Sensory coding in supragranular cells of the vibrissal cortex in anesthetized and awake mice
Date
2017
Authors
Ranjbar-Salmloo, Yadollah
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Abstract
Sensory perception entails reliable representation of the
external stimuli as impulse activity of individual neurons (i.e.
spikes) and neuronal populations in the sensory area. An ongoing
challenge in neuroscience is to identify and characterize the
features of the stimuli which are relevant to a specific sensory
modality and neuronal strategies to effectively and efficiently
encode those features. It is widely hypothesized that the
neuronal populations employ “sparse coding” strategies to
optimize the stimulus representations with a low energetic cost
(i.e. low impulse activity). In the past two decades, a wealth of
experimental evidence has supported this hypothesis by showing
spatiotemporally sparse activity in sensory area. Despite
numerous studies, the extent of sparse coding and its underlying
mechanisms are not fully understood, especially in primary
vibrissal somatosensory cortex (vS1), which is a key model system
in sensory neuroscience. Importantly, it is not clear yet whether
sparse activation of supragranular vS1 is due to insufficient
synaptic input to the majority of the cells or the absence of
effective stimulus features.
In this thesis, first we asked how the choice of stimulus could
affect the degree of sparseness and/or the overall fraction of
the responsive vS1 neurons. We presented whisker deflections
spanning a broad range of intensities, including “standard
stimuli” and a high-velocity, “sharp” stimulus, which
simulated the fast slip events that occur during whisker mediated
object palpation. We used whole-cell and cell-attached recording
and calcium imaging to characterize the neuronal responses to
these stimuli. Consistent with previous literature, whole-cell
recording revealed a sparse response to the standard range of
velocities: although all recorded cells showed tuning to velocity
in their postsynaptic potentials, only a small fraction produced
stimulus-evoked spikes. In contrast, the sharp stimulus evoked
reliable spiking in a large fraction of regular spiking neurons
in the supragranular vS1. Spiking responses to the sharp stimulus
were binary and precisely timed, with minimum trial-to-trial
variability. Interestingly, we also observed that the sharp
stimulus produced a consistent and significant reduction in
action potential threshold.
In the second step we asked whether the stimulus dependent sparse
and dense activations we found in anesthetized condition would
generalize to the awake condition. We employed cell-attached
recordings in head-fixed awake mice to explore the degree of
sparseness in awake cortex. Although, stimuli delivered by a
piezo-electric actuator evoked significant response in a small
fraction of regular spiking supragranular neurons (16%-29%), we
observed that a majority of neurons (84%) were driven by manual
probing of whiskers. Our results demonstrate that despite sparse
activity, the majority of neurons in the superficial layers of
vS1 contribute to coding by representing a specific feature of
the tactile stimulus.
Thesis outline: Chapter 1 provides a review of the current
knowledge on sparse coding and an overview of the whisker-sensory
pathway. Chapter 2 represents our published results regarding
sparse and dense coding in vS1 of anesthetized mice
(Ranjbar-Slamloo and Arabzadeh 2017). Chapter 3 represents our
pending manuscript with results obtained with piezo and manual
stimulation in awake mice. Finally, in Chapter 4 we discuss and
conclude our findings in the context of the literature. The
appendix provides unpublished results related to Chapter 2. This
section is referenced in the final chapter for further
discussion.
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Keywords
sparse coding, post synaptic potentials, Fano factor, intrinsic-signal optical imaging, two-photon imaging, somatosensory cortex, AP threshold, whisker velocity, sparseness, whisker tracking, feature selectivity
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