Memory in repeat sports-related concussive injury and single-impact traumatic brain injury.

Background: Repeat sports-related concussive/subconcussive injury (RC/SCI) is related to memory impairment. Objective & Methods: We sought to determine memory differences between persons with RC/SCI, moderate-to-severe single-impact traumatic brain injury (SI-TBI), and healthy controls. MRI scans from a subsample of participants with SI-TBI were used to identify the neuroanatomical correlates of observed memory process differences between the brain injury groups. Results: Both brain injury groups evidenced worse learning and recall in contrast to controls, although SI-TBI group had poorer memory than the RC/SCI group. Regarding memory process differences, in contrast to controls, the SI-TBI group evidenced difficulties with encoding, consolidation, and retrieval, while the RC/SCI group showed deficits in consolidation and retrieval. Delayed recall was predicted by encoding, with consolidation as a secondary predictor in the SI-TBI group. In the RC/SCI group, delayed recall was only predicted by consolidation. MRI data showed that the consolidation index we used mapped onto hippocampal atrophy. Conclusions: RC/SCI is primarily associated with consolidation deficits, which differs from SI-TBI. Given the role of the hippocampus in memory consolidation and the fact that hyperphosphorylated tau tends to accumulate in the medial temporal lobe in RC/SCI, consolidation deficits may be a cognitive marker of chronic traumatic encephalopathy in athletes.

Differential roles of delay-period neural activity in the monkey dorsolateral prefrontal cortex in visual-haptic crossmodal working memory.

Previous studies have shown that neurons of monkey dorsolateral prefrontal cortex (DLPFC) integrate information across modalities and maintain it throughout the delay period of working-memory (WM) tasks. However, the mechanisms of this temporal integration in the DLPFC are still poorly understood. In the present study, to further elucidate the role of the DLPFC in crossmodal WM, we trained monkeys to perform visuo-haptic (VH) crossmodal and haptic-haptic (HH) unimodal WM tasks. The neuronal activity recorded in the DLPFC in the delay period of both tasks indicates that the early-delay differential activity probably is related to the encoding of sample information with different strengths depending on task modality, that the late-delay differential activity reflects the associated (modality-independent) action component of haptic choice in both tasks (that is, the anticipation of the behavioral choice and/or active recall and maintenance of sample information for subsequent action), and that the sustained whole-delay differential activity likely bridges and integrates the sensory and action components. In addition, the VH late-delay differential activity was significantly diminished when the haptic choice was not required. Taken together, the results show that, in addition to the whole-delay differential activity, DLPFC neurons also show early- and late-delay differential activities. These previously unidentified findings indicate that DLPFC is capable of (i) holding the coded sample information (e.g., visual or tactile information) in the early-delay activity, (ii) retrieving the abstract information (orientations) of the sample (whether the sample has been haptic or visual) and holding it in the late-delay activity, and (iii) preparing for behavioral choice acting on that abstract information.

Modulation of Frontoparietal Neurovascular Dynamics in Working Memory.

Our perception of the world is represented in widespread, overlapping, and interactive neuronal networks of the cerebral cortex. A majority of physiological studies on the subject have focused on oscillatory synchrony as the binding mechanism for representation and transmission of neural information. Little is known, however, about the stability of that synchrony during prolonged cognitive operations that span more than just a few seconds. The present research, in primates, investigated the dynamic patterns of oscillatory synchrony by two complementary recording methods, surface field potentials (SFPs) and near-infrared spectroscopy (NIRS). The signals were first recorded during the resting state to examine intrinsic functional connectivity. The temporal modulation of coactivation was then examined on both signals during performance of working memory (WM) tasks with long delays (memory retention epochs). In both signals, the peristimulus period exhibited characteristic features in frontal and parietal regions. Examination of SFP signals over delays lasting tens of seconds, however, revealed alternations of synchronization and desynchronization. These alternations occurred within the same frequency bands observed in the peristimulus epoch, without a specific correspondence between any definite cognitive process (e.g., WM) and synchrony within a given frequency band. What emerged instead was a correlation between the degree of SFP signal fragmentation (in time, frequency, and brain space) and the complexity and efficiency of the task being performed. In other words, the incidence and extent of SFP transitions between synchronization and desynchronization-rather than the absolute degree of synchrony-augmented in correct task performance compared with incorrect performance or in a control task without WM demand. An opposite relationship was found in NIRS: increasing task complexity induced more uniform, rather than fragmented, NIRS coactivations. These findings indicate that the particular features of neural oscillations cannot be linearly mapped to cognitive functions. Rather, information and the cognitive operations performed on it are primarily reflected in their modulations over time. The increased complexity and fragmentation of electrical frequencies in WM may reflect the activation of hierarchically diverse cognits (cognitive networks) in that condition. Conversely, the homogeneity in coherence of NIRS responses may reflect the cumulative vascular reactions that accompany that neuroelectrical proliferation of frequencies and the longer time constant of the NIRS signal. These findings are directly relevant to the mechanisms mediating cognitive processes and to physiologically based interpretations of functional brain imaging.

Past makes future: role of pFC in prediction.

The pFC enables the essential human capacities for predicting future events and preadapting to them. These capacities rest on both the structure and dynamics of the human pFC. Structurally, pFC, together with posterior association cortex, is at the highest hierarchical level of cortical organization, harboring neural networks that represent complex goal-directed actions. Dynamically, pFC is at the highest level of the perception-action cycle, the circular processing loop through the cortex that interfaces the organism with the environment in the pursuit of goals. In its predictive and preadaptive roles, pFC supports cognitive functions that are critical for the temporal organization of future behavior, including planning, attentional set, working memory, decision-making, and error monitoring. These functions have a common future perspective and are dynamically intertwined in goal-directed action. They all utilize the same neural infrastructure: a vast array of widely distributed, overlapping, and interactive cortical networks of personal memory and semantic knowledge, named cognits, which are formed by synaptic reinforcement in learning and memory acquisition. From this cortex-wide reservoir of memory and knowledge, pFC generates purposeful, goal-directed actions that are preadapted to predicted future events.

Cognitive Networks (Cognits) Process and Maintain Working Memory.

Ever since it was discovered in the monkey's prefrontal cortex, persistent neuronal activity during the delay period of delay tasks has been considered a phenomenon of working memory. Operationally, this interpretation is correct, because during that delay those tasks require the memorization of a sensory cue, commonly visual. What is incorrect is the assumption that the persistent activity during the delay is caused exclusively by the retention of the sensory cue. In this brief review, the author takes the position that the neural substrate of working memory is an array of long-term memory networks, that is, of cognitive networks (cognits), updated and orderly activated for the attainment of a behavioral goal. In the case of a behavioral task, that activated array of cognits has been previously formed in long-term memory (throughout this text, the expression "long-term memory" refers to all experiences acquired after birth, including habits and so-called procedural memory, such as the learning of a behavioral task). The learning of a task is the forming of synaptic associations between neural representations of three cognitive components of the task: perceptual, motor, and reward-related. Thereafter, when needed, the composite cognit of the task is activated in an orderly fashion to serve working memory in the perception-action cycle. To make his points on a complex issue, which has been the focus of his work, and to delineate a frontier for future research, the author refers to several of his own publications and previously published reviews.

The prefrontal cortex in the neurology clinic.

Throughout the nervous system, posterior structures are mainly devoted to receptive functions-sensation and perception-while anterior structures are devoted to motor functions. In the cortex, that dichotomy is unclear because perception and action are intertwined in the perception-action cycle, the biocybernetic cycle that adapts the organism to its environment. All neural systems store information (memory), which they enact in behavior and language. There are no "systems of memory" but the memory of systems. The cortex of the frontal lobe is a hierarchical system: motor cortex at the bottom for coordination of simple movements, and prefrontal cortex at the top for complex goal-directed actions. In the coordination of such actions, the frontal hierarchy engages the posterior (perceptual) cortex in the perception-action cycle. Inputs to the cycle come to prefrontal cortex from sensory-evoked perceptual memory and biologic (phyletic) memory. The first comes from neocortex, the second from limbic structures-through orbitomedial cortex. Outputs flow to pyramidal and diencephalic structures. Feedback inputs for monitoring and correction operate at all levels of the cycle. All prefrontal functions-planning, executive attention, working memory, decision-making, and inhibitory controls-are prospective, i.e., have a future perspective for the cycle to reach its goal. Damage to lateral prefrontal cortex impairs all of them. Orbitofrontal damage impairs the exclusionary aspect of attention and often leads to poor impulse control, excessive risk taking, unstable mood, and antisocial behavior. Medial prefrontal damage leads to poor monitoring of behavioral outcome for prevention of errors.

Jackson and the frontal executive hierarchy.

Executive actions are represented and hierarchically organized in the cortex of the frontal lobe. The representation and coordination of an action or series of actions have the same anatomical substrate: an executive neuronal network (cognit) in forntal cortex. That network interacts structurally and dynamically with perceptual networks of posterior cortex at the highest levels of the perception-action cycle.

The cognit: a network model of cortical representation.

The prevalent concept in modular models is that there are discrete cortical domains dedicated more or less exclusively to such cognitive functions as visual discrimination, language, spatial attention, face recognition, motor programming, memory retrieval, and working memory. Most of these models have failed or languished for lack of conclusive evidence. In their stead, network models are emerging as more suitable and productive alternatives. Network models are predicated on the basic tenet that cognitive representations consist of widely distributed networks of cortical neurons. Cognitive functions, namely perception, attention, memory, language, and intelligence, consist of neural transactions within and between these networks. The present model postulates that memory and knowledge are represented by distributed, interactive, and overlapping networks of neurons in association cortex. Such networks, named cognits, constitute the basic units of memory or knowledge. The association cortex of posterior-post-rolandic-regions contains perceptual cognits: cognitive networks made of neurons associated by information acquired through the senses. Conversely, frontal association cortex contains executive cognits, made of neurons associated by information related to action. In both posterior and frontal cortex, cognits are hierarchically organized. At the bottom of that organization-that is, in parasensory and premotor cortex-cognits are small and relatively simple, representing simple percepts or motor acts. At the top of the organization-in temporo-parietal and prefrontal cortex-cognits are wider and represent complex and abstract information of perceptual or executive character. Posterior and frontal networks are associated by long reciprocal cortico-cortical connections. These connections support the dynamics of the perception-action cycle in sequential behavior, speech, and reasoning.

Upper processing stages of the perception-action cycle.

The neural substrate for behavioral, cognitive and linguistic actions is hierarchically organized in the cortex of the frontal lobe. In their methodologically impeccable study, Koechlin et al. reveal the neural dynamics of the frontal hierarchy in behavioral action. Progressively higher areas control the performance of actions requiring the integration of progressively more complex and temporally dispersed information. The study substantiates the crucial role of the prefrontal cortex in the temporal organization of behavior.

Somatosensory cell response to an auditory cue in a haptic memory task.

Neurons in the monkey's anterior parietal cortex (Brodmann's areas 3a, 3b, 1, and 2) have been reported to retain information from a visual cue that has been associated with a tactile stimulus in a haptic memory task. This cross-modal transfer indicates that neurons in somatosensory cortex can respond to non-tactile stimuli if they are associated with tactile information needed for performance of the task. We hypothesized that neurons in somatosensory cortex would be activated by other non-tactile stimuli signaling the haptic movements--of arm and hand--that the task required. We found such cells in anterior parietal areas. They reacted with short-latency activity changes to an auditory signal (a click) that prompted those movements. Further, some of those cells changed their discharge in temporal correlation with the movements themselves, with the touch of the test objects, and with the short-term memory of those objects for subsequent tactile discrimination. These findings suggest that cells in the somatosensory cortex participate in the behavioral integration of auditory stimuli with other sensory stimuli and with motor acts that are associated with those stimuli.

Cajal in Barcelona: From yesterday to today, from the neuron to the network

The synaptic and network theories of memory, which Cajal first advanced in Barcelona around 1890, have been firmly established and elaborated by three generations of neuroscientists. This article outlines a corollary model of memory in the cerebral cortex that derives from those theories and is empirically supported by modern functional methods. The model posits that the elementary unit of memory or knowledge is a network of neurons of the cerebral cortex associated by life experience according to Hebbian principles of synaptic modulation (a cognit). Networks or cognits of perceptual memory are hierarchically organized and distributed in posterior association cortex; those of executive memory, also hierarchically organized, are distributed in frontal association cortex. In the course of goal-directed behavior and language, perceptual and executive cognits engage in the perception-action cycle, the cybernetic cycle that dynamically links the cortical cognitive networks with the environment in the pursuit of goals,. The prefrontal cortex, at the summit of that cycle, and interacting with cortical and subcortical structures, guides behavior and language to their goals by means of its executive functions of planning, executive attention, working memory, decision-making, and inhibitory control

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Memory for performed and observed activities following traumatic brain injury.

Traumatic brain injury (TBI) is associated with deficits in memory for the content of completed activities. However, TBI groups have shown variable memory for the temporal order of activities. We sought to clarify the conditions under which temporal order memory for activities is intact following TBI. Additionally, we evaluated activity source memory and the relationship between activity memory and functional outcome in TBI participants. Thus, we completed a study of activity memory with 18 severe TBI survivors and 18 healthy age- and education-matched comparison participants. Both groups performed eight activities and observed eight activities that were fashioned after routine daily tasks. Incidental encoding conditions for activities were utilized. The activities were drawn from two counterbalanced lists, and both performance and observation were randomly determined and interspersed. After all of the activities were completed, content memory (recall and recognition), source memory (conditional source identification), and temporal order memory (correlation between order reconstruction and actual order) for the activities were assessed. Functional ability was assessed via the Community Integration Questionnaire (CIQ). In terms of content memory, TBI participants recalled and recognized fewer activities than comparison participants. Recognition of performed and observed activities was strongly associated with social integration on the CIQ. There were no between- or within-group differences in temporal order or source memory, although source memory performances were near ceiling. The findings were interpreted as suggesting that temporal order memory following TBI is intact under conditions of both purposeful activity completion and incidental encoding, and that activity memory is related to functional outcomes following TBI.

Cognit activation: a mechanism enabling temporal integration in working memory.

Working memory is critical to the integration of information across time in goal-directed behavior, reasoning and language, yet its neural substrate is unknown. Based on recent research, we propose a mechanism by which the brain can retain working memory for prospective use, thereby bridging time in the perception/action cycle. The essence of the mechanism is the activation of 'cognits', which consist of distributed, overlapping and interactive cortical networks that in the aggregate encode the long-term memory of the subject. Working memory depends on the excitatory reentry between perceptual and executive cognits of posterior and frontal cortices, respectively. Given the pervasive role of working memory in the structuring of purposeful cognitive sequences, its mechanism looms essential to the foundation of behavior, reasoning and language.

Cortex and memory: emergence of a new paradigm.

Converging evidence from humans and nonhuman primates is obliging us to abandon conventional models in favor of a radically different, distributed-network paradigm of cortical memory. Central to the new paradigm is the concept of memory network or cognit--that is, a memory or an item of knowledge defined by a pattern of connections between neuron populations associated by experience. Cognits are hierarchically organized in terms of semantic abstraction and complexity. Complex cognits link neurons in noncontiguous cortical areas of prefrontal and posterior association cortex. Cognits overlap and interconnect profusely, even across hierarchical levels (heterarchically), whereby a neuron can be part of many memory networks and thus many memories or items of knowledge.

Working memory cells' behavior may be explained by cross-regional networks with synaptic facilitation.

Neurons in the cortex exhibit a number of patterns that correlate with working memory. Specifically, averaged across trials of working memory tasks, neurons exhibit different firing rate patterns during the delay of those tasks. These patterns include: 1) persistent fixed-frequency elevated rates above baseline, 2) elevated rates that decay throughout the tasks memory period, 3) rates that accelerate throughout the delay, and 4) patterns of inhibited firing (below baseline) analogous to each of the preceding excitatory patterns. Persistent elevated rate patterns are believed to be the neural correlate of working memory retention and preparation for execution of behavioral/motor responses as required in working memory tasks. Models have proposed that such activity corresponds to stable attractors in cortical neural networks with fixed synaptic weights. However, the variability in patterned behavior and the firing statistics of real neurons across the entire range of those behaviors across and within trials of working memory tasks are typical not reproduced. Here we examine the effect of dynamic synapses and network architectures with multiple cortical areas on the states and dynamics of working memory networks. The analysis indicates that the multiple pattern types exhibited by cells in working memory networks are inherent in networks with dynamic synapses, and that the variability and firing statistics in such networks with distributed architectures agree with that observed in the cortex.

Distributed and associative working memory.

This study explores the cortical cell dynamics of unimodal and cross-modal working memory (WM). Neuronal activity was recorded from parietal areas of monkeys performing delayed match-to-sample tasks with tactile or visual samples. Tactile memoranda (haptic samples) consisted of rods with differing surface features (texture or orientation of ridges) perceived by active touch. Visual memoranda (icons) consisted of striped patterns of differing orientation. In a haptic-haptic task, the animal had to retain through a period of delay the surface feature of the sample rod to select a rod that matched it. In a visual-haptic task, the animal had to retain the icon for the haptic choice of a rod with ridges of the same orientation as the icon's stripes. Units in all areas responded with firing change to one or more task events. Also in all areas, cells responded differently to different sample memoranda. Differential sample coherent firing was present in most areas during the memory period (delay). It is concluded that neurons in somatosensory and association areas of parietal cortex participate in broad networks that represent various task events and stimuli (auditory, motor, proprioceptive, tactile, and visual). Neurons in the same networks take part in retaining in WM the memorandum for each trial, whether it is encoded haptically or visually. The VH association by parietal cells in WM is analogous to the auditory-visual association previously observed in prefrontal cortex. Both illustrate the capacity of cortical neurons to associate sensory information across time and across modalities in accord with the rules of a behavioral task.

Frontal lobe and cognitive development.

In phylogeny as in ontogeny, the association cortex of the frontal lobe, also known as the prefrontal cortex, is a late-developing region of the neocortex. It is also one of the cortical regions to undergo the greatest expansion in the course of both evolution and individual maturation. In the human adult, the prefrontal cortex constitutes as much as nearly one-third of the totality of the neocortex. The protracted, relatively large, development of the prefrontal cortex is manifest in gross morphology as well as fine structure. In the developing individual, its late maturation is made most apparent by the late myelination of its axonal connections. This and other indices of morphological development of the prefrontal cortex correlate with the development of cognitive functions that neuropsychological studies in animals and humans have ascribed to this cortex. In broad outline, the ventromedial areas of the prefrontal cortex, which with respect to other prefrontal areas develop relatively early, are involved in the expression and control of emotional and instinctual behaviors. On the other hand, the late maturing areas of the lateral prefrontal convexity are principally involved in higher executive functions. The most general executive function of the lateral prefrontal cortex is the temporal organization of goal-directed actions in the domains of behavior, cognition, and language. In all three domains, that global function is supported by a fundamental role of the lateral prefrontal cortex in temporal integration, that is, the integration of temporally discontinuous percepts and neural inputs into coherent structures of action. Temporal integration is in turn served by at least three cognitive functions of somewhat different prefrontal topography: working memory, preparatory set, and inhibitory control. These functions engage the prefrontal cortex in interactive cooperation with other neocortical regions. The development of language epitomizes the development of temporal integrative cognitive functions and their underlying neural substrate, notably the lateral prefrontal cortex and other late-developing cortical regions.

El paradigma reticular de la memoria cortical / The reticular paradigm of cortical memory

Introducción. Los avances de la neurociencia cognitiva en los últimos años nos obligan a cambiar radicalmente el modelo tradicional de representación de memoria en la corteza cerebral. El viejo modelo (modular) postulaba un área distinta para cada forma de representación cognitiva (memoria operante, visual, auditiva, táctil, fisonómica, semántica, etc.). En el nuevo paradigma, las memorias y objetos mentales de conocimiento están constituidos por amplias redes de neuronas corticales ligadas sinápticamente por la experiencia. Desarrollo. Se presentan los principios fundamentales de este paradigma, con énfasis en sus aspectos estructurales, clínicos y de desarrollo. A partir del nacimiento, y con cada nueva experiencia, estas redes o cógnitos se van formando o reformando por medio de procesos asociativos sinápticos que siguen gradientes filogenéticos, ontogenéticos y conectivos, desde las áreas sensoriales y motoras hacia las cortezas asociativas. Los cógnitos nuevos se van autoorganizando en dos jerarquías de redes, con base sensorial y motora. La jerarquía perceptual, en la corteza posterior, representa cógnitos definidos por parámetros sensoriales en áreas sensoriales primarias, y los cógnitos perceptivos individuales (por ejemplo, memoria autobiográfica y episódica, conocimiento semántico), en áreas asociativas posteriores. La jerarquía ejecutiva, por otra parte, representa movimientos concretos en las áreas motoras frontales, y acciones más complejas (p. ej., planes de conducta) en la corteza prefrontal. Conclusiones. La investigación reciente nos obliga a abandonar los modelos tradicionales, ‘modulares’ o ‘geográficos’, de la memoria cortical. En su lugar, se impone con creciente vigor su paradigma reticular, el cual tiene importantes implicaciones con respecto al desarrollo cognitivo del individuo, la clínica de las lesiones corticales y la rehabilitación del enfermo con tales lesiones (AU)

Introduction. Recent advances in cognitive neuroscience oblige us to change radically the traditional model of representation of memory in the cerebral cortex. The old –modular– model postulates a separate area for each form of memory (working memory, episodic memory, visual memory, auditory memory, tactile memory, etc.). In the new –reticular– paradigm, memories and items of knowledge are made of widely distributed networks of neuron populations synaptically connected by experience. Development. Memory networks overlap and interact profusely; a neuron or group of neurons can be part of many networks, thus many memories or items of knowledge. After birth and throughout life, each new experience is etched in the form of those networks or cognits by synaptic associative processes that course from area to area along phylogenetic, ontogenetic, and connective gradients, from sensory and motor areas into associative areas. By self-organization, new cognits distribute themselves within two cortical hierarchies with a sensory and motor base, respectively. The perceptual hierarchy, in posterior cortex, houses cognits defined by sensory parameters in sensory areas and perceptual memories in associative areas. The executive hierarchy, on the other hand, represents concrete movements in frontal motor areas and more complex actions (e.g., plans) in prefrontal cortex. Conclusions. The reticular memory paradigm has important implications with regard to the cognitive development of the individual, cortical clinical syndromes, and cognitive rehabilitation (AU)