The neural substrate of spatial memory stabilization depends on the distribution of the training sessions.

SignificanceDistributed training has long been known to lead to more robust memory formation as compared to massed training. Using the water maze, a well-established task for assessing memory in laboratory rodents, we found that distributed and massed training differentially engage the dorsolateral and dorsomedial striatum, and optogenetic priming of dorsolateral striatum can artificially increase the robustness of massed training to the level of distributed training. Overall, our findings demonstrate that spatial memory consolidation engages different neural substrates depending on the training regimen, identifying a therapeutic avenue for memory enhancement.

Why do individuals with Williams syndrome or Down syndrome fail the Weather Prediction Task?

Williams syndrome (WS) and Down syndrome (DS) are two neurodevelopmental disorders with distinct genetic origins characterized by mild to moderate intellectual disability. Individuals with WS or DS exhibit impaired hippocampus-dependent place learning and enhanced striatum-dependent spatial response learning. Here, we used the Weather Prediction Task (WPT), which can be solved using hippocampus- or striatum-dependent learning strategies, to determine whether individuals with WS or DS exhibit similar profiles outside the spatial domain. Only 10% of individuals with WS or DS solved the WPT. We further assessed whether a concurrent memory task could promote reliance on procedural learning to solve the WPT in individuals with WS but found that the concurrent task did not improve performance. To understand how the probabilistic cue-outcome associations influences WPT performance, and whether individuals with WS or DS can ignore distractors, we assessed performance using a visual learning task with differing reward contingencies, and a modified WPT with unpredictive cues. Both probabilistic feedback and distractors negatively impacted the performance of individuals with WS or DS. These findings are consistent with deficits in hippocampus-dependent learning and executive functions, and reveal the importance of congruent feedback and the minimization of distractors to optimize learning in these two populations.

Memory Systems, the Epistemic Arrow of Time, and the Second Law.

The epistemic arrow of time is the fact that our knowledge of the past seems to be both of a different kind and more detailed than our knowledge of the future. Just like with the other arrows of time, it has often been speculated that the epistemic arrow arises due to the second law of thermodynamics. In this paper, we investigate the epistemic arrow of time using a fully formal framework. We begin by defining a memory system as any physical system whose present state can provide information about the state of the external world at some time other than the present. We then identify two types of memory systems in our universe, along with an important special case of the first type, which we distinguish as a third type of memory system. We show that two of these types of memory systems are time-symmetric, able to provide knowledge about both the past and the future. However, the third type of memory systems exploits the second law of thermodynamics, at least in all of its instances in our universe that we are aware of. The result is that in our universe, this type of memory system only ever provides information about the past. We also argue that human memory is of this third type, completing the argument. We end by scrutinizing the basis of the second law itself. This uncovers a previously unappreciated formal problem for common arguments that try to derive the second law from the "Past Hypothesis", i.e., from the claim that the very early universe was in a state of extremely low entropy. Our analysis is indebted to prior work by one of us but expands and improves upon this work in several respects.

Impaired and Spared Auditory Category Learning in Developmental Dyslexia.

Categorization has a deep impact on behavior, but whether category learning is served by a single system or multiple systems remains debated. Here, we designed two well-equated nonspeech auditory category learning challenges to draw on putative procedural (information-integration) versus declarative (rule-based) learning systems among adult Hebrew-speaking control participants and individuals with dyslexia, a language disorder that has been linked to a selective disruption in the procedural memory system and in which phonological deficits are ubiquitous. We observed impaired information-integration category learning and spared rule-based category learning in the dyslexia group compared with the neurotypical group. Quantitative model-based analyses revealed reduced use of, and slower shifting to, optimal procedural-based strategies in dyslexia with hypothesis-testing strategy use on par with control participants. The dissociation is consistent with multiple category learning systems and points to the possibility that procedural learning inefficiencies across categories defined by complex, multidimensional exemplars may result in difficulty in phonetic category acquisition in dyslexia.

Benefits of collaborative remembering in older and younger couples: the role of conversation dynamics and gender.

We experimentally explored whether and how conversation dynamics would benefit collaborative remembering in intimate couples over time. To this end, we ran a study with a three-factor mixed design with relationship type (couples vs. strangers) and age (older adults vs. younger adults) as between-participants variables, and remembering condition (collaborative vs. individual) as a within-participants variable. Thirty pairs of intimate couples (fifteen long-term relationship older couples, fifteen short-term relationship younger couples) and thirty pairs of corresponding stranger-pairs (including older strangers and younger strangers) were compared with respect to recall accuracy and conversation dynamics, specifically considering the role of gender. Results revealed significant collaborative facilitation only in older couples. Also, females' communication behaviours facilitated males' collaborative remembering performance only in older (vs. younger) couples. In addition, a gender-specific pattern of shifts from the individual to collaborative context emerged only in older couple (vs. strangers). The findings are consistent with the notion that a longer experience of collaboration and more effective conversation dynamics allow older (vs. younger) couples to perform better at collaborative remembering. We discuss processes underlying the observed gender differences, and the social and motivational implications of collaborative remembering.

When and how do children solve the Weather Prediction Task?

The Weather Prediction Task (WPT) can be solved by adults using several strategies dependent on different memory systems. One developmental study reported that 8-12-year-old children can solve WPT-like tasks but, because of inadequate analyses, the cognitive processes involved in solving the task have not been established. The present study aimed to determine at what age children can first solve the WPT and identify the strategies used by children of different ages. We tested 3-12-year-old typically developing children and 20-30-year-old adults on a modified WPT. We performed detailed analyses of performance for each pattern of cue-outcome associations to decipher the strategies used by participants. None of the 3-5.5-year-old children solved the task. About one third of 5.5-7.5-year-old children performed above chance, relying only on the two most predictive cues. In contrast, about 80% of 7.5-12-year-old children performed above chance, relying on a conditional hierarchical strategy. Similar to 20-30-year-old adults, 7.5-12-year-old children considered the highly predictive cues primarily and the less predictive cues secondarily. These findings indicate that the learning strategies used to solve the WPT evolve from middle to late childhood and reflect an increasing ability to use a conditional strategy concomitant with the development of the hippocampus-dependent memory system.

Stress-induced cortisol modulates the control of memory retrieval towards the dorsal striatum.

Stress can modulate the recruitment of multiple memory systems during learning, favouring dorsal striatal "habit" learning over hippocampal "cognitive" learning. Here, we tested whether stress may also bias the engagement of "cognitive" and "habit" systems during retrieval and thereby affect the nature of remembering. To this end, participants first performed a probabilistic classification learning task that can be solved by both the "cognitive" and the "habit" system. Twenty-four hours later, participants underwent either a stress manipulation or a non-stressful control procedure before they completed a retention test for the previously learned task in the MRI scanner. During this retention test, stress-induced cortisol levels were linked to a relative bias towards behavioural strategies indicative for the "habit" system. At the neural level, stress led to increased dorsal striatal activity during retrieval. Elevated cortisol levels were directly correlated with increased activity in the dorsal striatum and further linked to reduced functional connectivity between the hippocampus and the amygdala, which is assumed to orchestrate the stress-related shift from "cognitive" to "habitual" control. Together, our data suggest that stress may bias the contributions of multiple memory systems also at retrieval, in a manner that promotes dorsal striatal "habit" processes and most likely driven by cortisol.

Neurobiological insights into twice-exceptionality: Circuits, cells, and molecules.

Twice-exceptional learners face a unique set of challenges arising from the intersection of extraordinary talent and disability. Neurobiology research has the capacity to complement pedagogical research and provide support for twice-exceptional learners. Very few studies have attempted to specifically address the neurobiological underpinnings of twice-exceptionality. However, neurobiologists have built a broad base of knowledge in nervous system function spanning from the level of neural circuits to the molecular basis of behavior. It is known that distinct neural circuits mediate different neural functions, which suggests that 2e learning may result from enhancement in one circuit and disruption in another. Neural circuits are known to adapt and change in response to experience, a cellular process known as neuroplasticity. Plasticity is controlled by a bidirectional connection between the synapse, where neural signals are received, and the nucleus, where regulated gene expression can return to alter synaptic function. Complex molecular mechanisms compose this connection in distinct neural circuits, and genetic alterations in these mechanisms are associated with both memory enhancements and psychiatric disorder. Understanding the consequences of these changes at the molecular, cellular, and circuit levels will provide critical insights into the neurobiological bases of twice-exceptionality.

Progesterone rapidly alters the use of place and response memory during spatial navigation in female rats.

17ß-Estradiol (E2) and progesterone (P) influence place and response memory in female rats in spatial navigation tasks. Use of these memory systems is associated with the hippocampus and the dorsal striatum, respectively. Injections of E2 result in a well-established bias to use place memory, while much less is understood about the role of P. A total of 120 ovariectomized female rats were tested within a dual-solution T-maze task and treated with either low E2 (n = 24), high E2 (10 µg/kg; n = 24), or high E2 in combination with P (500 µg/kg) at three time points before testing: 15 min (n = 24), 1 h (n = 24), and 4 h (n = 24). Given alone, high E2 biases rats to the use of place memory, but this effect is reversed when P is given 1 h or 4 h before testing. This indicates that P may be playing an inhibitory role in the hippocampus during spatial tasks, which is consistent with past findings. Our findings show that P acts rapidly (within an hour) to affect performance during spatial tasks.

The role of transactive memory systems, psychological safety and interpersonal conflict in hospital team performance.

Safe patient care in hospitals relies on teamwork. Transactive Memory Systems (TMS), are shared cognitive systems that have been linked to team performance in other domains, but have received limited attention in healthcare. This study investigated the role of TMS, psychological safety and interpersonal conflict in predicting team performance in hospital ward teams where team membership is dynamic and often loosely defined. Hospital staff (n = 106) in four wards completed a battery of instruments assessing team performance, TMS, psychological safety and interpersonal conflict. TMS was a weak predictor of team performance, but the relationship was mediated by psychological safety. Overall, team performance was predicted by high psychological safety, low interpersonal conflict and low reliance on team members' knowledge (i.e. TMS credibility). These findings suggest that, in hospital teams, TMS is not a strong predictor of team performance but team culture is critical to ensure the quality and safety of patient care. Practitioner summary: This study investigated the role of Transactive Memory Systems (TMS) and cultural factors in hospital team performance. Team performance was predicted by psychological safety, low interpersonal conflict and low reliance on team members' untested knowledge. This highlights the importance of a supportive and psychologically safe team culture for safe care in hospitals. Abbreviations: TMS: transactive memory systems; HCA: health care assistant.

The Hippocampus and Dorsolateral Striatum Integrate Distinct Types of Memories through Time and Space, Respectively.

Several decades of research have established that different kinds of memories result from the activity of discrete neural networks. Studying how these networks process information in experiments that target specific types of mnemonic representations has provided deep insights into memory architecture and its neural underpinnings. However, in natural settings reality confronts organisms with problems that are not neatly compartmentalized. Thus, a critical problem in memory research that still needs to be addressed is how distinct types of memories are ultimately integrated. Here we demonstrate how two memory networks, the hippocampus and dorsolateral striatum, may accomplish such a goal. The hippocampus supports memory for facts and events, collectively known as declarative memory and often studied as spatial memory in rodents. The dorsolateral striatum provides the basis for habits that are assessed in stimulus-response types of tasks. Expanding previous findings, the current work revealed that in male Long-Evans rats, the hippocampus and dorsolateral striatum use time and space in distinct and largely complementary ways to integrate spatial and habitual representations. Specifically, the hippocampus supported both types of memories when they were formed in temporal juxtaposition, even if the learning took place in different environments. In contrast, the lateral striatum supported both types of memories if they were formed in the same environment, even at temporally distinct points. These results reveal for the first time that by using fundamental aspects of experience in specific ways, the hippocampus and dorsolateral striatum can transcend their attributed roles in information storage.SIGNIFICANCE STATEMENT The current paradigm in memory research postulates that different types of memories reflected in separate types of behavioral strategies result from activity in distinct neural circuits. However, recent data have shown that when rats concurrently acquired in the same environment of hippocampal-dependent spatial navigation and striatal-dependent approach of a visual cue, each of the two types of memories became dependent on both the hippocampus and dorsolateral striatum. The current work reveals that the hippocampus and dorsolateral striatum use distinct and complementary principles to integrate different types of memories in time and space: the hippocampus integrates memories formed in temporal proximity, while the lateral striatum integrates memories formed in the same space.

Time-dependent changes in hippocampal and striatal glycogen long after maze training in male rats.

Long-lasting biological changes reflecting past experience have been studied in and typically attributed to neurons in the brain. Astrocytes, which are also present in large number in the brain, have recently been found to contribute critically to learning and memory processing. In the brain, glycogen is primarily found in astrocytes and is metabolized to lactate, which can be released from astrocytes. Here we report that astrocytes themselves have intrinsic neurochemical plasticity that alters the availability and provision of metabolic substrates long after an experience. Rats were trained to find food on one of two versions of a 4-arm maze: a hippocampus-sensitive place task and a striatum-sensitive response task. Remarkably, hippocampal glycogen content increased while striatal levels decreased during the 30 days after rats were trained to find food in the place version, but not the response version, of the maze tasks. A long-term consequence of the durable changes in glycogen stores was seen in task-by-site differences in extracellular lactate responses activated by testing on a working memory task administered 30 days after initial training, the time when differences in glycogen content were most robust. These results suggest that astrocytic plasticity initiated by a single experience may augment future availability of energy reserves, perhaps priming brain areas to process learning of subsequent experiences more effectively.

Amygdala-hippocampal interactions in synaptic plasticity and memory formation.

Memories of emotionally arousing events tend to endure longer than other memories. This review compiles findings from several decades of research investigating the role of the amygdala in modulating memories of emotional experiences. Episodic memory is a kind of declarative memory that depends upon the hippocampus, and studies suggest that the basolateral complex of the amygdala (BLA) modulates episodic memory consolidation through interactions with the hippocampus. Although many studies in rodents and imaging studies in humans indicate that the amygdala modulates memory consolidation and plasticity processes in the hippocampus, the anatomical pathways through which the amygdala affects hippocampal regions that are important for episodic memories were unresolved until recent optogenetic advances made it possible to visualize and manipulate specific BLA efferent pathways during memory consolidation. Findings indicate that the BLA influences hippocampal-dependent memories, as well as synaptic plasticity, histone modifications, gene expression, and translation of synaptic plasticity associated proteins in the hippocampus. More recent findings from optogenetic studies suggest that the BLA modulates spatial memory via projections to the medial entorhinal cortex, and that the frequency of activity in this pathway is a critical element of this modulation.

Mechanisms of context conditioning in the developing rat.

The article reviews our studies of contextual fear conditioning (CFC) in rats during a period of development---Postnatal Day (PND) 17-33---that represents the late-infant, juvenile, and early-adolescent stages. These studies seek to acquire 'systems level' knowledge of brain and memory development and apply it to a rodent model of Fetal Alcohol Spectrum Disorder (FASD). This rodent model focuses on alcohol exposure from PND4-9, a period of brain development equivalent to the human third trimester, when neocortex, hippocampus, and cerebellum are especially vulnerable to adverse effects of alcohol. Our research emphasizes a variant of CFC, termed the Context Preexposure Facilitation Effect (CPFE, Fanselow, 1990), in which context representations incidentally learned on one occasion are retrieved and associated with immediate shock on a subsequent occasion. These representations can be encoded at the earliest developmental stage but seem not to be retained or retrieved until the juvenile period. This is associated with developmental differences in context-elicited expression, in prefrontal cortex, hippocampus, and amygdala, of immediate early genes (IEGs) that are implicated in long-term memory. Loss-of-function studies establish a functional role for these regions as soon as the CPFE emerges during ontogeny. In our rodent model of FASD, the CPFE is much more sensitive to alcohol dose than other commonly used cognitive tasks. This impairment can be reversed by acute administration during behavioral testing of drugs that enhance cholinergic function. This effect is associated with normalized IEG expression in prefrontal cortex during incidental context learning. In summary, our findings suggest that long-term memory of incidentally-learned context representations depends on prefrontal-hippocampal circuitry that is important both for the normative development of context conditioning and for its disruption by developmental alcohol exposure.

Interindividual differences in memory system local field potential activity predict behavioral strategy on a dual-solution T-maze.

Individuals can use diverse behavioral strategies to navigate their environment including hippocampal-dependent place strategies reliant upon cognitive maps and striatal-dependent response strategies reliant upon egocentric body turns. The existence of multiple memory systems appears to facilitate successful navigation across a wide range of environmental and physiological conditions. The mechanisms by which these systems interact to ultimately generate a unitary behavioral response, however, remain unclear. We trained 20 male, Sprague-Dawley rats on a dual-solution T-maze while simultaneously recording local field potentials that were targeted to the dorsolateral striatum and dorsal hippocampus. Eight rats spontaneously exhibited a place strategy while the remaining 12 rats exhibited a response strategy. Interindividual differences in behavioral strategy were associated with distinct patterns of LFP activity between the dorsolateral striatum and dorsal hippocampus. Specifically, striatal-hippocampal theta activity was in-phase in response rats and out-of-phase in place rats and response rats exhibited elevated striatal-hippocampal coherence across a wide range of frequency bands. These contrasting striatal-hippocampal activity regimes were (a) present during both maze-learning and a 30 min premaze habituation period and (b) could be used to train support vector machines to reliably predict behavioral strategy. Distinct patterns of neuronal activity across multiple memory systems, therefore, appear to bias behavioral strategy selection and thereby contribute to interindividual differences in behavior.

Resistance, vulnerability and resilience: A review of the cognitive cerebellum in aging and neurodegenerative diseases.

In the context of neurodegeneration and aging, the cerebellum is an enigma. Genetic markers of cellular aging in cerebellum accumulate more slowly than in the rest of the brain, and it generates unknown factors that may slow or even reverse neurodegenerative pathology in animal models of Alzheimer's Disease (AD). Cerebellum shows increased activity in early AD and Parkinson's disease (PD), suggesting a compensatory function that may mitigate early symptoms of neurodegenerative pathophysiology. Perhaps most notably, different parts of the brain accumulate neuropathological markers of AD in a recognized progression and generally, cerebellum is the last brain region to do so. Taken together, these data suggest that cerebellum may be resistant to certain neurodegenerative mechanisms. On the other hand, in some contexts of accelerated neurodegeneration, such as that seen in chronic traumatic encephalopathy (CTE) following repeated traumatic brain injury (TBI), the cerebellum appears to be one of the most susceptible brain regions to injury and one of the first to exhibit signs of pathology. Cerebellar pathology in neurodegenerative disorders is strongly associated with cognitive dysfunction. In neurodegenerative or neurological disorders associated with cerebellar pathology, such as spinocerebellar ataxia, cerebellar cortical atrophy, and essential tremor, rates of cognitive dysfunction, dementia and neuropsychiatric symptoms increase. When the cerebellum shows AD pathology, such as in familial AD, it is associated with earlier onset and greater severity of disease. These data suggest that when neurodegenerative processes are active in the cerebellum, it may contribute to pathological behavioral outcomes. The cerebellum is well known for comparing internal representations of information with observed outcomes and providing real-time feedback to cortical regions, a critical function that is disturbed in neuropsychiatric disorders such as intellectual disability, schizophrenia, dementia, and autism, and required for cognitive domains such as working memory. While cerebellum has reciprocal connections with non-motor brain regions and likely plays a role in complex, goal-directed behaviors, it has proven difficult to establish what it does mechanistically to modulate these behaviors. Due to this lack of understanding, it's not surprising to see the cerebellum reflexively dismissed or even ignored in basic and translational neuropsychiatric literature. The overarching goals of this review are to answer the following questions from primary literature: When the cerebellum is affected by pathology, is it associated with decreased cognitive function? When it is intact, does it play a compensatory or protective role in maintaining cognitive function? Are there theoretical frameworks for understanding the role of cerebellum in cognition, and perhaps, illnesses characterized by cognitive dysfunction? Understanding the role of the cognitive cerebellum in neurodegenerative diseases has the potential to offer insight into origins of cognitive deficits in other neuropsychiatric disorders, which are often underappreciated, poorly understood, and not often treated.

Ventromedial Prefrontal Cortex Is Necessary for Normal Associative Inference and Memory Integration.

The ability to flexibly combine existing knowledge in response to novel circumstances is highly adaptive. However, the neural correlates of flexible associative inference are not well characterized. Laboratory tests of associative inference have measured memory for overlapping pairs of studied items (e.g., AB, BC) and for nonstudied pairs with common associates (i.e., AC). Findings from functional neuroimaging and neuropsychology suggest the ventromedial prefrontal cortex (vmPFC) may be necessary for associative inference. Here, we used a neuropsychological approach to test the necessity of vmPFC for successful memory-guided associative inference in humans using an overlapping pairs associative memory task. We predicted that individuals with focal vmPFC damage (n = 5; 3F, 2M) would show impaired inferential memory but intact non-inferential memory. Performance was compared with normal comparison participants (n = 10; 6F, 4M). Participants studied pairs of visually presented objects including overlapping pairs (AB, BC) and nonoverlapping pairs (XY). Participants later completed a three-alternative forced-choice recognition task for studied pairs (AB, BC, XY) and inference pairs (AC). As predicted, the vmPFC group had intact memory for studied pairs but significantly impaired memory for inferential pairs. These results are consistent with the perspective that the vmPFC is necessary for memory-guided associative inference, indicating that the vmPFC is critical for adaptive abilities that require application of existing knowledge to novel circumstances. Additionally, vmPFC damage was associated with unexpectedly reduced memory for AB pairs post-inference, which could potentially reflect retroactive interference. Together, these results reinforce an emerging understanding of a role for the vmPFC in brain networks supporting associative memory processes.SIGNIFICANCE STATEMENT We live in a constantly changing environment, so the ability to adapt our knowledge to support understanding of new circumstances is essential. One important adaptive ability is associative inference which allows us to extract shared features from distinct experiences and relate them. For example, if we see a woman holding a baby, and later see a man holding the same baby, then we might infer that the two adults are a couple. Despite the importance of associative inference, the brain systems necessary for this ability are not known. Here, we report that damage to human ventromedial prefrontal cortex (vmPFC) disproportionately impairs associative inference. Our findings show the necessity of the vmPFC for normal associative inference and memory integration.

Aberrant memory system connectivity and working memory performance in subjective cognitive decline.

Subjective cognitive decline, a perceived worsening of cognitive functioning without objective deficit on assessment, could indicate incipient dementia. However, the neural correlates of subjective cognitive decline as assessed by magnetic resonance imaging remain somewhat unclear. Here, we evaluated differences in functional connectivity across memory regions, and cognitive performance, between healthy older adults aged 50 to 85 with (n = 35, Age = 68.5 ±â€¯7.7, 22 female), and without (n = 48, Age = 67.0 ±â€¯8.8, 29 female) subjective cognitive decline. We also evaluated neurite density, fractional anisotropy, and mean diffusivity of the parahippocampal cingulum, cingulate gyrus cingulum, and uncinate fiber bundles in a subsample of participants (n = 37). Participants with subjective cognitive decline displayed lower average functional connectivity across regions of a putative posterior memory system, and lower retrosplenial-precuneus functional connectivity specifically, than those without memory complaints. Furthermore, participants with subjective cognitive decline performed poorer than controls on visual working memory. However, groups did not differ in cingulum or uncinate diffusion measures. Our results show differences in functional connectivity and visual working memory in participants with subjective cognitive decline that could indicate potential incipient dementia.

Memory systems 2018 - Towards a new paradigm.

The multiple memory systems theory (MMS) postulates that the brain stores information based on the independent and parallel activity of a number of modules, each with distinct properties, dynamics, and neural basis. Much of the evidence for this theory comes from dissociation studies indicating that damage to restricted brain areas cause selective types of memory deficits. MMS has been the prevalent paradigm in memory research for more than thirty years, even as it has been adjusted several times to accommodate new data. However, recent empirical results indicating that the memory systems are not always dissociable constitute a challenge to fundamental tenets of the current theory because they suggest that representations formed by individual memory systems can contribute to more than one type of memory-driven behavioral strategy. This problem can be addressed by applying a dynamic network perspective to memory architecture. According to this view, memory networks can reconfigure or transiently couple in response to environmental demands. Within this context, the neural network underlying a specific memory system can act as an independent unit or as an integrated component of a higher order meta-network. This dynamic network model proposes a way in which empirical evidence that challenges the idea of distinct memory systems can be incorporated within a modular memory architecture. The model also provides a framework to account for the complex interactions among memory systems demonstrated at the behavioral level. Advances in the study of dynamic networks can generate new ideas to experimentally manipulate and control memory in basic or clinical research.

The role of context in animal memory.

Past research has shown that testing memory in the same context in which the memory was encoded leads to improved retention relative to testing memory in a new context. Context-dependent memory is directly related to the extent to which the encoding context can be reproduced. An experiment with pigeons is reported in which the context was a colored house-light that completely enveloped the learning and testing contexts. Under this condition, perfect retention of a visual discrimination that reversed at midsession was shown. Beyond reactivation of memory, new research with pigeons suggests that context provides access to different working and reference memory systems. Finally, experiments are reported that suggest context may selectively access information about features from the different dimensions of place, color, and time.