Jc. Clarey et al., FUNCTIONAL-ORGANIZATION OF SOUND DIRECTION AND SOUND PRESSURE LEVEL IN PRIMARY AUDITORY-CORTEX OF THE CAT, Journal of neurophysiology, 72(5), 1994, pp. 2383-2405
1. The functional organization of neuronal tuning to the azimuthal loc
ation and sound pressure level (SPL) of noise bursts was studied in hi
gh-frequency primary auditory cortex (AI) of barbiturate-anesthetized
cats. Three data collection strategies were used to map neural respons
es: 1) electrode penetrations oriented normal to the cortical surface
provided information on the radial organization of neurons' responses;
2) neurons' responses were examined at a few points in the middle cor
tical layers in multiple normal penetrations across AI to produce fine
-grain maps of azimuth and level selectivity; and 3) electrode penetra
tions oriented tangential to the cortical surface provided information
on neurons' responses along the isofrequency dimension. 2. An azimuth
-level data set was obtained for each single- or multiple- (multi-) un
it recording; this consisted of responses to noise bursts at five SPLs
(0-80 dB in 20-dB steps) from seven azimuthal locations in the fronta
l hemifield (-90 to +90 degrees in 30 degrees steps; 0 degrees elevati
on). An azimuth function was derived from these data by averaging resp
onse magnitude over all SPLs at each azimuth tested. A preferred azimu
th range (PAR; range of azimuths over which the response was greater t
han or equal to 75% of maximum) was calculated from the azimuth functi
on and provided a level-independent measure of azimuth selectivity. Ea
ch PAR was assigned to one of four azimuth preference categories (cont
ralateral-, midline-, ipsilateral-preferring, or broad/multipeaked) ac
cording to its location and extent. A level function obtained from the
data set (responsiveness averaged over all azimuths) was classified a
s monotonic if it showed a decrease of less than or equal to 25% (rela
tive to maximum) at the highest SPL tested (usually 80 dB), and nonmon
otonic if it showed a decrease of >25%. The percentage reduction in re
sponsiveness, relative to maximum, at the highest lever tested (termed
nonmonotonic strength) and the preferred level range (PLR; range of S
PLs over which responsiveness was greater than or equal to 75% of maxi
mum) of each response was also determined. 3. Normal penetrations typi
cally showed a predominance of one azimuth preference category and/or
level function type. The majority of penetrations (26/36: 72.2%) showe
d statistically significant azimuth preference homogeneity, and approx
imately one-half (17/36: 47.2%) showed significant level function type
homogeneity. Over one-third (13/36) showed significant homogeneity fo
r both azimuth preference and level function type. 4. Mapping experime
nts (n = 4) sampled the azimuth and level response functions at two or
more depths in closely spaced normal penetrations that covered severa
l square millimeters of AI. Cortical sites containing neurons with the
same azimuth preference and/or level function type were grouped toget
her. The most extensive map revealed a contralateral-preferring band o
f cortex that was elongated parallel to the frequency gradient and cov
ered a frequency range from 8 to 20 kHz. Immediately ventral to this w
as a midline-preferring region that covered the same frequency range.
5. Azimuth and level response functions were obtained at closely space
d sites along 20 tangential penetrations oriented parallel to isofrequ
ency contours, and most penetrations showed sequential responses that
exhibited similar azimuth preferences and/or level function types. The
majority (15/20: 75%) of penetrations showed significant clustering o
f neurons with the same azimuth preference. There were often multiple,
discontinuous groups of neurons with similar azimuth preferences (usu
ally contralateral) along isofrequency bands. Systematic shifts in the
azimuth preference of successive neurons or groups of neurons were no
t commonly seen. Over one-half (12/20: 60%) the penetrations showed si
gnificant clustering of monotonic and nonmonotonic response types. 6.
The organization of neurons with differing degrees of azimuth sensitiv
ity, as measured by azimuth function modulation, was also examined. Ne
urons' responses were categorized as either high directional (HD; grea
ter than or equal to 75% modulation) or low directional (LD; <75% modu
lation). Only a minority of normal and tangential penetrations showed
significant homogeneity or clustering of neurons with HD or LD respons
es. 7. In an attempt to validate the use of the multiunit technique to
study functional organization, comparisons of the azimuth and level r
esponse properties of single and multiunits were made. This was achiev
ed by 1) comparing the responses of multi/single-unit pairs (n = 114)
recorded at the same location and single-unit pairs (n = 16) recorded
at the same location; and 2) comparing the distribution of responses f
alling into the different azimuth and level categories from a large mu
ltiunit sample and a comparable single-unit sample collected during th
ese experiments. Multi- and single-unit recordings showed differences
in nonmonotonic strength and azimuth function modulation. However, mea
sures of azimuth preference and selectivity (best azimuth) or level se
lectivity (best SPL) that were derived from the range of maximum respo
nsiveness of the azimuth or level function showed significant correlat
ions.