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Abstract

Integrating sentence meaning over time requires memory ranging from milliseconds (words) to seconds (sentences) and minutes (discourse). How do transient events like action potentials in the human language system support memory at these different temporal scales? Here we investigate the nature of processing memory in a neurobiologically motivated model of sentence comprehension. The model was a recurrent, sparsely connected network of spiking neurons. Synaptic weights were created randomly and there was no adaptation or learning. As input the network received word sequences generated from construction grammar templates and their syntactic alternations (e.g., active/passive transitives, transfer datives, caused motion). The language environment had various features such as tense, aspect, noun/verb number agreement, and pronouns which created positional variation in the input. Similar to natural speech, word durations varied between 50ms and 0.5s of real, physical time depending on their length. The model's task was to incrementally interpret these word sequences in terms of semantic roles. There were 8 target roles (e.g., Agent, Patient, Recipient) and the language generated roughly 1,2m distinct utterances from which a sequence of 10,000 words was randomly selected and filtered through the network. A set of readout neurons was then calibrated by means of logistic regression to decode the internal network dynamics onto the target semantic roles. In order to accomplish the role assignment task, network states had to encode and maintain past information from multiple cues that could occur several words apart. To probe the circuit's memory capacity, we compared models where network connectivity, the shape of synaptic currents, and properties of neuronal adaptation were systematically manipulated. We found that task-relevant memory could be derived from a mechanism of neuronal spike-rate adaptation, modelled as a conductance that hyperpolarized the membrane following a spike and relaxed to baseline exponentially with a fixed time-constant. By acting directly on the membrane potential it provided processing memory that allowed the system to successfully interpret its sentence input. Near optimal performance was also observed when an exponential decay model of post-synaptic currents was added into the circuit, with time-constants approximating excitatory NMDA and inhibitory GABA-B receptor dynamics. Thus, the information flow was extended over time, creating memory characteristics comparable to spike-rate adaptation. Recurrent connectivity, in contrast, only played a limited role in maintaining information; an acyclic version of the recurrent circuit achieved similar accuracy. This indicates that random recurrent connectivity at the modelled spatial scale did not contribute additional processing memory to the task. Taken together, these results suggest that memory for language might be provided by activity-silent dynamic processes rather than the active replay of past input as in storage-and-retrieval models of working memory. Furthermore, memory in biological networks can take multiple forms on a continuum of time-scales. Therefore, the development of neurobiologically realistic, causal models will be critical for our understanding of the role of memory in language processing.