ORGANIZATION OF AUDITORY

CORTEX
Once auditory information has ascended
through subcortical paths, it arrives at
auditory cortex, just beneath the Sylvian
fissure in the temporal lobe
The layout of auditory cortex has been studied
in other mammals including primates through
single-cell recording
NEUROIMAGING STUDIES
Neuroimaging studies in humans have also
addressed the organization of auditory cortex
Although studying auditory perception in a
neuroimaging environment poses some
challenges
For example, the machinery required to conduct
fMRI studies can generate a lot of noise. In
addition, it can be difficult to create naturalistic
sounds (e.g. sounds perceived as coming from
various distances) within the constraints of a
magnetic resonance imaging system.

DIVISION OF AUDITORY CORTEX
Auditory cortex can be subdivided into a few
regions,
Core ---it can be further subdivided into areas A1
(primary auditory cortex) and regions anterior to
A1, referred to as the rostral and rostrotemporal
fields. The core region receives the input from
medial geniculate nucleus
belt (which surrounds the core)---belt region
receives most of input from core
parabelt (which surrounds the belt)---receives
input from the belt
There appears to be a rough hierarchy
extending from the medial geniculate to the
core, then to the belt, then to the parabelt.
However the belt and parabelt regions also
receive some direct input from the medial
geniculate, so the hierarchy is not completely
rigid
All three of the areas within the core contain
tonotopic maps.
Figure illustrates a tonotopic map within area
A1.
Cells that are maximally responsive to same
frequency are localized in similar bands or
regions
Cells maximally responsive to lower
frequencies are located more rostrally
Whereas those responsive to higher
frequencies are located more caudally

TONOTOPIC MAP
A tonotopic map is somewhat analogous to retinotopic
map in the visual cortex, in the sense that they both map
out certain features of sensory world in a systematic way.
However whereas a retinotopic map is essentially a map of
space (that is a map of retina) the tonotopic map is not a
map of space. Rather it is a map of sound frequencies.
Frequency of sound
The frequency of sound refers to how fast sound waves
oscillate, and roughly corresponds to our perceptual
experience of pitch;
high frequency sounds are perceived as high pitches
low frequency as low pitches
In primary auditory cortex each cell responds best to a
certain frequency of sound, and neighboring cells
respond to similar frequencies. In a sense the
tonotopic map is map of cochlea because the cochlear
cells are also organized by frequency.
Cells in a tonotopic map can be thought of as having
receptive fields, but these receptive fields are not
spetialy defined as in visual system
Rather individual cells have preferred sound
frequencies such that only frequencies within a
particular range will excite the cell; other frequencies
will produce no response from the cell.
TUNING CURVE
For each cell, we can create a graph that shows the
cells sensitivity across different sound frequencies.
This graph is often referred to as the cells tuning
curve.
The curve represents the sound intensity that is
needed to provoke a response from each cell, as a
function of sound frequency
The low point on the curve indicates the sound
frequency to which the cell is most responsive. Cell A is
more responsive to lower sound frequencies than cell
B.
Cell A is more responsive to lower sound
frequencies than cell B.
TYPES OF TUNING CURVES
Cells in the different parts of the auditory cortex have different
kinds of tuning curves.
Within area A1, cells are sharply tuned to specific frequencies. In A1
cells also tend to respond best to pure tones (tones composed of
only one frequency) whereas in the surrounding regions cells tend
to respond best to more complex stimuli that incorporate several
individual sound frequencies
In the belt and parabelt regions, cells are more broadly tuned,
responding to a variety of stimulus frequencies. If the belt and
parabelt areas are damaged in monkeys while the core is left intact,
the monkeys are able to detect individual sounds, but they have
difficulty recognizing complex patterns, such as tone sequences.
Other studies in monkeys have shown that cells within lateral belt
area are especially responsive to the sounds of monkey
vocalizations. Thus as in the visual system, there is a hierarchy in
which early cortical stages seems to represent simpler features,
whereas later cortical stages represents more complex
combinations of features.
In humans lateral belt and parabelt regions are thought
to correspond to the planum temporale, an anatomical
region that is known to be especially important in
speech perception.
The planum temporale on the left side of the brain is
activated by the speech sounds, while the region of the
brain in both the left and right hemispheres is also
activated in response to other complex auditory stimuli
like sound patterns, music and environmental sounds.
Thus this region seems crucial in auditory pattern
recognition.
One hypothesis about the organization of auditory cortex proposes
that spatial and nonspatial features of a stimulus are handled in
somewhat separate streams.
Information about spatial location is processed in more
caudal/posterior regions of the auditory cortex;
Whereas nonspatial features (such as sound patterns) are
represented in more rostral/anterior sectors of auditory cortex.
This organization is analogous to the what and where
segregation of function in the visual system. Some evidence for this
view is found in the anatomical projections of auditory cortex.
Caudal auditory cortex projects to areas known to be important in
spatial localization, such as the parietal cortex and frontal eye fields
Whereas rostral auditory cortex projects to association areas to the
temporal lobe and orbitofrontal regions.
In addition recordings from single cells in the
monkey offer some support for this notion.
Researchers recorded the activity of cells in the
lateral belt region of auditory cortex while
monkeys listened to communication calls from
other monkeys.
Cells in the anterior portion of the belt were
more responsive to the specific type of call,
Whereas cells in the posterior region of the belt
were more sensitive to the spatial location
Some imaging studies in humans have also provided evidence for
the idea of dissociation between auditory spatial localization versus
auditory object recognition.
For example one study found that auditory localization task
activated the inferior parietal lobe and middle and inferior frontal
gyrus,
whereas an auditory recognition task activated middle temporal
gyrus and the precuneus area.
Although not all imaging studies consistently support a clear
dissociation between spatial localization and object recognition.
Recent evidence combining fMRI and MEG results provides some
strong support for such a distinction. Participants heard a vowel
emanating from a specific location in space. Next they heard a
target that was either an identical item, a different vowel from
same location
Posterior regions of both Heschels gyrus and the
superior temporal gyrus responded more when the
items differed in spatial location than when they were
different vowels, suggesting that these regions are
sensitive to where an auditory item is located.
In contrast anterior region of both Heschels gyrus and
superior temporal gyrus responded more when the
items differed in the vowel presented rather than
spatial location, suggesting that these regions are more
sensitive to what an auditory item is.
These responses occur very quickly within 70
to 150 milliseconds of stimulus onset, with
information in the where pathway activated
approximately 30 milliseconds before that in
the what pathway.
The researchers suggested that perhaps
information from where pathway can be fed
to what pathway to help direct attention to
auditory objects that need to be identified