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Switching probability B infection z imdb 400 mg ofloxacin order with visa, We consider here, for example, a toy brain of two coupled model brain regions X and Y, undergoing sparsely synchronized oscillations. In each of the two possible states, infor mation conveyed by spiking code words emitted by source neurons in the phaseleading area can be decoded from code words emitted by target neurons in the phaselaggard area (70% of shared information). However, decoding efficiency does not rise above chance level (·) in the opposite laggardto leader direction. Switching between phaselocking modes can be induced by precisely phased pulse perturbations, applied within a specific control phase range (correctly predicted by theory, highlighted range). Furthermore, all of them can be applied to the analysis of both empirical and simulated time series. When dealing with oscillatory neural signals, their functional coupling can vary as a function of fre quency. The coupling between neural oscillations can also be quantified using phase synchronization (Rosenblum, Pikovsky, & Kurths, 1996), defined as the entrainment of phases irrespectively of amplitude correlations, or phaselocking value (Lachaux et al. Beyond autoregressive modeling, Granger (1980) formalized a general condition of "Granger non causality" between two time series X and Y as p(Yi + 1 Y (i), X (i)) = p(Yi + 1 Y (i)), (62. Accordingly, causality can be defined as a deviation from this condi tion of "noncausality" and quantified by calculating the information theoretical Kullback Leibler divergence (MacKay, 2003) between the two conditional probabili ties in equation (62. Tools for the non parametric estimation of spectrally decomposed Granger causality directly from Fourier and wavelet transforms of time series data are available (Dhamala, Rangara jan, & Ding, 2008). In the frame work of autoregressive models, this allows the estima tion of model coefficients across trials on short time windows for the computation of coherence and Granger causality spectra with high temporal precision (see. Neural coupling, however, may vary across trials and reflect behavioral modulations occur ring during learning and adaptive behav iors. Such an approach can be used to estimate singletrial phase synchrony (Lachaux Battaglia and Brovelli: Functional Connectivity and Neuronal Dynamics 741 et al. Functional connectivity dynamics along a task Ultimately, cognition necessarily unrolls in time, and mental oper ations are built out of successive steps, which assemble into a cognitive architecture mixing serial and mas sively parallel information processing, also dubbed a human Turing machine (Zylberberg et al. This phenomenon is epitomized by simple toy models involving only a small number of coupled areas. Such an abstract structural motif serves as a metaphor for canonical cortical circuits in which the relative weights of top down and bottomup functional influences must be dynamically adjusted. Every brain region is modeled as a local network of thousands of excitatory and inhibitory spiking neurons, connected by random recurrent connectivity. Three different partially overlapping networks (right) activate and deactivate with a characteristic recruitment sched ule (left). Since firing is Poissonlike, spike trains have a high entropy, and a large amount of infor mation can be conveyed by the oscillating population within every oscillation cycle. In other words, the oscil lations themselves are not likely to encode information but act as carriers for general code words encoded in detailed spiking patterns "surfing on the wave. For sufficiently strong inhibition, a multiplicity of out ofphase locking modes tend to emerge, in which one of the two regions leads in phase over the other, despite the reciprocity of coupling. Besides the unidirectional transfer of information, other functional motifs can be implemented by our toy brain (bidirectional, either anisotropic or symmetric; effective disconnection; and so on). Self organized control of information routing Under the effect of an arbitrary perturbation, the system will be transiently destabilized, but its dynamics will then con verge back to one of the available intrinsic modes. If the applied perturbation kicks the system out of the phase space basin of attraction of the current dynamic state- a valley in an idealized landscape-the system will converge toward a dif ferent state within its dyn ome. Various mechanisms could force the system to leave its current state and then be used for implementing routing control. A first possibility would be to modulate the relative attractiveness of dif ferent states (in the landscape metaphor of figure 62. In the presence of multistabil ity between multiple dynamic configurations, it would be enough to apply a steady input bias to one of the two populations to automatically enhance its probability of becoming phase leader and thus act as an effective information sender (Palmigiano et al. An unspe cific, weak bias would be enough because its role would just be to favor the other wise self organized selection of a specific routing state from a preexisting repertoire. Therefore, no additional circuitry for routing control would be required besides the one already responsible for the generation of collective oscillations themselves, in contrast with other proposed mechanisms. Such a steady bias could be provided by some top down modu latory signal, neuromodulation, or even stimulus saliency itself. Such theoretical prediction has not yet been confirmed but could be experimentally validated using, for example, closedloop optogenetic stimulation (Witt et al. We have also generalized our findings to larger net works with arbitrarily complex modular topologies (Kirst, Timme, & Battaglia, 2016). In our study we indeed predicted that the dom inant direction of connectivity between two regions X and Y could be reverted by applying a drive bias to a third "remote controller" region Z. Such a mechanism, emphasizing the nonlocality of the effects of a local ized system perturbation, is robust since connectivity patterns would be stable over broad ranges of par ameters, and switching would occur- suddenly and Battaglia and Brovelli: Functional Connectivity and Neuronal Dynamics 743 "everywhere"- only in the proximity of specific, criti cal working points of operation. Selforganized routing with transient and stochastic oscilla tions the toy models considered in figure 62. In reality, oscillatory episodes in vivo are usually transient, arising at stochastic timings and with an inconsistently volatile frequency (Ray & Maunsell, 2015; Xing et al. Different filters can be defined to track the stochastic manifestation of dif ferent routing states. Via stateresolved analyses, we can thus conclude that the transient and stochastic nature of oscillations is not an obstacle to the flexible and controllable selective routing of input signals, thanks to collective self organization. A key prediction of the model-that calls for an experimental confirmation-is that directed information transfer between coupled regions should be intermittent-that is, strongly enhanced during co occurring oscillatory bursts and reduced to baseline, or even actively suppressed, between these oscillatory events (Palmigiano et al. However, if a rich repertoire of states is sampled, either spontaneously as an effect of noise or in a way guided by exogenous (sensory) or endogenous (cognitive) bias, every averaging procedure is going to destroy precious information (Hutchison et al. This is true even for averaging over time aligned trials since we cannot a priori guarantee that transitions between internal states are really so tightly linked to taskrelated events. In reality, matching oscillatory burst ing events with different phase relations are stochasti cally occurring along each trial and at different timings for different trials.
In the future antibiotic groups 200 mg ofloxacin for sale, validating models on unseen data can help identify the most reliable predictors of working memory at the level of single individuals. Precision Although the majority of individual differences studies of working memory have focused on capacity, people also differ in their working memory precision. Increases in set size were accompanied by per for mance decrements and lower pattern classification accuracy for the remembered stimuli, a measure of representational precision. Estimates of orientation selectivity in visual cortex were correlated with differences in representational acuity across participants, also suggesting links between working memory precision and sustained neural activity in sensory cortex. Finally, Galeano Weber, Peters, Hahn, Bledowski, and Fiebach (2016) reported that participants with more stable working memory per formance. Predictive Models of Working Memory To date, predictive network models have characterized individual differences in the precision, but not capacity, of working memory. Asking whether interactions between perceptual and attentional systems affect working memory precision, Galeano Weber, Hahn, Hilger, and Fiebach (2017) scanned participants while they performed a visual working memory and a visual attention task. For each participant, they also calculated functional connectivity between the occipital and parietal regions activated during both tasks. Participants with better working memory precision showed higher connectivity between occipital and parietal regions during encoding. Mirroring findings with attention, these results suggest that engaging memory-related circuits magnifies individual differences in memory-related functional connections. Nonetheless, these results leave open the possibility that models based on whole-brain functional connectivity, rather than a circumscribed set of regions of interest, could predict individual differences in working memory capacity. Attention- memory interactions Although attention and working memory are often studied in isolation, they are intimately intertwined (Engle, 2002). For example, attentional mechanisms can gate entry into our capacity-limited working memory (Awh, Vogel, & Oh, 2006) and manipulate stored information (Myers, Stokes, & Nobre, 2017), the contents of working memory can influence how we focus our attention and resist distraction (de Fockert, Rees, Frith, & Lavie, 2001; Downing, 2000), and working memory itself can be considered a form of internally directed attention (Chun, Golomb, & Turk-Browne, 2011). Interactions between attention and memory are also evident at the level of large- scale brain networks. The model significantly predicted memory-test per formance, such that individuals with stronger highattention networks and weaker low-attention networks during reading better comprehended and remembered what they had read (Jangraw et al. These results demonstrate links between sustained attention and short-term memory and suggest that cross-task prediction approaches can elucidate relationships between the constituent processes of attention and working memory. Current work explores relationships between aspects of attentional control and memory. These models generalized to predict visual and verbal memory in 157 older adults from a Samsung Medical Center data set, highlighting relationships between processes underlying attention, working memory, and short-term memory across the lifespan (Avery et al. Limitations of Predictive Network Models Although this chapter has focused on the benefits of predictive network models, there are several limitations associated with the approach. First, individual differences studies provide correlational (rather than causal) evidence of brain-behavior relationships and are limited by sample size and composition, the reliability of single- subject data, and the degree to which data reflect state-like versus trait-like influences (Braver, Cole, & Yarkoni, 2010). Confounds such as head motion can also induce spurious relationships between functional connectivity and behav ior, undermining model validity if not appropriately controlled. Finally, translating brain-based predictive models to clinical settings requires the careful consideration of issues related to implementation and patient privacy (Rosenberg, Casey, & Holmes, 2018). Conclusions A driving question in psychology is how the mind is organized into distinct processes. Proposed taxonomies of attention and working memory have suggested that attention comprises three independent systems (alerting, orienting, and executive control), that these components vary along a number of dimensions. Thus, moving forward, cognitive network neuroscientific approaches may not only shed light on the functional organization of the brain, but may also inform the organization of the mind. Whole-brain functional connectivity predicts working memory per for mance in novel healthy and memory-impaired individuals. Proceedings of the National Academy of Sciences of the United States of America, 106(21), 87198724. The segregation and integration of distinct brain networks and their relationship to cognition. The hubs of the human connectome are generally implicated in the anatomy of brain disorders. Distributed patterns of activity in sensory cortex reflect the precision of multiple items maintained in visual short-term memory. Functional connectome fingerprinting: Identifying individuals using patterns of brain connectivity. Spontaneous neuronal activity distinguishes human dorsal and ventral attention systems. Proceedings of the National Academy of Sciences of the United States of America, 103(26), 1004610051. The human brain is intrinsically organized into dynamic, anticorrelated functional networks. Proceedings of the National Academy of Sciences of the United States of Amer ica, 102(27), 96739678. Quantity not quality: the relationship between fluid intelligence and working memory capacity. Prediction as a humanitarian and pragmatic contribution from human cognitive neuroscience. Distributed patterns of occipito-parietal functional connectivity predict the precision of visual working memory. Superior intraparietal sulcus controls the variability of visual working memory precision.
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The ubiquity of rewards and the fundamental behaviors they support underscore the importance of neuronal reward processing bacteria 1710 purchase ofloxacin amex. The proper appreciation and use of rewards requires significant neural resources because reward processing poses a difficult neurocomputational problem. For example, common, everyday rewards include delicious wine, good company, and excellent music. There is no common biophysical receptor that can detect all of these rewards, and there is no physical ruler that can compare them. Nevertheless, individuals instantaneously recognize rewards, rank them according to preference, and even exchange one for another. In fact, particular brain areas appear specialized for processing rewards and reward-related behav iors. These (and possibly other) regions define a loosely organized brain network that we describe as the reward system. These neurons reside in the midbrain and send axons to the basal ganglia and frontal cortex (figure 49. Dopamine neurons emit action potentials at low background frequencies (08 imp/s) and respond to behaviorally relevant events with phasic responses- short duration (150300 ms) bursts or pauses of action potentials. These phasic responses code for reward prediction errors, the difference between received and predicted rewards. Prediction errors are a necessary condition for associative learning, and therefore dopamine reward prediction error responses are natural teaching signals. Increases in activity indicate the outcome was better than predicted, and the preceding behav ior should be repeated or invigorated, whereas reductions in activity indicate that the outcome was worse than predicted and the preceding behav ior should be lessened or avoided. Crucially, dopamine activations and suppressions indicate not just what direction to update, positive or negative. The magnitude of the response also indicates by what degree behav iors should be updated. The magnitudes of dopamine responses correlate positively with reward amount and expected value (reward amount multiplied by the probability of delivery). These factors are important determinants of what decision-makers are most likely to choose. Reward Prediction Error Responses 3 mm Some of the earliest indications that value is a quantity coded for by single neurons in the brain arose from dopamine neuron recordings. Unpredicted rewards briefly activated dopamine neurons, whereas unpredicted reward omissions caused dopamine activity to pause (Schultz, Apicella, & Ljungberg, 1993). This pattern of dopamine activity resembles the prediction error in the Rescorla-Wagner (R-W) reinforcement-learning rule, defined as reward received- reward predicted. Moreover, dopamine neurons are more readily activated by rewards than by motivationally equivalent aversive outcomes (Mirenowicz & Schultz, 1996). Accordingly, phasic dopamine responses are known as reward prediction error responses. Similarly, dopamine responses code for the prediction errors associated with higher- order conditional stimuli (Schultz, Apicella, & Ljungberg, 1993) and are sensitive to small temporal deviations in learned intervals (Hollerman & Schultz, 1998). Bright regions are stained with an antibody for tyrosine hydroxylase, an enzyme specific to dopamine neurons in the midbrain. Utility is defined in economics as a mathematical description of subjective value inferred from observed choice behavior. As we will see, psychophysical measurements of utility combined with neurophysiology have shown that dopamine teaching signals reflect the same values used for economic decisions. Thus, dopamine neurons constitute a critical neural interface between learning and decision-making. The purpose of this chapter is Amount, Probability, and Expected Value Biological evidence for value coding is found in the magnitude of dopamine responses to different amounts of unpredicted rewards. Reward prediction error responses scale positively with the amount of juice delivered. Dopamine neurons respond with the largest activation to the largest reward and the smallest activation to the 588 Reward and Decision-Making smallest reward (Tobler, Fiorillo, & Schultz, 2005). Thus, dopamine responses show a positive monotonic relationship to reward amount, a primary factor that determines reward value. The predicted probability of reward delivery affects value in a manner proportional to the chance of getting the reward. The dopamine response to a particular reward is larger when the predicted probability of its delivery is smaller, and the reward responses are diminished as the predicted probability gets larger. These facts reveal that objective factors including amount and probability and their mathematical expectation are only partially responsible for decisions. For example, individuals will often consume too much of a highly valued reward, and the resulting state of satiety temporarily devalues that same reward. Thus, the same physical reward takes on different values based on the individual decision-maker. Two reward parameters in particular, timing and risk, are critical for separating objective factors, such as amount, from subjective values. With regard to time, individuals often prefer to receive rewards sooner rather than later. A $10 reward is more valuable when it is delivered immediately, compared to when it is delivered in one week, despite the physical amount of reward being identical. This phenomenon is called temporal discounting and can be used to separate the coding of subjective value from the coding of the physical amount. Temporal discounting has been used to demonstrate that dopamine neurons, basal ganglia, and frontal cortex regions code for the subjective value of rewards. Hyperbolicdiscounting models-rather than exponentialdiscounting models that use a constant discount rate-predict well human and monkey preferences.