Neuronal assembly dynamics in the rat auditory cortex during reorganization induced by intracortical microstimulation

Single neurons, acting alone, cannot account for the complex and rapid computations that are routinely accomplished by the behaving nervous system. Recent studies with separable multineuron recordings are showing that neuronal assemblies can indeed be detected and that their organization is very dynamic, depending on variables such as time, physical stimulus, and context. Here we explore both single-neuron and assembly properties in the rat's auditory cortex. Acoustic stimuli are used as a normal, physiological input, and weak electrical intracortical microstimulation (ICMS) as a perturbation that forces a rapid cortical reorganization. In this setting, various aspects of neuronal interactions are changed by the ICMS. We found that cortical neurons exhibited highly synchronous oscillatory firing patterns that were enhanced by ICMS. Cross-correlation studies between two spike trains showed that statistically significant correlations depended on the anatomical distance between the two neurons. ICMS changed the strength and the local number of such correlations. Joint petristimulus analysis and gravity analysis showed that the correlation between neuronal activities varied dynamically at several time scales. We have identified neuronal assemblies in two ways, defined through similarity of receptive field properties and defined through correlated firing. Close anatomical spacing between neurons was conducive to, but not sufficient for membership in, the same assembly with either definition. ICMS changed cortical organization by altering assembly membership. Our data show that neuronal assemblies in the rat auditory cortex can be established transiently in time and that their membership is dynamic.     

Histopathologic findings in 37 cases of functional hemispherectomy.

Hemispherectomy procedures are performed in patients for whom focal cortical resection would be predicted to produce a significant reduction in seizures. The functional hemispherectomy procedure consists of disconnecting the hemispheres while attempting, in some cases, to preserve parenchyma. This study retrospectively reviews the histopathologic findings in 37 cases of functional hemispherectomy performed between 1990 and 1998 at a major epilepsy center. Procedures were performed in 20 males and 17 females who ranged in age from 3 months to 37 years (mean age, 9.6 years). In all but two cases, more than half or all the material submitted for pathologic testing was examined histologically. Cortical dysplasias or hemimegalencephaly were identified in 14 patients. The most common patterns of dysplasia observed included architectural disorganization (n = 13), increased molecular layer neurons (n = 11), and neuronal cytomegaly (n = 11). One patient was known to have epidermal nevus syndrome. Six patients had Sturge-Weber syndrome. Remote infarct/ischemic damage was identified as the etiology of seizures in six patients; four of these patients had mild associated secondary cortical architectural abnormalities. Three patients demonstrated pathology consistent with Rasmussen's encephalitis; one additional patient had chronic encephalitis changes, not otherwise specified. In two cases, changes consistent with hippocampal sclerosis were identified; additionally, hippocampal neuronal loss and gliosis was focally identified in three patients. Most of these patients had coexistent cortical dysplasia or radiographic evidence of remote infarct. One specimen demonstrated areas of infarct following resection of an arteriovenous malformation. In two specimens, significant histopathologic findings were not identified; both of these patients had radiographic evidence of remote infarct. The spectrum of pathologic conditions that may be encountered in the setting of a functional hemispherectomy is varied and in this study most frequently included cortical dysplasia, Sturge-Weber syndrome remote infarct, and Rasmussen's encephalitis.     

The organization of serotonergic projections to cerebral cortex in primates: regional distribution of axon terminals.

Serotonergic axons are widely distributed in the primate forebrain and represent the most abundant ascending projection from the reticular formation. Immunocytochemical methods have been utilized to examine the density, laminar distribution and morphology of serotonergic axons in both primary projection (motor, somatosensory) and association areas (dorsolateral prefrontal, area 5) as well as in the hippocampus and in cingulate cortex of rhesus and cynomolgus macaques. Serotonergic axons are present in all areas of cortex examined, and all cortical layers receive serotonergic afferents. However, the intracortical distribution of serotonergic axon terminals is not uniform; rather, different regions of cortex exhibit dissimilarities in both the density and laminar distribution of serotonergic axons. Thus, there are local patterns of serotonin innervation that are characteristic of each cortical area. Highly diverse patterns of serotonin innervation are found in heterotypical areas of cortex; more subtle variations are present among homotypical areas. Two morphologic types of serotonergic axon terminals, fine and beaded, are present in all cortical areas, and they typically exhibit different laminar distributions: in most areas of neocortex, beaded axons predominate in layer I while fine axons predominate in layers II-VI. However, exceptions to this pattern were observed in primary visual cortex and in the hippocampal formation. The distinctive local patterns of serotonin innervation observed in this study indicate that raphe-cortical projections are likely to have differential influences on particular cytoarchitectonic areas of cerebral cortex in the primate. Moreover, the discrete laminar distribution of serotonin axons suggests that serotonergic projections selectively innervate particular neuronal elements in cerebral cortex. The present findings suggest that the two classes of serotonergic axons, fine and beaded, which have different patterns of termination, affect different sets of cortical neurons. In addition, these two serotonergic projections may be associated with different sets of serotonergic receptors and thus produce selective effects on cortical function.     

Differential dynamics of transient neuronal assemblies in visual compared to auditory cortex

Large-scale, coherent, but highly transient networks of neurons, ‘neuronal assemblies’, operate over a sub-second time frame. Such assemblies of brain cells need not necessarily respect well-defined anatomical compartmentalisation, but represent an intermediate level of brain organisation between identified brain regions and individual neurons dependent on the activity status of the synaptic connections and axonal projections. To study neuronal assemblies both in slices and in the living brain, optical imaging using voltage-sensitive dyes (VSDI) offers the highest spatial and temporal resolution in real-time. Applying VSDI technique to compare assemblies in visual versus auditory cortices under standardised experimental protocols, we observed no significant variations in the basic parameters of fluorescence signal and assembly size: such results might be predicted from the canonical invariance of cortical structures across modalities. However, further analysis revealed less obvious yet significant differences in the assembly dynamics of the two regions. The neural assemblies spread widely across layers in the two cortices following paired-pulse stimulation of putative layer 4. The respective patterns of activity started to differentiate within a specific time frame (250–300 ms). The signal was predominant near the point of stimulation in the visual cortex, whereas in the auditory cortex the signal was stronger in the superficial layers. This modality-specific divergence in assembly dynamics highlights a previously under-appreciated level of neuronal processing. Additionally, these findings could prompt a new approach to the understanding of how information from different senses, transmitted as action potentials with identical electrochemical characteristics across different cortices, be it visual or auditory, can eventually yield, nonetheless, the qualitatively distinct experiences of seeing or hearing.     

Neuronal loss in different parts of the nucleus basalis is related to neuritic plaque formation in cortical target areas in Alzheimer's disease

In order to substantiate the hypothesis of a cholinergic pathogenesis of neuritic plaques in Alzheimer's disease the relationship between the loss of cholinergic neurons in six subdivisions of the nucleus basalis of Meynert and density of neuritic plaques in five neocortical target areas and hippocampus was studied in five cases with Alzheimer's disease. Distribution of plaques in different cortical areas as well as degeneration pattern of neurons within the subpopulations of the nucleus basalis were markedly different in the cases of Alzheimer's disease. Quantitative evaluation of the number of neuritic plaques in the five cortical areas revealed a strong correlation with the loss of neurons in those subpopulations of the nucleus basalis which give rise to the cholinergic innervation of the affected cortical areas. The nonlinearity of this correlation may reflect two different modes of plaque formation. Either plaque formation is a self-perpetuating process with an increasing rate depending on the number of plaques already formed or additional mechanisms, with an increasing rate of influence during plaque formation are induced. The shape of the regression function is different for the various cortical regions and their corresponding subpopulations of the nucleus basalis suggesting a different dependency of neuritic plaque formation on the neuronal loss in the nucleus basalis. This might reflect a different density of cholinergic fibers within these areas, a different degree of collateralization of the fibers or other factors not yet known. The findings indicate that degeneration of cortical cholinergic afferents from the neurons of the nucleus basalis is an important feature in the pathogenesis of neuritic plaques.     

The neurobiology of language and verbal memory: observations from awake neurosurgery.

Neurosurgical operations under local anesthesia provide a unique opportunity to investigate the neurobiology of human cognition. We have studied the cortical organization of language and verbal memory in this setting, using two different techniques: electrical stimulation mapping and extracellular microelectrode recording of activity of individual neurons. The two techniques provide very different perspectives. Stimulation mapping identifies brain areas that are essential for a behavior, while changes in neuronal activity can occur in non-essential regions. Stimulation mapping identifies multiple discrete areas in perisylvian cortex of the dominant hemisphere as essential for a function, with separation of areas for different aspects of language including naming in two languages, different semantic classes, naming compared to reading, and language from verbal memory. There is substantial individual variation in the location of these essential areas, variability that in part relates to subjects age, gender and verbal abilities. Neurons changing activity with language or verbal memory are widely distributed, in both hemispheres. However, individual neurons usually change activity with only one function, including naming in only one of two languages, only naming or reading, or with recent verbal memory encoding but not identification of similar items. A few lateralized changes in neuronal activity have been identified, including a predominance of inhibition in dominant hemisphere with naming, and polymodal memory responses in dominant hemisphere, unimodal in nondominant. Specific neuronal populations have been identified that are related to different aspects of memory, that differentiate correct from incorrect identification or memory performance and differentiate learned from unlearned associations, with some evidence of differences in neuronal activity related to subjects' ability.     

Sex dimorphism of neuron-glia correlations in the frontal areas of the human brain

In the series of frontal 20 microm thick paraffin sections, stained using Nissl's cresyl violet method, frontal areas 8, 10, 12, 47 were studied. The important parameters of neuronal functional and metabolic activity--total density of pyramidal neuron distribution (N), the density of satellite gliocyte distribution (SG), the density of the distribution of the neurons, surrounded by satellite glia (NS) as well as the neuron-glia index (NS/N)--were calculated in the cortical layers III and V, areas 8, 10, 12, 47 in both left and right hemispheres in 0.001 mm3 of brain substance. It was found that in women the parameters studied (SG, NS and NS/N) were higher in the cortical areas 12, 47, 10 as compared to those in men, and were lower in area 8. It is concluded that in women, cortical neurons had higher functional and metabolic activity in the areas, associated with emotions, will and intellectual activity while in men they were higher in the areas involved in motor activity.     

The developmental characteristics of speech-motor areas 44 and 45 in the left and right hemispheres of the human brain in early postnatal development

Cytoarchitecture of speech areas 44 and 45 of human brain was studied on frontal series of sections of brain of a newborn, 6 months, 1 year and 2 years old child. Sections were stained with cresyl violet after Nissl. Density of neurons and gliocytes location as well as fraction of satellite gliocytes and surrounded by the latter, glia index were determined by quantitative methods. General principles of cortical areas 44 and 45 were distinguished as well as peculiarities of their formation if left and right hemispheres of human brain in postnatal ontogenesis. Heterochronia of postnatal development of areas 44 and 45 was demonstrated. A hypothesis on different activity of right and left hemispheres speech areas in different periods of speech development of the child was discussed.     

Cross-correlation studies of movement-related cortical potentials during unilateral and bilateral muscle contractions in humans

A useful method of studying the degree of association between two signals of varying amplitude in the time domain is to use cross-correlation analysis. We applied this to the movement-related cortical potentials digitally filtered so as to eliminate the low frequency component before applying it during maximal unilateral left (UL L), unilateral right (UL R) and bilateral (BL) contractions in I I right-handed subjects. The recording electrode sites were over the right and left motor cortex areas (C3 and C4). The BL condition revealed higher cross-correlation levels of cortical activities between the two hemispheres than in UL L or UL R contraction [UL L, r = 0.68 (SEM 0.05); UL R, r = 0.73 (SEM 0.03); BL, r = 0.76 (SEM 0.02)]. The UL R revealed a positive phase difference [5 (SEM 2) ms] when the maximal cross-correlation coefficient was shown and UL L showed a negative phase difference [5 (SEM 3) ms]. However, BL revealed a smaller phase difference [2 (SEM 1) ms] than that for UL. It was concluded that during maximal BL contraction cortical cellular activities in both hemispheres was more synchronized in amplitude and time course compared with maximal UL contractions. Our data suggested that central common drive existed between the right and left motor areas during the maximal BL handgrip contractions and the amplitude of potentials of both hemispheres was modified by the interhemispheric inhibition mechanism as reported in other studies.     

Delayed cell migration in the developing rat brain following maternal omega 3 alpha linolenic acid dietary deficiency

Diminished levels of docosahexaenoic acid (DHA, 22:6n-3), the major polyunsaturated fatty acid (FA) synthesized from alpha linolenic acid (ALA, 18:3n-3), have been implicated in changes in neurotransmitter production, ion channel disruption and impairments of a variety of cognitive, behavioral and motor functions in the perinatal and adult mammal. Neuronal migration in the cortex and hippocampus of newborn and postnatal rats after ALA-deficiency, beginning on the 2nd day after conception and continuing for three weeks, was investigated. A marked decrease in the migration of bromodeoxyuridine⁽⁺⁾/neuronal nuclei⁽⁺⁾/neurofilament⁽⁺⁾ and glia fibrillary acidic protein^(–) neuronal cells to the dense cortical plate was accompanied by a corresponding abundance of non-migrating cells in several regions such as cortical layers IV–VI, corpus callosum and the sub-ventricular zone of ALA-deficient newborns. Similarly, a delayed migration of cells to CA1 and dentate gyrus areas was noticed while most cells were retained in the subicular area adjacent to the hippocampus. The reversibility of delay in migration in the hippocampus and cortex, after one and two weeks respectively, may be attributed to a temporary reelin disorganization or partial deficiency. Transient obstruction of neuronal cell migration may have long-lasting consequences on the organization of neuronal assemblies, on the connection between neurons (lateral connections) and acquisition of function in the adult brain.     

Hebb's concept of cell assemblies an the psychophysiology of word processing

Hebb's brain-theoretical approach suggests that tightly connected networks of neurons, Hebbian cell assemblies, are the building blocks of cognitive functions. These assemblies are not necessarily restricted to a small cortical locus but may be dispersed over distant cortical areas. Assemblies with different topographies can be postulated for different kinds of words, such as meaningful content versus grammatical function words or words eliciting motor versus visual associations. Evidence from evoked potentials and gamma-band electrocortical responses elicited by lexical material supports a cell assembly model of language and other higher cognitive functions.     

Structural bases of individual variability in the human brain

The paper shows considerable differences in the cytoarchitectonical organization of cortical fields 44, 45, and 4, 6 in the brain of different persons. The profile fields of pyramidal neurons in layers III and V substantially differ from each other for the cortical fields under investigation. The specific features of the total fractions of neurons and glia in the cortical fields in different brains are outlined. The density of neuronal location is an important criterion for individual variability. The magnitude of individual variability manifests itself the right and left hemispheres by different ways.     

完全性生长激素缺乏患儿大脑皮质形态变化的3.0 T MRI研究

目的 探讨完全性生长激素缺乏(CGHD)患儿与特发性矮小(ISS)患儿大脑结构的灰质体积、皮层表面积及皮层厚度的差异。方法 回顾性队列研究。纳入2015年1月—2019年1月山东省立医院确诊的12例CGHD患儿(男8例、女4例,年龄5~14岁,生长激素刺激释放试验峰值小于5.0 μg/L)与12例ISS患儿(男9例、女3例,年龄5~14岁,生长激素刺激释放试验峰值大于10.0 μg/L)。在首次就诊时采集患者高分辨率、高信噪比三维T₁WI MRI。利用FreeSurfer软件图像后处理方法获得两组患儿左右大脑半球及全脑灰质体积、皮层表面积与皮层厚度值,统计分析组间的差异情况。将每组患儿的上述形态学参数值分别进行平均以获得该组患儿的均值分布图。将两组患儿的测量均值进行相减,即获得两组患儿该测量均值的差异值分布图。结果 CGHD患儿左、右侧大脑半球的灰质体积分别为(228.50±36.72)cm³、 (229.10±34.95)cm³;皮层表面积分别为(737.02±140.48)cm²、(738.68±135.26)cm²;皮层厚度分别为(2.43±0.09)mm、(2.44±0.09)mm。ISS患儿左、右侧大脑半球的灰质体积分别为(272.36±34.77)cm³、 (272.54±32.76)cm³;皮层表面积分别为(841.88±141.75)cm²、(839.98±135.69)cm²;皮层厚度分别为(2.55±0.18)mm、(2.57±0.17)mm。CGHD患儿与ISS患儿的双侧半球与全脑的灰质体积(F_左=17.884, F_右=20.115, F_全=19.009)、皮层表面积(F_左=11.105, F_右=11.453, F_全=11.337)及皮层厚度(F_左=5.907, F_右=6.109, F_全=6.066)的差异均有统计学意义(P值均<0.05)。测量值的均值分布图与均值的差异值分布图显示两组患儿的灰质体积、皮层表面积与皮层厚度在诸多脑区存在差异。结论 CGHD患儿较ISS患儿的大脑灰质体积、皮层表面积与皮层厚度均较小,这可能与其智力、运动或其它功能发育相对落后相关。     

Dynamic spatiotemporal synaptic integration in cortical neurons: neuronal gain, revisited

Gain modulation is a ubiquitous phenomenon in cortical neurons, providing flexibility to operate under changing conditions. The prevailing view is that this modulation reflects a change in the relationship between mean input and output firing rate brought about by variation in neuronal membrane characteristics. An alternative mechanism is proposed for neuronal gain modulation that takes into account the capability of cortical neurons to process spatiotemporal synaptic correlations. Through the use of numerical simulations, it is shown that voltage-gated and leak conductances, membrane potential, noise, and input firing rate modify the sensitivity of cortical neurons to the degree of temporal correlation between their synaptic inputs. These changes are expressed in a change of the temporal window for synaptic integration and the range of input correlation over which response probability is graded. The study also demonstrates that temporal integration depends on the distance between the inputs and that this interplay of space and time is modulated by voltage-gated and leak conductances. Thus, gain modulation may reflect a change in the relationship between spatiotemporal synaptic correlations and output firing probability. It is further proposed that by acting synergistically with the network, dynamic spatiotemporal synaptic integration in cortical neurons may serve a functional role in the formation of dynamic cell assemblies.     

Natural rhythm: evidence for occult 40 Hz gamma oscillation in resting motor cortex

Fast gamma oscillations, often at 40 Hz, have been demonstrated throughout the brain including the thalamus, auditory, visual and motor cortices. The function of gamma rhythms is elusive, but several authors have hypothesized that they contribute to the "binding" of diverse information into a single coherent percept, and to the synchronization of movement. In skeletal muscle a "Piper rhythm" around 40 Hz is commonly observed during maximal voluntary contraction, and has been shown to correlate with activity of similar frequency in a limited area of contralateral motor cortex. Gamma rhythms are detected primarily during complex cortical activity, and are seldom recorded at rest or coherently over wide areas. Here we use bihemispheric transcranial magnetic stimulation (TMS) to study time-dependent correlations between evoked motor potentials from non-homologous muscles in opposite limbs of normal volunteers. The results suggest the presence of an occult, synchronous 40 Hz rhythm across broad areas of resting motor cortex in both hemispheres.     

Phase coupling between different motor areas during tongue-movement imagery

Motor imagery can be accompanied by an enhancement of brain oscillations (event-related synchronization, ERS) within specific frequency bands. To characterize the neuronal couplings involved during these prominent power changes, we have chosen a certain coupling measure that bears directly on the issue of transient cortical connections. Specifically, we applied for the first time the phase-locking value to investigate the phase coupling of sensorimotor rhythms in different motor areas during tongue-movement imagery. Most interesting, we showed that robust neuronal couplings within the alpha frequency range are established between the midcentral position and bilateral central electrode positions, overlying the supplementary motor area (SMA) and the right and left primary sensorimotor area, respectively. In contrast, no direct linkage was present between sensorimotor rhythms in both hemispheres. We suggest that the coupling results point at a separate interplay between neural networks within the SMA and lateralized networks in primary sensorimotor areas of each hemisphere during motor imagery.     

fMRI signal increases and decreases in cortical areas during small-field optokinetic stimulation and central fixation

Small-field optokinetic nystagmus (OKN) was performed in seven healthy volunteers in order to analyze the activation and deactivation patterns of visual motion, ocular motor, and multisensory vestibular cortex areas by means of fMRI during coherent visual motion stimulation. BOLD signal decreases (deactivations) were found in the first and second long insular gyri and retroinsular areas (the human homologue of the parietoinsular vestibular cortex and the visual posterior sylvian area in the monkey) of both hemispheres, extending into the transverse temporal gyrus and inferior-anterior parts of the superior temporal gyrus (BA 22), and the precentral gyri at two separate sites (BA 4 and 6). Further deactivations were found in cranioposterior parts of the superior temporal gyrus (BA 22) and the adjacent inferior parietal lobule (BA 40), anterior cingulate gyrus, hippocampus, and corpus callosum. Most of these BOLD signal decreases involved parts of the "multisensory vestibular cortical circuit". These findings support the concept of a reciprocally inhibitory visual–vestibular interaction that has now been demonstrated not only for large-field visual motion stimulation that induces vection (without eye movements) but also for optokinetically induced eye movements (without vection). The functional significance of this concept may be related to the perception of self-motion, since both large-field visual motion stimulation and optokinetic nystagmus are linked to the visual control of self-motion. With respect to activation of the cortical ocular motor system two separate and distinct areas of activations were delineated in the precentral sulcus of both hemispheres, one ventrolaterally (in BA 9) and the other dorsomedially at the junction of the superior frontal sulcus with the precentral sulcus (in BA 6). Both probably correspond to different subregions of the frontal eye field and the premotor cortex for the ocular motor performance of OKN. Electronic Publication     

Kindling phenomena induced by the repeated short-term high potassium stimuli in the ventral hippocampus of rats: on-line monitoring of extracellular glutamate overflow

Recent research indicates that areas of the primary somatosensory (SI) and primary motor cortex show massive cortical reorganization after amputation of the upper arm, forearm or fingers. Most of these studies were carried out months or several years after amputation. In the present study, we describe cortical reorganization of areas in the SI of a patient who underwent amputation of the traumatized middle and ring fingers of his right hand 10 days before cortical magnetic source imaging data were obtained. Somatosensory-evoked magnetic fields (SEF) to mechanical stimuli to the finger tips were recorded and single moving dipoles were calculated using a realistic volume conductor model. Results reveal that the dipoles representing the second and fifth fingers of the affected hand were closer together than the comparable dipoles of the unaffected hand. Our findings demonstrate that neural cell assemblies in SI which formerly represented the right middle and ring fingers of this amputee became reorganized and invaded by neighbouring cell assemblies of the index and little finger of the same hand. These results indicate that functional plasticity occurs within a period of 10 days after amputation.     

Prolonged wakefulness alters neuronal responsiveness to local electrical stimulation of the neocortex in awake rats

Prolonged wakefulness or a lack of sleep lead to cognitive deficits, but little is known about the underlying cellular mechanisms. We recently found that sleep deprivation affects spontaneous neuronal activity in the neocortex of sleeping and awake rats. While it is well known that synaptic responses are modulated by ongoing cortical activity, it remains unclear whether prolonged waking affects responsiveness of cortical neurons to incoming stimuli. By applying local electrical microstimulation to the frontal area of the neocortex, we found that after a 4 h period of waking the initial neuronal response in the contralateral frontal cortex was stronger and more synchronous, and was followed by a more profound inhibition of neuronal spiking as compared with the control condition. These changes in evoked activity suggest increased neuronal excitability and indicate that, after staying awake, cortical neurons become transiently bistable. We propose that some of the detrimental effects of sleep deprivation may be a result of altered neuronal responsiveness to incoming intrinsic and extrinsic inputs.     

Correlated discharges among putative pyramidal neurons and interneurons in the primate prefrontal cortex

Neurophysiological recordings have revealed that the discharges of nearby cortical cells are positively correlated in time scales that range from millisecond synchronization of action potentials to much slower firing rate co-variations, evident in rates averaged over hundreds of milliseconds. The presence of correlated firing can offer insights into the patterns of connectivity between neurons; however, few models of population coding have taken account of the neuronal diversity present in cerebral cortex, notably a distinction between inhibitory and excitatory cells. We addressed this question in the monkey dorsolateral prefrontal cortex by recording neuronal activity from multiple micro-electrodes, typically spaced 0.2-0.3 mm apart. Putative excitatory and inhibitory neurons were distinguished based on their action potential waveform and baseline discharge rate. We tested each pair of simultaneously recorded neurons for presence of significant cross-correlation peaks and measured the correlation of their averaged firing rates in successive trials. When observed, cross-correlation peaks were centered at time 0, indicating synchronous firing consistent with two neurons receiving common input. Discharges in pairs of putative inhibitory interneurons were found to be significantly more strongly correlated than in pairs of putative excitatory cells. The degree of correlated firing was also higher for neurons with similar spatial receptive fields and neurons active in the same epochs of the behavioral task. These factors were important in predicting the strength of both short time scale (<5 ms) correlations and of trial-to-trial discharge rate covariations. Correlated firing was only marginally accounted for by motor and behavioral variations between trials. Our findings suggest that nearby inhibitory neurons are more tightly synchronized than excitatory ones and account for much of the correlated discharges commonly observed in undifferentiated cortical networks. In contrast, the discharge of pyramidal neurons, the sole projection cells of the cerebral cortex, appears largely independent, suggesting that correlated firing may be a property confined within local circuits and only to a lesser degree propagated to distant cortical areas and modules.