Barrel Cortex Research Articles

Correlation between Cortical State and Locus Coeruleus Activity: Implications for Sensory Coding in Rat Barrel Cortex.

Cortical state modulates the background activity of cortical neurons, and their evoked response to sensory stimulation. Multiple mechanisms are involved in switching between cortical states including various neuromodulatory systems. Locus Coeruleus (LC) is one of the major neuromodulatory nuclei in the brainstem with widespread projections throughout the brain and modulates the activity of cells and networks. Here, we quantified the link between the LC spontaneous activity, cortical state and sensory processing in the rat vibrissal somatosensory “barrel” cortex (BC). We simultaneously recorded unit activity from LC and BC along with prefrontal electroencephalogram (EEG) while presenting brief whisker deflections under urethane anesthesia. The ratio of low to high frequency components of EEG (referred to as the L/H ratio) was employed to identify cortical state. We found that the spontaneous activity of LC units exhibited a negative correlation with the L/H ratio. Cross-correlation analysis revealed that changes in LC firing preceded changes in the cortical state: the correlation of the LC firing profile with the L/H ratio was maximal at an average lag of −1.2 s. We further quantified BC neuronal responses to whisker stimulation during the synchronized and desynchronized states. In the desynchronized state, BC neurons showed lower stimulus detection threshold, higher response fidelity, and shorter response latency. The most prominent change was observed in the late phase of BC evoked activity (100–400 ms post stimulus onset): almost every BC unit exhibited a greater late response during the desynchronized state. Categorization of the BC evoked responses based on LC activity (into high and low LC discharge rates) resulted in highly similar response profiles compared to categorization based on the cortical state (low and high L/H ratios). These findings provide evidence for the involvement of the LC neuromodulatory system in desynchronization of cortical state and the consequent enhancement of sensory coding efficiency.

Variable Temporal Integration of Stimulus Patterns in the Mouse Barrel Cortex.

Making sense of the world requires distinguishing temporal patterns and sequences lasting hundreds of milliseconds or more. How cortical circuits integrate over time to represent specific sensory sequences remains elusive. Here we assessed whether neurons in the barrel cortex (BC) integrate information about temporal patterns of whisker movements. We performed cell-attached recordings in anesthetized mice while delivering whisker deflections at variable intervals and compared the information carried by neurons about the latest interstimulus interval (reflecting sensitivity to instantaneous frequency) and earlier intervals (reflecting integration over timescales up to several hundred milliseconds). Neurons carried more information about the latest interval than earlier ones. The amount of temporal integration varied with neuronal responsiveness and with the cortical depth of the recording site, that is, with laminar location. A subset of neurons in the upper layers displayed the strongest integration. Highly responsive neurons in the deeper layers encoded the latest interval but integrated particularly weakly. Under these conditions, BC neurons act primarily as encoders of current stimulation parameters; however, our results suggest that temporal integration over hundreds of milliseconds can emerge in some neurons within BC.

A novel turning behavior control method for rat-robot through the stimulation of ventral posteromedial thalamic nucleus

The concept of a rat-robot was initially introduced in 2002, bringing to the field, a novel area of research using modern research into neuroscience and robotics. This paper brings to the table, a study into the method best used for navigation systems in a rat-robot. Current research is epitomized by the use of reward-based spatial navigation, combining the concept of an induced reward sensation as well as a ‘virtual touch’ sensation to control the movement of the rat-robot. However, such methods are plagued by limitations affecting the success rate as well as preparation procedures which may have varying effects on different rats, even under similar conditions. Hence, this paper studies the stimulation of two different portions of the brain to induce a turning motion within the rat, namely the Ventral Posteromedial (VPM) thalamic nucleus as well as the Barrel-Field (BF) cortex and demonstrates the preferential usage of VPM as the choice use of navigational control in a rat-robot.

LFP-guided targeting of a cortical barrel column for in vivo two-photon calcium imaging.

Two-photon microscopy of bulk-loaded functional dyes is an outstanding physiological technique that enables simultaneous functional mapping of hundreds of brain cells in vivo at single-cell resolution. However, precise targeting of a specific cortical location is not easy due to its fine dimensionality. To enable precise targeting, intrinsic-signal optical imaging is often additionally performed. However, the intrinsic-signal optical imaging is not only time-consuming but also ineffective in ensuring precision. Here, we propose an alternative method for precise targeting based on local field potential (LFP) recording, a conventional electrophysiological method. The heart of this method lies in use of the same glass pipette to record LFPs and to eject calcium dye. After confirming the target area by LFP using a glass pipette, the calcium dye is ejected from the same pipette without a time delay or spatial adjustment. As a result, the calcium dye is loaded into the same ensemble of brain cells from which the LFP was obtained. As a validation of the proposed LFP-based method, we targeted and successfully loaded calcium dye into layer 2/3 of a mouse barrel column.

Efficient population coding of naturalistic whisker motion in the ventro-posterior medial thalamus based on precise spike timing

The rodent whisker-associated thalamic nucleus (VPM) contains a somatotopic map where whisker representation is divided into distinct neuronal sub-populations, called “barreloids”. Each barreloid projects to its associated cortical barrel column and so forms a gateway for incoming sensory stimuli to the barrel cortex. We aimed to determine how the population of neurons within one barreloid encodes naturalistic whisker motion. In rats, we recorded the extracellular activity of up to nine single neurons within a single barreloid, by implanting silicon probes parallel to the longitudinal axis of the barreloids. We found that play-back of texture-induced whisker motion evoked sparse responses, timed with millisecond precision. At the population level, there was synchronous activity: however, different subsets of neurons were synchronously active at different times. Mutual information between population responses and whisker motion increased near linearly with population size. When normalized to factor out firing rate differences, we found that texture was encoded with greater informational-efficiency than white noise. These results indicate that, within each VPM barreloid, there is a rich and efficient population code for naturalistic whisker motion based on precisely timed, population spike patterns.

Experience of a First, “Whisker-Dependent,” Skill Affects the Induction of c-Fos Expression in Somatosensory Cortex Barrel Field Neurons in Rats on Training to a Second Skill

We report here studies in which a controlled history of sequential learning of two skills was created with the aim of identifying the pattern of activation of neurons for the first skill on formation of the second. Animals were initially trained to an operant drinking behavior requiring use of the whiskers on the left or right side (a “whisker-dependent” skill), which was followed by training to a food-procuring pedal-pressing skill not requiring use of the whiskers. The results showed that training to a food-procuring skill induced c-Fos expression in a significantly larger number of neurons in the barrel field in animals previously trained to the operant drinking (whisker-dependent) skill than in the analogous area of control animals previously trained to a non-operant drinking skill. Our data suggest that activation of c-Fos expression on repeat training also occurred in those neurons which had already become specialized in relation to the first, whisker-dependent, skill.

Calretinin-positive L5a pyramidal neurons in the development of the paralemniscal pathway in the barrel cortex.

BackgroundThe rodent barrel cortex has been established as an ideal model for studying the development and plasticity of a neuronal circuit. The barrel cortex consists of barrel and septa columns, which receive various input signals through distinct pathways. The lemniscal pathway transmits whisker-specific signals to homologous barrel columns, and the paralemniscal pathway transmits multi-whisker signals to both barrel and septa columns. The integration of information from both lemniscal and paralemniscal pathways in the barrel cortex is critical for precise object recognition. As the main target of the posterior medial nucleus (POm) in the paralemniscal pathway, layer 5a (L5a) pyramidal neurons are involved in both barrel and septa circuits and are considered an important site of information integration. However, information on L5a neurons is very limited. This study aims to explore the cellular features of L5a neurons and to provide a morphological basis for studying their roles in the development of the paralemniscal pathway and in information integration.Results1. We found that the calcium-binding protein calretinin (CR) is dynamically expressed in L5a excitatory pyramidal neurons of the barrel cortex, and L5a neurons form a unique serrated pattern similar to the distributions of their presynaptic POm axon terminals.2. Infraorbital nerve transection disrupts this unique alignment, indicating that it is input dependent.3. The formation of the L5a neuronal alignment develops synchronously with barrels, which suggests that the lemniscal and paralemniscal pathways may interact with each other to regulate pattern formation and refinement in the barrel cortex.4. CR is specifically expressed in the paralemniscal pathway, and CR deletion disrupts the unique L5a neuronal pattern, which indicates that CR may be required for the development of the paralemniscal pathway.ConclusionsOur results demonstrate that L5a neurons form a unique, input-dependent serrated alignment during the development of cortical barrels and that CR may play an important role in the development of the paralemniscal pathway. Our data provide a morphological basis for studying the role of L5a pyramidal neurons in information integration within the lemniscal and paralemniscal pathways.

Quantitative activity-induced manganese-dependent MRI for characterizing cortical layers in the primary somatosensory cortex of the rat.

The ability of Mn(2+) to follow Ca(2+) pathways upon stimulation transform them into remarkable surrogate markers of neuronal activity using activity-induced manganese-dependent MRI (AIM-MRI). In the present study, a precise follow-up of physiological parameters during MnCl2 and mannitol infusions improved the reproducibility of AIM-MRI allowing in-depth evaluation of the technique. Pixel-by-pixel T1 data were investigated using histogram distributions in the barrel cortex (BC) and the thalamus before and after Mn(2+) infusion, after blood brain barrier opening and after BC activation. Mean BC T1 values dropped significantly upon trigeminal nerve (TGN) stimulation (-38%, P=0.02) in accordance with previous literature findings. T1 histogram distributions showed that 34% of T1s in the range 600-1500ms after Mn(2+)+mannitol infusions shifted to 50-350ms after TGN stimulation corresponding to a twofold increase of the percentage of pixels with the lowest T1s in BC. Moreover, T1 changes in response to stimulation increased significantly from superficial cortical layers (I-III) to deeper layers (V-VI). Cortical cytoarchitecture detection during a functional paradigm was performed extending the potential of AIM-MRI. Quantitative AIM-MRI could thus offer a means to interpret local neural activity across cortical layers while identification of the role of calcium dynamics in vivo during brain activation could play a key role in resolving neurovascular coupling mechanisms.

Interhemispheric plasticity protects the deafferented somatosensory cortex from functional takeover after nerve injury.

Functional changes across brain hemispheres have been reported after unilateral cortical or peripheral nerve injury. Interhemispheric callosal connections usually underlie this cortico-cortical plasticity. However, the effect of the altered callosal inputs on local cortical plasticity in the adult brain is not well studied. Ipsilateral functional magnetic resonance imaging (fMRI) activation has been reliably detected in the deafferented barrel cortex (BC) at 2 weeks after unilateral infraorbital denervation (IO) in adult rats. The ipsilateral fMRI signal relies on callosal-mediated interhemispheric plasticity. This form of interhemispheric plasticity provides a good chronic model to study the interaction between callosal inputs and local cortical plasticity. The receptive field of forepaw in the primary somatosensory cortex (S1), which is adjacent to the BC, was mapped with fMRI. The S1 receptive field expanded to take over a portion of the BC in 2 weeks after both ascending inputs and callosal inputs were removed in IO rats with ablated contralateral BC (IO+ablation). This expansion, estimated specifically by fMRI mapping, is significantly larger than what has been observed in the IO rats with intact callosal connectivity, as well as in the rats with sham surgery. This work indicates that altered callosal inputs prevent the functional takeover of the deafferented BC from adjacent cortices and may help preserve the functional identity of the BC.

Imprecise Whisker Map in the Neonatal Rat Barrel Cortex.

The somatosensory barrel cortex in rodents contains a topographic map of the facial whiskers where each cortical barrel is tuned to a corresponding whisker. However, exactly when this correspondence is established during development and how precise the functional topography of the whisker protomap is at birth, before the anatomical formation of barrels, are questions that remain unresolved. Here, using extracellular and whole-cell recordings from the barrel cortex of 0- to 7-day-old (P0-7; P0 = day of birth) rat pups in vivo, we report a low level of tuning to the principal whisker at P0-1, with multiple adjacent whiskers evoking large multi- and single-unit responses and excitatory postsynaptic currents in cortical neurons. Additionally, we found broad and largely overlapping projection fields (PFs) for neighboring whiskers in the barrel cortex at P0-1. Starting from P2-3, a segregated whisker map emerged, characterized by preferential single whisker tuning and segregated whisker PFs. These results indicate that the functional whisker protomap in the somatosensory cortex is imprecise at birth, that for 2-3 days after birth, whiskers compete for the cortical target territories, and that formation of a segregated functional whisker map coincides with emergence of the anatomical barrel map.

Connection Changes in Somatosensory Cortex Induced by Different Doses of Propofol

BackgroundThe mechanism by which general anesthetics, widely used in clinical practice for over 160 years, effects on sensory responsiveness has been unclear until now. In the present study, the authors sought to explore the effect of different doses of propofol on somatosensory cortex by whisker stimulation in rats.MethodsIn a fixed cage, rats were anesthetized with propofol 80 mg/kg intraperitoneally and then cathetered tail vein with 23-gauge metal needle connected with a pump. Two holes (2 mm diameter) were drilled and recording electrodes implantated in the primary somatosensory cortex barrel field (S1BF) and secondary somatosensory cortex (S2). The extracellular (20 rats) and intracellular (8 rats) recordings were used to test the neuron activity in both cortices at different doses of propofol (20, 40 and 80 mg/kg/h) through tail vein by pump. Meantime, vibrissal, olfactory, corneal responses (VOCR, sedation), and tail-pinch response (TRP, analgesia) were tested every 10 min during the doses of propofol 20, 40 and 80 mg/kg/h.ResultsVOCR and TRP were depressed by propofol in a dose-dependent manner. The amplitude by whisker stimulation in S1BF was stronger and the peak latency was shorter compared with that of in S2. The response latency of S1BF and S2 was increased by raising infusion rate of propofol with the response latency in S2 being longer than that in S1BF at the same doses of propofol. The cross-correlation between S1BF and S2 decreased as the propofol infusion rate increased. The input resistance was higher by increasing infusion rate of propofol.ConclusionThe sedation and analgesia effects of propofol were dose-dependent. Both the connectivity and instinctive oscillation between S1BF and S2 were proportionally modulated by the different doses of propofol.

Effects of General Anesthesia with Propofol on Thalamocortical Sensory Processing in Rats

The effects of anesthetics on the transmission and processing of sensory information within the thalamocortical pathway and the underlying mechanism are not fully understood. Using the extracellular recording technique, we investigated the changes of spontaneous and stimulation-evoked activities within and between the ventral posteromedial nucleus (VPM) and primary somatosensory cortex barrel field (S1BF) of the rat in vivo during propofol anesthesia. Spontaneous local field potentials, whiskers deflection–elicited somatosensory evoked potentials, and multi-unit activities in VPM/S1BF were assessed at different depths of propofol anesthesia. In VPM and S1BF, powers of spontaneous and stimulation-evoked activities, coupled with stimulation-evoked multi-unit, were decreased with increasing of propofol anesthesia. Cortical onset latency increased during intermediate/deep level propofol anesthesia, whereas thalamic onset latencies were not changed even at different depths of anesthesia. In addition, spontaneous and whisker deflectionevoked alpha oscillations were observed during propofol anesthesia, which is similar to sleep spindles, These data suggest that propofol affects processing of sensory information by 1) attenuating respective neuronal activities in VPM and S1BF, 2) delaying the ascending signal transmission from VPM to S1BF, and 3) inducing a natural-sleep type of anesthesia.

Distribution of Corticosteroid Receptors in Mature Oligodendrocytes and Oligodendrocyte Progenitors of the Adult Mouse Brain

The expression of glucocorticoid receptors (GRs) was investigated immunohistochemically in two different lineages of oligodendrocytes, using carbonic anhydrase (CA) II and neuron glial antigen (NG) 2 as markers of mature oligodendrocytes and oligodendrocyte progenitors, respectively. We focused on the gray matter regions, including CA1, CA3 and the dentate gyrus of the hippocampus, the primary somatosensory cortex barrel field and the basolateral amygdala, and the white matter regions, including the corpus callosum, external capsule and fimbria of the hippocampus. More than 80% of CAII-immunoreactive (IR) cells and more than 95% of NG2-IR cells expressed GRs in various regions of the brain. In contrast, neither CAII-IR cells nor NG2-IR cells expressed mineralocorticoid receptors (MRs) in the same regions. The intensity of GR expression was drastically reduced in CA II-IR cells and NG2-IR cells in the same regions in adrenalectomized mice. Finally, steroid receptor co-activator (SRC)-1 and p300, both of which are cofactors for GR, were expressed in the gray and white matter regions in NG2-IR cells, but not in CAII-IR cells. These results suggest that the expression of GRs in oligodendrocytes and their progenitor cells mediates several functions in vivo, including differentiation and myelination, as a major target of glucocorticoids and their cofactors.

Lhx2 regulates a cortex-specific mechanism for barrel formation

LIM homeodomain transcription factors are critical regulators of early development in multiple systems but have yet to be examined for a role in circuit formation. The LIM homeobox gene Lhx2 is expressed in cortical progenitors during development and also in the superficial layers of the neocortex in maturity. However, analysis of Lhx2 function at later stages of cortical development has been hampered by severe phenotypes associated with early loss of function. We identified a particular Cre-recombinase line that acts in the cortical primordium after its specification is complete, permitting an analysis of Lhx2 function in neocortical lamination, regionalization, and circuit formation by selective elimination of Lhx2 in the dorsal telencephalon. We report a profound disruption of cortical neuroanatomical and molecular features upon loss of Lhx2 in the cortex from embryonic day 11.5. A unique feature of cortical circuitry, the somatosensory barrels, is undetectable, and molecular patterning of cortical regions appears disrupted. Surprisingly, thalamocortical afferents innervate the mutant cortex with apparently normal regional specificity. Electrophysiological recordings reveal a loss of responses evoked by stimulation of individual whiskers, but responses to simultaneous stimulation of multiple whiskers were present, suggesting that thalamic afferents are unable to organize the neurocircuitry for barrel formation because of a cortex-specific requirement of Lhx2. We report that Lhx2 is required for the expression of transcription factor paired box gene 6, axon guidance molecule Ephrin A5, and the receptor NMDA receptor 1. These genes may mediate Lhx2 function in the formation of specialized neurocircuitry necessary for neocortical function.

Sizing up the columns

A new study shows that the number of neurons in cortical barrel columns varies markedly within individual rats.

Cellular organization of cortical barrel columns is whisker-specific

The cellular organization of the cortex is of fundamental importance for elucidating the structural principles that underlie its functions. It has been suggested that reconstructing the structure and synaptic wiring of the elementary functional building block of mammalian cortices, the cortical column, might suffice to reverse engineer and simulate the functions of entire cortices. In the vibrissal area of rodent somatosensory cortex, whisker-related "barrel" columns have been referred to as potential cytoarchitectonic equivalents of functional cortical columns. Here, we investigated the structural stereotypy of cortical barrel columns by measuring the 3D neuronal composition of the entire vibrissal area in rat somatosensory cortex and thalamus. We found that the number of neurons per cortical barrel column and thalamic "barreloid" varied substantially within individual animals, increasing by ∼2.5-fold from dorsal to ventral whiskers. As a result, the ratio between whisker-specific thalamic and cortical neurons was remarkably constant. Thus, we hypothesize that the cellular architecture of sensory cortices reflects the degree of similarity in sensory input and not columnar and/or cortical uniformity principles.

Phase-of-firing coding of dynamical whisker stimuli and the thalamocortical code in barrel cortex

In the phase-of-firing (PoF) code the information carried by spikes is boosted when their firing is measured against the angular phase of the local extracellular signal. This coding strategy has been verified so far in visual and auditory cortices of monkeys at low frequencies [1,2]. Here we present the first evidence of this code in whisker modality. In particular, we characterise the PoF encoding of the white-noise whisker deflections in the barrel cortex responses of urethane anaesthetized rats. A novel aspect of our results is identifying the high frequency components of the PoF responses at certain cortical layers, with implications regarding the thalamo-cortical code and cortical information processing. The results indicate that the amount of information encoded using a PoF code was on average 100% greater than a spike rate code using MUA (Figure ​(Figure1A),1A), and this was up to 250% in deeper channels of the cortical columns. Contrary to the previous findings [1,2], the extra information in PoF using LFP was peaked at very high frequencies (100 Hz). The effect extended to >200 Hz bands of LFP. When CSD was used for labelling spikes, the effect was more localized with respect to depth, and was peaked at different frequencies at different cortical layers (Figure ​(Figure1C).1C). The depth-frequency profile was consistent across the multi-channel electrode penetrations. Why the PoF information was maximal at such high-frequency bands? ( >100 Hz) We hypothesised that the high-frequency PoF components originated from the high-frequency components in the white-noise stimuli. The depth-frequency profile and CSD analysis of layers showed association of the high-frequency components of PoF with the cortical layers that are known to receive direct thalamo-cortical input from VPM. A similar pattern was previously suggested in [5]. The high-frequency components of CSD phase responses were weak or absent in layers that are not associated with such projections (Figure ​(Figure1C1C). Figure 1 A. Phase-of-firing information and and spike rate information averaged across channels. B. Depth-frequency profile of the CSD phase information across the cortical layers (for two electrodes in two animals). C. same as B, but normalised to reveal the ... We then examined whether the pure spiking activity contributes in the information manifested in the high frequency PoF components. The temporal precision of spikes was quantified using a novel measure, information in the phase of the band-bass filtered spike trains. In all depths, the spikes lacked any temporal-resolution larger than 30 Hz (Figure ​(Figure1D).1D). We conclude that the output of the local computational processes in a cortical column lacked the temporal precision that they received in input (i.e., which was represented by CSD). This suggests a transformation of the information encoding from a high-precision input code relayed from thalamus, with fast and temporally precise dynamics (see [4]), into an output spiking code with a lower temporal resolution. Nevertheless, information about the high-frequency and fast varying components of the stimulus may still be encoded within the low resolution and irregular cortical spikes. In fact previous studies have shown that a spike-count code in the cortex can encode such stimulus features [3]. The computational demands of the whisker system (such as fine texture discrimination, and perception of subtle vibrations in the air) require that the thalamus relays the high-frequency components of the whisker deflections into the cortical barrels, which are capable of performing more complex neural computation.

Barrelettes without Barrels in the American Water Shrew

Water shrews (Sorex palustris) depend heavily on their elaborate whiskers to navigate their environment and locate prey. They have small eyes and ears with correspondingly small optic and auditory nerves. Previous investigations have shown that water shrew neocortex is dominated by large representations of the whiskers in primary and secondary somatosensory cortex (S1 and S2). Flattened sections of juvenile cortex processed for cytochrome oxidase revealed clear borders of the whisker pad representation in S1, but no cortical barrels. We were therefore surprised to discover prominent barrelettes in brainstem of juvenile water shrews in the present investigation. These distinctive modules were found in the principal trigeminal nucleus (PrV), and in two of the three spinal trigeminal subnuclei (interpolaris – SpVi and caudalis – SpVc). Analysis of the shrew's whisker pad revealed the likely relationship between whiskers and barrelettes. Barrelettes persisted in adult water shrew PrV, but barrels were also absent from adult cortex. Thus in contrast to mice and rats, which have obvious barrels in primary somatosensory cortex and less clear barrelettes in the principal nucleus, water shrews have clear barrelettes in the brainstem and no barrels in the neocortex. These results highlight the diverse ways that similar mechanoreceptors can be represented in the central nervous systems of different species.

Glucose and lactate metabolism in the awake and stimulated rat: a 13C-NMR study

Glucose is the major energetic substrate for the brain but evidence has accumulated during the last 20 years that lactate produced by astrocytes could be an additional substrate for neurons. However, little information exists about this lactate shuttle in vivo in activated and awake animals. We designed an experiment in which the cortical barrel field (S1BF) was unilaterally activated during infusion of both glucose and lactate (alternatively labeled with 13C) in rats. At the end of stimulation (1 h) both S1BF areas were removed and analyzed by HR-MAS NMR spectroscopy to compare glucose and lactate metabolism in the activated area vs. the non-activated one. In combination with microwave irradiation HR-MAS spectroscopy is a powerful technical approach to study brain lactate metabolism in vivo. Using in vivo 14C-2-deoxyglucose and autoradiography we confirmed that whisker stimulation was effective since we observed a 40% increase in glucose uptake in the activated S1BF area compared to the ipsilateral one. We first determined that lactate observed on spectra of biopsies did not arise from post-mortem metabolism. 1H-NMR data indicated that during brain activation there was an average 2.4-fold increase in lactate content in the activated area. When [1-13C]glucose + lactate were infused 13C-NMR data showed an increase in 13C-labeled lactate during brain activation as well as an increase in lactate C3-specific enrichment. This result demonstrates that the increase in lactate observed on 1H-NMR spectra originates from newly synthesized lactate from the labeled precursor ([1-13C]glucose). It also shows that this additional lactate does not arise from an increase in blood lactate uptake since it would otherwise be unlabeled. These results are in favor of intracerebral lactate production during brain activation in vivo which could be a supplementary fuel for neurons.

Increased physical activity is not enough to recover astrocytic population from dark-rearing. Synergy with multisensory enrichment is required

Elimination of sensory inputs (deprivation) modifies the properties of the sensory cortex and serves as a model for studying plasticity during postnatal development. Many studies on the effects of deprivation have been performed in the visual cortex using dark-rearing as a visual deprivation model. It induces changes in all cellular and molecular components, including astrocytes, which play an important role in the development, maintenance, and plasticity of the cortex, mediated by cytokines which have been termed angioglioneurins. When one sense is deprived, a compensatory mechanism called cross-modal plasticity increases performance in the remaining senses. Environmental enrichment is so far the best-known method to compensate sensorial deprivation. The aim of this work is to study the effects of exercise alone, and of an enriched environment combined with exercise, on astroglial population in order to observe the effects of exercise by itself, or the potential synergistic effect during the rat visual system development. Pregnant Sprague-Dawley rats were raised in one of the following rearing conditions: in total darkness and enriched environment conditions with physical exercise, and in total darkness with voluntary physical exercise. Astrocytic density was estimated by immunohistochemistry for S-100β protein and quantifications were performed in layer IV. The somatosensorial cortex barrel field was also studied as control. Our main result shows that an enriched environment combined with voluntary physical exercise manages to reverse the negative effects induced by darkness over the astroglial population of both the visual and the somatosensory cortices. On the other hand, exercise alone only produces effects upon the astroglial population of the somatosensory cortex, and less so when combined with an enriched environment.