Cortical Mechanisms of

13 Sound Localization and

Plasticity in Primates
Gregg H. Recanzone

CONTENTS

I.
Introduction
II.
Sound Localization Cues and Perception
III.
The Primate Auditory Cortex
IV.Lesion Studies
V.Electrophysiological Studies
VI.Plasticity of Sound Localization
A. Auditory Cortical Plasticity
B. Cortical Plasticity and Sound Localization
VII. Summary
References

I. INTRODUCTION
Sound localization has been studied extensively for many decades at the perceptual
level, particularly in humans1–3 but also in non-human primates4–6 and a variety of
other species (e.g., see References 7 to 9). However, at the neurophysiological level,
particularly in the cerebral cortex, relatively few studies have been conducted in
either monkeys or humans. The non-human primate, particularly the macaque mon-
key, has several advantages as an animal model of human auditory processing in
studies correlating neurophysiological responses to behavior and perception. This is
particularly true for studies on auditory cortical function because (1) this species
has perceptual abilities similar to humans in a variety of auditory discrimination
tasks that are dependent on normal cortical activity (for example, sound localization);
(2) it is possible to routinely record single-cell responses in awake monkeys actively
participating in discrimination experiments; and, therefore, (3) it is possible to
directly relate neuronal activity at the level of the single neuron to the auditory
perceptions of the animal. This chapter will concentrate on comparing both the
perceptual abilities and the neurophysiological correlates of sound localization in
non-human primates to those in humans, with some references to other species when

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considering areas where little is currently known in primates. It will also explore
the more general phenomenon of plasticity of the auditory cortex with reference to
how changes in the neuronal representations of acoustic stimuli can be related to
changes in perception.

II. SOUND LOCALIZATION CUES AND PERCEPTION

Because the auditory receptors, the hair cells within the cochlea, contain no spatial
information individually, the spatial location of acoustic stimuli must be computed
by the nervous system. Three principal cues are used in this computation.2 The first
two are based on differences in the sound waves that reach the two ears, known as
binaural cues. Interaural intensity differences result when a sound source is laterally
placed and the shadowing of the sound waves by the head and torso decreases the
intensity of the sound at the far ear with respect to the intensity at the near ear. This
cue is most effective for the higher frequency components of the sound, generally
above 4 kHz. The time of arrival and the phase of the sounds will also differ between
the two ears as a function of the laterality of the stimulus. This cue, based on
interaural time differences, is most effective for the low-frequency components,
generally below 1 kHz. These two interaural cues are used for determining the sound
location along the horizontal plane (azimuth) but are essentially ineffective for
stimuli that vary in elevation. Elevation cues are believed to arise from the differences
in the filtering of the acoustic stimulus by the pinnae, head, and torso.10,11 These
filtering properties generate spectral peaks and notches, referred to as the head-
related transfer function (HRTF), with the frequency of these peaks and particularly
the notches varying systematically as a function of stimulus elevation. Thus, all three
localization cues are available only for stimuli with a broad spectral bandwidth.
A comparison of studies across different paradigms and acoustic stimuli indicate
that the sound localization ability of humans and macaque monkeys are similar. For
example, monkeys and humans localize broadband noise stimuli best and band-
passed stimuli less well, and exhibit the poorest localization performance for tonal
stimuli (e.g., see Recanzone et al.6,12). Monkeys and humans also need the high-
frequency spectral cues to localize sounds in elevation.4–6,12 These observations
indicate that monkeys and humans take advantage of the same localization cues and
that the two species presumably have similar neuronal mechanisms subserving their
sound localization ability.

III. THE PRIMATE AUDITORY CORTEX

While the sound localization ability of primates and other species has been well
documented, the neuronal mechanisms of sound location perception are poorly
understood. Information from both ears is processed early in the ascending auditory
pathway, and several brainstem and midbrain structures, particularly the superior
olivary complex, are believed to be integral in processing binaural cues (for a review,
see Phillips and Brugge13). However, lesion studies indicate that the auditory divi-
sions of the cerebral cortex are critical for the perception of acoustic space. The
auditory cortex of the primate is composed of multiple cortical fields organized into

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a “core” region including the primary auditory cortex (AI), a surrounding “belt”
region including the caudomedial field (CM) and caudolateral field (CL), and a more
lateral “parabelt” region. This delineation of cortical fields is based on histochemical
and cytoarchitectonic analysis, the cortico-cortical and thalamo-cortical anatomical
connectivity, reversals in the frequency response properties to tones and band-passed
noise, and differences in physiological responses such as frequency tuning, intensity
tuning, and latency.14–25
It is currently unclear how spatial information is processed throughout the
auditory cortex, although neurons located caudally and laterally in the belt areas
(e.g., area CL) have sharper spatial tuning than neurons located more rostrally.25
This is consistent with the notion that auditory cortical fields are better tuned for
spatial processing in the more caudal regions of auditory cortex (i.e., CL and CM)
than neurons in the more rostral regions (see Woods et al.26) which can potentially
better encode spectral or temporal features of sound stimuli (see Rauschecker27 and
Kass et al.28). This is similar to the dorsal “what” and ventral “where” processing
proposed in the visual cortex of both monkeys29 and humans.30

IV. LESION STUDIES

The results from lesion studies in primates present strong evidence that the auditory
cortex plays a pivotal role in sound localization; however, the effects of lesions
restricted to one or even a few of the primate auditory cortical areas have yet to be
tested. Large bilateral lesions that likely incorporated all auditory cortical areas
resulted in sound localization deficits in macaque monkeys.31 An earlier study in
the squirrel monkey also showed sound localization deficits following unilateral
lesions.32 In this study, the lesions incorporated only part of the primary auditory
cortex and included parts of the belt regions as well, although the extent of the
lesions with respect to the cortical fields described above was not defined. Similarly,
large bilateral lesions that incorporate most or all of auditory cortex were also shown
to give rise to localization deficits in ferrets.33 In cats, small lesions restricted to the
representation of a few frequencies in the primary auditory cortex result in sound
localization deficits restricted to the frequencies represented in the lesioned zone.34
Thus, in multiple non-human species, auditory cortical lesions result in pronounced
deficits in the perception of acoustic space.
In humans, similar results have been found, although the extent of the lesions
are less well documented and are inconsistent across patients and studies. An early
study documented sound localization ability in normal subjects and patients with a
variety of cortical and subcortical lesions.35 This study found that patients with
temporal lobe lesions, presumably incorporating part or all of auditory cortex,
showed the most consistent and profound localization deficits in contralateral acous-
tic space. Patients with lesions in other, nonauditory, cortical areas generally had no
sound localization deficits. These findings were extended more recently in patients
who had undergone a hemispherectomy between the ages of 8 and 16 years and
were tested 6 months to 25 years after surgery.36 These patients had a deficit in
localizing sounds in contralateral space, but not in ipsilateral space. In a more recent
study, patients with partial auditory cortical lesions were examined with respect to

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sound localization as well as auditory recognition.37 In this study, the two patients
with more rostral auditory cortical lesions showed normal sound localization ability
but were impaired in their ability to recognize environmental sounds. In contrast,
the two patients with more caudal lesions were impaired in their ability to localize
both static and moving sound stimuli but were able to recognize environmental
sounds. These results indicate that different auditory cortical areas subserve these
two auditory discrimination processes.
Taken together, these data show that the auditory cortex plays an essential role
in sound location perception. The experiments in cats indicate that AI is essential
for this discrimination,34 which is consistent with the notion that auditory information
is first processed in AI and then spatial information is further processed in other,
more caudal, auditory cortical areas. More recent studies in humans show that the
caudal regions of auditory cortex are critical for this perception, consistent with the
anatomical and electrophysiological evidence (see below) observed in the macaque
monkey. Thus, it is likely that the macaque monkey will serve as an invaluable
animal model in studies of human sound localization ability.

V. ELECTROPHYSIOLOGICAL STUDIES
Experiments attempting to elucidate the cortical representation of acoustic space
have concentrated mainly on anesthetized preparations, particularly in cats.38–43
Fewer studies have investigated the spatial tuning properties of auditory cortical
neurons in the awake monkey.6,44,45 The results of these studies are converging on
the notion that the acoustic space in auditory cortex does not have a spatially
topographic organization. This is contrary to experimental evidence for spatially
topographic representations of acoustic space in noncortical areas, the best example
being the midbrain of the barn owl.46–48 Spatially topographic representations are
also seen in the superior colliculus of cats,49 ferrets,50 and monkeys.51,52 Thus,
topographical spatial representations, or “maps,” can be constructed in the brain but
are not constructed at the level of the auditory cortex.
A more recent study has taken advantage of the awake, behaving primate to
investigate how spatial information is processed in the macaque auditory cortex.6 In
this study, the activity of single neurons in the AI and CM were recorded while
monkeys performed a simple sound localization task. A restricted set of two different
stimuli was presented to all recorded neurons: a tone that was near the characteristic
frequency (the frequency that the neuron responded to at the lowest intensity) and
either a band-passed noise that contained the characteristic frequency within its
spectrum or a broadband noise. Representative examples of neuronal responses are
shown in Figures 13.1A and B for stimulus locations presented directly ahead and
at 15- and 30-degree eccentricity along both cardinal and oblique axes. Most neurons
in both AI and CM were responsive to all stimulus locations, although neurons in
CM generally had sharper spatial tuning compared to neurons in AI. The spatial
tuning of both AI and CM neurons also showed the same stimulus dependence as
the sound localization ability in the same monkeys. The spatial tuning of individual
neurons in azimuth was weaker for tone stimuli compared to noise stimuli, consistent
with the better localization performance for noise stimuli compared to tone stimuli

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 A. AI Neuron B. CM Neuron

C. Behavior D. Single Neuron E. Population

Prediction Predictions
15

predicted / measured
spikes / stimulus

1.0
4
Performance

0.5
2

0 0
-30 0 30 -30 0 30 0
Azimuth (deg) Azimuth (deg) AI CM

FIGURE 13.1 Responses from representative single neurons in the AI (A) and CM (B) to
noise stimuli presented from 17 different locations in frontal space. The center PSTH shows
the response of the neuron for stimuli presented directly in front of the monkey; the other
PSTHs are positioned at 15- and 30-degree eccentricity along both cardinal and oblique
axes. (C) Representative psychometric function showing localization in azimuth to a band-
passed stimulus during a go/no-go paradigm. Threshold was taken at the point where the
monkey detected this change at a 50% performance level (dashed lines). (D) Mean and
standard deviation of activity of a single neuron as a function of stimulus eccentricity in
azimuth. The neuronal threshold was taken as the eccentricity where the response differed
from the response to the center location by one standard deviation (dashed line). (E) Pooled
responses across neurons comparing the threshold predicted by the neuronal response to
the behaviorally measured threshold across all stimulus types (tones and noise) in both
azimuth and elevation. Only neurons that had statistically significant spatial tuning are
shown. (Adapted from Reference 6.)

measured behaviorally (see above). Also, very few neurons were spatially tuned in
elevation for tone stimuli or band-passed noise stimuli that contained only low
frequencies, again consistent with the poor sound localization acuity in elevation for
these stimuli.
Further analysis indicated that the population of neurons in CM, but not AI, had
response properties that were consistent with the sound localization acuity for each
animal. The eccentricity of a stimulus necessary for the neural response to be statistically
significantly different from the response when the stimulus was directly in front of the
monkey was calculated for each neuron (Figure 13.1D). This distance was then

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compared to the distance necessary for the monkey to correctly detect a change in
location on 50% of the trials (Figure 13.1C). Overall, neurons in CM more accurately
predicted the sound localization performance than neurons in AI. All neurons recorded
in sessions using the same tone, band-passed, and/or broadband stimulus were then
pooled and the analysis was repeated. The results (Figure 13.1E) showed that the pooled
neuronal responses in CM were not significantly different than the behavioral measures,
thus demonstrating that the firing rates of populations of cells in this cortical area, but
not in AI, contain sufficient information to account for the monkey’s behavior.
Less is currently known about the physiology of the human cerebral cortex
underlying sound localization. Although functional imaging studies have provided
valuable information about visual and somatosensory processing in humans, the
auditory system has not been as extensively studied. This is undoubtedly due to
the technical difficulties of overcoming the loud sounds (and constrained space)
generated by the functional magnetic resonance imaging (fMRI) technique. How-
ever, several imaging studies have shown that regions of the auditory cortex are
activated under different listening conditions, and a few studies have investigated
sound localization in particular. Results from positron emission tomography indi-
cate that the core and belt auditory areas generally do not show different levels
of activation during sound localization tasks compared to passive listening to the
same sounds (e.g., see Bushara et al.53 and Weeks et al.54,55). However, there is
significant activation in the inferior parietal lobule during sound localization tasks,
with greater activation in the right hemisphere compared to the left.54 Activation
of the primary auditory cortex during active sound localization was revealed using
magnetoencephalography, and again activation in the right hemisphere was greater
than that measured in the left hemisphere.56 This observation is consistent with
some previous reports of greater deficits in sound localization following right
hemisphere lesions compared to left hemisphere lesions in humans (e.g., see
Haeske-Dewick et al.57), although other studies have reported no hemisphere
difference (e.g., see Sanachez-Longo and Forster35 and Poirier et al.58). Direct
comparisons of deficits following left or right hemisphere lesions have not yet
been performed on macaque monkeys, although no obvious differences were
observed in squirrel monkeys.32 A hemispheric dominance or lack thereof has yet
to be definitively demonstrated in any non-human species.

VI. PLASTICITY OF SOUND LOCALIZATION

A. AUDITORY CORTICAL PLASTICITY
Several studies have shown that cortical activity can be altered in adults following
peripheral or central lesions, behavioral training, or manipulation of neuromodula-
tors.59–62 In the auditory system, early studies showed that classical conditioning could
induce changes in the responses of cortical neurons.63,64 These changes can occur
within only tens or scores of trials and can be specific to the paired frequency,65
suggesting that such a mechanism could influence a widespread change in the tono-
topic organization of auditory cortical fields.66 Such changes in the tonotopic repre-
sentation of the cochlea have been observed in AI following restricted cortical

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lesions67,68 and after pairing acoustic stimuli with basal forebrain stimulation,69,70
which can also alter temporal processing of auditory cortical neurons.70,71 The ability
of central neural structures to alter their response properties is consistent with the
ability to acquire new skills and behaviors, as well as the ability to adapt to changes
in the animal’s physical condition or in the environment.59,72
Plasticity of the primary auditory cortex has been directly related to changes
in perception in owl monkeys.73 Monkeys were trained to perform a frequency
discrimination task using the same base frequency on consecutive days. Through-
out several weeks of training, a steady improvement in performance at this task
was observed, as reflected in progressively decreasing thresholds measured during
each session (Figure 13.2A). Following this training, the animals were anesthe-
tized, and the responses of neurons throughout AI were investigated. Figures 13.2B
and C show the AI isofrequency bands encompassing neurons that have their
thresholds in the 2- to 4-kHz range from two different monkeys. Figure 13.2B
shows a normal monkey that was never trained and only a modest representation
(black areas) of this narrow frequency range (approximately 2500 to 2700 Hz).
In contrast, Figure 13.2C shows the representation of the same frequency range
in a monkey trained to discriminate frequencies within this narrow range. This
type of result was typical; monkeys trained at a particular frequency range had an
enlarged representation of those frequencies within AI. Further, the behaviorally
measured thresholds in the trained monkeys were significantly correlated with the
area of representation of that frequency range. These results suggest that changes
in perception, which commonly occur both during training and following periods
without training, are likely manifest as changes in cortical representations of the
stimulus features of the particular task.

FIGURE 13.2 Plasticity of AI following behavioral training. Panel A shows the progressive
improvement in performance with training. Each circle shows the threshold measured in the
same monkey trained to discriminate changes in the frequency of a 2.5-kHz standard tone.
Panel B shows the 2- to 4-kHz isofrequency band in the primary auditory cortex of a normal
monkey that had not been trained at any task. The black region shows recording locations
where the neurons responded to the narrow range of frequencies used in the training paradigm.
Panel C shows the same isofrequency region in a monkey trained to perform the discrimination
task. In this case, a much larger representation of the training frequencies was observed.
(Adapted from Reference 73.)

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B. CORTICAL PLASTICITY AND SOUND LOCALIZATION
Given the robust changes in cortical representations observed across sensory systems,
it seems likely that similar changes in the cortical representation of acoustic space
would take place during development and throughout adulthood. For example,
increasing head size during development can cause changes in the interaural time
and intensity cues. Studies in ferrets have shown that the spatial responses of auditory
neurons are influenced during development in a manner consistent with the changes
in both interaural and spectral cues.7,74
Studies on the ability of primates to adapt to changes in sound localization cues
have been largely confined to humans. The sound localization acuity of humans that
have been monaurally deaf since birth has been compared to normal humans, as
well as humans wearing sound-attenuating ear plugs in one ear.75 Two of five patients
that were monaurally congenitally deaf had profound sound localization deficits
similar to normal controls wearing a monaural earplug. Three patients, however,
showed localization performance that was equivalent in the two hemifields and
approached the level of the unplugged control subjects. In similar types of experi-
ments, it has been found that raising owls with a frequency-specific monaural plug
induces changes in the alignment of the auditory and visual receptive fields in the
midbrain,76 and ferrets raised with a monaural plug have sound localization ability
similar to unplugged controls.74 These results indicate that, at least in some instances,
subjects forced to rely almost exclusively on monaural (spectral) cues can learn to
accurately localize acoustic stimuli.
Humans can also adapt quickly to changes in sound localization cues. For example,
changing the interaural timing differences initially results in sound localization errors,
but subjects are able to compensate for these inappropriate cues within approximately
one week.77 Even more rapid adjustments have been observed when visual stimuli are
simultaneously presented with acoustic stimuli, but at a spatially disparate location.78
In these experiments, a visual stimulus is presented simultaneously with an acoustic
stimulus, but at a consistent spatial disparity (e.g., 4 degrees to the left). Exposure to
these disparities for as little as 20 minutes causes subjects to consistently localize stimuli
incorrectly by approximately the same amount as the previous visual–auditory disparity
(in this case, by 4 degrees to the left). This change in the internal representation of
acoustic space has recently been shown to also occur in macaque monkeys.79 Finally,
manipulation of the localization cues has been shown to influence localization percep-
tion both within tens of minutes80 and over the course of several days to weeks.81 For
example, Hofman et al.81 showed that changing the spectral cues produced an initial
deficit in localization in elevation, but most subjects slowly recovered to normal sound
localization ability within several weeks. Interestingly, once the normal spectral cues
were restored, the subjects were still able to localize sounds equally well, suggesting
the presence of two independent representations of acoustic space in these subjects.

VII. SUMMARY
Humans and macaque monkeys have similar sound localization abilities, indicating
that they likely share the same neuronal mechanisms to process acoustic space.

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Electrophysiological and neuroanatomical experiments in monkeys have led to the
hypothesis that spatial information is serially processed from the core areas through
the caudal belt areas toward the parietal lobe. Imaging studies in humans are largely
consistent with this idea, particularly with respect to the role of the parietal lobe in
processing spatial information. Studies in humans that manipulate either the binaural
or spectral cues, or both, have revealed some capacity to adapt to changing input
conditions, suggesting a parallel neuronal plasticity that underlies this adaptation.
Future studies in non-human primates may well reveal the neuronal locus of these
plastic changes, providing further evidence of how auditory spatial information is
processed in the primate cerebral cortex.

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