Models of cortical neuronal circuits commonly depend on inhibitory feedback to control gain, provide signal normalization, and to selectively amplify signals using winner-take-all (WTA) dynamics. Such models generally assume that excitatory and inhibitory neurons are able to interact easily, because their axons and dendrites are co-localized in the same small volume. However, quantitative neuroanatomical studies of the dimensions of axonal and dendritic trees of neurons in the neocortex show that this co-localization assumption is not valid. In this paper we describe a simple modification to the WTA circuit design that permits the effects of distributed inhibitory neurons to be coupled through synchronization, and so allows a single WTA to be distributed widely in cortical space, well beyond the arborization of any single inhibitory neuron, and even across different cortical areas. We prove by non-linear contraction analysis, and demonstrate by simulation that distributed WTA sub-systems combined by such inhibitory synchrony are inherently stable. We show analytically that synchronization is substantially faster than winner selection. This circuit mechanism allows networks of independent WTAs to fully or partially compete with each other.
The neocortex has a remarkably uniform neuronal organization, suggesting that common principles of processing are employed throughout its extent. In particular, the patterns of connectivity observed in the superficial layers of the visual cortex are consistent with the recurrent excitation and inhibitory feedback required for cooperative-competitive circuits such as the soft winner-take-all (WTA). WTA circuits offer interesting computational properties such as selective amplification, signal restoration, and decision making. But, these properties depend on the signal gain derived from positive feedback, and so there is a critical trade-off between providing feedback strong enough to support the sophisticated computations, while maintaining overall circuit stability. We consider the question of how to reason about stability in very large distributed networks of such circuits. We approach this problem by approximating the regular cortical architecture as many interconnected cooperative-competitive modules. We demonstrate that by properly understanding the behavior of this small computational module, one can reason over the stability and convergence of very large networks composed of these modules. We obtain parameter ranges in which the WTA circuit operates in a high-gain regime, is stable, and can be aggregated arbitrarily to form large stable networks. We use nonlinear Contraction Theory to establish conditions for stability in the fully nonlinear case, and verify these solutions using numerical simulations. The derived bounds allow modes of operation in which the WTA network is multi-stable and exhibits state-dependent persistent activities. Our approach is sufficiently general to reason systematically about the stability of any network, biological or technological, composed of networks of small modules that express competition through shared inhibition.
Although conditional branching between possible behavioural states is a hallmark of intelligent behavior, very little is known about the neuronal mechanisms that support this processing. In a step toward solving this problem we demonstrate by theoretical analysis and simulation how networks of richly inter-connected neurons, such as those observed in the superficial layers of the neocortex, can embed reliable robust finite state machines. We show how a multi-stable neuronal network containing a number of states can be created very simply, by coupling two recurrent networks whose synaptic weights have been configured for soft winner-take-all (sWTA) performance. These two sWTAs have simple, homogenous locally recurrent connectivity except for a small fraction of recurrent cross-connections between them, which are used to embed the required states. This coupling between the maps allows the network to continue to express the current state even after the input that elicted that state is withdrawn. In addition, a small number of 'transition neurons' implement the necessary input-driven transitions between the embedded states. We provide simple rules to systematically design and construct neuronal state machines of this kind. The significance of our finding is that it offers a method whereby the cortex could construct networks supporting a broad range of sophisticated processing by applying only small specializations to the same generic neuronal circuit.