Mechanisms underlying contrast-dependent orientation selectivity in mouse V1.

Recent experiments have shown that mouse primary visual cortex (V1) is very different from that of cat or monkey, including response properties-one of which is that contrast invariance in the orientation selectivity (OS) of the neurons' firing rates is replaced in mouse with contrast-dependent sharpening (broadening) of OS in excitatory (inhibitory) neurons. These differences indicate a different circuit design for mouse V1 than that of cat or monkey. Here we develop a large-scale computational model of an effective input layer of mouse V1. Constrained by experiment data, the model successfully reproduces experimentally observed response properties-for example, distributions of firing rates, orientation tuning widths, and response modulations of simple and complex neurons, including the contrast dependence of orientation tuning curves. Analysis of the model shows that strong feedback inhibition and strong orientation-preferential cortical excitation to the excitatory population are the predominant mechanisms underlying the contrast-sharpening of OS in excitatory neurons, while the contrast-broadening of OS in inhibitory neurons results from a strong but nonpreferential cortical excitation to these inhibitory neurons, with the resulting contrast-broadened inhibition producing a secondary enhancement on the contrast-sharpened OS of excitatory neurons. Finally, based on these mechanisms, we show that adjusting the detailed balances between the predominant mechanisms can lead to contrast invariance-providing insights for future studies on contrast dependence (invariance).

Balanced Active Core in Heterogeneous Neuronal Networks.

It is hypothesized that cortical neuronal circuits operate in a global balanced state, i.e., the majority of neurons fire irregularly by receiving balanced inputs of excitation and inhibition. Meanwhile, it has been observed in experiments that sensory information is often sparsely encoded by only a small set of firing neurons, while neurons in the rest of the network are silent. The phenomenon of sparse coding challenges the hypothesis of a global balanced state in the brain. To reconcile this, here we address the issue of whether a balanced state can exist in a small number of firing neurons by taking account of the heterogeneity of network structure such as scale-free and small-world networks. We propose necessary conditions and show that, under these conditions, for sparsely but strongly connected heterogeneous networks with various types of single-neuron dynamics, despite the fact that the whole network receives external inputs, there is a small active subnetwork (active core) inherently embedded within it. The neurons in this active core have relatively high firing rates while the neurons in the rest of the network are quiescent. Surprisingly, although the whole network is heterogeneous and unbalanced, the active core possesses a balanced state and its connectivity structure is close to a homogeneous Erdös-Rényi network. The dynamics of the active core can be well-predicted using the Fokker-Planck equation. Our results suggest that the balanced state may be maintained by a small group of spiking neurons embedded in a large heterogeneous network in the brain. The existence of the small active core reconciles the balanced state and the sparse coding, and also provides a potential dynamical scenario underlying sparse coding in neuronal networks.