Gain Modulation by an Urgency Signal Controls the Speed–Accuracy Trade-Off in a Network Model of a Cortical Decision Circuit

The speed–accuracy trade-off (SAT) is ubiquitous in decision tasks. While the neural mechanisms underlying decisions are generally well characterized, the application of decision-theoretic methods to the SAT has been difficult to reconcile with experimental data suggesting that decision thresholds are inflexible. Using a network model of a cortical decision circuit, we demonstrate the SAT in a manner consistent with neural and behavioral data and with mathematical models that optimize speed and accuracy with respect to one another. In simulations of a reaction time task, we modulate the gain of the network with a signal encoding the urgency to respond. As the urgency signal builds up, the network progresses through a series of processing stages supporting noise filtering, integration of evidence, amplification of integrated evidence, and choice selection. Analysis of the network's dynamics formally characterizes this progression. Slower buildup of urgency increases accuracy by slowing down the progression. Faster buildup has the opposite effect. Because the network always progresses through the same stages, decision-selective firing rates are stereotyped at decision time

Dynamic excitatory and inhibitory gain modulation can produce flexible, robust and optimal decision-making

<div><p>Behavioural and neurophysiological studies in primates have increasingly shown the involvement of urgency signals during the temporal integration of sensory evidence in perceptual decision-making. Neuronal correlates of such signals have been found in the parietal cortex, and in separate studies, demonstrated attention-induced gain modulation of both excitatory and inhibitory neurons. Although previous computational models of decision-making have incorporated gain modulation, their abstract forms do not permit an understanding of the contribution of inhibitory gain modulation. Thus, the effects of co-modulating both excitatory and inhibitory neuronal gains on decision-making dynamics and behavioural performance remain unclear. In this work, we incorporate time-dependent co-modulation of the gains of both excitatory and inhibitory neurons into our previous biologically based decision circuit model. We base our computational study in the context of two classic motion-discrimination tasks performed in animals. Our model shows that by simultaneously increasing the gains of both excitatory and inhibitory neurons, a variety of the observed dynamic neuronal firing activities can be replicated. In particular, the model can exhibit winner-take-all decision-making behaviour with higher firing rates and within a significantly more robust model parameter range. It also exhibits short-tailed reaction time distributions even when operating near a dynamical bifurcation point. The model further shows that neuronal gain modulation can compensate for weaker recurrent excitation in a decision neural circuit, and support decision formation and storage. Higher neuronal gain is also suggested in the more cognitively demanding reaction time than in the fixed delay version of the task. Using the exact temporal delays from the animal experiments, fast recruitment of gain co-modulation is shown to maximize reward rate, with a timescale that is surprisingly near the experimentally fitted value. Our work provides insights into the simultaneous and rapid modulation of excitatory and inhibitory neuronal gains, which enables flexible, robust, and optimal decision-making.</p></div

Categorical evidence, confidence and urgency during the integration of multi-feature information

Includes bibliographical references.2015 Summer.The present experiment utilized a temporally-extended categorization task to investigate the neural substrates underlying our ability to integrate information over time and across multiple stimulus features. Importantly, the design allowed differentiation of three important decision functions: 1) categorical evidence, 2) decisional confidence (the choice-independent probability that a decision will lead to a desirable state), and 3) urgency (a hypothetical signal representing a growing pressure to produce a behavioral response within each trial). In conjunction with model-based fMRI, the temporal evolution of these variables were tracked as participants deliberated about impending choices. The approach allowed investigation of the independent effects of urgency across the brain, and also the investigation of how urgency might modulate representations of categorical evidence and confidence. Representations associated with prediction errors during feedback were also investigated. Many cortical and striatal somatomotor regions tracked the dynamical evolution of categorical evidence, while many regions of the dorsal and ventral attention networks (Corbetta and Shulman, 2002) tracked decisional confidence and uncertainty. Urgency influenced activity in regions known to be associated with flexible control of the speed-accuracy trade-off (particularly the pre- SMA and striatum), and additionally modulated representations of categorical evidence and confidence. The results, therefore, link the urgency signal to two hypothetical mechanisms underling flexible control of decision thresholding (Bogacz et al., 2010): gain modulation of the striatal thresholding circuitry, and gain modulation of the integrated categorical evidence

Temporal Dynamics of Decision-Making during Motion Perception in the Visual Cortex

How does the brain make decisions? Speed and accuracy of perceptual decisions covary with certainty in the input, and correlate with the rate of evidence accumulation in parietal and frontal cortical "decision neurons." A biophysically realistic model of interactions within and between Retina/LGN and cortical areas V1, MT, MST, and LIP, gated by basal ganglia, simulates dynamic properties of decision-making in response to ambiguous visual motion stimuli used by Newsome, Shadlen, and colleagues in their neurophysiological experiments. The model clarifies how brain circuits that solve the aperture problem interact with a recurrent competitive network with self-normalizing choice properties to carry out probablistic decisions in real time. Some scientists claim that perception and decision-making can be described using Bayesian inference or related general statistical ideas, that estimate the optimal interpretation of the stimulus given priors and likelihoods. However, such concepts do not propose the neocortical mechanisms that enable perception, and make decisions. The present model explains behavioral and neurophysiological decision-making data without an appeal to Bayesian concepts and, unlike other existing models of these data, generates perceptual representations and choice dynamics in response to the experimental visual stimuli. Quantitative model simulations include the time course of LIP neuronal dynamics, as well as behavioral accuracy and reaction time properties, during both correct and error trials at different levels of input ambiguity in both fixed duration and reaction time tasks. Model MT/MST interactions compute the global direction of random dot motion stimuli, while model LIP computes the stochastic perceptual decision that leads to a saccadic eye movement.National Science Foundation (SBE-0354378, IIS-02-05271); Office of Naval Research (N00014-01-1-0624); National Institutes of Health (R01-DC-02852

Neural dynamics implement a flexible decision bound with a fixed firing rate for choice: a model-based hypothesis

Decisions are faster and less accurate when conditions favour speed, and are slower and more accurate when they favour accuracy. This speed-accuracy trade-off (SAT) can be explained by the principles of bounded integration, where noisy evidence is integrated until it reaches a bound. Higher bounds reduce the impact of noise by increasing integration times, supporting higher accuracy (vice versa for speed). These computations are hypothesized to be implemented by feedback inhibition between neural populations selective for the decision alternatives, each of which corresponds to an attractor in the space of network states. Since decision-correlated neural activity typically reaches a fixed rate at the time of commitment to a choice, it has been hypothesized that the neural implementation of the bound is fixed, and that the SAT is supported by a common input to the populations integrating evidence. According to this hypothesis, a stronger common input reduces the difference between a baseline firing rate and a threshold rate for enacting a choice. In simulations of a two-choice decision task, we use a reduced version of a biophysically-based network model (Wong & Wang, 2006) to show that a common input can control the SAT, but that changes to the threshold-baseline difference are epiphenomenal. Rather, the SAT is controlled by changes to network dynamics. A stronger common input decreases the model’s effective time constant of integration and changes the shape of the attractor landscape, so the initial state is in a more error-prone position. Thus, a stronger common input reduces decision time and lowers accuracy. The change in dynamics also renders firing rates higher under speed conditions at the time that an ideal observer can make a decision from network activity. The difference between this rate and the baseline rate is actually greater under speed conditions than accuracy conditions, suggesting that the bound is not implemented by firing rates per se

Of rodents and primates: Time-variant gain in drift-diffusion decision models

Sequential sampling models of decision-making involve evidence accumulation over time and have been successful in capturing choice behaviour. A popular model is the drift-diffusion model (DDM). To capture the finer aspects of choice reaction times (RTs), time-variant gain features representing urgency signals have been implemented in DDM that can exhibit slower error RTs than correct RTs. However, time-variant gain is often implemented on both DDM’s signal and noise features, with the assumption that increasing gain on the drift rate (due to urgency) is similar to DDM with collapsing decision bounds. Hence, it is unclear whether gain effects on just the signal or noise feature can lead to different choice behaviour. This work presents an alternative DDM variant, focusing on the implications of time-variant gain mechanisms, constrained by model parsimony. Specifically, using computational modelling of choice behaviour of rats, monkeys and humans, we systematically showed that time-variant gain only on the DDM’s noise was sufficient to produce slower error RTs, as in monkeys, while time-variant gain only on drift rate leads to faster error RTs, as in rodents. We also found minimal effects of time-variant gain in humans. By highlighting these patterns, this study underscores the utility of group-level modelling in capturing general trends and effects consistent across species. Thus, time-variant gain on DDM’s different components can lead to different choice behaviour, shed light on the underlying time-variant gain mechanisms for different species, and can be used for systematic data fitting

Global gain modulation generates time-dependent urgency during perceptual choice in humans

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A biologically motivated synthesis of accumulator and reinforcement-learning models for describing adaptive decision-making

Cognitive process models, such as reinforcement learning (RL) and accumulator models of decision-making, have proven to be highly insightful tools for studying adaptive behaviors as well as their underlying neural substrates. Currently, however, two major barriers exist preventing these models from being applied in more complex settings: 1) the assumptions of most accumulator models break down for decisions involving more than two alternatives; 2) RL and accumulator models currently exist as separate frameworks, with no clear mapping between trial-to-trial learning and the dynamics of the decision process. Recently I showed how a modified accumulator model, premised off of the architecture of cortico-basal ganglia pathways, both predicts human decisions in uncertain situations and evoked activity in cortical and subcortical control circuits. Here I present a synthesis of RL and accumulator models that is motivated by recent evidence that the basal ganglia acts as a site for integrating trial-wise feedback from midbrain dopaminergic neurons with accumulating evidence from sensory and associative cortices. I show how this hybrid model can explain both adaptive go/no-go decisions and multi-alternative decisions in a computationally efficient manner. More importantly, by parameterizing the model to conform to various underlying assumptions about the architecture and physiology of basal ganglia pathways, model predictions can be rigorously tested against observed patterns in behavior as well as neural recordings. The result is a biologically-constrained and behaviorally tractable description of trial-to-trial learning effects on decision-making among multiple alternatives