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Accurate interaction with the environment relies on the integration of external information about the spatial layout of potential actions and knowledge of their costs and benefits. Previous studies have shown that when given a choice between voluntary reaching movements, humans tend to prefer actions with lower biomechanical costs. However, these studies primarily focused on decisions made before the onset of movement ("decide-then-act" scenarios), and it is not known to what extent their conclusions generalize to many real-life situations, in which decisions occur during ongoing actions ("decide-while-acting"). For example, one recent study found that biomechanical costs did not influence decisions to switch from a continuous manual tracking movement to a point-to-point movement, suggesting that biomechanical costs may be disregarded in decide-while-acting scenarios. To better understand this surprising result, we designed an experiment in which participants were faced with the decision between continuing to track a target moving along a straight path or changing paths to track a new target that gradually moved along a direction that deviated from the initial one. We manipulated tracking direction, angular deviation rate, and side of deviation, allowing us to compare scenarios where biomechanical costs favored either continuing or changing the path. Crucially, here the choice was always between two continuous tracking actions. Our results show that in this situation decisions clearly took biomechanical costs into account. Thus we conclude that biomechanics are not disregarded during decide-while-acting scenarios but rather that cost comparisons can only be made between similar types of actions. NEW & NOTEWORTHY In this study, we aim to shed light on how biomechanical factors influence decisions made during ongoing actions. Previous work suggested that decisions made during actions disregard biomechanical costs, in contrast to decisions made before movement. Our results challenge that proposal and suggest instead that the effect of biomechanical factors is dependent on the types of actions being compared (e.g., continuous tracking vs. point-to-point reaching). These findings contribute to our understanding of the dynamic interplay between biomechanical considerations and action choices during ongoing interactions with the environment.

One of the most exciting new developments in systems neuroscience is the progress being made toward neurophysiological experiments that move beyond simplified laboratory settings and address the richness of natural behavior. This is enabled by technological advances such as wireless recording in freely moving animals, automated quantification of behavior, and new methods for analyzing large data sets. Beyond new empirical methods and data, however, there is also a need for new theories and concepts to interpret that data. Such theories need to address the particular challenges of natural behavior, which often differ significantly from the scenarios studied in traditional laboratory settings. Here, we discuss some strategies for developing such novel theories and concepts and some example hypotheses being proposed., Competing Interests: Declaration of competing interest The authors declare that no competing interests exist., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Prominent accounts of sentient behavior depict brains as generative models of organismic interaction with the world, evincing intriguing similarities with current advances in generative artificial intelligence (AI). However, because they contend with the control of purposive, life-sustaining sensorimotor interactions, the generative models of living organisms are inextricably anchored to the body and world. Unlike the passive models learned by generative AI systems, they must capture and control the sensory consequences of action. This allows embodied agents to intervene upon their worlds in ways that constantly put their best models to the test, thus providing a solid bedrock that is - we argue - essential to the development of genuine understanding. We review the resulting implications and consider future directions for generative AI., Competing Interests: Declaration of interests K.F. holds a chief scientific adviser position at VERSES AI. The other authors declare no conflicts of interest., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

When choosing between options with multiple attributes, do we integrate all attributes into a unified measure for comparison, or does the comparison also occur at the level of each attribute, involving parallel processes that can dynamically influence each other? What happens when independent sensory features all carry information about the same decision factor, such as reward value? To investigate these questions, we asked human participants to perform a two-alternative forced choice reaching task in which the reward value of a target was indicated by two visual attributes-its brightness ("bottom-up," BU feature) and its orientation ("top-down," TD feature). If decisions always occur after the integration of both features, there should be no difference in the reaction time (RT) regardless of the attribute combinations that drove the choice. Counter to that prediction, RT distributions depended on the attribute combinations of given targets and the choices made by participants. RTs were shortest when both attributes were congruent or when the choice was based on the bottom-up feature, and longer when the attributes were in conflict (favoring opposite options). In conflict trials, nearly two-thirds of participants made faster decisions when choosing the option favored by the bottom-up feature than when choosing the top-down-favored option. We also observed mid-reach changes-of-mind in a subset of conflict trials, mostly changing from the bottom-up to the top-down-favored target. These data suggest that multi-attribute value-based decisions are better explained by a distributed process including competition among different features than by a competition based on a single, integrated estimate of value. NEW & NOTEWORTHY We show that during value-based decisions, humans do not always use all reward-related information to make their choice, but instead can "jump the gun" using partial information. In particular, when different sources of information were in conflict, early decisions were mostly based on fast bottom-up information, and sometimes followed by corrective changes-of-mind based on slower top-down information. This supports parallel decision processes among different information sources, as opposed to a single integrated "common currency."

Psychology and neuroscience are concerned with the study of behavior, of internal cognitive processes, and their neural foundations. However, most laboratory studies use constrained experimental settings that greatly limit the range of behaviors that can be expressed. While focusing on restricted settings ensures methodological control, it risks impoverishing the object of study: by restricting behavior, we might miss key aspects of cognitive and neural functions. In this article, we argue that psychology and neuroscience should increasingly adopt innovative experimental designs, measurement methods, analysis techniques and sophisticated computational models to probe rich, ecologically valid forms of behavior, including social behavior. We discuss the challenges of studying rich forms of behavior as well as the novel opportunities offered by state-of-the-art methodologies and new sensing technologies, and we highlight the importance of developing sophisticated formal models. We exemplify our arguments by reviewing some recent streams of research in psychology, neuroscience and other fields (e.g., sports analytics, ethology and robotics) that have addressed rich forms of behavior in a model-based manner. We hope that these "success cases" will encourage psychologists and neuroscientists to extend their toolbox of techniques with sophisticated behavioral models - and to use them to study rich forms of behavior as well as the cognitive and neural processes that they engage., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Author(s). Published by Elsevier B.V. All rights reserved.)

Recent theoretical models suggest that deciding about actions and executing them are not implemented by completely distinct neural mechanisms but are instead two modes of an integrated dynamical system. Here, we investigate this proposal by examining how neural activity unfolds during a dynamic decision-making task within the high-dimensional space defined by the activity of cells in monkey dorsal premotor (PMd), primary motor (M1), and dorsolateral prefrontal cortex (dlPFC) as well as the external and internal segments of the globus pallidus (GPe, GPi). Dimensionality reduction shows that the four strongest components of neural activity are functionally interpretable, reflecting a state transition between deliberation and commitment, the transformation of sensory evidence into a choice, and the baseline and slope of the rising urgency to decide. Analysis of the contribution of each population to these components shows meaningful differences between regions but no distinct clusters within each region, consistent with an integrated dynamical system. During deliberation, cortical activity unfolds on a two-dimensional "decision manifold" defined by sensory evidence and urgency and falls off this manifold at the moment of commitment into a choice-dependent trajectory leading to movement initiation. The structure of the manifold varies between regions: In PMd, it is curved; in M1, it is nearly perfectly flat; and in dlPFC, it is almost entirely confined to the sensory evidence dimension. In contrast, pallidal activity during deliberation is primarily defined by urgency. We suggest that these findings reveal the distinct functional contributions of different brain regions to an integrated dynamical system governing action selection and execution., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2022 Thura et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Finding the right amount of deliberation, between insufficient and excessive, is a hard decision making problem that depends on the value we place on our time. Average-reward, putatively encoded by tonic dopamine, serves in existing reinforcement learning theory as the opportunity cost of time, including deliberation time. Importantly, this cost can itself vary with the environmental context and is not trivial to estimate. Here, we propose how the opportunity cost of deliberation can be estimated adaptively on multiple timescales to account for non-stationary contextual factors. We use it in a simple decision-making heuristic based on average-reward reinforcement learning (AR-RL) that we call Performance-Gated Deliberation (PGD). We propose PGD as a strategy used by animals wherein deliberation cost is implemented directly as urgency, a previously characterized neural signal effectively controlling the speed of the decision-making process. We show PGD outperforms AR-RL solutions in explaining behaviour and urgency of non-human primates in a context-varying random walk prediction task and is consistent with relative performance and urgency in a context-varying random dot motion task. We make readily testable predictions for both neural activity and behaviour., Competing Interests: The authors have declared that no competing interests exist.

Humans and other animals are able to adjust their speed-accuracy trade-off (SAT) at will depending on the urge to act, favoring either cautious or hasty decision policies in different contexts. An emerging view is that SAT regulation relies on influences exerting broad changes on the motor system, tuning its activity up globally when hastiness is at premium. The present study aimed to test this hypothesis. A total of 50 participants performed a task involving choices between left and right index fingers, in which incorrect choices led either to a high or to a low penalty in 2 contexts, inciting them to emphasize either cautious or hasty policies. We applied transcranial magnetic stimulation (TMS) on multiple motor representations, eliciting motor-evoked potentials (MEPs) in 9 finger and leg muscles. MEP amplitudes allowed us to probe activity changes in the corresponding finger and leg representations, while participants were deliberating about which index to choose. Our data indicate that hastiness entails a broad amplification of motor activity, although this amplification was limited to the chosen side. On top of this effect, we identified a local suppression of motor activity, surrounding the chosen index representation. Hence, a decision policy favoring speed over accuracy appears to rely on overlapping processes producing a broad (but not global) amplification and a surround suppression of motor activity. The latter effect may help to increase the signal-to-noise ratio of the chosen representation, as supported by single-trial correlation analyses indicating a stronger differentiation of activity changes in finger representations in the hasty context., Competing Interests: The authors have declared that no competing interests exist.

Studies of neural population dynamics of cell activity from monkey motor areas during reaching show that it mostly represents the generation and timing of motor behavior. We compared neural dynamics in dorsal premotor cortex (PMd) during the performance of a visuomotor task executed individually or cooperatively and during an observation task. In the visuomotor conditions, monkeys applied isometric forces on a joystick to guide a visual cursor in different directions, either alone or jointly with a conspecific. In the observation condition, they observed the cursor's motion guided by the partner. We found that in PMd neural dynamics were widely shared across action execution and observation, with cursor motion directions more accurately discriminated than task types. This suggests that PMd encodes spatial aspects irrespective of specific behavioral demands. Furthermore, our results suggest that largest components of premotor population dynamics, which have previously been suggested to reflect a transformation from planning to movement execution, may rather reflect higher cognitive-motor processes, such as the covert representation of actions and goals shared across tasks that require movement and those that do not., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

The nervous system is a product of evolution. That is, it was constructed through a long series of modifications, within the strong constraints of heredity, and continuously subjected to intense selection pressures. As a result, the organization and functions of the brain are shaped by its history. We believe that this fact, underappreciated in contemporary systems neuroscience, offers an invaluable aid for helping us resolve the brain's mysteries. Indeed, we think that the consideration of evolutionary history ought to take its place alongside other intellectual tools used to understand the brain, such as behavioural experiments, studies of anatomical structure and functional characterization based on recordings of neural activity. In this introduction, we argue for the importance of evolution by highlighting specific examples of ways that evolutionary theory can enhance neuroscience. The rest of the theme issue elaborates this point, emphasizing the conservative nature of neural evolution, the important consequences of specific transitions that occurred in our history, and the ways in which considerations of evolution can shed light on issues ranging from specific mechanisms to fundamental principles of brain organization. This article is part of the theme issue 'Systems neuroscience through the lens of evolutionary theory'.

This article outlines a hypothetical sequence of evolutionary innovations, along the lineage that produced humans, which extended behavioural control from simple feedback loops to sophisticated control of diverse species-typical actions. I begin with basic feedback mechanisms of ancient mobile animals and follow the major niche transitions from aquatic to terrestrial life, the retreat into nocturnality in early mammals, the transition to arboreal life and the return to diurnality. Along the way, I propose a sequence of elaboration and diversification of the behavioural repertoire and associated neuroanatomical substrates. This includes midbrain control of approach versus escape actions, telencephalic control of local versus long-range foraging, detection of affordances by the dorsal pallium, diversified control of nocturnal foraging in the mammalian neocortex and expansion of primate frontal, temporal and parietal cortex to support a wide variety of primate-specific behavioural strategies. The result is a proposed functional architecture consisting of parallel control systems, each dedicated to specifying the affordances for guiding particular species-typical actions, which compete against each other through a hierarchy of selection mechanisms. This article is part of the theme issue 'Systems neuroscience through the lens of evolutionary theory'.

Decision-making is traditionally described as a cognitive process of deliberation followed by commitment to an action choice, preceding the planning and execution of the chosen action. However, this is challenged by recent data suggesting that during situated decisions, multiple options are specified simultaneously and compete in premotor cortical areas for selection and execution. Previous studies focused on the competition during planning and left unaddressed the dynamics of decisions during movement. Does deliberation extend into the execution phase? Are nonselected options still considered? Here we studied a decision-making task in which human participants were instructed to select a reaching path trajectory from an origin to a rectangular target, where reward was distributed nonuniformly at the target. Critically, we applied mechanical perturbations to the arm during movement to study under which conditions such perturbations produce changes of mind. Our results show that participants initially selected the direction of movement toward the highest reward region and changed their mind most frequently when the two choices offered the same reward, showing that deliberation continues and follows cost-benefit considerations during movement. Furthermore, changes of mind were dependent on the intensity of the perturbation and the current state of the motor system, including velocity and distance to targets. Although reward remains most relevant, our results indicate that the state of the motor system when the perturbation occurs is a crucial determinant of changes of mind., (Copyright © 2021 Cos et al.)

Most current decision-making research focuses on classical economic scenarios, where choice offers are prespecified and where action dynamics play no role in the decision. However, our brains evolved to deal with different choice situations: "embodied decisions". As examples of embodied decisions, consider a lion that has to decide which gazelle to chase in the savannah or a person who has to select the next stone to jump on when crossing a river. Embodied decision settings raise novel questions, such as how people select from time-varying choice options and how they track the most relevant choice attributes; but they have long remained challenging to study empirically. Here, we summarize recent progress in the study of embodied decisions in sports analytics and experimental psychology. Furthermore, we introduce a formal methodology to identify the relevant dimensions of embodied choices (present and future affordances) and to map them into the attributes of classical economic decisions (probabilities and utilities), hence aligning them. Studying embodied decisions will greatly expand our understanding of what decision-making is., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Humans and other animals often need to balance the desire to gather sensory information (to make the best choice) with the urgency to act, facing a speed-accuracy tradeoff (SAT). Given the ubiquity of SAT across species, extensive research has been devoted to understanding the computational mechanisms allowing its regulation at different timescales, including from one context to another, and from one decision to another. However, animals must frequently change their SAT on even shorter timescales-that is, over the course of an ongoing decision-and little is known about the mechanisms that allow such rapid adaptations. The present study aimed at addressing this issue. Human subjects performed a decision task with changing evidence. In this task, subjects received rewards for correct answers but incurred penalties for mistakes. An increase or a decrease in penalty occurring halfway through the trial promoted rapid SAT shifts, favoring speeded decisions either in the early or in the late stage of the trial. Importantly, these shifts were associated with stage-specific adjustments in the accuracy criterion exploited for committing to a choice. Those subjects who decreased the most their accuracy criterion at a given decision stage exhibited the highest gain in speed, but also the highest cost in terms of performance accuracy at that time. Altogether, the current findings offer a unique extension of previous work, by suggesting that dynamic cha*nges in accuracy criterion allow the regulation of the SAT within the timescale of a single decision. NEW & NOTEWORTHY Extensive research has been devoted to understanding the mechanisms allowing the regulation of the speed-accuracy tradeoff (SAT) from one context to another and from one decision to another. Here, we show that humans can voluntarily change their SAT on even shorter timescales-that is, over the course of a decision. These rapid SAT shifts are associated with dynamic adjustments in the accuracy criterion exploited for committing to a choice.

A successful class of models link decision-making to brain signals by assuming that evidence accumulates to a decision threshold. These evidence accumulation models have identified neuronal activity that appears to reflect sensory evidence and decision variables that drive behavior. More recently, an additional evidence-independent and time-variant signal, called urgency, has been hypothesized to accelerate decisions in the face of insufficient evidence. However, most decision-making paradigms tested with fMRI or EEG in humans have not been designed to disentangle evidence accumulation from urgency. Here we use a face-morphing decision-making task in combination with EEG and a hierarchical Bayesian model to identify neural signals related to sensory and decision variables, and to test the urgency-gating model. Forty females and 34 males took part (mean age, 23.4 years). We find that an evoked potential time locked to the decision, the centroparietal positivity, reflects the decision variable from the computational model. We further show that the unfolding of this signal throughout the decision process best reflects the product of sensory evidence and an evidence-independent urgency signal. Urgency varied across subjects, suggesting that it may represent an individual trait. Our results show that it is possible to use EEG to distinguish neural signals related to sensory evidence accumulation, decision variables, and urgency. These mechanisms expose principles of cognitive function in general and may have applications to the study of pathologic decision-making such as in impulse control and addictive disorders. SIGNIFICANCE STATEMENT Perceptual decisions are often described by a class of models that assumes that sensory evidence accumulates gradually over time until a decision threshold is reached. In the present study, we demonstrate that an additional urgency signal impacts how decisions are formed. This endogenous signal encourages one to respond as time elapses. We found that neural decision signals measured by EEG reflect the product of sensory evidence and an evidence-independent urgency signal. A nuanced understanding of human decisions, and the neural mechanisms that support it, can improve decision-making in many situations and potentially ameliorate dysfunction when it has gone awry., (Copyright © 2021 the authors.)

Neurophysiological studies suggest that when decisions are made between concrete actions, the selection process involves a competition between potential action representations in the same sensorimotor structures involved in executing those actions. However, it is unclear how such models can explain situations, often encountered during natural behavior, in which we make decisions while were are already engaged in performing an action. Does the process of deliberation characterized in classical studies of decision-making proceed the same way when subjects are deciding while already acting? In the present study, human subjects continuously tracked a target moving in the horizontal plane and were occasionally presented with a new target to which they could freely choose to switch at any time, whereupon it became the new tracked target. We found that the probability of choosing to switch increased with decreasing distance to the new target and increasing size of the new target relative to the tracked target, as well as when the direction to the new target was aligned (either toward or opposite) to the current tracking direction. However, contrary to our expectations, subjects did not choose targets that minimized the energetic costs of execution, as calculated by a biomechanical model of the arm. When the constraints of continuous tracking were removed in variants of the task involving point-to-point movements, the expected preference for lower cost choices was seen. These results are discussed in the context of current theories of nested feedback control, internal models of forward dynamics, and high-dimensional neural spaces. NEW & NOTEWORTHY Current theories of decision-making primarily address how subjects make decisions before executing selected actions. However, in our daily lives we often make decisions while already performing some action (e.g., while playing a sport or navigating through a crowd). To gain insight into how current theories can be extended to such "decide-while-acting" scenarios, we examined human decisions during continuous manual tracking and found some intriguing departures from how decisions are made in classical "decide-then-act" paradigms.

Humans and other animals are faced with decisions about actions on a daily basis. These typically include a period of deliberation that ends with the commitment to a choice, which then leads to the overt expression of that choice through action. Previous studies with monkeys have demonstrated that neural activity in sensorimotor areas correlates with the deliberation process and reflects the moment of commitment before movement initiation, but the causal roles of these regions are challenging to establish. Here, we tested whether dorsal premotor (PMd) and primary motor cortex (M1) are causally involved in the volitional commitment to a reaching choice. We found that brief subthreshold microstimulation in PMd or M1 delayed commitment to an action but not the initiation of the action itself. Importantly, microstimulation only had a significant effect when it was delivered close to and before commitment time. These results are consistent with the proposal that PMd and M1 participate in the commitment process, which occurs when a critical firing rate difference is reached between cells voting for the selected option and those voting for the competing one. NEW & NOTEWORTHY The neural substrates of decisions between actions are typically investigated by correlating neural activity and subjects' decision behavior, but this does not establish causality. In a reaching decision task, we demonstrate that subthreshold microstimulation of the monkey dorsal premotor cortex or primary motor cortex delays the deliberation duration if applied shortly before choice commitment. This result suggests a causal role of the sensorimotor cortex in the determination of decisions between actions.

If we abandon the coding metaphor in favor of models of the full behavioral loop, we need a way to dissect that loop into understandable pieces. I suggest that evolutionary data provide a solution. We can subdivide behavior into parallel sensorimotor subsystems by following the phylogenetic history of how those systems differentiated and specialized during our evolution, leading to promising ways of re-interpreting neural activity within the context of its pragmatic role in mediating interaction.

Decisions about actions typically involve a period of deliberation that ends with the commitment to a choice and the motor processes overtly expressing that choice. Previous studies have shown that neural activity in sensorimotor areas, including the primary motor cortex (M1), correlates with deliberation features during action selection. However, the causal contribution of these areas to the decision process remains unclear. Here, we investigated whether M1 determines choice commitment or whether it simply reflects decision signals coming from upstream structures and instead mainly contributes to the motor processes that follow commitment. To do so, we tested the impact of a disruption of M1 activity, induced by continuous theta burst stimulation (cTBS), on the behavior of human subjects in 1 ) a simple reaction time (SRT) task allowing us to estimate the duration of the motor processes and 2 ) a modified version of the tokens task (Cisek P, Puskas GA, El-Murr S. J Neurosci 29: 11560-11571, 2009), which allowed us to estimate subjects' time of commitment as well as accuracy criterion. The efficiency of cTBS was attested by a reduction in motor evoked potential amplitudes following M1 disruption compared with those following a sham stimulation. Furthermore, M1 cTBS lengthened SRTs, indicating that motor processes were perturbed by the intervention. Importantly, all of the behavioral results in the tokens task were similar following M1 disruption and sham stimulation, suggesting that the contribution of M1 to the deliberation process is potentially negligible. Taken together, these findings favor the view that M1 contribution is downstream of the decision process. NEW & NOTEWORTHY Decisions between actions are ubiquitous in the animal realm. Deliberation during action choices entails changes in the activity of the sensorimotor areas controlling those actions, but the causal role of these areas is still often debated. With the use of continuous theta burst stimulation, we show that disrupting the primary motor cortex (M1) delays the motor processes that follow instructed commitment but does not alter volitional deliberation, suggesting that M1 contribution may be downstream of the decision process.

This article proposes that biologically plausible theories of behavior can be constructed by following a method of "phylogenetic refinement," whereby they are progressively elaborated from simple to complex according to phylogenetic data on the sequence of changes that occurred over the course of evolution. It is argued that sufficient data exist to make this approach possible, and that the result can more effectively delineate the true biological categories of neurophysiological mechanisms than do approaches based on definitions of putative functions inherited from psychological traditions. As an example, the approach is used to sketch a theoretical framework of how basic feedback control of interaction with the world was elaborated during vertebrate evolution, to give rise to the functional architecture of the mammalian brain. The results provide a conceptual taxonomy of mechanisms that naturally map to neurophysiological and neuroanatomical data and that offer a context for defining putative functions that, it is argued, are better grounded in biology than are some of the traditional concepts of cognitive science.

Humans and other animals are motivated to act so as to maximize their subjective reward rate. Here, we propose that reward rate maximization is accomplished by adjusting a context-dependent "urgency signal," which influences both the commitment to a developing action choice and the vigor with which the ensuing action is performed. We review behavioral and neurophysiological data suggesting that urgency is controlled by projections from the basal ganglia to cerebral cortical regions, influencing neural activity related to decision making as well as activity related to action execution. We also review evidence suggesting that different individuals possess specific policies for adjusting their urgency signal to particular contextual variables, such that urgency constitutes an individual trait which jointly influences a wide range of behavioral measures commonly related to the overall quality and hastiness of one's decisions and actions. Consequently, we argue that a central mechanism for reward rate maximization provides a potential link between personality traits such as impulsivity, as well as some of the motivation-related symptomology of clinical disorders such as depression and Parkinson's disease.

In this article, we challenge the usefulness of "attention" as a unitary construct and/or neural system. We point out that the concept has too many meanings to justify a single term, and that "attention" is used to refer to both the explanandum (the set of phenomena in need of explanation) and the explanans (the set of processes doing the explaining). To illustrate these points, we focus our discussion on visual selective attention. It is argued that selectivity in processing has emerged through evolution as a design feature of a complex multi-channel sensorimotor system, which generates selective phenomena of "attention" as one of many by-products. Instead of the traditional analytic approach to attention, we suggest a synthetic approach that starts with well-understood mechanisms that do not need to be dedicated to attention, and yet account for the selectivity phenomena under investigation. We conclude that what would serve scientific progress best would be to drop the term "attention" as a label for a specific functional or neural system and instead focus on behaviorally relevant selection processes and the many systems that implement them.

Prominent theories of decision making suggest that the basal ganglia (BG) play a causal role in deliberation between action choices. An alternative hypothesis is that deliberation occurs in cortical regions, while the BG control the speed-accuracy trade-off (SAT) between committing to a choice versus continuing to deliberate. Here, we test these hypotheses by recording activity in the internal and external segments of the globus pallidus (GPi/GPe) while monkeys perform a task dissociating the process of deliberation, the moment of commitment, and adjustment of the SAT. Our data suggest that unlike premotor and motor cortical regions, pallidal output does not contribute to the process of deliberation but instead provides a time-varying signal that controls the SAT and reflects the growing urgency to commit to a choice. Once a target is selected by cortical regions, GP activity confirms commitment to the decision and invigorates the subsequent movement., (Copyright © 2017 Elsevier Inc. All rights reserved.)

We report a case of a symptomatic and rapidly expanding aneurysm of an in situ saphenous vein graft in a 70-year-old man with extensive prior open and endovascular procedures for aneurysmal disease. He was found to have full-length aneurysmal dilation with rapid progression over the course of 6 months. Successful ligation and exclusion with subtotal excision of the aneurysmal segment was performed, and revision bypass was foregone because of adequate distal perfusion via collateralization. This rare complication of autologous vein bypass graft typically occurs at the anastomoses and merits aneurysmorrhaphy or ligation. Far less common is the scenario of nonanastomotic, true aneurysmal dilation of the entire conduit such as the current patient. Surgical intervention via open, endovascular, or a combination thereof is warranted for rapidly growing or symptomatic defects and those yielding distal malperfusion., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Recent studies have shown that activity in sensorimotor structures varies depending on the speed-accuracy trade-off (SAT) context in which a decision is made. Here we tested the hypothesis that the same areas also reflect a more local adjustment of SAT established between individual trials, based on the outcome of the previous decision. Two monkeys performed a reaching decision task in which sensory evidence continuously evolves during the time course of a trial. In two SAT contexts, we compared neural activity in trials following a correct choice vs. those following an error. In dorsal premotor cortex (PMd), we found that 23% of cells exhibited significantly weaker baseline activity after error trials, and for ∼30% of these this effect persisted into the deliberation epoch. These cells also contributed to the process of combining sensory evidence with the growing urgency to commit to a choice. We also found that the activity of 22% of PMd cells was increased after error trials. These neurons appeared to carry less information about sensory evidence and time-dependent urgency. For most of these modulated cells, the effect was independent of whether the previous error was expected or unexpected. We found similar phenomena in primary motor cortex (M1), with 25% of cells decreasing and 34% increasing activity after error trials, but unlike PMd, these neurons showed less clear differences in their response properties. These findings suggest that PMd and M1 belong to a network of brain areas involved in SAT adjustments established using the recent history of reinforcement., New & Noteworthy: Setting the speed-accuracy trade-off (SAT) is crucial for efficient decision making. Previous studies have reported that subjects adjust their SAT after individual decisions, usually choosing more conservatively after errors, but the neural correlates of this phenomenon are only partially known. Here, we show that neurons in PMd and M1 of monkeys performing a reach decision task support this mechanism by adequately modulating their firing rate as a function of the outcome of the previous decision., (Copyright © 2017 the American Physiological Society.)

We discuss how cybernetic principles of feedback control, used to explain sensorimotor behavior, can be extended to provide a foundation for understanding cognition. In particular, we describe behavior as parallel processes of competition and selection among potential action opportunities ('affordances') expressed at multiple levels of abstraction. Adaptive selection among currently available affordances is biased not only by predictions of their immediate outcomes and payoffs but also by predictions of what new affordances they will make available. This allows animals to purposively create new affordances that they can later exploit to achieve high-level goals, resulting in intentional action that links across multiple levels of control. Finally, we discuss how such a 'hierarchical affordance competition' process can be mapped to brain structure., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Perceptual decision making is often modeled as perfect integration of sequential sensory samples until the accumulated total reaches a fixed decision bound. In that view, the buildup of neural activity during perceptual decision making is attributed to temporal integration. However, an alternative explanation is that sensory estimates are computed quickly with a low-pass filter and combined with a growing signal reflecting the urgency to respond and it is the latter that is primarily responsible for neural activity buildup. These models are difficult to distinguish empirically because they make similar predictions for tasks in which sensory information is constant within a trial, as in most previous studies. Here we presented subjects with a variant of the classic constant-coherence motion discrimination (CMD) task in which we inserted brief motion pulses. We examined the effect of these pulses on reaction times (RTs) in two conditions: 1) when the CMD trials were blocked and subjects responded quickly and 2) when the same CMD trials were interleaved among trials of a variable-motion coherence task that motivated slower decisions. In the blocked condition, early pulses had a strong effect on RTs but late pulses did not, consistent with both models. However, when subjects slowed their decision policy in the interleaved condition, later pulses now became effective while early pulses lost their efficacy. This last result contradicts models based on perfect integration of sensory evidence and implies that motion signals are processed with a strong leak, equivalent to a low-pass filter with a short time constant., (Copyright © 2016 the American Physiological Society.)

Recent work suggests that while animals decide between reaching actions, neurons in dorsal premotor (PMd) and primary motor (M1) cortex reflect a dynamic competition between motor plans and determine when commitment to a choice is made. This competition is biased by at least two sources of information: the changing sensory evidence for one choice versus another, and an urgency signal that grows over time. Here, we test the hypothesis that the urgency signal adjusts the trade-off between speed and accuracy during both decision-making and movement execution. Two monkeys performed a reaching decision task in which sensory evidence continuously evolves over the course of each trial. In different blocks, task timing parameters encouraged monkeys to voluntarily adapt their behavior to be either hasty or conservative. Consistent with our hypothesis, during the deliberation process the baseline and gain of neural activity in decision-related PMd (29%) and M1 cells (45%) was higher when monkeys applied a hasty policy than when they behaved conservatively, but at the time of commitment the population activity was similar across blocks. Other cells (30% in PMd, 30% in M1) showed activity that increased or decreased with elapsing time until the moment of commitment. Movement-related neurons were also more active after longer decisions, as if they were influenced by the same urgency signal controlling the gain of decision-related activity. Together, these results suggest that the arm motor system receives an urgency/vigor signal that adjusts the speed-accuracy trade-off for decision-making and movement execution. Significance statement: This work addresses the neural mechanisms that control the speed-accuracy trade-off in both decisions and movements, in the kinds of dynamic situations that are typical of natural animal behavior. We found that many "decision-related" premotor and motor neurons are modulated in a time-dependent manner compatible with an "urgency" signal that changes between hasty and conservative decision policies. We also found that such modulation influenced cells related to the speed of the reaching movements executed by the animals to report their decisions. These results suggest that a unified mechanism determines speed-accuracy trade-off adjustments during decision-making and movement execution, potentially influencing both the cognitive and motor aspects of reward-oriented behavior., (Copyright © 2016 the authors 0270-6474/16/360938-19$15.00/0.)

In a recent report, Winkel, Keuken, van Maanen, Wagenmakers & Forstmann (Psychonomics Bulletin and Review 21(3): 777-784, 2014) show that during a random-dot motion discrimination task, early differences in motion evidence can influence reaction times (RTs) and error rates in human subjects. They use this as an argument in favor of the drift-diffusion model and against the urgency-gating model. However, their implementation of the urgency-gating model is incomplete, as it lacks the low-pass filter that is necessary to deal with noisy input such as the motion signal used in their experimental task. Furthermore, by focusing analyses solely on comparison of mean RTs they overestimate how long early information influences individual trials. Here, we show that if the urgency-gating model is correctly implemented, including a low-pass filter with a 250 ms time constant, it can successfully reproduce the results of the Winkel et al. experiment.

Speed-accuracy tradeoffs (SATs) exist in both decision-making and movement control, and are generally studied separately. However, in natural behavior animals are free to adjust the time invested in deciding and moving so as to maximize their reward rate. Here, we investigate whether shared mechanisms exist for SAT adjustment in both decisions and actions. Two monkeys performed a reach decision task in which they watched 15 tokens jump, one every 200 ms, from a central circle to one of two peripheral targets, and had to guess which target would ultimately receive the majority of tokens. The monkeys could decide at any time, and once a target was reached, the remaining token movements accelerated to either 50 ms ("fast" block) or 150 ms ("slow" block). Decisions were generally earlier and less accurate in fast than slow blocks, and in both blocks, the criterion of accuracy decreased over time within each trial. This could be explained by a simple model in which sensory information is combined with a linearly growing urgency signal. Remarkably, the duration of the reaching movements produced after the decision decreased over time in a similar block-dependent manner as the criterion of accuracy estimated by the model. This suggests that SATs for deciding and acting are influenced by a shared urgency/vigor signal. Consistent with this, we observed that the vigor of saccades performed during the decision process was higher in fast than in slow blocks, suggesting the influence of a context-dependent global arousal., (Copyright © 2014 the authors 0270-6474/14/3416442-13$15.00/0.)

Neurophysiological studies of decision-making have focused primarily on elucidating the mechanisms of classic economic decisions, for which the relevant variables are the values of expected outcomes and action is simply the means of reporting the selected choice. By contrast, here we focus on the particular challenges of embodied decision-making faced by animals interacting with their environment in real time. In such scenarios, the choices themselves as well as their relative costs and benefits are defined by the momentary geometry of the immediate environment and change continuously during ongoing activity. To deal with the demands of embodied activity, animals require an architecture in which the sensorimotor specification of potential actions, their valuation, selection and even execution can all take place in parallel. Here, we review behavioural and neurophysiological data supporting a proposed brain architecture for dealing with such scenarios, which we argue set the evolutionary foundation for the organization of the mammalian brain., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

When given a choice between actions that yield the same reward, we tend to prefer the one that requires the least effort. Recent studies have shown that humans are remarkably accurate at evaluating the effort of potential reaching actions and can predict the subtle energetic demand caused by the nonisotropic biomechanical properties of the arm. In the present study, we investigated the time course over which such information is computed and comes to influence decisions. Two independent approaches were used. First, subjects performed a reach decision task in which the time interval for deciding between two candidate reaching actions was varied from 200 to 800 ms. Second, we measured motor-evoked potential (MEPs) to single-pulse transcranial magnetic stimulation (TMS) over the primary motor cortex (M1) to probe the evolving decision at different times after stimulus presentation. Both studies yielded a consistent conclusion: that a prediction of the effort associated with candidate movements is computed very quickly and influences decisions within 200 ms after presentation of the candidate actions. Furthermore, whereas the MEPs measured 150 ms after stimulus presentation were well correlated with the choices that subjects ultimately made, later in the trial the MEP amplitudes were primarily related to the muscular requirements of the chosen movement. This suggests that corticospinal excitability (CSE) initially reflects a competition between candidate actions and later changes to reflect the processes of preparing to implement the winning action choice., (Copyright © 2014 the American Physiological Society.)

Spatial patterns of functional connectivity derived from resting brain activity may be used to elucidate the topological properties of brain networks. Such networks are amenable to study using graph theory, which shows that they possess small world properties and can be used to differentiate healthy subjects and patient populations. Of particular interest is the possibility that some of these differences are related to alterations in the dopamine system. To investigate the role of dopamine in the topological organization of brain networks at rest, we tested the effects of reducing dopamine synthesis in 13 healthy subjects undergoing functional magnetic resonance imaging. All subjects were scanned twice, in a resting state, following ingestion of one of two amino acid drinks in a randomized, double-blind manner. One drink was a nutritionally balanced amino acid mixture, and the other was tyrosine and phenylalanine deficient. Functional connectivity between 90 cortical and subcortical regions was estimated for each individual subject under each dopaminergic condition. The lowered dopamine state caused the following network changes: reduced global and local efficiency of the whole brain network, reduced regional efficiency in limbic areas, reduced modularity of brain networks, and greater connection between the normally anti-correlated task-positive and default-mode networks. We conclude that dopamine plays a role in maintaining the efficient small-world properties and high modularity of functional brain networks, and in segregating the task-positive and default-mode networks. This article is part of the Special Issue Section entitled 'Neuroimaging in Neuropharmacology'., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Neurophysiological studies of decision making have primarily focused on decisions about information that is stable over time. However, during natural behavior, animals make decisions in a constantly changing environment. To investigate the neural mechanisms of such dynamic choices, we recorded activity in dorsal premotor (PMd) and primary motor cortex (M1) while monkeys performed a two-choice reaching task in which sensory information about the correct choice was changing within each trial and the decision could be made at any time. During deliberation, activity in both areas did not integrate sensory information but instead tracked it and combined it with a growing urgency signal. Approximately 280 ms before movement onset, PMd activity tuned to the selected target reached a consistent peak while M1 activity tuned to the unselected target was suppressed. We propose that this reflects the resolution of a competition between the potential responses and constitutes the volitional commitment to an action choice., (Copyright © 2014 Elsevier Inc. All rights reserved.)

How does the brain decide between actions? Is it through comparisons of abstract representations of outcomes or through a competition in a sensorimotor map defining the actions themselves? Here, I review strengths and limitations of both of these proposals, and suggest that decisions emerge through a distributed consensus across many levels of representation., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

It is often suggested that decisions are made when accumulated sensory information reaches a fixed accuracy criterion. This is supported by many studies showing a gradual build up of neural activity to a threshold. However, the proposal that this build up is caused by sensory accumulation is challenged by findings that decisions are based on information from a time window much shorter than the build-up process. Here, we propose that in natural conditions where the environment can suddenly change, the policy that maximizes reward rate is to estimate evidence by accumulating only novel information and then compare the result to a decreasing accuracy criterion. We suggest that the brain approximates this policy by multiplying an estimate of sensory evidence with a motor-related urgency signal and that the latter is primarily responsible for neural activity build up. We support this hypothesis using human behavioral data from a modified random-dot motion task in which motion coherence changes during each trial.

Recent work has shown that human subjects are able to predict the biomechanical ease of potential reaching movements and use these predictions to influence their choices. Here, we examined how reach decisions are influenced by specific biomechanical factors related to the control of end-point stability, such as aiming accuracy or stopping control. Human subjects made free choices between two potential reaching movements that varied in terms of path distance and biomechanical cost in four separate blocks that additionally varied two constraints: the width of the targets (narrow or wide) and the requirement of stopping in them. When movements were unconstrained (very wide targets and no requirement of stopping), subjects' choices were strongly biased toward directions aligned with the direction of maximal mobility. However, as the movements became progressively constrained, factors related to the control of the end point gained relevance, thus reducing this bias. This demonstrates that, before movement onset, constraints such as stopping and aiming participate in a remarkably adaptive and flexible action selection process that trades off the advantage of moving along directions of maximal mobility for unconstrained movements against exploiting biomechanical anisotropies to facilitate control of end-point stability whenever the movement constraints require it. These results support a view of decision making between motor actions as a highly context-dependent gradual process in which the subjective desirability of potential actions is influenced by their dynamic properties in relation to the intrinsic properties of the motor apparatus.

During simple sensorimotor decision making, neurons in the parietal cortex extract evidence from sensory information provided by visual areas until a decision is reached. Contextual information can bias parietal activity during the task and change the decision-making parameters. One type of contextual information is the availability of reward for correct decisions. We tested the hypothesis that the frontal lobes and basal ganglia use contextual information to bias decision making to maximize reward. Human volunteers underwent functional MRI while making decisions about the motion of dots on a computer monitor. On rewarded trials, subjects responded more slowly by increasing the threshold to decision. Rewarded trials were associated with activation in the ventral striatum and prefrontal cortex in the period preceding coherent dot motion, and the degree of activation predicted the increased decision threshold. Decreasing dopamine transmission, using a tyrosine-depleting amino acid mixture, abolished the reward-related corticostriatal activation and eliminated the correlation between striatal activity and decision threshold. These observations provide direct evidence that some reward-related functional MRI signals in the striatum are the result of dopamine neuron activity and demonstrate that mesolimbic dopamine transmission can influence perceptual and decision-making neural processes engaged to maximize reward harvest.

Previous studies have shown that neural activity in primate dorsal premotor cortex (PMd) can simultaneously represent multiple potential movement plans, and that activity related to these movement options is modulated by their relative subjective desirability. These findings support the hypothesis that decisions about actions are made through a competition within the same circuits that guide the actions themselves. This hypothesis further predicts that the very same cells that guide initial decisions will continue to update their activities if an animal changes its mind. For example, if a previously selected movement option suddenly becomes unavailable, the correction will be performed by the same cells that selected the initial movement, as opposed to some different group of cells responsible for online guidance. We tested this prediction by recording neural activity in the PMd of a monkey performing an instructed-delay reach selection task. In the task, two targets were simultaneously presented and their border styles indicated whether each would be worth 1, 2, or 3 juice drops. In a random subset of trials (FREE), the monkey was allowed a choice while in the remaining trials (FORCED) one of the targets disappeared at the time of the GO signal. In FORCED-LOW trials the monkey was forced to move to the less valuable target and started moving either toward the new target (Direct) or toward the target that vanished and then curved to reach the remaining one (Curved). Prior to the GO signal, PMd activity clearly reflected the monkey's subjective preference, predicting his choices in FREE trials even with equally valued options. In FORCED-LOW trials, PMd activity reflected the switch of the monkey's plan as early as 100 ms after the GO signal, well before movement onset (MO). This confirms that the activity is not related to feedback from the movement itself, and suggests that PMd continues to participate in action selection even when the animal changes its mind on-line. These findings were reproduced by a computational model suggesting that switches between action plans can be explained by the same competition process responsible for initial decisions.

To make an accurate movement, the CNS has to overcome the inherent complexities of the multijoint limb. For example, interaction torques arise when motion of individual arm segments propagates to adjacent segments causing their movement without any muscle contractions. Since these passive joint torques significantly add to the overall torques generated by active muscular contractions, they must be taken into account during planning or execution of goal-directed movements. We investigated the role of the corticospinal tract in compensating for the interaction torques during arm movements in humans. Twelve subjects reached to visual targets with their arm supported by a robotic exoskeleton. Reaching to one target was accompanied by interaction torques that assisted the movement, while reaching to the other target was accompanied by interaction torques that resisted the movement. Corticospinal excitability was assessed at different times during movement using single-pulse transcranial magnetic stimulation (TMS) over the upper-arm region of M1 (primary motor cortex). We found that TMS responses in shoulder monoarticular and elbow-shoulder biarticular muscles changed together with the interaction torques during movements in which the interaction torques were resistive. In contrast, TMS responses did not correlate with assistive interaction torques or with co-contraction. This suggests that the descending motor command includes compensation for passive limb dynamics. Furthermore, our results suggest that compensation for interaction torques involves the biarticular muscles, which span both shoulder and elbow joints and are in a biomechanically advantageous position to provide such compensation.

There is considerable debate on the extent to which biomechanical properties of movements are taken into account before and during voluntary movements. For example, while several models have described reach planning as primarily kinematic, some studies have suggested that implicit knowledge about biomechanics may also exert some influence on the planning of reaching movements. Here, we investigated whether decisions about reaching movements are influenced by biomechanical factors and whether these factors are taken into account before movement onset. To this end, we designed an experimental paradigm in which humans made free choices between two potential reaching movements where the options varied in path distance as well as biomechanical factors related to movement energy and stability. Our results suggest that the biomechanical properties of potential actions strongly influence the selection between them. In particular, in our task, subjects preferred movements whose final trajectory was better aligned with the major axis of the arm's mobility ellipse, even when the launching properties were very similar. This reveals that the nervous system can predict biomechanical properties of potential actions before movement onset and that these predictions, in addition to purely abstract criteria, may influence the decision-making process.

Detecting the temporal relationship among events in the environment is a fundamental goal of the brain. Following pulses of rhythmic stimuli, neurons of the retina and cortex produce activity that closely approximates the timing of an omitted pulse. This omitted stimulus response (OSR) is generally interpreted as a transient response to rhythmic input and is thought to form a basis of short-term perceptual memories. Despite its ubiquity across species and experimental protocols, the mechanisms underlying OSRs remain poorly understood. In particular, the highly transient nature of OSRs, typically limited to a single cycle after stimulation, cannot be explained by a simple mechanism that would remain locked to the frequency of stimulation. Here, we describe a set of realistic simulations that capture OSRs over a range of stimulation frequencies matching experimental work. The model does not require an explicit mechanism for learning temporal sequences. Instead, it relies on spike timing-dependent plasticity (STDP), a form of synaptic modification that is sensitive to the timing of pre- and post-synaptic action potentials. In the model, the transient nature of OSRs is attributed to the heterogeneous nature of neural properties and connections, creating intricate forms of activity that are continuously changing over time. Combined with STDP, neural heterogeneity enabled OSRs to complex rhythmic patterns as well as OSRs following a delay period. These results link the response of neurons to rhythmic patterns with the capacity of heterogeneous circuits to produce transient and highly flexible forms of neural activity.

A new study by Klaes et al. in this issue of Neuron shows that the brain can simultaneously apply two rules to the same sensory information in order to specify two parallel potential action goals, which then compete for execution in the sensorimotor system., (Copyright © 2011 Elsevier Inc. All rights reserved.)

It has been proposed that whenever an animal faces several action choices, their neural representations are processed in parallel in frontoparietal cortex and compete in a manner biased by any factor relevant to the decision. We tested this hypothesis by recording single-unit activity in dorsal premotor cortex (PMd) while a monkey performed two delayed center-out reaching tasks. In the one-target task, a single target was presented and its border style indicated its reward value. The two-target task was the same except two targets were presented and the value of each was varied. During the delay period of the one-target task, directionally tuned PMd activity showed no modulation with value. In contrast, during the two-target task, the same neurons showed strong effects of the value associated with their preferred target, always in relation to the value of the other target. Furthermore, the competition between action choices was strongest when targets were furthest apart. This angular distance effect appeared in neural activity as soon as cells became tuned, while modulation by relative value appeared much later. All of these findings can be reproduced by a computational model which suggests that decisions between actions are made through a biased competition taking place within a sensorimotor map of potential actions.

It has been suggested that, during decisions about actions, multiple options are initially specified in parallel and then gradually eliminated in a competition for overt execution. To further test this hypothesis, we studied the modulation of human corticospinal excitability during the reaction time of the Eriksen flanker task. In the task, subjects responded with finger flexion or extension to a central arrow while ignoring congruent or incongruent flanker arrows. Single-pulse transcranial magnetic stimulation (TMS) was applied over primary motor cortex (M1) at one of five different latencies after stimulus onset, and motor-evoked potentials (MEPs) were measured in the contralateral index finger. During the control (no flankers) and congruent conditions, MEP size in the agonist increased gradually over the course of reaction time, indicating an increase in corticospinal excitability. Conversely, when the same muscle acted as an antagonist, MEP size decreased, suggesting inhibition. Critically, in the incongruent condition, MEPs briefly increased in the muscle corresponding to an initial default response to the flanker arrows and were later replaced by MEPs corresponding to the correct response to the central arrow. Finally, we found that the gradually growing MEPs for the three conditions reached a constant maximum level just before movement initiation. We propose that this dynamic modulation in corticospinal excitability reflects the competition process, leading to the selection of one response and the rejection of the other. Our results suggest that response competition influences activity in primary motor cortex and that its timing directly influences motor output latency.

The neural bases of behavior are often discussed in terms of perceptual, cognitive, and motor stages, defined within an information processing framework that was originally inspired by models of human abstract problem solving. Here, we review a growing body of neurophysiological data that is difficult to reconcile with this influential theoretical perspective. As an alternative foundation for interpreting neural data, we consider frameworks borrowed from ethology, which emphasize the kinds of real-time interactive behaviors that animals have engaged in for millions of years. In particular, we discuss an ethologically-inspired view of interactive behavior as simultaneous processes that specify potential motor actions and select between them. We review how recent neurophysiological data from diverse cortical and subcortical regions appear more compatible with this parallel view than with the classical view of serial information processing stages.

Several widely accepted models of decision making suggest that, during simple decision tasks, neural activity builds up until a threshold is reached and a decision is made. These models explain error rates and reaction time distributions in a variety of tasks and are supported by neurophysiological studies showing that neural activity in several cortical and subcortical regions gradually builds up at a rate related to task difficulty and reaches a relatively constant level of discharge at a time that predicts movement initiation. The mechanism responsible for this buildup is believed to be related to the temporal integration of sequential samples of sensory information. However, an alternative mechanism that may explain the neural and behavioral data is one in which the buildup of activity is instead attributable to a growing signal related to the urgency to respond, which multiplicatively modulates updated estimates of sensory evidence. These models are difficult to distinguish when, as in previous studies, subjects are presented with constant sensory evidence throughout each trial. To distinguish the models, we presented human subjects with a task in which evidence changed over the course of each trial. Our results are more consistent with "urgency gating" than with temporal integration of sensory samples and suggest a simple mechanism for implementing trade-offs between the speed and accuracy of decisions.

Neural synchronization is of wide interest in neuroscience and has been argued to form the substrate for conscious attention to stimuli, movement preparation, and the maintenance of task-relevant representations in active memory. Despite a wealth of possible functions, the mechanisms underlying synchrony are still poorly understood. In particular, in vitro preparations have demonstrated synchronization with no apparent periodicity, which cannot be explained by simple oscillatory mechanisms. Here, we investigate the possible origins of nonperiodic synchronization through biophysical simulations. We show that such aperiodic synchronization arises naturally under a simple set of plausible assumptions, depending crucially on heterogeneous cell properties. In addition, nonperiodicity occurs even in the absence of stochastic fluctuation in membrane potential, suggesting that it may represent an intrinsic property of interconnected networks. Simulations capture some of the key aspects of population-level synchronization in spontaneous network spikes (NSs) and suggest that the intrinsic nonperiodicity of NSs observed in reduced cell preparations is a phenomenon that is highly robust and can be reproduced in simulations that involve a minimal set of realistic assumptions. In addition, a model with spike timing-dependent plasticity can overcome a natural tendency to exhibit nonperiodic behavior. After rhythmic stimulation, the model does not automatically fall back to a state of nonperiodic behavior, but keeps replaying the pattern of evoked NSs for a few cycles. A cluster analysis of synaptic strengths highlights the importance of population-wide interactions in generating this result and describes a possible route for encoding temporal patterns in networks of neurons.