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OPEN SCIENCE, VERSION 3.0: Breaking down barriers for equitable and efficient research communication
(2022)
Two generic mechanisms for emergence of direction selectivity coexist in recurrent neural networks
(2013)
Poster presentation: Twenty Second Annual Computational Neuroscience Meeting: CNS*2013. Paris, France. 13-18 July 2013.
In the mammalian visual cortex, the time-averaged response of many neurons is maximal for stimuli moving in a particular direction. Such a direction selective response is not found in LGN, upstream of the visual processing pathway, suggesting that cortical networks play a strong role in the generation of direction selectivity. Here we investigate the mechanisms for the emergence of direction selectivity in the recurrent networks of nonlinear firing rate neurons in layer 4 of V1 receiving the input from LGN. In the model the LGN inputs are characterized by different receptive field positions, and their relative temporal phase shifts are reversed for the stimuli moving in the opposite direction. We propose that two distinct mechanisms result in the neuronal direction selective response in these recurrent networks. The first one is a result of nonlinear feed-forward summation of several time-shifted inputs. The second mechanism is based on the competition between neurons for firing in a winner-take-all regime. Both mechanisms rely on inhibitory interactions in the connectivity matrix of lateral connections, but the second one involves inhibitory loops. Typically, the first mechanism results in lower selectivity values than the second, but the time-course of acquiring direction selective response is faster for the first mechanism. Importantly, the two mechanisms have different input frequency tuning. The first mechanism, based on the nonlinear summation, result in a relatively narrow tuning curve around the preferred frequency of the stimulus in the case of the moving grating. In contrast, the direction selectivity arising from the second mechanism depends only weakly on the input frequency, i.e. has a broader tuning curve. These differences allow us to provide the recipe for identifying in experiment which of the two mechanisms is used by a given direction selective neuron. We then analyze how the statistics of the connections in the random recurrent networks affect the relative contributions from these two mechanisms and determine the distributions of the direction selectivity values. We identify the motifs in the connectivity matrix, which are required for each mechanism and show that the minimal conditions for both mechanisms are met in a very broad set of random recurrent networks with sufficiently strong inhibitory connections. Thus, we propose that these mechanisms coexist in generic recurrent networks with inhibition. Our results may account for the recent experimental observations that direction selectivity is present in dark-reared mice and ferrets [1,2]. It can also explain the emergence of direction selectivity in species lacking a spatially organized direction selectivity map.
This paper considers the logic FOcard, i.e., first-order logic with cardinality predicates that can specify the size of a structure modulo some number. We study the expressive power of FOcard on the class of languages of ranked, finite, labelled trees with successor relations. Our first main result characterises the class of FOcard-definable tree languages in terms of algebraic closure properties of the tree languages. As it can be effectively checked whether the language of a given tree automaton satisfies these closure properties, we obtain a decidable characterisation of the class of regular tree languages definable in FOcard. Our second main result considers first-order logic with unary relations, successor relations, and two additional designated symbols < and + that must be interpreted as a linear order and its associated addition. Such a formula is called addition-invariant if, for each fixed interpretation of the unary relations and successor relations, its result is independent of the particular interpretation of < and +. We show that the FOcard-definable tree languages are exactly the regular tree languages definable in addition-invariant first-order logic. Our proof techniques involve tools from algebraic automata theory, reasoning with locality arguments, and the use of logical interpretations. We combine and extend methods developed by Benedikt and Segoufin (ACM ToCL, 2009) and Schweikardt and Segoufin (LICS, 2010).