Why Neurons Have Thousands of Synapses, A Theory of Sequence Memory in Neocortex

Neocortical neurons have thousands of excitatory synapses. It is a mystery how neurons integrate the input from so many synapses and what kind of large-scale network behavior this enables. It has been previously proposed that non-linear properties of dendrites enable neurons to recognize multiple patterns. In this paper we extend this idea by showing that a neuron with several thousand synapses arranged along active dendrites can learn to accurately and robustly recognize hundreds of unique patterns of cellular activity, even in the presence of large amounts of noise and pattern variation. We then propose a neuron model where some of the patterns recognized by a neuron lead to action potentials and define the classic receptive field of the neuron, whereas the majority of the patterns recognized by a neuron act as predictions by slightly depolarizing the neuron without immediately generating an action potential. We then present a network model based on neurons with these properties and show that the network learns a robust model of time-based sequences. Given the similarity of excitatory neurons throughout the neocortex and the importance of sequence memory in inference and behavior, we propose that this form of sequence memory is a universal property of neocortical tissue. We further propose that cellular layers in the neocortex implement variations of the same sequence memory algorithm to achieve different aspects of inference and behavior. The neuron and network models we introduce are robust over a wide range of parameters as long as the network uses a sparse distributed code of cellular activations. The sequence capacity of the network scales linearly with the number of synapses on each neuron. Thus neurons need thousands of synapses to learn the many temporal patterns in sensory stimuli and motor sequences.Comment: Submitted for publicatio

Evolving Dendritic Morphologies Highlight the Impact of Structured Synaptic Inputs on Neuronal Performance

Acknowledgements I would like to express my sincere gratitude to Dr. Rene te Boekhorst for his valued support and guidance extended to me.Postprin

Comparison Between Supervised and Unsupervised Classifications of Neuronal Cell Types: A Case Study

In the study of neural circuits, it becomes essential to discern the different neuronal cell types that build the circuit. Traditionally, neuronal cell types have been classified using qualitative descriptors. More recently, several attempts have been made to classify neurons quantitatively, using unsupervised clustering methods. While useful, these algorithms do not take advantage of previous information known to the investigator, which could improve the classification task. For neocortical GABAergic interneurons, the problem to discern among different cell types is particularly difficult and better methods are needed to perform objective classifications. Here we explore the use of supervised classification algorithms to classify neurons based on their morphological features, using a database of 128 pyramidal cells and 199 interneurons from mouse neocortex. To evaluate the performance of different algorithms we used, as a “benchmark,” the test to automatically distinguish between pyramidal cells and interneurons, defining “ground truth” by the presence or absence of an apical dendrite. We compared hierarchical clustering with a battery of different supervised classification algorithms, finding that supervised classifications outperformed hierarchical clustering. In addition, the selection of subsets of distinguishing features enhanced the classification accuracy for both sets of algorithms. The analysis of selected variables indicates that dendritic features were most useful to distinguish pyramidal cells from interneurons when compared with somatic and axonal morphological variables. We conclude that supervised classification algorithms are better matched to the general problem of distinguishing neuronal cell types when some information on these cell groups, in our case being pyramidal or interneuron, is known a priori. As a spin-off of this methodological study, we provide several methods to automatically distinguish neocortical pyramidal cells from interneurons, based on their morphologies