SENSITIVITY TO INTERAURAL INTENSITY DIFFERENCES OF NEURONS IN PRIMARY
AUDITORY CORTEX OF THE CAT: I. TYPES OF SENSITIVITY AND EFFECTS OF VARIATIONS
IN SOUND PRESSURE LEVEL.
Irvine, D. R. F., R. Rajan and L. M. Aitkin.
Departments of Psychology and Physiology, Monash University, Clayton, Vic.
3168, Australia.
APStracts 2:0249N, 1995.
SUMMARY AND CONCLUSIONS
1. Interaural intensity differences (IIDs) provide the major cue to the
azimuthal location of high-frequency narrow-band sounds. In recent studies of
the azimuthal sensitivity of high-frequency neurons in the primary auditory
cortex (field AI) of the cat, a number of different types of azimuthal
sensitivity have been described and the azimuthal sensitivity of many neurons
was found to vary as a function of changes in stimulus intensity. The extent
to which the shape and the intensity-dependence of the azimuthal sensitivity
of AI neurons reflects features of their IID sensitivity was investigated by
obtaining data on IID sensitivity from a large sample of neurons with
characteristic frequency (CF) >5.5 ? kHz in AI of anesthetized cats. IID-
sensitivity functions were classified in a manner that facilitated comparison
with previously obtained data on azimuthal sensitivity, and the effects of
changes in the base intensity at which IIDs were introduced were examined. 2.
IID-sensitivity functions for CF tonal stimuli were obtained at one or more
intensities for a total of 294 neurons, in most cases by a method of
generating IIDs that kept the average binaural intensity (ABI) of the stimuli
at the two ears constant. In the standard ABI range at which a function was
obtained for each unit, 5 types of IID sensitivity were distinguished. Contra-
max neurons (50% of the sample) had maximum response (a peak or a plateau) at
IIDs corresponding to contralateral azimuths, while ipsi-max neurons (17%) had
the mirror-image form of sensitivity. Near-zero-max neurons (18%) had a
clearly defined maximum response (peak) in the range +/-10 ? dB IID, while a
small group of trough neurons (2%) had a restricted range of minimal
responsiveness with near maximal responses at IIDs on either side. A final 18%
of AI neurons were classified as insensitive to IIDs. The proportions of
neurons exhibiting the various types of sensitivity corresponded closely to
the proportions found to exhibit corresponding types of azimuthal sensitivity
in a previous study. 3. There was a strong correlation between a neuron's
binaural interaction characteristics and the form of its IID-sensitivity
function. Thus, neurons excited by monaural stimulation of only one ear but
with either inhibitory, facilitatory, or mixed facilitatory/inhibitory effects
of stimulation of the other ear had predominantly contra-max IID sensitivity
(if contralateral monaural stimulation was excitatory) or ipsi-max sensitivity
(if ipsilateral monaural stimulation was excitatory). Neurons driven weakly or
not at all by monaural stimulation but facilitated binaurally almost all
exhibited near-zero-max IID sensitivity. The exception to this tight
association between binaural input and IID sensitivity was provided by neurons
excited by monaural stimulation of either ear (EE neurons). Although EE
neurons have frequently been considered to be insensitive to IIDs, our data
were in agreement with two recent reports indicating that they can exhibit
various forms of IID sensitivity: only 23 of 75 EE neurons were classified as
insensitive and the remainder exhibited diverse types of sensitivity. 4. IID
sensitivity was examined at two or more intensities (3-5 in most cases) for 84
neurons. The form of the IID-sensitivity function (defined in terms of both
shape and position along the IID axis) was invariant with changes in ABI for
only a small proportion of IID-sensitive neurons (approximately 15% if a
strict criterion of invariance was employed), and for many of these neurons
the spike counts associated with a given IID varied with ABI, particularly at
near-threshold levels. When the patterns of variation in the form of IID
sensitivity produced by changes in ABI were classified in a manner equivalent
to that used previously to classify the effects of intensity on azimuthal
sensitivity, there was a close correspondence between the effects of intensity
on corresponding types of azimuthal and IID sensitivity. 5. In agreement with
the report of Semple and Kitzes (1993a,b), the responses of a significant
proportion of high-frequency AI neurons (30-40%) varied non-monotonically with
both IID and ABI such that a restricted focus of maximal responsiveness was
associated with a particular combination of limited IID and ABI ranges. 6. The
close correspondence between the types of IID and azimuthal sensitivity and
the effects of variation in intensity on the two types of sensitivity supports
the view that the azimuthal sensitivity of high-frequency AI neurons is shaped
by their IID sensitivity. The fact that IID sensitivity is invariant with
changes in stimulus intensity (at suprathreshold levels) for only a small
proportion of AI neurons has implications for the manner in which IID (and
azimuthal location) are represented by populations of neurons in AI. If IID is
represented by the distributed pattern of activity across a population of
neurons, then the pattern of activity associated with a given IID would vary
with intensity unless the population were restricted to the small proportion
of neurons with intensity-invariant IID sensitivity. 7. Comparison of the
types of IID (and azimuthal) sensitivity exhibited by high-frequency AI
neurons with those seen in the inferior colliculus, and of the binaural
interaction characteristics of high-frequency neurons at these two levels,
indicates far greater diversity in cortex. Significant transformations in the
binaural properties of high-frequency neurons occur in the projections from
the midbrain to the cortex and/or as a consequence of mechanisms intrinsic to
the cortex.
Received 13 June 1994; accepted in final form 2 August 1995.
APS Manuscript Number J338-4.
Article publication pending J. Neurophysiol.
ISSN 1080-4757 Copyright 1995 The American Physiological Society.
Published in APStracts on 24 August 1995.