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1.
Fried I  Mukamel R  Kreiman G 《Neuron》2011,69(3):548-562
Understanding how self-initiated behavior is encoded by neuronal circuits in the human brain remains elusive. We recorded the activity of 1019 neurons while twelve subjects performed self-initiated finger movement. We report progressive neuronal recruitment over ~1500 ms before subjects report making the decision to move. We observed progressive increase or decrease in neuronal firing rate, particularly in the supplementary motor area (SMA), as the reported time of decision was approached. A population of 256 SMA neurons is sufficient to predict in single trials the impending decision to move with accuracy greater than 80% already 700 ms prior to subjects' awareness. Furthermore, we predict, with a precision of a few hundred ms, the actual time point of this voluntary decision to move. We implement a computational model whereby volition emerges once a change in internally generated firing rate of neuronal assemblies crosses a threshold.  相似文献   

2.
Neurotropin (Nippon Zoki Co, Ltd) effects on firing patterns of the CA 3 hippocampus neurons in rabbits under a 24-h food deprivation was studied. Neuronal firing was recorded in a state of hunger (40 neurons) and under neurotropin (27 neurons) injected through a catheter implanted into the lateral brain ventricles. Initially, in 20% of neurons histograms had a bimodal interspike interval distribution: 1.5-25 ms and 100-400 ms. Neurotropin (50 microliters) increased the baseline spike rate regularity of hippocampus neurons. Histograms had a monomodal interspike interval distribution. Neurotropin (70 microliters) inhibited firing patterns and histograms, had trimodal interspike interval distribution: 1.5-5 ms; 250-400 ms and 1000 ms. Thus, these data suggested the involvement of neuro-immuno-modulator mechanisms in organization of firing patterns in rabbits.  相似文献   

3.
In the sensorimotor cortex of rabbits with a hidden focus of excitation in the CNS, the firing rate of neurons that responded to light was significantly lower (p = 0.01) than the firing rate of neurons that did not respond to light. The same phenomenon was observed in the visual cortex of intact rabbits. Both in intact rabbits and animals with the hidden focus of excitation, 36% of neurons in the sensorimotor responded to a nonspecific for them light stimulation. In the sensorimotor cortex of rabbits with the hidden focus of excitation, more (p = 0.01) neurons responded to light with the latency lower that 100 ms and less (p = 0.02) neurons responded to light with the latency from 200 to 300 ms as compared to intact animals. In the visual cortex of rabbits with the hidden excitation focus, less (p = 0.01) neurons responded to light stimulation with the latency from 50 to 100 ms as compared to intact rabbits.  相似文献   

4.
RV Florian 《PloS one》2012,7(8):e40233
In many cases, neurons process information carried by the precise timings of spikes. Here we show how neurons can learn to generate specific temporally precise output spikes in response to input patterns of spikes having precise timings, thus processing and memorizing information that is entirely temporally coded, both as input and as output. We introduce two new supervised learning rules for spiking neurons with temporal coding of information (chronotrons), one that provides high memory capacity (E-learning), and one that has a higher biological plausibility (I-learning). With I-learning, the neuron learns to fire the target spike trains through synaptic changes that are proportional to the synaptic currents at the timings of real and target output spikes. We study these learning rules in computer simulations where we train integrate-and-fire neurons. Both learning rules allow neurons to fire at the desired timings, with sub-millisecond precision. We show how chronotrons can learn to classify their inputs, by firing identical, temporally precise spike trains for different inputs belonging to the same class. When the input is noisy, the classification also leads to noise reduction. We compute lower bounds for the memory capacity of chronotrons and explore the influence of various parameters on chronotrons' performance. The chronotrons can model neurons that encode information in the time of the first spike relative to the onset of salient stimuli or neurons in oscillatory networks that encode information in the phases of spikes relative to the background oscillation. Our results show that firing one spike per cycle optimizes memory capacity in neurons encoding information in the phase of firing relative to a background rhythm.  相似文献   

5.
Reward-modulated spike-timing-dependent plasticity (STDP) has recently emerged as a candidate for a learning rule that could explain how behaviorally relevant adaptive changes in complex networks of spiking neurons could be achieved in a self-organizing manner through local synaptic plasticity. However, the capabilities and limitations of this learning rule could so far only be tested through computer simulations. This article provides tools for an analytic treatment of reward-modulated STDP, which allows us to predict under which conditions reward-modulated STDP will achieve a desired learning effect. These analytical results imply that neurons can learn through reward-modulated STDP to classify not only spatial but also temporal firing patterns of presynaptic neurons. They also can learn to respond to specific presynaptic firing patterns with particular spike patterns. Finally, the resulting learning theory predicts that even difficult credit-assignment problems, where it is very hard to tell which synaptic weights should be modified in order to increase the global reward for the system, can be solved in a self-organizing manner through reward-modulated STDP. This yields an explanation for a fundamental experimental result on biofeedback in monkeys by Fetz and Baker. In this experiment monkeys were rewarded for increasing the firing rate of a particular neuron in the cortex and were able to solve this extremely difficult credit assignment problem. Our model for this experiment relies on a combination of reward-modulated STDP with variable spontaneous firing activity. Hence it also provides a possible functional explanation for trial-to-trial variability, which is characteristic for cortical networks of neurons but has no analogue in currently existing artificial computing systems. In addition our model demonstrates that reward-modulated STDP can be applied to all synapses in a large recurrent neural network without endangering the stability of the network dynamics.  相似文献   

6.
Ca(v)1.2 L-type calcium channels support hippocampal synaptic plasticity, likely by facilitating dendritic Ca2+ influx evoked by action potentials (AP) back-propagated from the soma. Ca2+ influx into hippocampal neurons during somatic APs is sufficient to activate signalling pathways associated with late phase LTP. Thus, mechanisms controlling AP firing of hippocampal neurons are of major functional relevance. We examined the excitability of CA1 pyramidal cells using somatic current-clamp recordings in brain slices from control type mice and mice with the Ca(v)1.2 gene inactivated in principal hippocampal neurons. Lack of the Ca(v)1.2 protein did not affect either affect basic characteristics, such as resting membrane potential and input resistance, or parameters of single action potentials (AP) induced by 5 ms depolarising current pulses. However, CA1 hippocampal neurons from control and mutant mice differed in their patterns of AP firing during 500 ms depolarising current pulses: threshold voltage for repetitive firing was shifted significantly by about 5 mV to more depolarised potentials in the mutant mice (p<0.01), and the latency until firing of the first AP was prolonged (73.2+/-6.6 ms versus 48.1+/- 7.8 ms in control; p<0.05). CA1 pyramidal cells from the mutant mice also showed a lowered initial spiking frequency within an AP train. In control cells, isradipine had matching effects, while BayK 8644 facilitated spiking. Our data demonstrate that Ca(v)1.2 channels are involved in regulating the intrinsic excitability of CA1 pyramidal neurons. This cellular mechanism may contribute to the known function of Ca(v)1.2 channels in supporting synaptic plasticity and memory.  相似文献   

7.
Neurons in the primary visual cortex typically reach their highest firing rate after an abrupt image transition. Since the mutual information between the firing rate and the currently presented image is largest during this early firing period it is tempting to conclude this early firing encodes the current image. This view is, however, made more complicated by the fact that the response to the current image is dependent on the preceding image. Therefore we hypothesize that neurons encode a combination of current and previous images, and that the strength of the current image relative to the previous image changes over time. The temporal encoding is interesting, first, because neurons are, at different time points, sensitive to different features such as luminance, edges and textures; second, because the temporal evolution provides temporal constraints for deciphering the instantaneous population activity. To study the temporal evolution of the encoding we presented a sequence of 250 ms stimulus patterns during multiunit recordings in areas 17 and 18 of the anaesthetized ferret. Using a novel method we decoded the pattern given the instantaneous population-firing rate. Following a stimulus transition from stimulus A to B the decoded stimulus during the first 90ms was more correlated with the difference between A and B (B-A) than with B alone. After 90ms the decoded stimulus was more correlated with stimulus B than with B-A. Finally we related our results to information measures of previous (B) and current stimulus (A). Despite that the initial transient conveys the majority of the stimulus-related information; we show that it actually encodes a difference image which can be independent of the stimulus. Only later on, spikes gradually encode the stimulus more exclusively.  相似文献   

8.
Star-nosed moles have a series of mechanosensory appendages surrounding each nostril. Each appendage is covered with sensory organs (Eimer's organs) containing both rapidly adapting and slowly adapting mechanoreceptors and each appendage is represented in primary somatosensory cortex (S1) by a single cortical module. When the skin surface of an appendage is depressed, neurons in the corresponding module in S1 respond in either a transient or sustained fashion. The aim of this study was to characterize and compare the responses of these two classes of neurons to both short (5 or 20 ms) and long (500 ms) mechanosensory stimulation. Activity from neurons in the representation of appendage 11, the somatosensory fovea, was recorded while delivering mechanosensory stimuli to the corresponding skin surface. Transient and sustained neurons had different levels of spontaneous activity and different responses to both short and long mechanosensory stimulation. Neurons with sustained responses had a significantly higher spontaneous firing rate than neurons with transient responses. Transient neurons responded to a 5 ms stimulus with excitation followed by suppression of discharge whereas sustained neurons did not exhibit post-excitatory suppression. Rather, responses of sustained neurons to 5 ms stimuli lasted several hundred milliseconds. Consequently sustained responses contained significantly more spikes than transient responses. These experiments suggest contact to the appendages causes two distinct firing patterns in cortex regardless of the duration of the stimulus. The sustained and transient responses could reflect either the activity of fundamentally different classes of neurons or activity in distinct subcortical and cortical networks.  相似文献   

9.
Most neurons in the primary visual cortex initially respond vigorously when a preferred stimulus is presented, but adapt as stimulation continues. The functional consequences of adaptation are unclear. Typically a reduction of firing rate would reduce single neuron accuracy as less spikes are available for decoding, but it has been suggested that on the population level, adaptation increases coding accuracy. This question requires careful analysis as adaptation not only changes the firing rates of neurons, but also the neural variability and correlations between neurons, which affect coding accuracy as well. We calculate the coding accuracy using a computational model that implements two forms of adaptation: spike frequency adaptation and synaptic adaptation in the form of short-term synaptic plasticity. We find that the net effect of adaptation is subtle and heterogeneous. Depending on adaptation mechanism and test stimulus, adaptation can either increase or decrease coding accuracy. We discuss the neurophysiological and psychophysical implications of the findings and relate it to published experimental data.  相似文献   

10.
1. The first part of this study looks at spontaneously active neurons located in the rostral ventrolateral medulla (RVLM) with projections to the thoracic spinal cord. Sixteen neurons were intracellularly recorded in vivo. Four out of 16 neurons were antidromically activated from the thoracic spinal cord (axonal conduction velocities varied from 1.8 m/s to 9.5 m/s).2. The simultaneous averages of the neuronal membrane potential and arterial blood pressure triggered by the pulsatile arterial wave or the EKG-R wave demonstrated changes in membrane potential (hyperpolarization or depolarization) locked to the cardiac cycle in four neurons in this group. These neurons (three of them bulbospinal) were further tested for barosensitivity by characterizing the responses to electrical stimulation of the aortic depressor nerve. Four neurons responded with inhibitory hyperpolarizing responses characterized as inhibitory postsynaptic potentials (IPSP) to aortic nerve stimulation (onset latency: 32.3 ± 5.0 ms; mean ± SEM).3. In two neurons in the RVLM, one of them characterized as barosensitive, electrical stimulation of the opposite RVLM (0.5 Hz, 1.0 ms pulse duration, 25–100 A) elicited excitatory postsynaptic potentials (EPSPs) with latencies of 9.07 and 10.5 ms. At resting membrane potential, the onset latency of the evoked EPSPs did not change with increasing stimulus intensities. Some of the recorded neurons were intracellularly labelled with biocytin for visualization. They were found in the RVLM.4. These experiments in vivo would support the idea of a functional commissural pathway between the RVLM of both sides.5. Anatomical data have shown that some of those commissural bundle fibers originate in the C1 adrenergic neuronal group in the RVLM. In the second part of this study, we used an intracellular recording technique in vitro to investigate the effects of the indirect adrenergic agonist tyramine on neurons in the RVLM with electrophysiological properties similar to premotor sympathetic neurons in vivo.6. Tyramine (0.5–1 mM) produced a pronounced inhibitory effect with hyperpolarization and increase in membrane input resistance on two neurons characterized as regularly firing (R), and on one neuron characterized as irregularly firing (I). This effect was preceded by a transient depolarization with increases in firing rate.7. These results would indicate that neurons in the RVLM recorded in vitro and with properties similar to premotor sympathetic neurons can be modulated by catecholamines released from terminals probably making synaptic contacts.  相似文献   

11.
Neuronal cortical spike trains contain precisely replicating patterns whose presence cannot be accounted for by chance production. A comparison of the number of triplets of spikes present two times with the number of doublets replicated three times in the same window duration gives a frequency-insensitive measure of this type of fine temporal organisation. By varying the tolerance with which such precisely replicating patterns are detected, one can evaluate the accuracy of spike timing in spike trains. In the sample of data here analysed, it was found that replicating patterns were best seen in the precision range 0.4–1.4 ms (a result evidently at variance with a simple ‘integrate and fire’ model of neurons). Surprisingly, the fine temporal structure of spike trains thus evidenced was stronger at relatively low firing rate discharges and was present in both the ‘spontaneous’ and ‘evoked’ responses.  相似文献   

12.
Odor perception depends on the odorant-evoked changes on Mitral/Tufted cell firing pattern within the olfactory bulb (OB). The OB exhibits a significant "ongoing" or spontaneous activity in the absence of sensory stimulation. We characterized this ongoing activity by simultaneously recording several single neurons in the mitral cell layer (MCL) of anesthetized rats and determined the extent of synchrony and oscillations under nasal and tracheal breathing. We recorded 115 neurons and found no significant differences in the mean firing rates between both breathing conditions. Surprisingly, nearly all single units exhibited a long refractory period averaging 14.4 ms during nasal respiration that was not different under tracheal breathing. We found a small incidence (2% of neurons) of gamma band oscillations and a low incidence (8.1%) of correlated firing between adjacent MCL cells. During nasal respiration, a significant oscillation at the respiratory rate was observed in 12% of cells that disappeared during tracheal breathing. Thus, in the absence of odorants, MCL cells exhibit a long refractory period, probably reflecting the intrinsic OB network properties. Furthermore, in the absence of sensory stimulation, MCL cell discharge does not oscillate in the gamma band and the respiratory cycle can modulate the firing of these cells.  相似文献   

13.
We used phase resetting methods to predict firing patterns of rat subthalamic nucleus (STN) neurons when their rhythmic firing was densely perturbed by noise. We applied sequences of contiguous brief (0.5–2 ms) current pulses with amplitudes drawn from a Gaussian distribution (10–100 pA standard deviation) to autonomously firing STN neurons in slices. Current noise sequences increased the variability of spike times with little or no effect on the average firing rate. We measured the infinitesimal phase resetting curve (PRC) for each neuron using a noise-based method. A phase model consisting of only a firing rate and PRC was very accurate at predicting spike timing, accounting for more than 80% of spike time variance and reliably reproducing the spike-to-spike pattern of irregular firing. An approximation for the evolution of phase was used to predict the effect of firing rate and noise parameters on spike timing variability. It quantitatively predicted changes in variability of interspike intervals with variation in noise amplitude, pulse duration and firing rate over the normal range of STN spontaneous rates. When constant current was used to drive the cells to higher rates, the PRC was altered in size and shape and accurate predictions of the effects of noise relied on incorporating these changes into the prediction. Application of rate-neutral changes in conductance showed that changes in PRC shape arise from conductance changes known to accompany rate increases in STN neurons, rather than the rate increases themselves. Our results show that firing patterns of densely perturbed oscillators cannot readily be distinguished from those of neurons randomly excited to fire from the rest state. The spike timing of repetitively firing neurons may be quantitatively predicted from the input and their PRCs, even when they are so densely perturbed that they no longer fire rhythmically.  相似文献   

14.
The activity patterns of rhythmically firing neurons in monkey primary somatosensory cortex (SI) were studied during trained wrist movements that were performed in response to palmar vibration. Of 1,222 neurons extracellularly recorded in SI, 129 cells (11%) discharged rhythmically (at 30 Hz) during maintained wrist position. During the initiation of vibratory-cued movements, neuronal activity usually decreased at 25 ms after vibration onset followed by an additional decrease in activity at 60 ms prior to movement onset. Rhythmically firing neurons are not likely to be integrate-and-fire neurons because, during activity changes, their rhythmic firing pattern was disrupted rather than modulated. The activity pattern of rhythmically firing neurons was complimentary to that of quickly adapting SI neurons recorded during the performance of this task (Nelson et al., 1991). Moreover, disruptions of rhythmic activity of individual SI neurons were similar to those reported previously for local field potential (LFP) oscillations in sensorimotor cortex during trained movements (Sanes and Donoghue, 1993). However, rhythmic activity of SI neurons did not wax and wane like LFP oscillations (Murthy and Fetz, 1992; Sanes and Donoghue, 1993). It has been suggested that fast (20–50 Hz) cortical oscillations may be initiated by inhibitory interneurons (Cowan and Wilson, 1994; Llinas et al., 1991; Stern and Wilson, 1994). We suggest that rhythmically firing neurons may tonically inhibit quickly adapting neurons and release them from the inhibition at go-cue onsets and prior to voluntary movements. It is possible that rhythmically active neurons may evoke intermittent oscillations in other cortical neurons and thus regulate cortical population oscillations.  相似文献   

15.
通常采用恒定电脉冲间隔的高频刺激(high-frequency stimulation,HFS),进行深部脑刺激治疗帕金森氏症等运动障碍疾病.为了开发适用于不同脑疾病治疗的新刺激模式,近年来脉冲间隔(inter-pulse-interval,IPI)变化的变频刺激模式受到关注.已有研究表明,即使具有相同的平均电脉冲频率,变频刺激与恒频刺激的治疗效果也不同.我们推测,变频刺激的短小IPI变化就足以改变HFS对于神经元的作用.为了验证此推测,本文在大鼠海马CA1区锥体神经元的输入轴突纤维上交替施加恒频刺激(100或133 Hz,即IPI=10 ms或7.5 ms)和随机变频刺激(100~200 Hz,即IPI=5~10 ms,平均频率为133 Hz),记录并分析刺激下游神经元群体的诱发电位,用于定量评价神经元对于恒频和变频刺激的响应.实验结果表明,持续的恒频刺激使得神经元的响应从最初的同步发放形成的群峰电位(population spike,PS)转变为非同步的动作电位发放(即单元锋电位).但是,当刺激切换为变频模式时,却又可以诱发神经元群体同步产生动作电位,重新形成PS波.并且,变频刺激诱发的PS幅值和神经元发放的同步程度可达基线的单脉冲刺激诱发波的水平.但是,PS的发生率只有脉冲刺激频率的7%左右,表明在持续的变频刺激时,多个脉冲累积的作用才能诱发这种同步的神经元发放.而且PS的出现与前导IPI的长度之间存在一定关系.神经元的轴突和突触等结构对于高频刺激的非线性响应可能是变频刺激诱发同步活动的原因.这些结果表明,变频刺激序列中短小的间隔变化可以产生与恒定间隔不同的调控作用.本文的结果对于揭示脑刺激的作用机制,促进新型刺激模式的开发及其在不同类型脑疾病治疗中的应用具有重要意义.  相似文献   

16.
Spike timing-dependent plasticity (STDP) has been shown to enable single neurons to detect repeatedly presented spatiotemporal spike patterns. This holds even when such patterns are embedded in equally dense random spiking activity, that is, in the absence of external reference times such as a stimulus onset. Here we demonstrate, both analytically and numerically, that STDP can also learn repeating rate-modulated patterns, which have received more experimental evidence, for example, through post-stimulus time histograms (PSTHs). Each input spike train is generated from a rate function using a stochastic sampling mechanism, chosen to be an inhomogeneous Poisson process here. Learning is feasible provided significant covarying rate modulations occur within the typical timescale of STDP (~10-20 ms) for sufficiently many inputs (~100 among 1000 in our simulations), a condition that is met by many experimental PSTHs. Repeated pattern presentations induce spike-time correlations that are captured by STDP. Despite imprecise input spike times and even variable spike counts, a single trained neuron robustly detects the pattern just a few milliseconds after its presentation. Therefore, temporal imprecision and Poisson-like firing variability are not an obstacle to fast temporal coding. STDP provides an appealing mechanism to learn such rate patterns, which, beyond sensory processing, may also be involved in many cognitive tasks.  相似文献   

17.
In the nucleus interpositus (IP) of the cat cerebellum the response patterns to peripheral stimulation were recorded, and the interactions between electrophysiologically identified neurons were studied with cross correlation techniques. The response patterns were composed of excitation appearing with latencies of about 8 to 22 ms., separated and succeeded by phases of inhibitions. Four basic types of interneuronal connectivities were observed: 1) intranuclear excitation, 2) shared input from a common source, 3) intranuclear inhibition, and 4) stimulus coordinated firing. Shared input appeared in all combinations of paired neurons and extended for a distance from 300 microns up to 1000 microns. Intranuclear excitation as well as inhibitory synaptic connection occurred mainly in combinations between interneurons and efferent neurons. Stimulus coordinated firing of paired neurons was found in almost the entire extent of the nucleus interpositus.  相似文献   

18.
We examined the kinetics of voltage-dependent sodium currents in cerebellar Purkinje neurons using whole-cell recording from dissociated neurons. Unlike sodium currents in other cells, recovery from inactivation in Purkinje neurons is accompanied by a sizeable ionic current. Additionally, the extent and speed of recovery depend markedly on the voltage and duration of the prepulse that produces inactivation. Recovery is faster after brief, large depolarizations (e.g., 5 ms at +30 mV) than after long, smaller depolarizations (e.g., 100 ms at -30 mV). On repolarization to -40 mV following brief, large depolarizations, a resurgent sodium current rises and decays in parallel with partial, nonmonotonic recovery from inactivation. These phenomena can be explained by a model that incorporates two mechanisms of inactivation: a conventional mechanism, from which channels recover without conducting current, and a second mechanism, favored by brief, large depolarizations, from which channels recover by passing transiently through the open state. The second mechanism is consistent with voltage-dependent block of channels by a particle that can enter and exit only when channels are open. The sodium current flowing during recovery from this blocked state may depolarize cells immediately after an action potential, promoting the high-frequency firing typical of Purkinje neurons.  相似文献   

19.
To study plasticity, we cultured cortical networks on multielectrode arrays, enabling simultaneous recording from multiple neurons. We used conditional firing probabilities to describe functional network connections by their strength and latency. These are abstract representations of neuronal pathways and may arise from direct pathways between two neurons or from a common input. Functional connections based on direct pathways should reflect synaptic properties. Therefore, we searched for long-term potentiation (this mechanism occurs in vivo when presynaptic action potentials precede postsynaptic ones with interspike intervals up to ∼20 ms) in vitro. To investigate if the strength of functional connections showed a similar latency-related development, we selected periods of monotonously increasing or decreasing strength. We observed increased incidence of short latencies (5-30 ms) during strengthening, whereas these rarely occurred during weakening. Furthermore, we saw an increased incidence of 40-65 ms latencies during weakening. Conversely, functional connections tended to strengthen in periods with short latency, whereas strengthening was significantly less than average during long latency. Our data suggest that functional connections contain information about synaptic connections, that conditional firing probability analysis is sensitive enough to detect it and that a substantial fraction of all functional connections is based on direct pathways.  相似文献   

20.
To study the effects of different durations of forward masker sound on neuronal firing and rate-intensity function(RIF)of mouse inferior collicular(IC)neurons,a tone relative to 5 dB above the minimum threshold(re MT+5 dB)of the best frequency of recorded neurons was used as forward masker sound under free field stimulation condition.The masker durations used were 40,60,80,and 100 ms.Results showed that as masker duration was increased,inhibition in neuronal firing was enhanced(P<0.0001,n=41)and the latency of neurons was lengthened(P<0.01,n=41).In addition,among 41 inhibited IC neurons,90.2%(37/41)exhibited narrowed dynamic range(DR)when masker sound duration was increased(P<0.0001),whereas the DR of 9.8%(4/41)became wider.These data suggest that masking effects of different durations of forward masker sound might be correlated with the amplitude and duration of inhibitory input to IC neurons elicited by the masker sound.  相似文献   

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