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We wished to test the hypothesis that there are modules in low-frequency AI that can be identified by their responsiveness to communication calls or particular regions of space. Units were recorded in anaesthetised guinea pig AI and stimulated with conspecific vocalizations and a virtual motion stimulus (binaural beats) presented via a closed sound system. Recording tracks were mainly oriented orthogonally to the cortical surface. Some of these contained units that were all time-locked to the structure of the chutter call (14/22 tracks) and/or the purr call (12/22 tracks) and/or that had a preference for stimuli from a particular region of space (8/20 tracks with four contralateral, two ipsilateral and two midline), or where there was a strong asymmetry in the response to beats of different direction (two tracks). We conclude that about half of low-frequency AI is organized into modules that are consistent with separate "what" and "where" pathways.

In visual and somatosensory cortex there are important functional differences between layers. Although it is difficult to identify laminar borders in the primary auditory cortex (AI) laminar differences in functional processing are still present. We have used electrodes inserted orthogonal to the cortical surface to compare the response properties of cells in all six layers of AI in anaesthetised guinea pigs. Cells were stimulated with short tone pips and two conspecific vocalizations. When frequency response areas were measured for 248 units the tuning bandwidth was broader for units in the deep layers. The mean Q (10) value for tuning in layers IV-VI was significantly smaller (Mann-Whitney test P < 0.001) than for layers I-III. When response latencies were measured, the shortest latencies were found in layer V and the mean latency in this layer was shorter than in any of the more superficial layers (I-IV) when compared with a Tukey analysis of variance (P < 0.005). There were also laminar differences in the best threshold with layer V having the highest mean value. The mean best threshold for layer V (32.7 dB SPL) was significantly different from the means for layers II (25.5 dB SPL) and III (26.3 dB SPL). The responses to two vocalizations also varied between layers: the response to the first phrase of a chutter was smaller and about 10 ms later in the deep layers than in layers II and III. By contrast, the response to an example of whistle was stronger in the deep layers. These results are consistent with a model of AI that involves separate inputs to different layers and descending outputs from layers V/VI (to thalamus and brainstem) that are different from the output from layers II/III (to ipsilateral cortex).

Stellate cells within the ventral cochlear nucleus (VCN) are a diverse cell group that have been classified according to their size and morphology. Some of these stellate cell types constitute major projection neurones into the brainstem and directly into the inferior colliculus, while others are implicated in more local processing. It is still not clear whether a specific physiological profile is uniquely associated with each distinct type of stellate cell. To investigate such associations, we have analysed 23 units with a battery of physiological stimuli in vivo and then examined their shape and outputs following juxtacellular labelling with biocytin. Five physiologically identified groups of cells were filled. These formed two major response classes: onset cells and chopper cells. The two classes could be separated purely on morphological grounds. The onset cells had large somata, large symmetrical dendritic trees and profuse axonal branches that were restricted to the cochlear nucleus on one (On-L) or both sides (On-C) of the brainstem. The chopper cells had smaller, asymmetric, dendritic trees, which were either planar or marginal, had smaller somata and an output axon that left via the trapezoid body. We have confirmed profuse projections into the dorsal cochlear nucleus from all onset cells, and more focal projections from some members of all three groups of chopper cells.

Despite numerous studies stretching over the last 100 years there is still no general agreement on the number of auditory areas in the human cortex or even how to define them by histological methods. Full definition of these areas will require a combination of functional and histological methods but, by using six complementary histological methods, of which most have been used in the monkey, we provide a clearer description of these areas. The primary auditory area was located on the posteromedial two-thirds of the first transverse temporal (Heschl's) gyrus and was distinguished by a dense band of cytochrome oxidase activity in layer IV and the base of layer III, as well as a relatively thick, pale layer V and VI. Layers V and VI together made up 40% of the cortical thickness. Acetylcholinesterase (AChE)-containing pyramidal cells were sparsely distributed within the primary auditory area. The anterolateral third of Heschl's gyrus did not have a clear band of high cytochrome oxidase activity but contained a moderately high density of AChE-containing pyramidal cells and thus appeared to be part of the auditory belt. Within Heschl's sulcus there was a third area, which had a band of high cytochrome oxidase activity and bands of high parvalbumin immunoreactivity and AChE activity in layer IV. This area appeared to be part of the auditory core. Thus the use of staining methods for cytochrome oxidase, AChE and parvalbumin provided additional information which allowed a clearer definition of auditory areas than Nissl or myelin staining alone. Our results suggest that there are two core areas surrounded by at least six belt areas in the human auditory region.

By studying the efferent projections of five auditory areas in the guinea pig cortex, we sought evidence that the larger fields can be divided into subareas based on unique patterns of cortical connections. Small extracellular injections of biocytin were made in combination with evoked potential mapping or single-unit analysis and histochemical determination of cortical landmarks. The two core fields, primary (AI) and dorsocaudal (DC), are partially surrounded by six adjacent belt areas, leaving two gaps: one at the rostral edge of AI and the other at the dorsal edge. All of the areas studied projected to their nearest neighbors, but AI was the only area to project to all seven of the other auditory areas. The caudal, high-frequency (more than 4 kHz) end of AI had different projections from the rostral, low-frequency (less than 1.5 kHz) end, and there was no evidence of connections between the two ends. Each end had separate dorsal and ventral projections. The two ends of AI may be working independently. By contrast, area DC had strong connections between its high- and low-frequency ends and it may be involved in auditory/visual integration. The dorsorostral belt (DRB) was subdivided into two zones on the basis of its projections: the more rostral part appears to overlap the second somatosensory area and be bimodal, while the caudal part has stronger auditory connections. The small belt area (area S) had separate physiological and anatomical properties from the rest of the rostral belt.

The organisation of guinea pig auditory cortex was studied by combining histological methods with microelectrode mapping. This allowed the location of seven auditory areas to be determined in relation to the visual and primary somatosensory areas. The auditory areas were identified by single-unit recordings and their borders defined by evoked potential mapping. The visual areas were identified by their relatively high densities of myelinated fibres, while the primary somatosensory cortex was identified by its characteristic barrels of high cytochrome oxidase (CYO) activity in layer IV. The auditory region had moderate levels of CYO and myelin staining. When staining was optimal, there was a clear edge to the moderate CYO activity, which apparently corresponds to the dorsal border of the primary auditory area (AI) and the other core field that lies dorsocaudal to it (DC). Thus the primary somatosensory area and the visual and auditory regions were separated from each other by a region with lower levels of CYO and myelin staining. The ventral borders of AI and DC could not be determined histologically as there were no sharp transitions in the levels of CYO or myelin staining. The two core areas were partially surrounded by belt areas. The dorsorostral belt and most of the belt around DC responded more strongly to broad-band stimuli than pure tones, while the ventrorostral belt, small field and a belt zone ventral to the rostral part of DC responded better to pure tones. Units in the small field (S) typically had higher thresholds and broader tuning to pure tones than AI, while units in the ventrorostral belt typically had longer onset latencies and gave more sustained responses than units in AI.

The dynamic ranges of cochlear nerve fibres in cats were determined automatically and were related to the fibres' rates of spontaneous activity, in both pooled data and data from individual cochlear nerves. The dynamic range represents the range of levels of a tone at the characteristic frequency of the fibre evoking mean discharge rates between spontaneous and saturated activity. In common with the findings of other investigators, the distribution of spontaneous discharge rates was bimodal. The total population could be divided into two sub-populations with spontaneous discharge rates above and below 15 spikes/s, respectively. The mean dynamic range of fibres having spontaneous discharge rates in excess of 15 spikes/s, was 41 dB (+/- 0.65 S.E.); that for fibres with rates below 15 spikes/s was 50 dB (+/- 1.2 S.E.). While the distributions of dynamic ranges of the two populations overlapped, they were significantly different, and dynamic ranges in excess of 60 dB were only found in substantial numbers (23%) in the population having low spontaneous discharge rates. Some of these were not saturated at the highest stimulus levels used.

We have investigated the responses of neurones in the guinea-pig superior colliculus to combinations of visual and auditory stimuli. When these stimuli were presented separately, some of these neurones responded only to one modality, others to both and a few neurones reliably to neither. To bimodal stimulation, many of these neurones exhibited some form of cross-modality interaction, the degree and nature of which depended on the relative timing and location of the two stimuli. Facilitatory and inhibitory interactions were observed and, occasionally, both effects were found in the same neurone at different inter-stimulus intervals. Neurones whose responses to visual stimuli were enhanced by an auditory stimulus were found in the superficial layers. Although visual-enhanced and visual-depressed auditory neurones were found throughout the deep layers, the majority of them were recorded in the stratum griseum profundum. Neurones that responded to both visual and auditory stimuli presented separately and gave enhanced or depressed responses to bimodal stimulation were found throughout the deep layers, but were concentrated in the stratum griseum intermediale and extended into the stratum opticum.