Interference between space and time estimations: from behavior to neurons

Influences between time and space can be found in our daily life in which we are surrounded by numerous spatial metaphors to refer to time. For instance, when we move files from one folder to another in our computer a horizontal line that grows from left to right informs us about the elapsed and remaining time to finish the procedure and, similarly, in our communication we use several spatial terms to refer to time. Although with some differences in the degree of interference, not only space has an influence on time but both magnitudes influence each other. Indeed, since our childhood our estimations of time are influenced by space even when space should be irrelevant and the same occurs when estimating space with time as distractor. Such interference between magnitudes has also been observed in monkeys even if they do not use language or computers, suggesting that the two magnitudes are tightly coupled beyond communication and technology. Imaging and lesion studies have indicated that same brain areas are involved during the processing of both magnitudes and have suggested that rather than coding the specific magnitude itself the brain represents them as abstract concepts. Recent neurophysiological studies in prefrontal cortex, however, have shown that the coding of absolute and relative space and time in this area is realized by independent groups of neurons. Interestingly, instead, a high overlap was observed in this same area in the coding of goal choices across tasks. These results suggest that rather than during perception or estimation of space and time the interference between the two magnitudes might occur, at least in the prefrontal cortex, in a subsequent phase in which the goal has to be chosen or the response provided

Outcome contingency selectively affects the neural coding of outcomes but not of tasks

Value-based decision-making is ubiquitous in every-day life, and critically depends on the contingency between choices and their outcomes. Only if outcomes are contingent on our choices can we make meaningful value-based decisions. Here, we investigate the effect of outcome contingency on the neural coding of rewards and tasks. Participants performed a reversal-learning paradigm in which reward outcomes were contingent on trial-by-trial choices, and performed a ‘free choice’ paradigm in which rewards were random and not contingent on choices. We hypothesized that contingent outcomes enhance the neural coding of rewards and tasks, which was tested using multivariate pattern analysis of fMRI data. Reward outcomes were encoded in a large network including the striatum, dmPFC and parietal cortex, and these representations were indeed amplified for contingent rewards. Tasks were encoded in the dmPFC at the time of decision-making, and in parietal cortex in a subsequent maintenance phase. We found no evidence for contingency-dependent modulations of task signals, demonstrating highly similar coding across contingency conditions. Our findings suggest selective effects of contingency on reward coding only, and further highlight the role of dmPFC and parietal cortex in value-based decision-making, as these were the only regions strongly involved in both reward and task coding

Rule learning enhances structural plasticity of long-range axons in frontal cortex.

Rules encompass cue-action-outcome associations used to guide decisions and strategies in a specific context. Subregions of the frontal cortex including the orbitofrontal cortex (OFC) and dorsomedial prefrontal cortex (dmPFC) are implicated in rule learning, although changes in structural connectivity underlying rule learning are poorly understood. We imaged OFC axonal projections to dmPFC during training in a multiple choice foraging task and used a reinforcement learning model to quantify explore-exploit strategy use and prediction error magnitude. Here we show that rule training, but not experience of reward alone, enhances OFC bouton plasticity. Baseline bouton density and gains during training correlate with rule exploitation, while bouton loss correlates with exploration and scales with the magnitude of experienced prediction errors. We conclude that rule learning sculpts frontal cortex interconnectivity and adjusts a thermostat for the explore-exploit balance

The evolution of brain architectures for predictive coding and active inference

This article considers the evolution of brain architectures for predictive processing. We argue that brain mechanisms for predictive perception and action are not late evolutionary additions of advanced creatures like us. Rather, they emerged gradually from simpler predictive loops (e.g. autonomic and motor reflexes) that were a legacy from our earlier evolutionary ancestors-and were key to solving their fundamental problems of adaptive regulation. We characterize simpler-to-more-complex brains formally, in terms of generative models that include predictive loops of increasing hierarchical breadth and depth. These may start from a simple homeostatic motif and be elaborated during evolution in four main ways: these include the multimodal expansion of predictive control into an allostatic loop; its duplication to form multiple sensorimotor loops that expand an animal's behavioural repertoire; and the gradual endowment of generative models with hierarchical depth (to deal with aspects of the world that unfold at different spatial scales) and temporal depth (to select plans in a future-oriented manner). In turn, these elaborations underwrite the solution to biological regulation problems faced by increasingly sophisticated animals. Our proposal aligns neuroscientific theorising-about predictive processing-with evolutionary and comparative data on brain architectures in different animal species. This article is part of the theme issue 'Systems neuroscience through the lens of evolutionary theory'

Foraging as an evidence accumulation process

A canonical foraging task is the patch-leaving problem, in which a forager must decide to leave a current resource in search for another. Theoretical work has derived optimal strategies for when to leave a patch, and experiments have tested for conditions where animals do or do not follow an optimal strategy. Nevertheless, models of patch-leaving decisions do not consider the imperfect and noisy sampling process through which an animal gathers information, and how this process is constrained by neurobiological mechanisms. In this theoretical study, we formulate an evidence accumulation model of patch-leaving decisions where the animal averages over noisy measurements to estimate the state of the current patch and the overall environment. Evidence accumulation models belong to the class of drift diffusion processes and have been used to model decision making in different contexts. We solve the model for conditions where foraging decisions are optimal and equivalent to the marginal value theorem, and perform simulations to analyze deviations from optimal when these conditions are not met. By adjusting the drift rate and decision threshold, the model can represent different strategies, for example an increment-decrement or counting strategy. These strategies yield identical decisions in the limiting case but differ in how patch residence times adapt when the foraging environment is uncertain. To account for sub-optimal decisions, we introduce an energy-dependent utility function that predicts longer than optimal patch residence times when food is plentiful. Our model provides a quantitative connection between ecological models of foraging behavior and evidence accumulation models of decision making. Moreover, it provides a theoretical framework for potential experiments which seek to identify neural circuits underlying patch leaving decisions