Hierarchical effects of task engagement on amplitude modulation encoding in auditory cortex.

We recorded from middle lateral belt (ML) and primary (A1) auditory cortical neurons while animals discriminated amplitude-modulated (AM) sounds and also while they sat passively. Engagement in AM discrimination improved ML and A1 neurons' ability to discriminate AM with both firing rate and phase-locking; however, task engagement affected neural AM discrimination differently in the two fields. The results suggest that these two areas utilize different AM coding schemes: a "single mode" in A1 that relies on increased activity for AM relative to unmodulated sounds and a "dual-polar mode" in ML that uses both increases and decreases in neural activity to encode modulation. In the dual-polar ML code, nonsynchronized responses might play a special role. The results are consistent with findings in the primary and secondary somatosensory cortices during discrimination of vibrotactile modulation frequency, implicating a common scheme in the hierarchical processing of temporal information among different modalities. The time course of activity differences between behaving and passive conditions was also distinct in A1 and ML and may have implications for auditory attention. At modulation depths ≥ 16% (approximately behavioral threshold), A1 neurons' improvement in distinguishing AM from unmodulated noise is relatively constant or improves slightly with increasing modulation depth. In ML, improvement during engagement is most pronounced near threshold and disappears at highly suprathreshold depths. This ML effect is evident later in the stimulus, and mainly in nonsynchronized responses. This suggests that attention-related increases in activity are stronger or longer-lasting for more difficult stimuli in ML.

Excitatory and inhibitory intensity tuning in auditory cortex: evidence for multiple inhibitory mechanisms.

The intensity tuning of excitatory and suppressive domain frequency response areas was investigated in 230 cat primary auditory cortical and 92 posterior auditory field neurons. Suppressive domains were explored using simultaneous 2-tone stimulation with one tone at the best excitatory frequency. The intensity tuning of excitatory and suppressive domains was negatively correlated, supporting the hypothesis that inhibitory sidebands are related to excitatory domain intensity tuning. To further test this hypothesis, we compared the slopes of the edges of suppressive bands to the intensity tuning of excitatory domains. Edges of suppressive bands next to excitatory domains had slopes significantly more slanted toward the excitatory area in neurons with intensity-tuned excitatory domains. This relationship was not observed for suppressive band edges not next to the excitatory domain (e.g., the lower edge of lower suppressive bands). This indicates that intensity tuning ultimately observed in the excitatory domain results from overlapping excitatory and inhibitory inputs. In combination with results using forward masking, our results suggest that there are separate early and late sources of inhibition contributing to cortical frequency response areas, and only the early-stage inhibition contributes to excitatory domain intensity tuning.

Spectrotemporal organization of excitatory and inhibitory receptive fields of cat posterior auditory field neurons.

The excitatory and inhibitory frequency/intensity response areas (FRAs) and spectrotemporal receptive fields (STRFs) of posterior auditory cortical field (PAF) single neurons were investigated in barbiturate anesthetized cats. PAF neurons' pure-tone excitatory FRAs (eFRAs) exhibited a diversity of shapes, including some with very broad frequency tuning and some with multiple distinct excitatory frequency ranges (i.e., multipeaked eFRAs). Excitatory FRAs were analyzed after selectively excluding spikes on the basis of spike response times relative to stimulus onset. This analysis indicated that spikes with shorter response times were confined to narrow regions of the eFRAs, while spikes with longer response times were more broadly distributed over the eFRA. First-spike latencies in higher threshold response peaks of multipeaked eFRAs were approximately 10 ms longer, on average, than latencies in lower threshold response peaks. STRFs were constructed to examine the dynamic frequency tuning of neurons. More than half of the neurons (51%) had STRFs with "sloped" response maxima, indicating that the excitatory frequency range shifted with time. A population analysis demonstrated that the median first-spike latency varied systematically as a function of frequency with a median slope of approximately 12 ms per octave. Inhibitory frequency response areas were determined by simultaneous two-tone stimulation. As in primary auditory cortex (A1), a diversity of inhibitory band structures was observed. The largest class of neurons (25%) had an inhibitory band flanking each eFRA edge, i.e., one lower and one upper inhibitory band in a "center-surround" organization. However, in comparison to a previous report of inhibitory structure in A1 neurons, PAF exhibited a higher incidence of neurons with more complex inhibitory band structure (for example, >2 inhibitory bands). As was the case with eFRAs, spikes with longer response times contributed to the complexity of inhibitory FRAs. These data indicate that PAF neurons integrate temporally varying excitatory and inhibitory inputs from a broad spectral extent and, compared with A1, may be suited to analyzing acoustic signals of greater spectrotemporal complexity than was previously thought.

Global processing of spectrally complex sounds in macaques (Macaca mullata) and humans.

How nonhuman species perceive the world is a biological question of fundamental importance, and has major significance for establishing the validity and possible limitations of animal models of human sensory function and perception. Studies in comparative hearing have revealed that almost all animals, including monkeys, are worse than humans at discriminating tone frequencies. Less is known, however, about comparative differences in discriminating more spectrally complex sounds. We compared the capacity of macaques and humans to discriminate complex sound patterns by measuring spectral-contrast sensitivity using stimuli having sine-modulated power spectra, analogous to sine-wave gratings used in visual studies. We found that the auditory system of the macaque is far less sensitive than the human system over the sine-profile frequency range tested (0.5-2.0 cycles/octave). These results indicate that rhesus macaques hear at least some spectrally complex sounds with less fidelity than do humans, and demonstrate large differences in primate species' abilities to process low-resolution spectral patterns. These results cannot be accounted for by traditional, narrowband peripheral filter models of spectral analysis, but instead, imply the involvement of a central, frequency integration process that may differ significantly across species.

Shapes and level tolerances of frequency tuning curves in primary auditory cortex: quantitative measures and population codes.

The shape and level tolerance of the excitatory frequency/intensity tuning curves (eFTCs) of 160 cat primary auditory cortical (A1) neurons were investigated. Overall, A1 cells were characterized by tremendous variety in eFTC shapes and symmetries; eFTCs were U-shaped ( approximately 20%), V-shaped ( approximately 20%), lower-tail-upper-sharp ( approximately 15%), upper-tail-lower-sharp (<2%), slant-lower ( approximately 10%), slant-upper (<3%), multipeaked ( approximately 10%), and circumscribed ( approximately 20%). Quantitative analysis suggests that eFTC are best thought of as forming a continuum of shapes, rather than falling into discrete categories. A1 eFTCs tended to be more level tolerant than eFTCs from earlier stations in the ascending auditory system as inferred from other studies. While individual peaks of multipeaked eFTCs were similar to single peaked eFTCs, the overall eFTC of multipeaked neurons (spanning the range of all peaks) tended to have high-frequency tails. Measurements of shape and symmetry indicate that A1 eFTCs, on average, tended to have greater area on the low-frequency side of characteristic frequency (CF) than on the high-frequency side. A1 cells showed a relationship between CF and the inverse slope of low-frequency edges of eFTCs, but not for high-frequency edges. These data demonstrate that frequency tuning, particularly along the eFTC low-frequency border, sharpens along the lemniscal pathway to A1. The results are consistent with studies in mustached bats (Suga 1997) and support the idea that spectral decomposition along the ascending lemniscal pathway up to A1 is a general organizing principle of mammalian auditory systems. Altogether, these data suggest that A1 neurons' eFTCs are shaped by complex patterns of inhibition and excitation accumulating along the auditory pathways, implying that central rather than peripheral filtering properties are responsible for certain psychophysical phenomena.

Auditory scene analysis in dyslexics.

It has been argued that dyslexics suffer from temporal sensory processing deficits which affect their ability to discriminate speech in quiet environments. The impact of auditory deficits on non-language aspects of perception, however, is poorly understood. In almost every natural-listening environment, one must constantly construct scenes of the auditory world by grouping and analyzing sounds generated by multiple sources. We investigated whether dyslexics have difficulties grouping sounds. The results demonstrate that dyslexics have an impairment in grouping auditory objects that depends both on the sounds' frequency and presentation rate (i.e. the spectrotemporal context of the sound). We conclude that dyslexics have difficulty constructing scenes of the auditory world, and that these deficits can contribute to learning impairments.

Modular organization of frequency integration in primary auditory cortex.

Two fundamental aspects of frequency analysis shape the functional organization of primary auditory cortex. For one, the decomposition of complex sounds into different frequency components is reflected in the tonotopic organization of auditory cortical fields. Second, recent findings suggest that this decomposition is carried out in parallel for a wide range of frequency resolutions by neurons with frequency receptive fields of different sizes (bandwidths). A systematic representation of the range of frequency resolution and, equivalently, spectral integration shapes the functional organization of the iso-frequency domain. Distinct subregions, or "modules," along the iso-frequency domain can be demonstrated with various measures of spectral integration, including pure-tone tuning curves, noise masking, and electrical cochlear stimulation. This modularity in the representation of spectral integration is expressed by intrinsic cortical connections. This organization has implications for our understanding of psychophysical spectral integration measures such as the critical band and general cortical coding strategies.

Global spectral and location effects in auditory perceptual grouping.

An important problem in cognitive and systems neuroscience concerns the extent to which perceptual organization can be explained by "local," peripheral physiological mechanisms, or rather by more "global," central, and higher-level processes. Though central in vision research, this issue has received little attention in the field of audition. One claim is that auditory-perceptual grouping mechanisms, possibly related to visual figure-from-ground segregation or "pop-out," are low level, resulting from local processing in the frequency domain. However, no experiments have been performed specifically to test this question. We examined the effects of perceptual grouping on detection for reversal of two repeated target tones, one constant in frequency (1030 Hz), the other free to vary between trials (1045-8580 Hz). Detection was examined in the presence of a 1000-Hz background tone that repeated between target presentations. By varying the frequency of the high-target tone, this task was designed to modulate grouping between the background and low-target tones, thereby affecting reversal detection. We predicted that at large target frequency differences (deltaf), the high-target tone would segregate from the background and low-target tones, and so render the background and low-target tones less distinct. We found that reversal detection declined from optimal levels with increasing deltaf, and that performance was improved by spatially separating the location of the target and background sounds by at least 32 degrees. These results demonstrate that global frequency integration over at least three octaves occurs through grouping, and that grouping is affected by source location. This implies that auditory-perceptual grouping involves global neural processing, i.e., the participation of neurons with very broad frequency input that are also sensitive to spatial location.

Organization of inhibitory frequency receptive fields in cat primary auditory cortex.

Based on properties of excitatory frequency (spectral) receptive fields (esRFs), previous studies have indicated that cat primary auditory cortex (A1) is composed of functionally distinct dorsal and ventral subdivisions. Dorsal A1 (A1d) has been suggested to be involved in analyzing complex spectral patterns, whereas ventral A1 (A1v) appears better suited for analyzing narrowband sounds. However, these studies were based on single-tone stimuli and did not consider how neuronal responses to tones are modulated when the tones are part of a more complex acoustic environment. In the visual and peripheral auditory systems, stimulus components outside of the esRF can exert strong modulatory effects on responses. We investigated the organization of inhibitory frequency regions outside of the pure-tone esRF in single neurons in cat A1. We found a high incidence of inhibitory response areas (in 95% of sampled neurons) and a wide variety in the structure of inhibitory bands ranging from a single band to more than four distinct inhibitory regions. Unlike the auditory nerve where most fibers possess two surrounding "lateral" suppression bands, only 38% of A1 cells had this simple structure. The word lateral is defined in this sense to be inhibition or suppression that extends beyond the low- and high-frequency borders of the esRF. Regional differences in the distribution of inhibitory RF structure across A1 were evident. In A1d, only 16% of the cells had simple two-banded lateral RF organization, whereas 50% of A1v cells had this organization. This nonhomogeneous topographic distribution of inhibitory properties is consistent with the hypothesis that A1 is composed of at least two functionally distinct subdivisions that may be part of different auditory cortical processing streams.

Functional organization of spectral receptive fields in the primary auditory cortex of the owl monkey.

Recent experiments in the cat have demonstrated that several response parameters, including frequency tuning, intensity tuning, and FM selectivity, are spatially segregated across the isofrequency axis. To investigate whether a similar functional organization exists in the primate, we have studied the spatial distribution of pure-tone receptive field parameters across the primary auditory cortex (AI) in six owl monkeys (Aotus trivirgatus). The distributions of binaural interaction types and onset latency were also examined. Consistent with previous studies, the primary auditory cortex contained a clear cochleotopic organization. We demonstrate here that several other properties of the responses to tonal stimuli also showed nonrandom spatial distributions that were largely independent from each other. In particular, the sharpness of frequency tuning to pure tones, intensity tuning and sensitivity, response latency, and binaural interaction types all showed spatial variations that were independent from the representation of characteristic frequency and from each other. Statistical analysis confirmed that these organizations did not reflect random distributions. The overall organizational pattern of overlaying but independent functional maps that emerged was quite similar to that seen in AI of cats and, in general, appears to reflect a fundamental organization principle of primary sensory cortical fields.

Auditory temporal integration in the rhesus macaque (Macaca mulatta).

Temporal integration for pure tones was examined in two rhesus macaques. The subjects were required to respond to a brief sound (a tone burst) that deviated from a previous series of sounds (noise bursts) on a trial (a deviant-stimulus detection paradigm). Psychometric functions and thresholds were determined from correct detections (hit proportions) alone, and from d' scores. Two models describing the decline in threshold as a function of stimulus duration, one a power function the other an exponential, were tested against the data. When the decline (slope) in threshold per log stimulus duration is used as a rate measure, our results yield a lower estimate of temporal integration rate in rhesus than did a previous study [Clack, J. Acoust. Soc. Am. 40, 1140-1146 (1966)]. Both studies, however, gave slope estimates of integration rate that were higher than in most other species. Comparison of the models using data from several species, revealed that the exponential, but not the power model, could account for two sources of variation in threshold measurement. One source is due to the range across threshold as a function of duration (the linear rate component), and is described by the constant of proportionality Ik in the model. The other source of variation arises from the rate of decline within this range (the nonlinear rate component), and is described by the time constant tau. In terms of this model, differences in rate estimates between Clack's study and ours (and between rhesus and other species) are primarily due to the linear component. The nonlinear rate component was about equal for our study and Clack's (tau = approximately 150 ms): a time constant that is just slightly larger (indicating a rate of temporal integration slightly slower) than for most other species examined.

Functional topography of cat primary auditory cortex: response latencies.

Minimum onset latency (Lmin) of single- and multiple-unit responses were mapped in the primary auditory cortex (AI) of barbiturate-anesthetized cats. Contralateral Lmin for multiple units was non-homogeneously distributed along the dorso-ventral/isofrequency axis of the AI. Responses with shorter latencies were more often located in the central, more sharply tuned region while longer latencies were more frequently encountered in the dorsal and ventral portions of the AI. For single units, a large scatter of Lmin values was found throughout the extent of the AI including cortical depth. The relationship between Lmin and previously reported spectral, intensity and temporal parameters was analyzed and revealed statistically significant correlations between minimum onset latency and the following response properties in some but not all studied animals: sharpness of tuning of a frequency response area 10 dB above threshold, broadband transient response, strongest response level, monotonicity of rate/level functions, dynamic range, and preferred frequency modulation sweep direction. This analysis suggests that Lmin is determined by several independent factors and that the prediction of Lmin based on relationships with other spectral and temporal response properties is inherently weak. The spatial distribution and the functional relationship between these response parameters may provide an important aspect of the time-based cortical representation of specific features in the animal's natural environment.

Topography of intensity tuning in cat primary auditory cortex: single-neuron versus multiple-neuron recordings.

1. We studied the spatial distributions of amplitude tuning (monotonicity of rate-level functions) and response threshold of single neurons along the dorsoventral extent of cat primary auditory cortex (AI). To pool data across animals, we used the multiple-unit map of monotonicity as a frame of reference. Amplitude selectivity of multiple units is known to vary systematically along isofrequency contours, which run roughly in the dorsoventral direction. Clusters sharply tuned for intensity (i.e., "nonmonotonic" clusters) are located near the center of the contour. A second nonmonotonic region can be found several millimeters dorsal to the center. We used the locations of these two nonmonotonic regions as reference points to normalize data across animals. Additionally, to compare this study to sharpness of frequency tuning results, we also used multiple-unit bandwidth (BW) maps as references to pool data. 2. The multiple-unit amplitude-related topographies recorded in previous studies were confirmed. Pooled multiple-unit maps closely approximated the previously reported individual case maps when the multiple-unit monotonicity or the map of bandwidth (in octaves) of pure tones to which a cell responds 40 dB above minimum threshold were used as the pooling reference. When the map of bandwidth (in octaves) of pure tones to which a cell responds 10 dB above minimum threshold map was used as part of the measure, the pooled spatial pattern of multiple-unit activity was degraded. 3. Single neurons exhibited nonmonotonic rate-level functions more frequently than multiple units. Although common in single-neuron recordings (28%), strongly nonmonotonic recordings (firing rates reduced by > 50% at high intensities) were uncommon (8%) in multiple-unit recordings. Intermediately nonmonotonic neurons (firing rates reduced between 20% and 50% at high intensities) occurred with nearly equal probability in single-neuron (28%) and multiple-unit (26%) recordings. The remaining recordings for multiple units (66%) and single units (44%) were monotonic (firing rates within 20% of the maximum at the highest tested intensity). 4. In ventral AI (AIv), the topography of monotonicity for single units was qualitatively similar to multiple units, although single units were on average more intensity selective. In dorsal AI (AId) we consistently found a spatial gradient for sharpness of intensity tuning for multiple units; however, for pooled single units in Aid there was no clear topographic gradient. 5. Response (intensity) thresholds of single neurons were not uniformly distributed across the dorsoventral extent of AI.(ABSTRACT TRUNCATED AT 400 WORDS)

Global synchronous response to autogenous song in zebra finch HVc.

1. The spatial distribution of neuronal responses to autogenous song (AS) was investigated in the HVc of urethan-anesthetized adult male zebra finches (Taeniopygia guttata). In seven birds, penetrations covered the entire mediolateral, rostrocaudal, or dorsoventral extents of HVc. In an eighth, control birth penetrations were made near to but outside of HVc. Reconstruction of recording sites from histological material indicated a good correspondence between sites that exhibited stronger responses to AS than to tone or noise bursts, and sites that were within HVc. 2. Within each experimental bird but not in the control, multiple-unit responses to AS were similar across the entire spatial extent of HVc (up to 1.3 mm). For each experimental bird, the strongest responses occurred within a narrow range of times. The middle of this range of times is called the time of maximum synchronization (TMS). Across birds, 34-75% of recording sites exhibited the same TMS. With the use of a criterion of > 33% of sites exhibiting their strongest responses at the TMS, the temporal scatter around the TMS varied between 6 and 138 ms across individuals. In six of the seven experimental birds, the position of the TMS was not affected by changing the window of integration from 10 to 150 ms. In two experimental birds, short windows of integration tended to emphasize beginning portions of the song. In one case this effect was sufficiently strong to change the TMS for short windows of integration. 3. Each TMS was associated with a syllable of maximum synchronization (SMS). The positions of the SMS varied considerably across birds. In four birds the SMS was one of the syllables of the first motif (a motif is a temporal sequence of syllables that can be repeated > or = 1 times to form a song), in two birds the SMS was the introductory note of song, and in one bird the SMS was the second syllable of the last (3rd) motif. Syllables of the same type as the SMS but occurring in other motifs typically elicited much weaker responses, in many cases weaker than other syllables in those motifs. Syllables that elicited strong responses in non-SMS motifs did not necessarily elicit strong responses in the SMS motif, even if they preceded the SMS. There were no apparent acoustical features of the SMS or the preceding syllable that could account for the global synchronous response to song.(ABSTRACT TRUNCATED AT 400 WORDS)

Distributed representation in the song system of oscines: evolutionary implications and functional consequences.

This paper reviews the organizational principles and implications that have emerged from the analysis of HVc, a forebrain nucleus that is a major site of sensory, motor, and sensorimotor integration in the song control system of oscine passerine birds (songbirds). Anatomical, physiological, and behavioral data support the conclusion that HVc exists within a hierarchically organized system with parallel pathways that converge onto HVc. The organization of HVc is distributed and redundant, and its outputs exhibit broad divergence. A similar pattern of connectivity exists for neostriatum adjacent to HVc. This and other data support the hypothesis that the song system arose from an elaboration or duplication of pathways generally present in all birds. Spontaneous and auditory response activity is strongly correlated throughout HVc, with auditory responses exhibiting strong temporal modulation in a synchronized fashion throughout the nucleus. This suggests that the auditory representation of song is encoded in the synchronized temporal patterns of activation, and that the predominant selectivity for the individual's own song that is observed for HVc neurons results from interactions of auditory input with central pattern generators for song. Most, or all HVc neurons are recruited during singing. The auditory response and motor recruitment properties of individual HVc neurons have no simple relationship, and the spontaneous activity in HVc may build up in the seconds preceding a song. To the extent HVc participates in perceptual phenomena associated with song, production and perception are not tightly linked in adults but may be linked by shared developmental processes during periods of sensorimotor learning.

Functional topography of cat primary auditory cortex: responses to frequency-modulated sweeps.

The spatial distribution of neuronal responses to frequency-modulated (FM) sweeps was mapped with microelectrodes in the primary auditory cortex (AI) of barbiturate-anesthetized cats. Increasing and decreasing FM sweeps (upward- and downward-directed FM sweeps, respectively) covering a range of 0.25-64.0 kHz were presented at three different rates of frequency change over time (i.e, sweep speed). Using multiunit recordings, the high-frequency domain (between 3.2 and 26.3 kHz) of AI was mapped over most of its dorsoventral extent (as determined by the distribution of the excitatory bandwidth, Q10dB) for all six cases studied. The spatial distributions of the preferred sweep speed and the preferred sweep direction were determined for each case. Neuronal responses for frequency sweeps of different speeds appeared to be systematically distributed along the dorsoventral axis of AI. In the dorsal region, cortical cells typically responded best to fast and/or medium FM sweeps, followed more ventrally by cells that responded best to medium--then slow--, then medium-speed FM sweeps. In the more ventral aspect of AI (which in some cases may also have included cells located in the dorsal region of the second auditory field, AII), neurons generally preferred fast FM sweeps. However, a comparison of maps from different animals showed that there was more variability in the distribution of preferred speed responses in the ventral region of the cortex. The directional preference of units for FM sweeps was determined for the sweep speed producing the strongest response. Direction selectivity appeared to be nonrandomly distributed along the dorsoventral axis of AI. In general, units that responded best to upward-directed FM sweeps were located in the more dorsal and ventral aspects of AI while units that responded best to downward-directed FM sweeps were usually located in the mid-region of AI. Direction selectivity was also determined for multiunit responses at each of the three FM sweep speeds. In general, there was a relatively close agreement between the spatial distributions of direction selectivity determined for the strongest response with those calculated for the fast and medium speeds. The spatial distribution of direction selectivity determined for slow FM sweeps deviated somewhat from that determined for the strongest response. Near the dorsoventral center of the mapped areas, the distribution of units that responded best to downward sweeps tended to overlay the distribution of units that responded best to slow speeds, suggesting some spatial covariance of the two parameters.(ABSTRACT TRUNCATED AT 400 WORDS)

Topography of excitatory bandwidth in cat primary auditory cortex: single-neuron versus multiple-neuron recordings.

1. The spatial distribution of the sharpness of tuning of single neurons along the dorsoventral extent of primary auditory cortex (AI) was studied. A sharpness of tuning gradient was initially obtained with multiple-unit recordings, and in combination with the cochleotopic organization, served as a frame of reference for the locations of single neurons. The frequency selectivity or "integrated excitatory bandwidth" of multiple units varied systematically along the dorsoventral extent of AI. The most sharply tuned unit clusters were found at the approximate center of the dorsoventral extent. A gradual broadening of the integrated excitatory bandwidth in both dorsal and ventral directions was consistently seen. 2. The multiple-unit measures of the bandwidth 10 (BW10) and 40 dB (BW40) above minimum threshold, pooled across several animals and expressed in octaves, were similar to those described within individual cases in cats. As in the individual animals, the bandwidth maps were V shaped with minima located at the approximate center of the dorsal-ventral extent of AI. The location of the minimum in the multiple-unit bandwidth map (i.e., the most sharply tuned area) was used as a reference point to pool single-neuron data across animals. 3. For single neurons, the dorsal half of the BW40 distribution showed a gradient paralleling that found for multiple units. For both single and multiple units, the average excitatory bandwidth increased at a rate of approximately 0.27 octaves/mm from the center of AI toward the dorsal fringe. Differing from the dorsal half of AI, the ventral half of AI showed no clear BW40 gradient for single units along its dorsoventral extent. At 40 dB above minimum threshold, most ventral neurons encountered were sharply tuned. By contrast, the multiple-unit BW40 showed a gradient similar to the dorsal half with 0.23 octaves/mm increasing from the center toward the ventral border of AI. 4. For single neurons, BW10 showed no clear systematic spatial distribution in AI. Neither the dorsal nor the ventral gradient was significantly different from zero slope, although the dorsal half showed a trend toward increasing BW10s. Contrasting single neurons, both dorsal and ventral halves of AI showed BW10 slopes for multiple units confirming a V-shaped map of the integrated excitatory bandwidth within the dorsoventral extent of AI. 5. On the basis of the distribution of the integrated (multiple-unit) excitatory bandwidth, AI was parceled into three regions: the dorsal gradient, the ventral gradient, and the central, narrowly tuned area.(ABSTRACT TRUNCATED AT 400 WORDS)

Functional topography of cat primary auditory cortex: representation of tone intensity.

The neuronal response to tones as a function of intensity was topographically studied with multiple-unit recordings in the primary auditory cortex (AI) of barbiturate-anesthetized cats. The spatial distribution of the characteristics of rate/level functions was determined in each of three intensely studied cases and their relationship to the distribution of spectral parameters (sharpness of tuning and responses to broadband transients) in the same animals was determined. The growth of the high-intensity portion of rate/level functions was estimated by linear regression. Locations with monotonically growing high-intensity portions were spatially segregated from locations with nonmonotonic rate/level functions. Two noncontiguous areas with a high degree of nonmonotonicity were observed. One was located at the dorsoventral center of AI, and a second in the dorsal third of AI. The more ventral aggregate of high nonmonotonicity coincided with the region of sharp frequency tuning. The stimulus levels that produced the highest firing rate (strongest response level, SRL) at any sampled location ranged from 10 to 80 dB sound pressure level (SPL). Several spatial aggregates with either high or low SRLs were observed in AI. The region of sharpest tuning was always associated with a region of low SRLs. The response threshold to contralateral tones at the characteristic frequency (CF) ranged from -10 dB SPL to 85 dB SPL with the majority between 0 and 40 dB SPL. The spatial distribution of response thresholds indicated several segregated areas containing clusters with either higher or lower response thresholds. The correlation of response threshold with integrated bandwidth and transient responses was only weak. Low- and high-intensity tones of the same frequency are represented at different locations in AI as judged by the amount of evoked neuronal activity and are largely independent of the frequency organization. The spatial distribution of locations with high monotonicity and low strongest response levels were aligned with the organization of the integrated excitatory bandwidth and covaried with the response strength to broadband stimuli.

Physiology and topography of neurons with multipeaked tuning curves in cat primary auditory cortex.

1. The physiology and topography of single neuron responses along the isofrequency domain of the middle- and high-frequency portions [characteristic frequencies (CFs) greater than 4 kHz] of the primary auditory cortex (AI) were investigated in the barbiturate-anesthetized cat. Single neurons were recorded at several locations along the extent of isofrequency contours, defined from initial multiple-unit mapping. For each neuron a high-resolution excitatory tuning curve was determined, and for some neurons high-resolution two-tone tuning curves were recorded to measure inhibitory/suppressive areas. 2. A physiologically distinct population of neurons was found in the dorsal part of cat AI. These neurons exhibited two or three distinct excitatory frequency ranges, whereas most neurons in AI responded with excitation to a single narrow frequency range. These were called multipeaked neurons because of the shape of their tuning curves. At frequencies between the excitatory regions, the multipeaked neurons were inhibited or unresponsive. 3. Multipeaked neurons exhibited several distinct threshold minima in their frequency tuning curves. Most of the multipeaked neurons (88%) displayed two frequency minima, whereas the rest exhibited three minima. 4. The frequency separation between threshold minima was less than 1 octave in 71% of the double-peaked neurons recorded. Occasionally, the frequency peaks of these neurons closely corresponded to a response to second and third harmonics without a response to the fundamental frequency. 5. Multipeaked neurons exhibited a wide range of total bandwidths (highest excitatory frequency minus lowest excitatory frequency expressed in octaves). Bandwidths of the isolated peaks within the same neuron were also quite variable. 6. Response latencies to tones with frequencies within each peak of a multipeaked neuron could vary considerably. In 71% (17) of the neurons, tones corresponding to the high-frequency peak (CFh) elicited a longer response latency (greater than 4 ms) than those corresponding to the low-frequency peak (CF1). 7. Inhibitory/suppressive bands, as demonstrated with a two-tone paradigm, were often present between the peaks. Typically, neurons with excitatory peaks of similar response latencies showed an inhibitory band located between the peaks. 8. Ninety percent of the topographically localized multipeaked neurons were in the dorsal part of AI (greater than 1 mm dorsal to the maximum in the sharpness-of-tuning map). Although these neurons were restricted to dorsal AI, only 35% of neurons in this region were multipeaked. 9. Multipeaked neurons could show decreased response latencies and thresholds to two-tone combinations.(ABSTRACT TRUNCATED AT 400 WORDS)

An interactive clinical data system.

A clinical data system is under development at the Upjohn Company that will reduce the time lag from patient visit to data availability, data entry errors, and the amount of training required for system use. The system design allows generality for both the study monitor and the biostatistician, editing of the data before entry, simplified forms design, direct entry of textual responses without coding, and a reduced need for special programs. Interactive subsystems allow a user to describe a study to the system, generate tailor-made entry/edit programs for each study, undate the date base, and produce ad hoc reports. The system is being implemented on top of a data base management system and runs on an IBM 370/148 under VM/CMS.