Talk abstract:
Specializations in the Auditory Midbrain
for Detecting
Temporal Parameters of Biologically Important Sounds
John H. Casseday
Department of Psychology
University of Washington
casseday@u.washington.edu
The auditory midbrain, or inferior colliculus (IC), is an
obligatory waystation for >90% of the neurons of the ascending
auditory pathways. Thus, nearly all auditory information destined
for the thalamus and cortex must pass through synapses in the
IC. Cells in the IC receive synaptic input from several different
auditory centers in the lower brainstem, each of which produces
its own transformation of auditory signals. Therefore, cells
in the IC integrate different sorts of auditory information,
and from many of them, new signal transformations emerge. Some
of the most biologically important of these emergent properties
are selective responses to time varying stimuli. These properties
emerge from the interaction of excitatory and inhibitory inputs,
temporally offset from one another. We have studied two kinds
of selectivity in the IC of the big brown bat-neural tuning
for sound duration and tuning for the rate of repetitive frequency
modulations. Duration tuning refers to the ability of some neurons
to respond to a narrow range of sound duration but not to shorter
or longer sounds. For example, a duration tuned neuron may give
a vigorous response to a sound 5 ms long but completely fail
to respond to sound <3 ms or >7 ms. This neuron has a best duration
of 5 ms; different duration tuned neurons have different best
durations. A fairly simple model, comprised of three inputs,
can account for duration tuning: 1) Leading inhibition--inhibitory
input has the shortest latency and arrives prior to the excitatory
input; it is a sustained response that lasts at least as long
as the duration of the sound. 2) Offset excitation--an excitatory
event is correlated with the offset of sound; its magnitude
is below the threshold for spike initiation. 3) Delayed excitation--excitatory
and transient input has a long latency relative to the onset
of sound; it is temporally locked to the onset of sound; it
arrives after the initial inhibitory input; its magnitude is
just at or below spike threshold. The neuron fires when the
two excitatory inputs coincide in time with one another. The
best duration of a neuron is determined by the time delay between
the onset of inhibitory input and the delayed excitation. Other
specialized neurons respond to repetitive frequency modulations
but do not respond to simple sounds such as pure tones or noises.
Moreover, these neurons are tuned for the rate of frequency
modulations. This sort of specialization can be accounted for
by a modification of the duration tuning model. The most important
modification is that the excitatory and inhibitory components
are linked to different frequency components of the sound. These
two types of neurons appear to be filters for the same range
of temporal parameters of sounds as encountered by the bat during
detection of prey or avoidance of predators.
Research supported by NIH grants DC-00287 and DC-00607.
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1998-1999
Mathematics in Biology