Zipser D, Kehoe B, Littlewort G, Fuster J
Department of Cognitive Science, University of California, San Diego, La Jolla 92093.
J Neurosci. 1993 Aug;13(8):3406-20. doi: 10.1523/JNEUROSCI.13-08-03406.1993.
Studies of cortical neurons in monkeys performing short-term memory tasks have shown that information about a stimulus can be maintained by persistent neuron firing for periods of many seconds after removal of the stimulus. The mechanism by which this sustained activity is initiated and maintained is unknown. In this article we present a spiking neural network model of short-term memory and use it to investigate the hypothesis that recurrent, or "re-entrant," networks with constant connection strengths are sufficient to store graded information temporarily. The synaptic weights that enable the network to mimic the input-output characteristics of an active memory module are computed using an optimization procedure for recurrent networks with non-spiking neurons. This network is then transformed into one with spiking neurons by interpreting the continuous output values of the nonspiking model neurons as spiking probabilities. The behavior of the model neurons in this spiking network is compared with that of 179 single units previously recorded in monkey inferotemporal (IT) cortex during the performance of a short-term memory task. The spiking patterns of almost every model neuron are found to resemble closely those of IT neurons. About 40% of the IT neuron firing patterns are also found to be of the same types as those of model neurons. A property of the spiking model is that the neurons cannot maintain precise graded activity levels indefinitely, but eventually relax to one of a few constant activities called fixed-point attractors. The noise introduced into the model by the randomness of spiking causes the network to jump between these attractors. This switching between attractor states generates spike trains with a characteristic statistical temporal structure. We found evidence for the same kind of structure in the spike trains from about half of the IT neurons in our test set. These results show that the behavior of many real cortical memory neurons is consistent with an active storage mechanism based on recurrent activity in networks with fixed synaptic strengths.
对执行短期记忆任务的猴子的皮层神经元研究表明,在刺激移除后,有关刺激的信息可通过神经元持续放电持续数秒来维持。这种持续活动启动和维持的机制尚不清楚。在本文中,我们提出了一个短期记忆的脉冲神经网络模型,并用它来研究这样一个假设:具有恒定连接强度的循环或“折返”网络足以临时存储分级信息。使用针对非脉冲神经元的循环网络的优化程序,计算出使网络能够模拟活跃记忆模块输入输出特性的突触权重。然后,通过将非脉冲模型神经元的连续输出值解释为脉冲发放概率,将该网络转换为具有脉冲神经元的网络。将这个脉冲网络中模型神经元的行为与之前在猴子颞下(IT)皮层执行短期记忆任务期间记录的179个单个神经元的行为进行比较。发现几乎每个模型神经元的脉冲发放模式都与IT神经元的模式非常相似。还发现约40%的IT神经元发放模式与模型神经元的模式属于同一类型。脉冲模型的一个特性是,神经元不能无限期地维持精确的分级活动水平,而是最终松弛到少数几个称为定点吸引子的恒定活动之一。脉冲发放的随机性引入模型的噪声导致网络在这些吸引子之间跳跃。吸引子状态之间的这种切换产生具有特征性统计时间结构的脉冲序列。我们在测试集中约一半的IT神经元的脉冲序列中发现了这种相同类型结构的证据。这些结果表明,许多真实皮层记忆神经元的行为与基于具有固定突触强度的网络中的循环活动的活跃存储机制一致。