Plasticity in the barrel cortex of the adult mouse: transient increase of GAD-immunoreactivity following sensory stimulation
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Barrel cortex
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findings reported by Berkowitz (1) -are extremely interesting, informative, and consistent with theoretical formulations and experimental results relating sensory deprivation to stimulus-seeking behaviors, (4, 5, 6, 7, 9, 10). It is the intention of this comment to suggest that Berkowitz's conclusion (1, p. 192) that The results as a whole do not support optimal-arousal theory; nor do they fall under the exclusive domain of any other extant theory of the phenomena is not entirely correct and that his remarks while the present data favor a deficit-oriented more than an optimal-arousal stance, unilateral application of the former to the infantile experience and/or sensory reinforcement areas is premature (1, p. 194) is, in fact, not consistent with the early deprivation and stimulation literature. In the references listed above a theoretical formulation has been offered that sensory deprivation during early development leads to stimulus-seeking behaviors that are related to the sensory system that has been deprived; further, that hyperactivity, hyperreactivity, and increased violent-aggressive behaviors commonly reported following maternal-social deprivation in mammals represent forms of stimulus-seeking behaviors which are attributable to somatosensory deprivation and not to deprivation of the other sensory systems. With respect to underlying mediating neural mechanisms, this writer has extended and emphasized the relevance of Cannon's Law of Denervation Supersensitivity (10) in accounting for the neural-behavior effects of early sensory deprivation, as initially observed by Riesen (14). Riesen (11, 12, 13, 14, 15) can be constructively consulted for a thorough analysis of the many aspects of early sensory deprivation upon neural-behavioral ontogeny. Additionally, Cannon and Rosenbleuth (2), Stavraky (17), and Sharpless (16) may be consulted for a thorough and systematic treatment of denervation supersensitivity phenomena. It is the contention of this writer that early experimental
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The normal development of sensory perception in mammals depends on appropriate sensory experience between birth and maturity. Numerous reports have shown that trimming some or all of the large mystacial vibrissa (whiskers) on one side of the face after birth has a detrimental effect on the maturation of cortical function. The objective of the present study was to understand the differences that occur after unilateral whisker trimming compared with those that occur after bilateral deprivation. Physiological deficits produced by bilateral trimming (BD) of all whiskers for 2 mo after birth were compared with the deficits produced by unilateral trimming (UD) for the same period of time using extracellular recording under urethan anesthesia from single cells in rat barrel cortex. Fast spiking (FSUs) and regular spiking (RSUs) units were separated and their properties compared in four subregions identified by histological reconstructions of the electrode penetrations, namely: layer IV barrel and septum, and layers II/III above a barrel and above a septum. UD upregulated responses in layer IV septa and in layers II/III above septa and perturbed the timing of responses to whisker stimuli. After BD, nearly all responses were decreased, and poststimulus latencies were increased. Circuit changes are proposed as an argument for how inputs arising from the spared whiskers project to the undeprived cortex and, via commissural fibers, could upregulate septal responses after UD. Following BD, more global neural deficits create a signature difference in the outcome of UD and BD in rat barrel cortex.
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Introduction: Barrel cortex of rats is a part of somatosensory cortex, which receives information from facial whiskers. Vibrisectomy by sensory deprivation leads to changes in the barrel cortex, which is known as experience dependent plasticity. On the other hand, Norepinephrine (NE) and locus coeruleus as the main source of NE, modulate response properties of cortical barrel neurons. In this study, the effect of NE depleted and sensory deprivation on induction of experience dependent plasticity was investigated. Methods: In this study sixty Wistar rats (250±25g) were used. Rats were divided into four groups: 1.Control group 2. NE depleted group (Norepinephrine was selectively depleted by IP injection of DSP4). 3. Sensory deprivation group (all whiskers except the whisker D2 on the left side were trimmed every other day). 4. NE depleted + sensory deprivation group. Excitatory (magnitude and latency) and inhibitatory (Conditioning Test Ratio, CTR index) receptive fields of barrel cortical neurons were assessed Using extracellular single unit recordings. Results: Sensory deprivation led to an increase both in the magnitude of response to principle whisker deflection (spared whisker) and in the CTR. In NE depleted + sensory deprivation group, the response magnitude and CTR index were the same as control group. Conclusion: The result showed that experience dependent plasticity has a facilitating effect on excitatory receptive field while decreasing the inhibitory circuits in the brain. When NE content of the brain was depleted before sensory deprivation, these changes were not seen. We conclude that NE depletion inhibits the plastic changes in the response properties of neurons following sensory deprivation.
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Experience-dependent plasticity (EDP) is essential for anatomical and functional maturation of sensory circuits during development. Although the principal synaptic and circuit mechanisms of EDP are increasingly well studied experimentally and computationally, its molecular mechanisms remain largely elusive. EDP can be readily studied in the rodent barrel cortex, where each "barrel column" preferentially represents deflections of its own principal whisker. Depriving select whiskers while sparing their neighbours introduces competition between barrel columns, ultimately leading to weakening of intracortical, translaminar (i.e., cortical layer (L)4-to-L2/3) feed-forward excitatory projections in the deprived columns. The same synapses are potentiated in the neighbouring spared columns. These experience-dependent alterations of synaptic strength are thought to underlie somatosensory map plasticity. We used RNA sequencing in this model system to uncover cortical-column and -layer specific changes on the transcriptome level that are induced by altered sensory experience. Column- and layer-specific barrel cortical tissues were collected from juvenile mice with all whiskers intact and mice that received 11-12 days of long whisker (C-row) deprivation before high-quality RNA was purified and sequenced. The current dataset entails an average of 50 million paired-end reads per sample, 75 base pairs in length. On average, 90.15% of reads could be uniquely mapped to the mm10 reference mouse genome. The current data reveal the transcriptional changes in gene expression in the barrel cortex upon altered sensory experience in juvenile mice and will help to molecularly map the mechanisms of cortical plasticity.
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Abstract We examined whether sensory deprivation during formation of the cortical circuitry influences the pattern of intracortical single‐cell connections in rat barrel cortex. Excitatory postsynaptic potentials (EPSPs) from layer 2/3 (L2/3) pyramidal neurons were recorded in vitro using patch‐clamp techniques. In order to evoke EPSPs, presynaptic neurons were stimulated by photolytically applied glutamate, thus generating action potentials. Synaptic connections between the stimulated and the recorded neuron were identified by the occurrence of PSPs following photostimulation. Sensory deprivation changed the pattern of projections from L4 and L2/3 neurons to L2/3 pyramidal cells. In slices of non‐deprived rats 86% of the total presynaptic neurons were located in the first and only 10% in the second barrel column. Deprivation changed these values to 67% and 26%, respectively. Therefore, the probability of presynaptic cells projecting to L2/3 neurons was shifted from adjacent to more remote barrel columns. These results indicate that deprivation of sensory input influences the pattern of intracortical connections.
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The capacity of adult barrel cortex to show experience-dependent plasticity after early restricted neonatal sensory deprivation was analyzed in barrel field cortex neurons. Selective sensory deprivation was induced by trimming two whiskers from postnatal day 0 (P0) to P21, namely, the principal D2 whisker plus one adjacent surround whisker (D3). At maturity (P90), responses of supragranular (layer II/III) and barrel (layer IV) neurons, all located in the D2 barrel column, were analyzed for modified responses to the deprived principal whisker (D2) and the nondeprived (D1) and deprived (D3) adjacent surround whiskers. For supragranular neurons, the responses to both principal and surround whiskers were reduced at maturity, whereas the barrel neurons showed mildly elevated responses to the principal whisker but a reduced response to the deprived surround whisker. In normal adult rats, trimming all but the principal D2 whisker and an adjacent D3 whisker for 3 d (whisker pairing) produced the expected bias: elevated responses from the intact D3 compared with the cut D1 whisker in both barrel and supragranular neurons. When the neonatally deprived D2 and D3 whiskers were paired at maturity, a similar D3/D1 bias was generated in barrel neurons, but no bias occurred in supragranular neuron responses. Pairing the maintained D1 and deprived D2 whiskers produced a much greater bias toward D1 compared with the deprived D3 whisker in barrel neurons than in supragranular neurons. There were minimal effects on response latencies in layer IV under any of the experimental conditions. These findings indicate that a restricted period of sensory deprivation in early postnatal life (1) impairs intracortical relay of deprived inputs from layer IV to layer II/III in barrel cortex at maturity and (2) degrades receptive field plasticity of the supragranular layer cells but not the thalamic-recipient barrel neurons.
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Each cerebral hemisphere processes sensory input from both sides of the body, but the impact of this convergence on shaping and modifying receptive field properties remains controversial. Here we investigated the effect of chronic deprivation of ipsilateral sensory whiskers on receptive field plasticity in primary somatosensory cortex. In the absence of ipsilateral whiskers, cortical receptive fields were significantly larger than control after 1 week. Removal of all but a single whisker from one side of the face [single-whisker experience (SWE)] has been shown to result in the expansion of the cortical area responding to the spared whisker. We compared the effects of SWE in the presence (SWE-unilateral) and absence (SWE-bilateral) of ipsilateral whiskers. SWE-bilateral deprivation results in a significant increase in neuronal responses to spared whisker stimulation both in its cognate barrel column and in adjacent, surrounding barrel columns compared with control and SWE-unilateral deprived animals. Surround receptive fields in deprived columns were maintained in SWE-bilateral treated animals but depressed in SWE-unilateral animals. The increase in spared whisker responses was progressive with longer deprivation periods. These data show that ipsilateral whiskers can constrain receptive field size in the barrel cortex.
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