, 39% were identifiable from visible electrolytic lesions or electrode tracks and were used to reconstruct the positions of all recording sites. A total of 226 cells were located in the superior temporal sulcus and 81 cells in area V1. The location and visuotopic organization of the foveal portion of the middle temporal (MT) visual area were determined in three hemispheres. MT was defined physiologically on the basis of direction-selectivity, receptive field size, and retinotopic organization. Of 170 MT neurons, most were motion sensitive, and 65% had a directionality index, (best — opposite)/best, of 0.6 or higher. MT was defined anatomically on the basis of myelin staining within the superior temporal sulcus (STS). On the posterior bank of the STS the physiologically defined border corresponded closely to a myelin border visible on our sections. Distinct myelin borders were not consistently identifiable on the anterior bank. The representation of the central fovea (eccentricities of 0–1 deg) was located partly on the floor, but mostly on the posterior bank of the STS at the extreme postero-lateral edge of MT. In all three hemispheres foveal MT extended onto the roof of a cleft formed between the posterior bank and a wide flattened area on the floor of the STS. This region lies 10–12 mm below the brain surface, measuring along a line normal to the surface at a point 2–3 mm antero-lateral to foveal V1. The area of MT was 6–9 mm² for the central fovea (0–1 deg), 15–24 mm² for the entire fovea (0–3 deg), and 28–40 mm² including the fovea and parafovea (0–10 deg). A visuotopic map of central foveal V1 (0–1 deg) was obtained in one animal. The measured area of this representation was 116 mm². Using published estimates of the total areas of cynomolgus MT and V1 (73 and 1200 mm² respectively) the ratio of central foveal to total area was calculated to be 0.10 for both MT (7.5/73) and V1 (116/1200), indicating that the relative magnification of the foveal versus the peripheral visual field is preserved in the mapping of V1 onto MT. A separate representation of the central visual field was found immediately adjacent to foveal MT. This region, the FST area (Ungerleider et al. 1982; Ungerleider and Desimone 1986a, b), was distinguishable from MT in three ways: 1) by the presence of occasional visually unresponsive cells, 2) by the presence of cells with very large receptive fields intermingled with cells whose receptive fields are comparable in size to those found in foveal MT, and 3) by an increased incidence of cells responding during tracking. Of 34 FST neurons, 53% had a directionality index of 0.6 or higher. An additional 22 cells recorded in the superior temporal sulcus were judged to be outside both MT and FST.     

Recovery of function after lesions in the superior temporal sulcus in the monkey.

1. Ibotenic acid lesions in the monkey's middle temporal area (MT) and the medial superior temporal area (MST) in the superior temporal sulcus (STS) have previously been shown to produce a deficit in initiation of smooth-pursuit eye movements to moving visual targets. The deficits, however, recovery within a few days. In the present experiments we investigated the factors that influence that recovery. 2. We tested two aspects of the monkey's ability to use motion information to acquire moving targets. We used eye-position error as a measure of the monkey's ability to make accurate initial saccades to the moving target. We measured eye speed within the first 100 ms after the saccade to evaluate the monkey's initial smooth pursuit. 3. We determined that pursuit recovery was not dependent specifically on the use of neurotoxic lesions. Although the rate of recovery was slightly altered by replacing the usual neurotoxin (ibotenic acid) with another neurotoxin [alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA)] or with an electrolytic lesion, pursuit recovery still occurred within a period of days to weeks. 4. There was a relationship between the size and location of the lesion and the recovery time. The time to recovery for eye-position error and initial eye speed increased with the fraction of MT removed. Whether the rate of recovery and size of lesions within regions on the anterior bank were related was unresolved. 5. We found that a large AMPA lesion within the STS that removed all of MT and nearly all of MST drastically altered the rate of recovery. Recovery was incomplete more than 7 mo after the lesion. Even with this lesion, however, the monkey's ability to use motion information for pursuit was not completely eliminated. 6. The large lesion also included parts of areas V1, V2, V3, and V4, but analysis of the visual fields associated with this lesion indicated that these areas probably did not have a substantial effect on recovery. 7. We tested whether visual motion experience of the monkey after a lesion was necessary for recovery by limiting the monkey's experience either by using a mask or by using 4-Hz stroboscopic illumination. In one monkey the eye-position error component of pursuit was prolonged to greater than 2 wk, but recovery of eye speed was not. Reduced motion experience had little effect on recovery in the other two monkeys. These results suggest that such visual motion experience is not necessary for the recovery of pursuit.(ABSTRACT TRUNCATED AT 400 WORDS)     

Polysensory properties of neurons in the anterior bank of the caudal superior temporal sulcus of the macaque monkey

1. We examined the sensory properties of cells in the anterior bank of the caudal part of the superior temporal sulcus (caudal STS) in anesthetized, paralyzed monkeys to visual, auditory, and somesthetic stimuli. 2. In the anterior bank of the caudal STS, there were three regions distinguishable from each other and also from the middle temporal area (MT) in the floor of the STS and area Tpt in the superior temporal gyrus. The three regions were located approximately in the respective inner, middle, and outer thirds of the anterior bank of the caudal STS. These three regions are referred to, from the inner to the outer, as the medial superior temporal region (MST), the mostly unresponsive region, and the caudal STS polysensory region (cSTP), respectively. 3. The extent of MST and its response properties agreed with previous studies. Cells in MST responded exclusively to visual stimuli, had large visual receptive fields (RFs), and nearly all (91%) showed directional selectivity. 4. In the mostly unresponsive region, three quarters of cells were unresponsive to any stimulus used in this study. A quarter of the cells responded to only visual stimuli and most did not show directional selectivity for moving stimuli. Several directionally selective cells responded to movements of three-dimensional objects, but not of projected stimuli. 5. The response properties of cells in the superficial cortex of the caudal superior temporal gyrus, a part of area Tpt, external to cSTP were different from those of cells in the three regions in the anterior bank of the STS. Cells in Tpt were exclusively auditory, and had much larger auditory RFs (mean = 271 degrees) than those of acoustically-driven cSTP cells (mean = 138 degrees). 6. The cSTP contained unimodal visual, auditory, and somesthetic cells as well as multimodal cells of two or all three modalities. The sensory properties of cSTP cells were as follows. 1) Out of 200 cells recorded, 102 (51%) cells were unimodal (59 visual, 33 auditory, and 10 somesthetic), 36 (18%) cells were bimodal (21 visual+auditory, 7 visual+somesthetic, and 8 auditory+somesthetic), and four (2%) cells were trimodal. Visual and auditory responses were more frequent than somesthetic responses: the ratio of the population of cells driven by visual: auditory: somesthetic stimuli was 3:2:1. 2) Visual RFs were large (mean diameter, 59 degrees), but two-thirds were limited to the contralateral visual hemifield. About half the cells showed directional selectivity for moving visual stimuli. None showed selectivity for particular visual shapes.(ABSTRACT TRUNCATED AT 400 WORDS)     

Directional tuning of motion-sensitive cells in the anterior superior temporal polysensory area of the macaque

An investigation was made into the directional sensitivity of cells in the macaque anterior superior temporal polysensory region (STPa) to the motion of objects. The cells studied were sensitive to the presence of motion but showed little or no selectivity for the form of the stimulus. Directional tuning was not continuously distributed about all possible directions. The majority of cells were most responsive to motion in a direction within 15° of one of the three cartesian axes (up/down, left/right, towards/away). Tuning to direction varied in sharpness. For most (34/37) cells the angular change in direction required to reduce response to half maximal was between 45 and 70° (for 3/37 cells it was > 90°). The estimates of the directionality (median I _d = 0.97) of STPa cells was similar to that reported for posterior motion processing areas (the middle temporal area, MT, and the medial superior temporal area, MST). The tuning for direction (sharpness, distribution and discrimination) of the motion-sensitive STPa cells were found to be similar to the tuning for perspective view of STPa cells selective for static form of the head and body. On average the STPa responses showed a 100- to 300-ms transient burst of activity followed by a tonic discharge maintained at approximately 20% of the peak firing rate for the duration of stimulation. The responses of motion-sensitive STPa cells occurred at an earlier latency (mean 91 ms) than responses of cells selective for static form (mean 119 ms), but the time course of responses of the two classes of cell were similar in many other respects. The early response latency and directional selectivity indicate that motion sensitivity in STPa cells derives from the dorsal visual pathway via MT/MST. The similarity of tuning for direction and perspective view within STPa may facilitate the integration of motion and form processing within this high-level brain area.     

Object-centered encoding by face-selective neurons in the cortex in the superior temporal sulcus of the monkey

Summary Neurophysiological studies have shown that some neurons in the cortex in the superior temporal sulcus and in the inferior temporal cortex respond to faces. To determine if some face responsive neurons encode stimuli in an object-centered coordinate system rather than a viewer-centered coordinate system, a large number of neurons were tested for sensitivity to head movement in 3 macaque monkeys. Ten neurons responded only when a head undergoing rotatory movements was shown. All of these responded to a particular movement independently of the orientation of the moving head in relation to the viewer, maintaining specificity even when the moving head was inverted or shown from the back, thereby reversing viewer-centered movement vectors. This was taken as evidence that the movement was encoded in object-centered coordinates. In tests of whether there are neurons in this area which respond differently to the faces of different individuals relatively independently of viewing angle, it was found that a further 18 neurons responded more to one static face than another across different views. However, for 16 of these 18 cells there was still some modulation of the neuronal response with viewing angle. These 16 neurons thus did not respond perfectly in relation to the object shown independently of viewing angle, and may represent an intermediate stage between a viewercentered and an object-centered representation. In the same area as these neurons, other cells were found which responded on the basis of viewercentered coordinates. These neurophysiological findings provide evidence that some neurons in the inferior temporal visual cortex respond to faces (or heads) on the basis of object-centered coordinates, and that others have responses which are intermediate between object-centered and viewer-centered representations. The results are consistent with the hypothesis that object-centered representations are built in the inferior temporal visual cortex.     

Differential characteristics of face neuron responses within the anterior superior temporal sulcus of macaques

The anterior superior temporal sulcus (STS) of macaque monkeys is thought to be involved in the analysis of incoming perceptual information for face recognition or identification; face neurons in the anterior STS show tuning to facial views and/or gaze direction in the faces of others. Although it is well known that both the anatomical architecture and the connectivity differ between the rostral and caudal regions of the anterior STS, the functional heterogeneity of these regions is not well understood. We recorded the activity of face neurons in the anterior STS of macaque monkeys during the performance of a face identification task, and we compared the characteristics of face neuron responses in the caudal and rostral regions of the anterior STS. In the caudal region, facial views that elicited optimal responses were distributed among all views tested; the majority of face neurons responded symmetrically to right and left views. In contrast, the face neurons in the rostral region responded optimally to a single oblique view; right-left symmetry among the responses of these neurons was less evident. Modulation of the face neuron responses according to gaze direction was more evident in the rostral region. Some of the face neuron responses were specific to a certain combination of a particular facial view and a particular gaze direction, whereas others were associated with the relative spatial relationship between facial view and gaze direction. Taken together, these results indicated the existence of a functional heterogeneity within the anterior STS and suggested a plausible hierarchical organization of facial information processing.     

Recovery of visual responses in foveal V1 neurons following bilateral foveal lesions in adult monkey

Summary Cells in the foveal representation of V1 cortex of adult primates became visually responsive after normal sensory input was removed. Immediately after foveae were lesioned bilaterally, a region was found where no cells' activity could be modulated by visual stimulation. Recordings made in that deafferented zone at > 2.5 months after lesions revealed that activity of over half of the cells could be modulated by visual stimuli presented to intact peripheral retina. Although response characteristics made cells with recovered driving quite unlike normal cells, the result suggests a level of visual cortical reorganization previously observed only in immature animals.     

Commissural afferents to the cortex surrounding the posterior part of the superior temporal sulcus in the monkey

Commissural afferents to the cortex surrounding the posterior part of the superior temporal sulcus were studied in Japanese monkeys by the horseradish peroxidase method. After injection of the enzyme, many callosal neurons were labeled contralaterally in the cortical area corresponding to the injection site (homotopical area) and in other regions (heterotopical area). Most of the callosal neurons were triangular in shape, occurring for the most part in layer III of both homotopical and heterotopical areas (about 75-90% of the total number of labeled cells). Mean diameters of the cell bodies were about 11-13 micron.     

Functional organization of a visual area in the posterior bank of the superior temporal sulcus of the rhesus monkey

1. Anatomical studies have shown the cortex of the posterior bank of the superior temporal sulcus to receive a projection from visual cortical areas, including areas 17, 18 and 19. In this paper the response of single neurones in this area to simple visual stimulation is reported. Ten monkeys were studied.2. A clear but relatively crude topographic representation of the visual field was found. There was a large variation in the size of the receptive fields of individual cells, even in a single penetration. Some cells, with the central parts of their receptive fields located from between 1 and 5 degrees from the centre of gaze had receptive fields averaging about 10 degrees x 10 degrees or even larger. Other cells with central receptive fields had much smaller field sizes.3. Two main types of neurones were encountered, with subdivisions within each type. The first type responded to movement irrespective of form. These could be subdivided into neurones which responded to movement in any direction within the receptive field and neurones which responded to movement in one direction only (directionally selective neurones). Another type of cell was responsive to both contour and movement, much like the complex and lower order hypercomplex cells. Almost all such neurones were directionally selective.4. In oblique penetrations through this cortical region, there tended frequently to be an orderly shift in preferred directions of motion, thus suggesting the possibility of a columnar organization for movement.5. Combined anatomical (degeneration) and electrophysiological experiments showed that these types of neurones are found in those regions of the posterior bank of the superior temporal sulcus receiving a direct projection from area 17.     

Recovery of saccadic dysmetria following localized lesions in monkey superior colliculus

Damage to the monkey superior colliculus (SC) produces deficits in the generation of saccadic eye movements. Recovery of the accuracy of saccades is rapid, but saccadic latency and peak velocity recover slowly or not at all. In the present experiments we revisited the issue of recovery of function following localized lesions of the SC using three methodological advances: implantation of wire recording electrodes into the SC for the duration of the experiment to ensure that we were recording from the same site on the SC map on successive days; quantification of changes in saccadic accuracy, latency, and velocity using a standard grid of target points in the visual field contralateral to the SC lesion; measurement of movement field size to quantitatively determine any changes following the lesion. We confirmed a decrease in saccadic accuracy following electrolytic lesions of the SC, and we found that this dysmetria recovered within about 4 days. Saccadic latency increased for saccades to the lesion area and this deficit persisted. Peak saccadic velocity decreased immediately after the lesion and decreased further during the 10 days to 2 weeks of the experiment. We found no indication of an expansion of the movement fields of neurons adjacent to the lesion area. This lack of reorganization suggests that movement field changes within the SC cannot mediate the recovery in accuracy of the saccade. The persistence of the latency and velocity deficits despite the recovery of amplitude deficits indicates that saccadic latency and peak velocity are dependent upon the SC whereas saccadic amplitude is not.     

Task-specific deficits in reversal performance following superior colliculus lesions in rats

Rats with lesions of the superior colliculus or control operations were tested in acquisition, extinction and reversal performance of operant visual tasks in four experiments. The tasks used in the experiments were: a simultaneous choice task (Experiment 1), a successive choice task (Experiment 2), a simultaneous go/no-go task (Experiment 3) and a successive go/no-go task (Experiment 4). Acquisition discrimination performance was affected by lesions only in Experiment 3, where in contrast with previous reports animals with lesions performed better than controls. Reversal performance of animals with lesions was impaired in Experiments 1 through 3, but not in Experiment 4. Effects on extinction performance were minimal in all experiments. The data are interpreted to indicate a role of the superior colliculus in the processing of spatial information.     

Corticothalamic connections of the superior temporal sulcus in rhesus monkeys

Summary The corticothalamic connections of the superior temporal sulcus (STS) were studied by means of the autoradiographic technique. The results indicate that corticothalamic connections of the STS in general reciprocate thalamocortical connections. The cortex of the upper bank of the STS-multimodal areas TPO and PGa-projects to four major thalamic targets: the pulvinar complex, the mediodorsal nucleus, the limitanssuprageniculate nucleus, as well as intralaminar nuclei. Within the pulvinar complex, the main projections of the upper bank of the STS are directed to the medial pulvinar (PM) nucleus. Rostral upper bank regions tend to project caudally and medially within the PM nucleus, caudal upper bank regions, more laterally and ventrally. The mid-portion of the upper bank tends to occupy the central sector of the PM nucleus. There are also relatively minor projections from upper bank regions to the lateral pulvinar (PL) and oral pulvinar (PO) nuclei. In contrast to the upper bank, the projections from the lower bank are directed primarily to the pulvinar complex, with only minor projections to intralaminar nuclei. The rostral portion of the lower bank projects mainly to caudal and medial regions of the PM nucleus, whereas the caudal lower bank projects predominantly to the lateral PM nucleus, and also to the PL, PO, and inferior pulvinar (PI) nuclei. The mid-portion of the lower bank projects mainly to central and lateral portions of the PM nucleus, and also to the PI and PL nuclei. The rostral depth of the STS projects mainly to the PM nucleus, with only minor connections to the PO, PI, and PL nuclei. The midportion of multimodal area TPO of the upper bank, areas TPO₂ and TPO₃, projects preferentially to the central sector of the PM nucleus. It is possible that this STS-thalamic connectivity has a role in behavior that is dependent upon more than one sensory modality.Abbreviations AM anterior medial nucleus - AS arcuate sulcus - AV anterior ventral nucleus - BSC brachium of the superior colliculus - Cd caudate nucleus - Cif nucleus centralis inferior - Cim nucleus centralis intermedialis - CL central lateral nucleus - CM centromedian nucleus - CM-Pf centromedian-parafascicular nucleus - Cs nucleus centralis superior - CS central sulcus - CSL nucleus centralis lateralis superior - GLd dorsal lateral geniculate nucleus - GM medial geniculate nucleus - Hb habenula - IOS inferior occipital sulcus - IPS intraparietal sulcus - LD lateral dorsal nucleus - LF lateral fissure - Li limitans nucleus - LP lateral posterior nucleus - LS lunate sulcus - MD mediodorsal nucleus - Pa paraventricular nucleus - Pen paracentral nucleus - Pf parafascicular nucleus - PI inferior pulvinar nucleus - PL lateral pulvinar nucleus - PM medial pulvinar nucleus - PO oral pulvinar nucleus - PS principal sulcus - Pt parataenial nucleus - R reticular nucleus - Re reuniens nucleus - SG suprageniculate nucleus - STN subthalamic nucleus - STS superior temporal sulcus - THI habenulo-interpeduncular tract - VLc nucleus ventralis lateralis, pars caudalis - VLm nucleus ventralis lateralis, pars medialis - VLo nucleus ventralis lateralis, pars oralis - VLps nucleus ventralis lateralis, pars postrema - VPI ventroposteroinferior nucleus - VPLc nucleus ventralis posterior lateralis, pars caudalis - VPLo nucleus ventralis posterior lateralis, pars oralis - VPM ventroposteromedial nucleus - VPMpc ventroposteromedial nucleus, parvocellular portion - X nucleus X     

Both striate cortex and superior colliculus contribute to visual properties of neurons in superior temporal polysensory area of macaque monkey

Although the tectofugal system projects to the primate cerebral cortex by way of the pulvinar, previous studies have failed to find any physiological evidence that the superior colliculus influences visual activity in the cortex. We studied the relative contributions of the tectofugal and geniculostriate systems to the visual properties of neurons in the superior temporal polysensory area (STP) by comparing the effects of unilateral removal of striate cortex, the superior colliculus, or of both structures. In the intact monkey, STP neurons have large, bilateral receptive fields. Complete unilateral removal of striate cortex did not eliminate visual responses of STP neurons in the contralateral visual hemifield; rather, nearly half the cells still responded to visual stimuli in the hemifield contralateral to the lesion. Thus the visual properties of STP neurons are not completely dependent on the geniculostriate system. Unilateral striate lesions did affect the response properties of STP neurons in three ways. Whereas most STP neurons in the intact monkey respond similarly to stimuli in the two visual hemifields, responses to stimuli in the hemifield contralateral to the striate lesion were usually weaker than responses in the ipsilateral hemifield. Whereas the responses of many STP neurons in the intact monkey were selective for the direction of stimulus motion or for stimulus form, responses in the hemifield contralateral to the striate lesion were not selective for either motion or form. Whereas the median receptive field in the intact monkey extended 80 degrees into the contralateral visual field, the receptive fields of cells with responses in the contralateral field that survived the striate lesions had a median border that extended only 50 degrees into the contralateral visual field. Removal of both striate cortex and the superior colliculus in the same hemisphere abolished the responses of STP neurons to visual stimuli in the hemifield contralateral to the combined lesion. Nearly 80% of the cells still responded to visual stimuli in the hemifield ipsilateral to the lesion. Unilateral removal of the superior colliculus alone had only small effects on visual responses in STP. Receptive-field size and visual response strength were slightly reduced in the hemifield contralateral to the collicular lesion. As in the intact monkey, selectivity for stimulus motion or form were similar in the two visual hemifields. We conclude that both striate cortex and the superior colliculus contribute to the visual responses of STP neurons. Striate cortex is crucial for the movement and stimulus specificity of neurons in STP.(ABSTRACT TRUNCATED AT 400 WORDS)     

Hyperactivity and deficits in problem solving following superior colliculus lesions in the rat

Animals with bilateral superior colliculus lesions or control operations were tested for problem solving ability in a Hebb-Williams closed field and for locomotor activity and emotionality in an open field. Experimental animals were significantly both hyperactive and deficient in maze performance. Activity and maze performances were significantly related in the experimental animals but not in controls. The 2 groups did not differ in emotionality as measured by defecation. The results were interpreted as supporting the current theory that the superior colliculus is functionally involved in attention and orientation.     

Individual differences in the theory of mind and superior temporal sulcus

We investigated the brain area with regard to individual differences in the theory of mind. Using functional magnetic resonance imaging, we examined the brain area in which signal intensity was apparently related to performance of a theory-of-mind task on multiple regression analysis. A significant relation was observed between performance of theory-of-mind task and activation in the left anterior superior temporal sulcus. We could not find such an activation in the superior temporal sulcus and the temporo-parietal junction area. The present findings provide new evidence that the anterior superior temporal sulcus might dictate individual differences in theory of mind.     

The effect of superior temporal lesions on the recognition of species-specific calls in the squirrel monkey

Summary Eleven squirrel monkeys (Saimiri sciureus) were trained to discriminate species-specific calls from non-species-specific complex sounds in a go, no-go procedure with social contact as positive reinforcement. The task required that the animals not only responded to a particular call but that this response should be generalized to any squirrel monkey call, whether or not it had been presented previously in training.After having reached a performance level of 75% correct responses in three consecutive sessions, seven animals received bilateral lesions of the auditory cortex; the other four animals served as controls. It was found that small lesions within the superior temporal gyrus did not interfere with the discrimination task. Lesions destroying about three quarters of the auditory cortex led to loss of retention; during retraining the animals did not reach criterion, but performed significantly above chance. These animals were able, however, to master a simplified version of the task where one species-specific call had to be discriminated from one non-species-specific sound. Animals with almost total ablation of the auditory cortex were capable of mastering neither the generalized task nor the simplified version.From these results, together with those of the literature, it is concluded 1) that recognition of complex sounds is not possible after complete auditory cortex ablation, probably because of interference with gestalt-formation processing, and 2) that species-specific calls are processed in the auditory system in the same way as other complex sounds.