Spatial and nonspatial working memory and visual search

It has been indicated that visual search is interfered with in spatial working memory (WM), although not in nonspatial WM. In this study, the effects on visual search of individual differences in spatial and nonspatial WM were examined. Two visual search conditions were used: a conjunction search condition comprising two features (color and shape) and a disjunction condition comprising only one feature (color or shape). 96 participants (42 men, 54 women, M age = 20.9 yr., SD = 3.5) participated in this study. The participants were divided into high and low WM groups based on their spatial and nonspatial WM test scores. As a result, statistically significant group differences in the conjunction search rate were observed in spatial WM but not in nonspatial WM. These results suggest there is a relationship between visual search and the individual spatial WM ability, but this does not hold for nonspatial WM.     

A cognitive map for object memory in the hippocampus

The hippocampus has been proposed to support a cognitive map, a mental representation of the spatial layout of an environment as well as the nonspatial items encountered in that environment. In the present study, we recorded simultaneously from 43 to 61 hippocampal pyramidal cells as rats performed an object recognition memory task in which novel and repeated objects were encountered in different locations on a circular track. Multivariate analyses of the neural data indicated that information about object identity was represented secondarily to the primary information dimension of object location. In addition, the neural data related to performance on the recognition memory task. The results suggested that objects were represented as points of interest on the hippocampal cognitive map and that this map was useful in remembering encounters with particular objects in specific locations.The hippocampus plays an important role in spatial memory for both humans and rodents (O''Keefe 1999; Burgess et al. 2002). Findings from many studies in rodents indicate that the hippocampus supports memory for locations referenced to external landmarks, a capacity that O''Keefe and Nadel (1978) described over 30 yr ago as a “cognitive map” (using a term they borrowed from Tolman 1948). In the time since that pioneering thesis, it has become clear that the rodent hippocampus is also important for nonspatial memory (Eichenbaum et al. 1999). Damage to the rat hippocampus (defined here as CA fields, dentate gyrus, and subiculum) leads to impairments on nonspatial tasks, including object recognition memory (Clark et al. 2000; Fortin et al. 2004), transitive odor associations (Bunsey and Eichenbaum 1996), memory for temporal order (Fortin et al. 2002; Kesner et al. 2002), and social transmission of food preference (Alvarez et al. 2001; Clark et al. 2002).The circuitry by which information arrives at and exits from the hippocampus is consistent with the idea that the hippocampus is important for both spatial and nonspatial memory. In both rats and macaques, detailed anatomical studies have indicated that spatial information arrives at the hippocampus via the postrhinal cortex (parahippocampal cortex in primates) and the medial entorhinal cortex, whereas nonspatial information takes a path largely through the perirhinal cortex and lateral entorhinal cortex (Witter and Amaral 1991; Suzuki and Amaral 1994; Witter et al. 2000). Thus, the hippocampus is ideally situated to combine spatial and nonspatial information in the service of remembering item–location associations (Manns and Eichenbaum 2006).Single-unit recording studies in the rat hippocampus have largely focused on the spatial correlates of hippocampal pyramidal neuron firing rates. Fewer studies have investigated nonspatial correlates of hippocampal activity during memory tasks for nonspatial items. However, in one such study, Wood et al. (1999) found that some individual hippocampal pyramidal neurons responded to particular odors and that others responded to particular odors in specific locations during an odor recognition memory task. Thus, activity of individual cells appeared to contain information about nonspatial items as well as spatial locations.An important question is how the activity of individual hippocampal neurons combine to represent item–location associations as a neural ensemble. In particular, how is an encounter with an object in a particular location represented in the pattern of spiking among many hippocampal pyramidal neurons? How might this representation relate to memory for the object or for the location? In the present study, we recorded simultaneously from 43 to 61 hippocampal pyramidal cells as rats performed an object recognition memory task in which novel and repeated objects were encountered in different locations on a circular track. Multivariate analyses of the neural data indicated that information about object identity was represented secondarily to the primary information dimension of object location. In addition, the analyses indicated that the neural data related to performance on the recognition memory task. The results suggest that objects were represented as points of interest on the hippocampal cognitive map and that this map was useful in remembering encounters with particular objects in specific locations.     

Impaired odor recognition memory in patients with hippocampal lesions

In humans, impaired recognition memory following lesions thought to be limited to the hippocampal region has been demonstrated for a wide variety of tasks. However, the importance of the human hippocampus for olfactory recognition memory has scarcely been explored. We evaluated the ability of memory-impaired patients with damage thought to be limited to the hippocampal region to recognize a list of odors. The patients were significantly impaired after a retention delay of 1 h. Olfactory sensitivity was intact. This finding is in agreement with earlier reports that rats with hippocampal lesions exhibited memory impairment on an odor delayed nonmatching to sample task (after 30 min and 1 h) and that patients with damage thought to be limited to the hippocampal region were impaired on an odor span memory task. Olfactory recognition memory, similar to recognition memory in other sensory modalities, depends on the integrity of the hippocampal region.     

加工速度和工作记忆在认知年老化过程中的作用

研究加工速度和工作记忆在认知能力年老化过程中的作用。被试为20-90岁的健康成人1350名,完成加工速度、工作记忆及一组认知能力测验,应用分层回归分析方法考查在年老化过程中加工速度和工作记忆对多项认知变量的作用量。结果表明,在年老化过程中,加工速度和工作记忆对认知变量的作用量分别约为87% 和76%;二者的综合作用量约为94%。说明加工速度和工作记忆在认知能力年老化过程中起决定作用     

Allocentric spatial learning by hippocampectomised rats: A further test of the “spatial mapping” and “working memory” theories of hippocampal function

This paper reports a series of three experiments that tested the “spatial-mapping” and “working-memory” theories of hippocampal function. The experimental designs incorporate separate reference- and working-memory procedures of a water-escape task, using both spatial and non-spatial learning. In Experiment 1 (Reference memory), rats with hippocampal (HC) or cortical (CC) lesions and unoperated (UNOP) rats learned to swim to a rigid visible escape platform while avoiding contact with a floating one. In the nonspatial task, the platforms each occupied any of 8 possible positions in the pool over successive trials but differed in appearance. In the spatial task, the platforms were of identical appearance but the safe one always occupied a single fixed location. The HC rats showed a highly specific spatial learning impairment but did learn to perform consistently above chance towards the end of training. In Experiment 2 (working memory), new groups of rats were trained on similar spatial and nonspatial tasks, but the platform designated correct-in terms of its visual appearance or its spatial location-was randomly changed each day. No animal learned the nonspatial task despite extensive training. Performance on the spatial version unexpectedly revealed an impairment in the CC as well as the HC group relative to the UNOP rats. However, the HCs again performed at above chance levels and demonstrated rapid (I-trial) spatial learning towards the end training. Experiment 3 used a place navigation matching-to-sample task examine spatial working memory further. Each day, an underwater platform was hidden at any of 4 possible locations, and the rats were given 2 trials to search for it. Both UNOP and CC rats located the platform faster on Trial 2 than on Trial 1, even when the inter-trial interval was long as 30min. HC rats were no faster on Trial 2 than on Trial 1. We conclude that hippocampal lesions (1) severely but partially impair spatial but not visual reference memory and (2) give rise to different patterns impairment in different working-mermory tasks. The results are a chal lenge to both the spatial-mapping and working-memory theories.     

Differential effects of damage within the hippocampal region on memory for a natural, nonspatial Odor-Odor Association

Debate continues on whether the role of rodent hippocampus in memory is limited to the spatial domain. Recently, this controversy has been addressed with studies on the social transmission of food preference, an odor-odor association task with no spatial requirements. Multiple reports have concluded that damage to the hippocampal region impairs memory in this task, but there remain questions about the extent of damage essential to produce an impairment. Furthermore, a recent study () found no effect of hippocampal lesions on memory in this task. We tested animals with complete lesions of the hippocampus (H) lesions of the hippocampus plus subiculum (HS), and lesions of the adjacent, anatomically related cortices of the parahippocampal region (PHR). H lesions produced an impairment on spatial delayed alternation, but not on memory for the social transmission of food preference, whereas HS and PHR lesions produced severe and equivalent impairments on memory for the socially acquired food preference. We discuss possible explanations for the discrepancy with the results of and conclude that the hippocampus and subiculum together play a critical role in the formation of this form of nonspatial, relational memory.     

Mental representations of spatial and nonspatial relations

Four experiments investigated the representation and integration in memory of spatial and nonspatial relations. Subjects learned two-dimensional spatial arrays in which critical pairs of object names were semantically related (Experiment 1), semantically and episodically related (Experiment 2), or just episodically related (Experiments 3a and 3b). Episodic relatedness was established in a paired-associate learning task that preceded array learning. After learning an array, subjects participated in two tasks: item recognition, in which the measure of interest was priming; and distance estimation. Priming in item recognition was sensitive to the Euclidean distance between object names and, for neighbouring locations, to nonspatial relations. Errors in distance estimations varied as a function of distance but were unaffected by nonspatial relations. These and other results indicated that nonspatial relations influenced the probability of encoding spatial relations between locations but did not lead to distorted spatial memories.     

Deletion of the GluA1 AMPA receptor subunit impairs recency-dependent object recognition memory

Deletion of the GluA1 AMPA receptor subunit impairs short-term spatial recognition memory. It has been suggested that short-term recognition depends upon memory caused by the recent presentation of a stimulus that is independent of contextual-retrieval processes. The aim of the present set of experiments was to test whether the role of GluA1 extends to nonspatial recognition memory. Wild-type and GluA1 knockout mice were tested on the standard object recognition task and a context-independent recognition task that required recency-dependent memory. In a first set of experiments it was found that GluA1 deletion failed to impair performance on either of the object recognition or recency-dependent tasks. However, GluA1 knockout mice displayed increased levels of exploration of the objects in both the sample and test phases compared to controls. In contrast, when the time that GluA1 knockout mice spent exploring the objects was yoked to control mice during the sample phase, it was found that GluA1 deletion now impaired performance on both the object recognition and the recency-dependent tasks. GluA1 deletion failed to impair performance on a context-dependent recognition task regardless of whether object exposure in knockout mice was yoked to controls or not. These results demonstrate that GluA1 is necessary for nonspatial as well as spatial recognition memory and plays an important role in recency-dependent memory processes.     

Differential age-related effects on conjunctive and relational visual short-term memory binding

An age-related associative deficit has been described in visual short-term binding memory tasks. However, separate studies have suggested that ageing disrupts relational binding (to associate distinct items or item and context) more than conjunctive binding (to integrate features within an object). The current study directly compared relational and conjunctive binding with a short-term memory task for object–colour associations in 30 young and 30 older adults. Participants studied a number of object–colour associations corresponding to their individual object span level in a relational task in which objects were associated to colour patches and a conjunctive task where colour was integrated into the object. Memory for individual items and for associations was tested with a recognition memory test. Evidence for an age-related associative deficit was observed in the relational binding task, but not in the conjunctive binding task. This differential impact of ageing on relational and conjunctive short-term binding is discussed by reference to two underlying age-related cognitive difficulties: diminished hippocampally dependent binding and attentional resources.     

Spatial relational memory in 9-month-old macaque monkeys

This experiment assesses spatial and nonspatial relational memory in freely moving 9-mo-old and adult (11-13-yr-old) macaque monkeys (Macaca mulatta). We tested the use of proximal landmarks, two different objects placed at the center of an open-field arena, as conditional cues allowing monkeys to predict the location of food rewards hidden in one of two sets of three distinct locations. Monkeys were tested in two different conditions: (1) when local visual cues marked the two sets of potentially baited locations, so that monkeys could use both local and spatial information to discriminate these locations from never-baited locations; and (2) when no local visual cues marked the two sets of potentially baited locations, so that monkeys had to rely on a spatial relational representation of the environment to discriminate these locations. No 9-mo-old or adult monkey associated the presence of the proximal landmarks, at the center of the arena, with the presence of food in one set of three distinct locations. All monkeys, however, discriminated the potentially baited locations in the presence of local visual cues, thus providing evidence of visual discrimination learning. More importantly, all 9-mo-old monkeys tested discriminated the potentially baited locations in absence of the local visual cues, thus exhibiting evidence of spatial relational learning. These findings indicate that spatial memory processes characterized by a relational representation of the environment are present as early as 9 mo of age in macaque monkeys.     

Hippocampal formation lesions impair performance in an odor-odor association task independently of spatial context

The rodent hippocampal system is known to play an important role in memory. Evidence that this role is not limited to spatial memory has come from studies using a variety of non-spatial memory tasks. One example is the social transmission of food preference paradigm, a task in which rats learn an odor-odor association with no explicit spatial memory component. However, because training and testing in this task typically take place in the same environment, it is possible that memory for the spatial context in which odors are experienced during training is critical to subsequent retention performance. If this is the case, it might be expected that lesions of the hippocampal system would impair memory performance by disrupting the establishment of a representation of the training environment. We addressed this issue by training rats in one spatial context and then testing them either in the same or a different spatial context. Normal control rats performed equally well when tested in an environment that was the same or different from that used during training, and the retention impairment exhibited by rats with hippocampus plus subiculum lesions was equivalent in the two test environments. These results support the view that the hippocampal system is necessary for the flexible expression of nonspatial memories even when the spatial context in which the memory is acquired is not critical to retrieval.     

Domain-dependent activation during spatial and nonspatial auditory working memory

Visual system has been proposed to be divided into two, the ventral and dorsal, processing streams. The ventral pathway is thought to be involved in object identification whereas the dorsal pathway processes information regarding the spatial locations of objects and the spatial relationships among objects. Several studies on working memory (WM) processing have further suggested that there is a dissociable domain-dependent functional organization within the prefrontal cortex for processing of spatial and nonspatial visual information. Also the auditory system is proposed to be organized into two domain-specific processing streams, similar to that seen in the visual system. Recent studies on auditory WM have further suggested that maintenance of nonspatial and spatial auditory information activates a distributed neural network including temporal, parietal, and frontal regions but the magnitude of activation within these activated areas shows a different functional topography depending on the type of information being maintained. The dorsal prefrontal cortex, specifically an area of the superior frontal sulcus (SFS), has been shown to exhibit greater activity for spatial than for nonspatial auditory tasks. Conversely, ventral frontal regions have been shown to be more recruited by nonspatial than by spatial auditory tasks. It has also been shown that the magnitude of this dissociation is dependent on the cognitive operations required during WM processing. Moreover, there is evidence that within the nonspatial domain in the ventral prefrontal cortex, there is an across-modality dissociation during maintenance of visual and auditory information. Taken together, human neuroimaging results on both visual and auditory sensory systems support the idea that the prefrontal cortex is organized according to the type of information being maintained in WM.     

Effects of environmental enrichment on spatial memory and neurochemistry in middle-aged mice

The present study compared the effects of environmental enrichment on spatial memory, glutamic acid decarboxylase (GAD) activity, and synaptophysin levels in middle-aged male and female mice. Prior to testing, a subset of 18-month-old male and female C57BL/6 mice was housed with two to three toys and a running wheel in the home cage for up to 29 d. Adult mice (7 mo) of both sexes and the remaining middle-aged mice were group (social) housed, but not exposed to enriching objects. After the enrichment period, all mice were tested in a 1-day version of the Morris water maze, in which both spatial and nonspatial memory were assessed. Immediately after testing, the hippocampus and frontoparietal cortex were dissected, and GAD activity and synaptophysin levels were measured. Environmental enrichment reduced the age-related impairment in spatial acquisition and retention; relative to adult social controls, middle-aged enriched mice were unimpaired, whereas middle-aged social controls were impaired. This reduction was similar in middle-aged males and females. Enrichment did not affect cued memory in either sex. Although hippocampal GAD activity was increased by enrichment in males, all other neurochemical measurements were unaffected by enrichment or aging in either sex. These data suggest that environmental enrichment initiated at middle age can reduce age-related impairments in spatial memory in males and females, although the underlying neurobiological mechanisms of this effect remain unknown.     

Drug-induced amnesia impairs implicit relational memory

Amnesic patients have difficulty consciously recollecting past events, but they perform normally on implicit memory measures that do not require conscious access. Now, an innovative study offers compelling evidence that even implicit memory can be impaired during amnesia, temporarily induced in healthy subjects using a neuropharmacological manipulation (midazolam). Importantly, midazolam impaired implicit, relational memory, whereas non-relational memory was preserved, supporting the characterization of conscious memory in terms of relational encoding.     

Transgenic mice expressing a truncated form of CREB-binding protein (CBP) exhibit deficits in hippocampal synaptic plasticity and memory storage

Deletions, translocations, or point mutations in the CREB-binding protein (CBP) gene have been associated with Rubinstein-Taybi Syndrome; a human developmental disorder characterized by retarded growth and reduced mental function. To examine the role of CBP in memory, transgenic mice were generated in which the CaMKII alpha promoter drives expression of an inhibitory truncated CBP protein in forebrain neurons. Examination of hippocampal long-term potentiation (LTP), a form of synaptic plasticity thought to underlie memory storage, revealed significantly reduced late-phase LTP induced by dopamine-regulated potentiation in hippocampal slices from CBP transgenic mice. However, four-train induced late-phase LTP is normal. Behaviorally, CBP transgenic mice exhibited memory deficits in spatial learning in the Morris water maze and deficits in long-term memory for contextual fear conditioning, two hippocampus-dependent tasks. Together, these results demonstrate that CBP is involved in specific forms of hippocampal synaptic plasticity and hippocampus-dependent long-term memory formation.     

The role of spatial working memory in visual search efficiency

Many theories have proposed that visual working memory plays an important role in visual search. In contrast, by showing that a nonspatial working memory load did not interfere with search efficiency, Woodman, Vogel, and Luck (2001) recently proposed that the role of working memory in visual search is insignificant. However, the visual search process may interfere with spatial working memory. In the present study, a visual search task was performed concurrently with either a spatial working memory task (Experiment 1) or a nonspatial working memory task (Experiment 2). We found that the visual search process interfered with a spatial working memory load, but not with a nonspatial working memory load. These results suggest that there is a distinction between spatial and nonspatial working memory in terms of interactions with visual search tasks. These results imply that the visual search process and spatial working memory storage require the same limited-capacity mechanisms.     

Spatial working memory load affects counting but not subitizing in enumeration

The present study investigated whether subitizing reflects capacity limitations associated with two types of working memory tasks. Under a dual-task situation, participants performed an enumeration task in conjunction with either a spatial (Experiment 1) or a nonspatial visual (Experiment 2) working memory task. Experiment 1 showed that spatial working memory load affected the slope of a counting function but did not affect subitizing performance or subitizing range. Experiment 2 showed that nonspatial visual working memory load affected neither enumeration efficiency nor subitizing range. Furthermore, in both spatial and nonspatial memory tasks, neither subitizing efficiency nor subitizing range was affected by amount of imposed memory load. In all the experiments, working memory load failed to influence slope, subitizing range, or overall reaction time. These findings suggest that subitizing is performed without either spatial or nonspatial working memory. A possible mechanism of subitizing with independent capacity of working memory is discussed.     

Age-related neural activity during allocentric spatial memory

Age-related decline in allocentric (viewer-independent) spatial memory is seen across species. We employed a virtual reality analogue of the Morris Water Maze to study the effect of healthy ageing on neural activity during allocentric spatial memory using functional magnetic resonance imaging. Voxel-based morphometry was used to ascertain hippocampal volumetric integrity. A widespread neural network comprising frontal, parietal, occipital, thalamic, and cerebellar regions was activated in young and older adults, but only young adults significantly activated bilateral hippocampus and left parahippocampus, as well as right frontal pole and dorso-lateral prefrontal cortex (DLPFC) during encoding and right DLPC during retrieval. Hippocampal grey matter volume was unchanged in older adults; however, prefrontal and parahippocampal functional attenuation was accompanied by volumetric reduction. We conclude that the decline in allocentric spatial memory with age is associated with attenuated hippocampal function, as well as compromised function and structure of prefrontal and parahippocampal regions.     

Evidence for the substantia nigra pars compacta as an essential component of a memory system independent of the hippocampal memory system

The aim of the present study was to test if the nigrostriatal pathway is an essential component for a water maze cued task learning and if it works independently of the hippocampal memory system. This hypothesis was tested using an animal model of Parkinson's disease in which male Wistar rats were lesioned in the substantia nigra pars compacta (SNc) by the intranigral infusion of 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP), thus causing a partial depletion of striatal dopamine. SNc-lesioned and sham-operated animals were implanted bilaterally with guide cannulae above the dorsal hippocampus in order to be tested after the administration of 0.4 microl 2% lidocaine or saline into this structure. The animals were tested in a spatial or in a cued version of the water maze, memory tasks previously reported to model hippocampal-dependent spatial/relational and striatal-dependent S-R learning, respectively. Hippocampal inactivation, but not SNc lesion, impaired learning and memory in the spatial version of the water maze. An opposite situation was observed with the cued version. No significant interaction was observed between the SNc lesion and hippocampal inactivation conditions affecting scores in the spatial or in the cued version of the water maze. These results suggest that the nigrostriatal pathway is an essential part of the memory system that processes S-R learning and that it works independently of the hippocampal memory system that processes spatial/relational memories.