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Journal articles on the topic 'Rat Barrel Cortex Dynamics'

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Author: Grafiati

Published: 10 March 2023

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1

Lustig, Brian R., Robert M. Friedman, Jeremy E. Winberry, Ford F. Ebner, and Anna W. Roe. "Voltage-sensitive dye imaging reveals shifting spatiotemporal spread of whisker-induced activity in rat barrel cortex." Journal of Neurophysiology 109, no. 9 (May 1, 2013): 2382–92. http://dx.doi.org/10.1152/jn.00430.2012.

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In rats, navigating through an environment requires continuous information about objects near the head. Sensory information such as object location and surface texture are encoded by spike firing patterns of single neurons within rat barrel cortex. Although there are many studies using single-unit electrophysiology, much less is known regarding the spatiotemporal pattern of activity of populations of neurons in barrel cortex in response to whisker stimulation. To examine cortical response at the population level, we used voltage-sensitive dye (VSD) imaging to examine ensemble spatiotemporal dynamics of barrel cortex in response to stimulation of single or two adjacent whiskers in urethane-anesthetized rats. Single whisker stimulation produced a poststimulus fluorescence response peak within 12–16 ms in the barrel corresponding to the stimulated whisker (principal whisker). This fluorescence subsequently propagated throughout the barrel field, spreading anisotropically preferentially along a barrel row. After paired whisker stimulation, the VSD signal showed sublinear summation (less than the sum of 2 single whisker stimulations), consistent with previous electrophysiological and imaging studies. Surprisingly, we observed a spatial shift in the center of activation occurring over a 10- to 20-ms period with shift magnitudes of 1–2 barrels. This shift occurred predominantly in the posteromedial direction within the barrel field. Our data thus reveal previously unreported spatiotemporal patterns of barrel cortex activation. We suggest that this nontopographical shift is consistent with known functional and anatomic asymmetries in barrel cortex and that it may provide an important insight for understanding barrel field activation during whisking behavior.

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2

Lee, Soo-Hyun, and Daniel J. Simons. "Angular Tuning and Velocity Sensitivity in Different Neuron Classes Within Layer 4 of Rat Barrel Cortex." Journal of Neurophysiology 91, no. 1 (January 2004): 223–29. http://dx.doi.org/10.1152/jn.00541.2003.

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Local circuitry within layer IV whisker-related barrels is preferentially sensitive to thalamic population firing synchrony, and neurons respond most vigorously to stimuli, such as high-velocity whisker deflections, that evoke it. Field potential recordings suggest that thalamic barreloid neurons having similar angular preferences fire synchronously. To examine whether angular tuning of cortical neurons might also be affected by thalamic firing synchrony, we characterized responses of layer IV units to whisker deflections that varied in angular direction and velocity. Barrel regular-spike units (RSUs) became more tuned for deflection angle with slower whisker movements. Deflection amplitude had no affect. Barrel fast-spike units (FSUs) were poorly tuned for deflection angle, and their responses remained constant with different deflection velocity. The dependence of angular tuning on deflection velocity among barrel RSUs appears to reflect the same underlying response dynamics that determine their velocity sensitivity and receptive field focus. Unexpectedly, septal RSUs and FSUs are largely similar to their barrel counterparts despite available evidence suggesting that they receive different afferent inputs and are embedded within different local circuits.

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3

Iwasaki, Naoko, Akihiro Karashima, Yuichi Tamakawa, Norihiro Katayama, and Mitsuyuki Nakao. "Sleep EEG dynamics in rat barrel cortex associated with sensory deprivation." NeuroReport 15, no. 17 (December 2004): 2681–84. http://dx.doi.org/10.1097/00001756-200412030-00026.

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4

Sheth, Bhavin R., Christopher I. Moore, and Mriganka Sur. "Temporal Modulation of Spatial Borders in Rat Barrel Cortex." Journal of Neurophysiology 79, no. 1 (January 1, 1998): 464–70. http://dx.doi.org/10.1152/jn.1998.79.1.464.

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Sheth, Bhavin R., Christopher I. Moore, and Mriganka Sur. Temporal modulation of spatial borders in rat barrel cortex. J. Neurophysiol. 79: 464–470, 1998. We examined the effects of varying vibrissa stimulation frequency on intrinsic signal and neuronal responses in rat barrel cortex. Optical imaging of intrinsic signals demonstrated that the region of cortex activated by deflection of a single vibrissa at 1 Hz is more diffuse and more widespread than the territory activated at 5 or 10 Hz. With the use of two different paradigms, constant time of stimulation and constant number of vibrissa deflections, we showed that the optically imaged spread of activity is more discrete at higher stimulation frequencies. We combined optical imaging with multiple electrode recording and confirmed that the neuronal response to individual vibrissa stimulation at the optically imaged center of activity is greater than the response away from the imaged center. Consistent with the imaging data, these recordings also showed no response to a second vibrissa deflection at 5 Hz at a peripheral recording site, though there was a significant response to a second vibrissa deflection at 1 Hz at the same peripheral site. These findings demonstrate that vibrissa stimulation at higher frequencies leads to more focused physiological responses in cortex. Thus the spread of activation in rat barrel cortex is modulated in a dynamic fashion by the frequency of vibrissa stimulation.

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5

Lefebvre, Veronique, Ying Zheng, Chris Martin, Ian M. Devonshire, Samuel Harris, and John E. Mayhew. "A Dynamic Causal Model of the Coupling Between Pulse Stimulation and Neural Activity." Neural Computation 21, no. 10 (October 2009): 2846–68. http://dx.doi.org/10.1162/neco.2009.07-08-820.

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We present a dynamic causal model that can explain context-dependent changes in neural responses, in the rat barrel cortex, to an electrical whisker stimulation at different frequencies. Neural responses were measured in terms of local field potentials. These were converted into current source density (CSD) data, and the time series of the CSD sink was extracted to provide a time series response train. The model structure consists of three layers (approximating the responses from the brain stem to the thalamus and then the barrel cortex), and the latter two layers contain nonlinearly coupled modules of linear second-order dynamic systems. The interaction of these modules forms a nonlinear regulatory system that determines the temporal structure of the neural response amplitude for the thalamic and cortical layers. The model is based on the measured population dynamics of neurons rather than the dynamics of a single neuron and was evaluated against CSD data from experiments with varying stimulation frequency (1–40 Hz), random pulse trains, and awake and anesthetized animals. The model parameters obtained by optimization for different physiological conditions (anesthetized or awake) were significantly different. Following Friston, Mechelli, Turner, and Price ( 2000 ), this work is part of a formal mathematical system currently being developed (Zheng et al., 2005 ) that links stimulation to the blood oxygen level dependent (BOLD) functional magnetic resonance imaging (fMRI) signal through neural activity and hemodynamic variables. The importance of the model described here is that it can be used to invert the hemodynamic measurements of changes in blood flow to estimate the underlying neural activity.

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6

Petersen, Carl C. H. "Short-Term Dynamics of Synaptic Transmission Within the Excitatory Neuronal Network of Rat Layer 4 Barrel Cortex." Journal of Neurophysiology 87, no. 6 (June 1, 2002): 2904–14. http://dx.doi.org/10.1152/jn.2002.87.6.2904.

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The short-term plasticity of synaptic transmission between excitatory neurons within a barrel of layer 4 rat somatosensory neocortex was investigated. Action potentials in presynaptic neurons at frequencies ranging from 1 to 100 Hz evoked depressing postsynaptic excitatory postsynaptic potentials (EPSPs). Recovery from synaptic depression followed an exponential time course with best-fit parameters that differed greatly between individual synaptic connections. The average maximal short-term depression was close to 0.5 with a recovery time constant of around 500 ms. Analysis of each individual sweep showed that there was a correlation between the amplitude of the response to the first and second action potentials such that large first EPSPs were followed by smaller than average second EPSPs and vice versa. Short-term depression between excitatory layer 4 neurons can thus be termed use dependent. A simple model describing use-dependent short-term plasticity was able to closely simulate the experimentally observed dynamic behavior of these synapses for regular spike trains. More complex irregular trains of 10 action potentials occurring within 500 ms were initially well described, but during the train errors increased. Thus for short periods of time the dynamic behavior of these synapses can be predicted accurately. In conjunction with data describing the connectivity, this forms a first step toward computational modeling of the excitatory neuronal network of layer 4 barrel cortex. Simulation of whisking-evoked activity suggests that short-term depression may provide a mechanism for enhancing the detection of objects within the whisker space.

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7

Reyes-Puerta, Vicente, Yael Amitai, Jyh-Jang Sun, Itamar Shani, Heiko J. Luhmann, and Maoz Shamir. "Long-range intralaminar noise correlations in the barrel cortex." Journal of Neurophysiology 113, no. 9 (May 2015): 3410–20. http://dx.doi.org/10.1152/jn.00981.2014.

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Identifying the properties of correlations in the firing of neocortical neurons is central to our understanding of cortical information processing. It has been generally assumed, by virtue of the columnar organization of the neocortex, that the firing of neurons residing in a certain vertical domain is highly correlated. On the other hand, firing correlations between neurons steeply decline with horizontal distance. Technical difficulties in sampling neurons with sufficient spatial information have precluded the critical evaluation of these notions. We used 128-channel “silicon probes” to examine the spike-count noise correlations during spontaneous activity between multiple neurons with identified laminar position and over large horizontal distances in the anesthetized rat barrel cortex. Eigen decomposition of correlation coefficient matrices revealed that the laminar position of a neuron is a significant determinant of these correlations, such that the fluctuations of layer 5B/6 neurons are in opposite direction to those of layers 5A and 4. Moreover, we found that within each experiment, the distribution of horizontal, intralaminar spike-count correlation coefficients, up to a distance of ∼1.5 mm, is practically identical to the distribution of vertical correlations. Taken together, these data reveal that the neuron's laminar position crucially affects its role in cortical processing. Moreover, our analyses reveal that this laminar effect extends over several functional columns. We propose that within the cortex the influence of the horizontal elements exists in a dynamic balance with the influence of the vertical domain and this balance is modulated with brain states to shape the network's behavior.

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8

IWASAKI, Naoko, Akihiro KARASHIMA, Mitsuyuki NAKAO, Norihiro KATAYAMA, and Mitsuaki YAMAMOTO. "Changes in the dynamics of sleep electroencephalogram in rat barrel cortex associated with long-term sensory deprivation." Sleep and Biological Rhythms 1, no. 2 (June 2003): 155–57. http://dx.doi.org/10.1046/j.1446-9235.2003.00030.x.

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9

Cowan, Anna I., and Christian Stricker. "Functional Connectivity in Layer IV Local Excitatory Circuits of Rat Somatosensory Cortex." Journal of Neurophysiology 92, no. 4 (October 2004): 2137–50. http://dx.doi.org/10.1152/jn.01262.2003.

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There are two types of excitatory neurons within layer IV of rat somatosensory cortex: star pyramidal (SP) and spiny stellate cells (SS). We examined the intrinsic properties and connectivity between these neurons to determine differences in function. Eighty-four whole cell recordings of pairs of neurons were examined in slices of rat barrel cortex at 36 ± 1°C. Only minimal differences in intrinsic properties were found; however, differences in synaptic strength could clearly be shown. Connections between homonymous pairs (SS–SS or SP–SP) had a higher efficacy than heteronymous connections. This difference was mainly a result of quantal content. In 42 pairs, synaptic dynamics were examined. Sequences of action potentials (3–20 Hz) in the presynaptic neuron consistently caused synaptic depression ( Ē2/ Ē1 = 0.53 ± 0.18). The dominant component of depression was release-independent; this depression occurred even when preceding action potentials had failed to cause a response. The release-dependence of depression was target specific; in addition, release-independence was greater for postsynaptic SPs. In a subset of connections formed only between SP and any other cell type (43%), synaptic efficacy was dependent on the presynaptic membrane potential ( Vm); at −55 mV, the connections were almost silent, whereas at −85 mV, transmission was very reliable. We suggest that, within layer IV, there is stronger efficacy between homonymous than between heteronymous excitatory connections. Under dynamic conditions, the functional connectivity is shaped by synaptic efficacy at individual connections, by Vm, and by the specificity in the types of synaptic depression.

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10

Weber, B., C. Burger, M. T. Wyss, G. K. von Schulthess, F. Scheffold, and A. Buck. "Optical imaging of the spatiotemporal dynamics of cerebral blood flow and oxidative metabolism in the rat barrel cortex." European Journal of Neuroscience 20, no. 10 (November 2004): 2664–70. http://dx.doi.org/10.1111/j.1460-9568.2004.03735.x.

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11

Bernardi, Davide, Guy Doron, Michael Brecht, and Benjamin Lindner. "A network model of the barrel cortex combined with a differentiator detector reproduces features of the behavioral response to single-neuron stimulation." PLOS Computational Biology 17, no. 2 (February 8, 2021): e1007831. http://dx.doi.org/10.1371/journal.pcbi.1007831.

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The stimulation of a single neuron in the rat somatosensory cortex can elicit a behavioral response. The probability of a behavioral response does not depend appreciably on the duration or intensity of a constant stimulation, whereas the response probability increases significantly upon injection of an irregular current. Biological mechanisms that can potentially suppress a constant input signal are present in the dynamics of both neurons and synapses and seem ideal candidates to explain these experimental findings. Here, we study a large network of integrate-and-fire neurons with several salient features of neuronal populations in the rat barrel cortex. The model includes cellular spike-frequency adaptation, experimentally constrained numbers and types of chemical synapses endowed with short-term plasticity, and gap junctions. Numerical simulations of this model indicate that cellular and synaptic adaptation mechanisms alone may not suffice to account for the experimental results if the local network activity is read out by an integrator. However, a circuit that approximates a differentiator can detect the single-cell stimulation with a reliability that barely depends on the length or intensity of the stimulus, but that increases when an irregular signal is used. This finding is in accordance with the experimental results obtained for the stimulation of a regularly-spiking excitatory cell.

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12

IWASAKI, Naoko, Akihiro KARASHIMA, Norihiro KATAYAMA, and Mitsuyuki NAKAO. "Progressive changes in sleep electroencephalogram dynamics in the rat barrel cortex associated with long-term alternation of sensory input activities." Sleep and Biological Rhythms 6, no. 4 (October 2008): 208–14. http://dx.doi.org/10.1111/j.1479-8425.2008.00363.x.

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13

Sonnay, Sarah, João M. N. Duarte, and Nathalie Just. "Lactate and glutamate dynamics during prolonged stimulation of the rat barrel cortex suggest adaptation of cerebral glucose and oxygen metabolism." Neuroscience 346 (March 2017): 337–48. http://dx.doi.org/10.1016/j.neuroscience.2017.01.034.

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14

Lippert, Michael T., Kentaroh Takagaki, Weifeng Xu, Xiaoying Huang, and Jian-Young Wu. "Methods for Voltage-Sensitive Dye Imaging of Rat Cortical Activity With High Signal-to-Noise Ratio." Journal of Neurophysiology 98, no. 1 (July 2007): 502–12. http://dx.doi.org/10.1152/jn.01169.2006.

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We describe methods to achieve high sensitivity in voltage-sensitive dye (VSD) imaging from rat barrel and visual cortices in vivo with the use of a blue dye RH1691 and a high dynamic range imaging device (photodiode array). With an improved staining protocol and an off-line procedure to remove pulsation artifact, the sensitivity of VSD recording is comparable with that of local field potential recording from the same location. With this sensitivity, one can record from ∼500 individual detectors, each covering an area of cortical tissue 160 μm in diameter (total imaging field ∼4 mm in diameter) and a temporal resolution of 1,600 frames/s, without multiple-trial averaging. We can record 80–100 trials of intermittent 10-s trials from each imaging field before the VSD signal reduces to one half of its initial amplitude because of bleaching and wash-out. Taken together, the methods described in this report provide a useful tool for visualizing evoked and spontaneous waves from rodent cortex.

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15

Beierlein, Michael, Jay R. Gibson, and Barry W. Connors. "Two Dynamically Distinct Inhibitory Networks in Layer 4 of the Neocortex." Journal of Neurophysiology 90, no. 5 (November 2003): 2987–3000. http://dx.doi.org/10.1152/jn.00283.2003.

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Normal operations of the neocortex depend critically on several types of inhibitory interneurons, but the specific function of each type is unknown. One possibility is that interneurons are differentially engaged by patterns of activity that vary in frequency and timing. To explore this, we studied the strength and short-term dynamics of chemical synapses interconnecting local excitatory neurons (regular-spiking, or RS, cells) with two types of inhibitory interneurons: fast-spiking (FS) cells, and low-threshold spiking (LTS) cells of layer 4 in the rat barrel cortex. We also tested two other pathways onto the interneurons: thalamocortical connections and recurrent collaterals from corticothalamic projection neurons of layer 6. The excitatory and inhibitory synapses interconnecting RS cells and FS cells were highly reliable in response to single stimuli and displayed strong short-term depression. In contrast, excitatory and inhibitory synapses interconnecting the RS and LTS cells were less reliable when initially activated. Excitatory synapses from RS cells onto LTS cells showed dramatic short-term facilitation, whereas inhibitory synapses made by LTS cells onto RS cells facilitated modestly or slightly depressed. Thalamocortical inputs strongly excited both RS and FS cells but rarely and only weakly contacted LTS cells. Both types of interneurons were strongly excited by facilitating synapses from axon collaterals of corticothalamic neurons. We conclude that there are two parallel but dynamically distinct systems of synaptic inhibition in layer 4 of neocortex, each defined by its intrinsic spiking properties, the short-term plasticity of its chemical synapses, and (as shown previously) an exclusive set of electrical synapses. Because of their unique dynamic properties, each inhibitory network will be recruited by different temporal patterns of cortical activity.

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16

Petersen, Carl C. H., Amiram Grinvald, and Bert Sakmann. "Spatiotemporal Dynamics of Sensory Responses in Layer 2/3 of Rat Barrel Cortex MeasuredIn Vivoby Voltage-Sensitive Dye Imaging Combined with Whole-Cell Voltage Recordings and Neuron Reconstructions." Journal of Neuroscience 23, no. 4 (February 15, 2003): 1298–309. http://dx.doi.org/10.1523/jneurosci.23-04-01298.2003.

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17

Panzeri, Stefano, and Simon R. Schultz. "A Unified Approach to the Study of Temporal, Correlational, and Rate Coding." Neural Computation 13, no. 6 (June 1, 2001): 1311–49. http://dx.doi.org/10.1162/08997660152002870.

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We demonstrate that the information contained in the spike occurrence times of a population of neurons can be broken up into a series of terms, each reflecting something about potential coding mechanisms. This is possible in the coding regime in which few spikes are emitted in the relevant time window. This approach allows us to study the additional information contributed by spike timing beyond that present in the spike counts and to examine the contributions to the whole information of different statistical properties of spike trains, such as firing rates and correlation functions. It thus forms the basis for a new quantitative procedure for analyzing simultaneous multiple neuron recordings and provides theoretical constraints on neural coding strategies. We find a transition between two coding regimes, depending on the size of the relevant observation timescale. For time windows shorter than the timescale of the stimulus-induced response fluctuations, there exists a spike count coding phase, in which the purely temporal information is of third order in time. For time windows much longer than the characteristic timescale, there can be additional timing information of first order, leading to a temporal coding phase in which timing information may affect the instantaneous information rate. In this new framework, we study the relative contributions of the dynamic firing rate and correlation variables to the full temporal information, the interaction of signal and noise correlations in temporal coding, synergy between spikes and between cells, and the effect of refractoriness. We illustrate the utility of the technique by analyzing a few cells from the rat barrel cortex.

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18

Wei, Ling, Carl M. Rovainen, and Thomas A. Woolsey. "Ministrokes in Rat Barrel Cortex." Stroke 26, no. 8 (August 1995): 1459–62. http://dx.doi.org/10.1161/01.str.26.8.1459.

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19

Erchova, I. A. "Rapid Fluctuations in Rat Barrel Cortex Plasticity." Journal of Neuroscience 24, no. 26 (June 30, 2004): 5931–41. http://dx.doi.org/10.1523/jneurosci.1202-04.2004.

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20

Pinto, David J., Joshua C. Brumberg, and Daniel J. Simons. "Circuit Dynamics and Coding Strategies in Rodent Somatosensory Cortex." Journal of Neurophysiology 83, no. 3 (March 1, 2000): 1158–66. http://dx.doi.org/10.1152/jn.2000.83.3.1158.

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Previous experimental studies of both cortical barrel and thalamic barreloid neuron responses in rodent somatosensory cortex have indicated an active role for barrel circuitry in processing thalamic signals. Previous modeling studies of the same system have suggested that a major function of the barrel circuit is to render the response magnitude of barrel neurons particularly sensitive to the temporal distribution of thalamic input. Specifically, thalamic inputs that are initially synchronous strongly engage recurrent excitatory connections in the barrel and generate a response that briefly withstands the strong damping effects of inhibitory circuitry. To test this experimentally, we recorded responses from 40 cortical barrel neurons and 63 thalamic barreloid neurons evoked by whisker deflections varying in velocity and amplitude. This stimulus evoked thalamic response profiles that varied in terms of both their magnitude and timing. The magnitude of the thalamic population response, measured as the average number of evoked spikes per stimulus, increased with both deflection velocity and amplitude. On the other hand, the degree of initial synchrony, measured from population peristimulus time histograms, was highly correlated with the velocity of whisker deflection, deflection amplitude having little or no effect on thalamic synchrony. Consistent with the predictions of the model, the cortical population response was determined largely by whisker velocity and was highly correlated with the degree of initial synchrony among thalamic neurons ( R 2 = 0.91), as compared with the average number of evoked thalamic spikes ( R 2 = 0.38). Individually, the response of nearly all cortical cells displayed a positive correlation with deflection velocity; this homogeneity is consistent with the dependence of the cortical response on local circuit interactions as proposed by the model. By contrast, the response of individual thalamic neurons varied widely. These findings validate the predictions of the modeling studies and, more importantly, demonstrate that the mechanism by which the cortex processes an afferent signal is inextricably linked with, and in fact determines, the saliency of neural codes embedded in the thalamic response.

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21

Lenschow, Constanze, and Michael Brecht. "Barrel Cortex Membrane Potential Dynamics in Social Touch." Neuron 85, no. 4 (February 2015): 718–25. http://dx.doi.org/10.1016/j.neuron.2014.12.059.

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22

Lenschow, Constanze, and Michael Brecht. "Barrel Cortex Membrane Potential Dynamics in Social Touch." Neuron 85, no. 5 (March 2015): 1145. http://dx.doi.org/10.1016/j.neuron.2015.02.030.

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23

Margolis, David J., Henry Lütcke, and Fritjof Helmchen. "Microcircuit dynamics of map plasticity in barrel cortex." Current Opinion in Neurobiology 24 (February 2014): 76–81. http://dx.doi.org/10.1016/j.conb.2013.08.019.

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24

Diamond, M. E., M. Armstrong-James, and F. F. Ebner. "Experience-dependent plasticity in adult rat barrel cortex." Proceedings of the National Academy of Sciences 90, no. 5 (March 1, 1993): 2082–86. http://dx.doi.org/10.1073/pnas.90.5.2082.

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25

Khatri, V., and D. J. Simons. "Angularly Nonspecific Response Suppression in Rat Barrel Cortex." Cerebral Cortex 17, no. 3 (March 31, 2006): 599–609. http://dx.doi.org/10.1093/cercor/bhk006.

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26

Han, Yong, Ming-De Huang, Man-Li Sun, Shumin Duan, and Yan-Qin Yu. "Long-Term Synaptic Plasticity in Rat Barrel Cortex." Cerebral Cortex 25, no. 9 (April 15, 2014): 2741–51. http://dx.doi.org/10.1093/cercor/bhu071.

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27

Melzer, P., R. N. S. Sachdev, N. Jenkinson, and F. F. Ebner. "Stimulus Frequency Processing in Awake Rat Barrel Cortex." Journal of Neuroscience 26, no. 47 (November 22, 2006): 12198–205. http://dx.doi.org/10.1523/jneurosci.2620-06.2006.

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28

Lebedev, M. A. "Experience-dependent Plasticity of Rat Barrel Cortex: Redistribution of Activity across Barrel-columns." Cerebral Cortex 10, no. 1 (January 1, 2000): 23–31. http://dx.doi.org/10.1093/cercor/10.1.23.

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29

Harrell, Evan R., Matías A. Goldin, Brice Bathellier, and Daniel E. Shulz. "An elaborate sweep-stick code in rat barrel cortex." Science Advances 6, no. 38 (September 2020): eabb7189. http://dx.doi.org/10.1126/sciadv.abb7189.

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In rat barrel cortex, feature encoding schemes uncovered during broadband whisker stimulation are hard to reconcile with the simple stick-slip code observed during natural tactile behaviors, and this has hindered the development of a generalized computational framework. By designing broadband artificial stimuli to sample the inputs encoded under natural conditions, we resolve this disparity while markedly increasing the percentage of deep layer neurons found to encode whisker movements, as well as the diversity of these encoded features. Deep layer neurons encode two main types of events, sticks and sweeps, corresponding to high angular velocity bumps and large angular displacements with high velocity, respectively. Neurons can exclusively encode sticks or sweeps, or they can encode both, with or without direction selectivity. Beyond unifying coding theories from naturalistic and artificial stimulation studies, these findings delineate a simple and generalizable set of whisker movement features that can support a range of perceptual processes.

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30

von Heimendahl, Moritz, Pavel M. Itskov, Ehsan Arabzadeh, and Mathew E. Diamond. "Neuronal Activity in Rat Barrel Cortex Underlying Texture Discrimination." PLoS Biology 5, no. 11 (November 13, 2007): e305. http://dx.doi.org/10.1371/journal.pbio.0050305.

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31

Mitrukhina, Olga, Dmitry Suchkov, Roustem Khazipov, and Marat Minlebaev. "Imprecise Whisker Map in the Neonatal Rat Barrel Cortex." Cerebral Cortex 25, no. 10 (August 6, 2014): 3458–67. http://dx.doi.org/10.1093/cercor/bhu169.

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32

Arabzadeh, E. "Whisker Vibration Information Carried by Rat Barrel Cortex Neurons." Journal of Neuroscience 24, no. 26 (June 30, 2004): 6011–20. http://dx.doi.org/10.1523/jneurosci.1389-04.2004.

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33

Land, Peter W., and Daniel J. Simons. "Cytochrome oxidase staining in the rat smI barrel cortex." Journal of Comparative Neurology 238, no. 2 (August 8, 1985): 225–35. http://dx.doi.org/10.1002/cne.902380209.

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34

Wright, Nicholas, and Kevin Fox. "Origins of Cortical Layer V Surround Receptive Fields in the Rat Barrel Cortex." Journal of Neurophysiology 103, no. 2 (February 2010): 709–24. http://dx.doi.org/10.1152/jn.00560.2009.

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Layer IV of the barrel cortex contains an anatomical map of the contralateral whisker pad, which serves as a useful reference in relating receptive field properties of cells to the cortical columns in which they reside. Recent studies have shown that the degree to which the surround receptive fields of layer IV cells are generated intracortically or subcortically depends on whether they lie in barrel or septal columns. To investigate whether this is true for layer V cells, we blocked intracortical activity in the barrel cortex by infusing muscimol from the cortical surface and measuring spike responses to sensory stimulation in the presence of locally iontophoresed bicuculline. Layer V cells beneath barrels had small subcortically generated single- or double-whisker center receptive fields and larger intracortically generated six to seven whisker surround receptive fields. Conversely, septally located cells received multiwhisker input from both subcortical and cortical sources. Most properties of layer Va and layer Vb cells were very similar. However, layer Vb barrel neurons showed a relative lack of phasic inhibition evoked from sensory input compared with layer Va cells. The direct thalamic input to the layer V cells was not sufficient to evoke a sensory response in the absence of input from superficial layers. These findings suggest that despite the apparent overt similarity of layer V receptive fields, barrel and septal subdivisions process different sources of information within the barrel cortex.

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35

Minlebaev, Marat, Yehezkel Ben-Ari, and Rustem Khazipov. "Network Mechanisms of Spindle-Burst Oscillations in the Neonatal Rat Barrel Cortex In Vivo." Journal of Neurophysiology 97, no. 1 (January 2007): 692–700. http://dx.doi.org/10.1152/jn.00759.2006.

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Early in development, cortical networks generate particular patterns of activity that participate in cortical development. The dominant pattern of electrical activity in the neonatal rat neocortex in vivo is a spatially confined spindle-burst. Here, we studied network mechanisms of generation of spindle-bursts in the barrel cortex of neonatal rats using a superfused cortex preparation in vivo. Both spontaneous and sensory-evoked spindle-bursts were present in the superfused barrel cortex. Pharmacological analysis revealed that spindle-bursts are driven by glutamatergic synapses with a major contribution of AMPA/kainate receptors, but slight participation of NMDA receptors and gap junctions. Although GABAergic synapses contributed minimally to the pacing the rhythm of spindle-burst oscillations, surround GABAergic inhibition appeared to be crucial for their compartmentalization. We propose that local spindle-burst oscillations, driven by glutamatergic synapses and spatially confined by GABAergic synapses, contribute to the development of barrel cortex during the critical period of developmental plasticity.

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Suzuki, Takashi, Yasuhiro Ooi, and Junji Seki. "Infrared thermal imaging of rat somatosensory cortex with whisker stimulation." Journal of Applied Physiology 112, no. 7 (April 1, 2012): 1215–22. http://dx.doi.org/10.1152/japplphysiol.00867.2011.

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The present study aims to validate the applicability of infrared (IR) thermal imaging for the study of brain function through experiments on the rat barrel cortex. Regional changes in neural activity within the brain produce alterations in local thermal equilibrium via increases in metabolic activity and blood flow. We studied the relationship between temperature change and neural activity in anesthetized rats using IR imaging to visualize stimulus-induced changes in the somatosensory cortex of the brain. Sensory stimulation of the vibrissae (whiskers) was given for 10 s using an oscillating whisker vibrator (5-mm deflection at 10, 5, and 1 Hz). The brain temperature in the observational region continued to increase significantly with whisker stimulation. The mean peak recorded temperature changes were 0.048 ± 0.028, 0.054 ± 0.036, and 0.097 ± 0.015°C at 10, 5, and 1 Hz, respectively. We also observed that the temperature increase occurred in a focal spot, radiating to encompass a larger region within the contralateral barrel cortex region during single-whisker stimulation. Whisker stimulation also produced ipsilateral cortex temperature increases, which were localized in the same region as the pial arterioles. Temperature increase in the barrel cortex was also observed in rats treated with a calcium channel blocker (nimodipine), which acts to suppress the hemodynamic response to neural activity. Thus the location and area of temperature increase were found to change in accordance with the region of neural activation. These results indicate that IR thermal imaging is viable as a functional quantitative neuroimaging technique.

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Huang, Wei, Michael Armstrong-James, V. Rema, Mathew E. Diamond, and Ford F. Ebner. "Contribution of Supragranular Layers to Sensory Processing and Plasticity in Adult Rat Barrel Cortex." Journal of Neurophysiology 80, no. 6 (December 1, 1998): 3261–71. http://dx.doi.org/10.1152/jn.1998.80.6.3261.

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Huang, Wei, Michael Armstrong-James, V. Rema, Mathew E. Diamond, and Ford F. Ebner. Contribution of supragranular layers to sensory processing and plasticity in adult rat barrel cortex. J. Neurophysiol. 80: 3261–3271, 1998. In mature rat primary somatic sensory cortical area (SI) barrel field cortex, the thalamic-recipient granular layer IV neurons project especially densely to layers I, II, III, and IV. A prior study showed that cells in the supragranular layers are the fastest to change their response properties to novel changes in sensory inputs. Here we examine the effect of removing supragranular circuitry on the responsiveness and synaptic plasticity of cells in the remaining layers. To remove the layer II + III (supragranular) neurons from the circuitry of barrel field cortex, N-methyl-d-aspartate (NMDA) was applied to the exposed dura over the barrel cortex, which destroys those neurons by excitotoxicity without detectable damage to blood vessels or axons of passage. Fifteen days after NMDA treatment, the first responsive cells encountered were 400–430 μm below the pial surface. In separate cases triphenyltetrazolium chloride (TTC), a vital dye taken up by living cells, was absent from the lesion area. Cytochrome oxidase (CO) activity was absent in the first few tangential sections through the barrel field in all cases before arriving at the CO-dense barrel domains. These findings indicate that the lesions were quite consistent from animal to animal. Controls consisted of applying vehicle without NMDA under similar conditions. Responses of D2 barrel cells were assessed for spontaneous activity and level of response to stimulation of the principal D2 whisker and four surround whiskers D1, D3, C2, and E2. In two additional groups of animals treated in the same way, sensory plasticity was assessed by trimming all whiskers except D2 and either D1 or D3 (called Dpaired) for 7 days before recording cortical responses. Such whisker pairing normally potentiates D2 barrel cell responses to stimulation of the two intact whiskers (D2 + Dpaired). After NMDA lesions, cortical cells still responded to all whiskers tested. Cells in lesioned cortex showed reduced response amplitude compared with sham-operated controls to all D-row whiskers. In-arc surround whisker (C2 or E2) responses were normal. Spontaneous activity did not change significantly in any remaining layer at the time tested. Modal latencies to stimulation of principal D2 or surround D1 or D3 whiskers showed no significant change after lesioning. These findings indicate that there is a reasonable preservation of the response properties of layer IV, V, VI neurons after removal of layer II–III neurons in this way. Whisker pairing plasticity in layer IV–VI D2 barrel column neurons occurred in both lesioned and sham animals but was reduced significantly in lesioned animals compared with controls. The response bias generated by whisker trimming (Dpaired/Dcut + Dpaired ratio) was less pronounced in NMDA-lesioned than sham-lesioned animals. Proportionately fewer neurons in layer IV (52 vs. 64%) and in the infragranular layers (55 vs. 68%) exhibited a clear response bias to paired whiskers. We conclude that receptive-field plasticity can occur in layers IV–VI of barrel cortex in the absence of the supragranular layer circuitry. However, layer I–III circuitry does play a role in normal receptive-field generation and is required for the full expression of whisker pairing plasticity in granular and infragranular layer cells.

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Hemelt, Marie E., Ernest E. Kwegyir-Afful, Randy M. Bruno, Daniel J. Simons, and Asaf Keller. "Consistency of Angular Tuning in the Rat Vibrissa System." Journal of Neurophysiology 104, no. 6 (December 2010): 3105–12. http://dx.doi.org/10.1152/jn.00697.2009.

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Each region along the rat mystacial vibrissa pathway contains neurons that respond preferentially to vibrissa deflections in a particular direction, a property called angular tuning. Angular tuning is normally defined using responses to deflections of the principal vibrissa, which evokes the largest response magnitude. However, neurons in most brain regions respond to multiple vibrissae and do not necessarily respond to different vibrissae with the same angular tuning. We tested the consistency of angular tuning across the receptive field in several stations along the vibrissa-to-cortex pathway: primary somatosensory (barrel) cortex, ventroposterior medial nucleus of the thalamus (VPM), second somatosensory cortex, and superior colliculus. We found that when averaged across the population, neurons in all of these regions have low (superior colliculus and second somatosensory cortex) or statistically insignificant (barrel cortex and VPM) angular tuning consistencies across vibrissae. Nevertheless, in each region there are a small number of neurons that display consistent angular tuning for at least some vibrissae. We discuss the relevance of these findings for the transformation of inputs along the vibrissa trigeminal pathway and for the detection of sensory cues by whisking animals.

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Roy, Noah C., Thomas Bessaih, and Diego Contreras. "Comprehensive mapping of whisker-evoked responses reveals broad, sharply tuned thalamocortical input to layer 4 of barrel cortex." Journal of Neurophysiology 105, no. 5 (May 2011): 2421–37. http://dx.doi.org/10.1152/jn.00939.2010.

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Cortical neurons are organized in columns, distinguishable by their physiological properties and input-output organization. Columns are thought to be the fundamental information-processing modules of the cortex. The barrel cortex of rats and mice is an attractive model system for the study of cortical columns, because each column is defined by a layer 4 (L4) structure called a barrel, which can be clearly visualized. A great deal of information has been collected regarding the connectivity of neurons in barrel cortex, but the nature of the input to a given L4 barrel remains unclear. We measured this input by making comprehensive maps of whisker-evoked activity in L4 of rat barrel cortex using recordings of multiunit activity and current source density analysis of local field potential recordings of animals under light isoflurane anesthesia. We found that a large number of whiskers evoked a detectable response in each barrel (mean of 13 suprathreshold, 18 subthreshold) even after cortical activity was abolished by application of muscimol, a GABAA agonist. We confirmed these findings with intracellular recordings and single-unit extracellular recordings in vivo. This constitutes the first direct confirmation of the hypothesis that subcortical mechanisms mediate a substantial multiwhisker input to a given cortical barrel.

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Petersen, Carl C. H., and Bert Sakmann. "The Excitatory Neuronal Network of Rat Layer 4 Barrel Cortex." Journal of Neuroscience 20, no. 20 (October 15, 2000): 7579–86. http://dx.doi.org/10.1523/jneurosci.20-20-07579.2000.

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Allitt, Benjamin J., Dasuni S. Alwis, and Ramesh Rajan. "Laminar-specific encoding of texture elements in rat barrel cortex." Journal of Physiology 595, no. 23 (October 15, 2017): 7223–47. http://dx.doi.org/10.1113/jp274865.

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Ajima, Ayako, Yoshitaka Matsuda, Kenichi Ohki, Dae-Shik Kim, and Shigeru Tanaka. "GABA-mediated representation of temporal information in rat barrel cortex." NeuroReport 10, no. 9 (June 1999): 1973–79. http://dx.doi.org/10.1097/00001756-199906230-00033.

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Filipkowski, R. K. "Tactile Experience Induces c-fos Expression in Rat Barrel Cortex." Learning & Memory 7, no. 2 (March 1, 2000): 116–22. http://dx.doi.org/10.1101/lm.7.2.116.

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Blue, Mary E., and Michael V. Johnston. "The ontogeny of glutamate receptors in rat barrel field cortex." Developmental Brain Research 84, no. 1 (January 1995): 11–25. http://dx.doi.org/10.1016/0165-3806(94)00147-r.

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Bobrov, Evgeny, Jason Wolfe, Rajnish P. Rao, and Michael Brecht. "The Representation of Social Facial Touch in Rat Barrel Cortex." Current Biology 24, no. 1 (January 2014): 109–15. http://dx.doi.org/10.1016/j.cub.2013.11.049.

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Safaai, H., M. von Heimendahl, J. M. Sorando, M. E. Diamond, and M. Maravall. "Coordinated Population Activity Underlying Texture Discrimination in Rat Barrel Cortex." Journal of Neuroscience 33, no. 13 (March 27, 2013): 5843–55. http://dx.doi.org/10.1523/jneurosci.3486-12.2013.

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Katz, Y., J. E. Heiss, and I. Lampl. "Cross-Whisker Adaptation of Neurons in the Rat Barrel Cortex." Journal of Neuroscience 26, no. 51 (December 20, 2006): 13363–72. http://dx.doi.org/10.1523/jneurosci.4056-06.2006.

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Ichikawa, Takehiko, Takafumi Akasaki, Satoshi Shimegi, Yumiko Yoshimura, and Hiromichi Sato. "1809 Neuronal connectivity among barrels in the rat barrel cortex." Neuroscience Research 28 (January 1997): S220. http://dx.doi.org/10.1016/s0168-0102(97)90599-x.

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Kim, Uhnoh, and Ford F. Ebner. "Barrels and septa: Separate circuits in rat barrel field cortex." Journal of Comparative Neurology 408, no. 4 (June 14, 1999): 489–505. http://dx.doi.org/10.1002/(sici)1096-9861(19990614)408:4<489::aid-cne4>3.0.co;2-e.

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Armstrong-James, Michael, and Kevin Fox. "Spatiotemporal convergence and divergence in the rat S1 ?Barrel? cortex." Journal of Comparative Neurology 263, no. 2 (September 8, 1987): 265–81. http://dx.doi.org/10.1002/cne.902630209.

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