Dissociable stages of problem solving (II): first evidence for process-contingent temporal order of activation in dorsolateral prefrontal cortex

In a companion study, eye-movement analyses in the Tower of London task (TOL) revealed independent indicators of functionally separable cognitive processes during problem solving, with processes of building up an internal representation of the problem preceding actual planning processes. These results imply that processes of internalization and planning should also be distinguishable in time and space with respect to concomitant brain activation patterns. To investigate this possibility, here we conducted analyses of fMRI data for left and right dorsolateral prefrontal cortex (dlPFC) during problem solving in the TOL task by accounting for the trial-by-trial variability of onsets and durations of the different cognitive processing stages. Comparisons between stimulus-locked and response-locked modeling approaches affirmed that activation in left dlPFC was elicited particularly during early processes of internalization, comprising the extraction of goal information and the generation of an internal problem representation, whereas activation in right dlPFC was predominantly attributable to later processes of mental transformations on this representation, that is planning proper. Thus, present data corroborate the proposal that often observed bilateral dlPFC activation patterns during complex cognitive tasks such as problem solving may reflect functionally and, to some extent, even temporally separable processes with opposing lateralizations.     

Hippocampal damage and exploratory preferences in rats: memory for objects, places, and contexts

Rats have a natural tendency to spend more time exploring novel objects than familiar objects, and this preference can be used as an index of object recognition. Rats also show an exploratory preference for objects in locations where they have not previously encountered objects (an index of place memory) and for familiar objects in contexts different from those in which the objects were originally encountered (an index of context memory). In this experiment, rats with cytotoxic lesions of the hippocampal formation were tested on all three versions of the novelty-preference paradigm, with a 5-min retention interval between the familiarization and test phases. Rats with sham lesions displayed a novelty preference on all three trial types, whereas the rats with hippocampal lesions displayed a novelty preference on Object trials but did not discriminate between the objects on Place trials or Context trials. The findings indicate that hippocampal damage impairs memory for contextual or spatial aspects of an experience, whereas memory for objects that were part of the same experience are left relatively intact.     

An analysis of independence and interactions of brain substrates that subserve multiple attributes, memory systems, and underlying processes

It is proposed that memory is organized into event-based, knowledge-based, and rule-based memory systems. Furthermore, each system is composed of the same set of multiple attributes and characterized by a set of process oriented operating characteristics that are mapped onto multiple neural regions and interconnected neural circuits. Based on this theoretical model of memory, it is possible to investigate the independence and interaction among brain regions between any two systems for any of the proposed attributes or processes. This applies also to the investigation of independence and interactions between any two attributes within a system and between processes associated with a system for any of the proposed attributes. In this article, research evidence is presented to suggest that there are both dissociations and interactions between the hippocampus and caudate nucleus in mediating spatial and response attributes within the event-based memory system, between the hippocampus and the parietal cortex in subserving the spatial attribute within the event-based and knowledge-based memory systems, and between the hippocampus and the prefrontal cortex in subserving the spatial attribute within the event-based and rule-based memory systems.     

Visual working memory for dynamic objects: Impaired binding between object feature and location

In daily life, people frequently need to observe dynamic objects and temporarily maintain their representations in visual working memory (VWM). The present study explored the mechanism underlying the binding between perceptual features and locations of dynamic objects in VWM. In three experiments, we measured and compared the memory performance for feature-location binding of multiple dynamic and static objects. The results showed that the feature-location binding was impaired for the dynamic objects compared with the static objects. The impairment persisted when the global spatial configuration of the objects remained intact during the motion, as well as when the binding task was relatively easy, such as binding between single-feature objects and coarse locations. The results indicate that object features and locations are not maintained in VWM as well-integrated object files; rather, the formation of feature-location binding may require additional processes, which are disrupted by the constant change of locations in dynamic circumstances. We propose a consolidation process as possible underlying mechanism, and discuss factors that may influence the strength of feature-location binding in dynamic circumstances.     

Preterm infant body composition,working memory,and temperament

Altered body composition in preterm infants is associated with risks to cognitive development, but the effect specific to prefrontal cortex (PFC) development is unknown. We were interested in the impact of fat mass (FM) and fat free mass (FFM) gains out to 4 months corrected gestational age (CGA) on PFC development, as indexed by working memory and temperament. This is a prospective observational pilot study recruiting 100 preterm (<33 weeks gestation), appropriate for gestational age, and very low birth weight infants, of which 49 infants met inclusion criteria. Body composition was measured using air displacement plethysmography at hospital discharge and 4 months CGA. Questionnaire based temperament assessments were completed at 12 and 24 months CGA and a working memory assessment was completed at 24 months CGA. Associations between developmental tests and body composition obtained at term and 4 months were analyzed. Increased FM at discharge was associated with increased fear and decreased soothability at 12 months. Increased FM at 4 months was associated with increased activity level, increased distress from limitations at 12 months and decreased attentional shifting, decreased frustration, and decreased inhibitory control at 24 months. Increased FFM at 4 months was associated with increased activity level at 12 months and increased impulsivity and decreased low intensity pleasure at 24 months. In this exploratory pilot study, increased FM out to 4 months and increased FFM after discharge are associated with negative markers of infant temperament. Infant temperament may be sensitive to body composition status at least to 4 months CGA.     

The hippocampus and prefrontal cortex are differentially involved in serial memory retrieval in non-stress and stress conditions

We previously showed that 24 h after learning, mice significantly remembered the first (D1) but not the second (D2) discrimination in a serial spatial task and that an acute stress delivered 5 min before the test phase reversed this memory retrieval pattern.A first experiment evaluated the effects of dorsal hippocampus (HPC) or prefrontal cortex (PFC) lesions, these two brain areas being well-known for their involvement in serial and spatial memory processes. For this purpose, six independent groups of mice were used: non-lesioned (controls), PFC or HPC-lesioned animals, submitted or not to an acute stress (electric footshocks; 0.9 mA). Results show that (i) non-stressed controls as well as PFC-lesioned mice (stressed or not) remembered D1 but not D2; (ii) stressed controls and HPC-lesioned mice (stressed or not) remembered D2 but not D1; (iii) stress significantly increased plasma corticosterone in controls and PFC-lesioned mice, but not in HPC-lesioned mice which already showed a significant plasma corticosterone increase in non-stressed condition.Since data from this first experiment showed that stress inhibited the hippocampal-dependent D1 memory retrieval, a second experiment evaluated the behavioral effect of intrahippocampal corticosterone injection in non-stressed mice. Results show that intrahippocampal corticosterone injection induced a reversal of serial memory retrieval pattern similar to that induced by acute stress.Overall, our study shows that (i) in non-stress condition, the emergence of D1 is HPC-dependent; (ii) in stress condition, the emergence of D2 requires the PFC integrity; moreover, intrahippocampal corticosterone injection mimicked the effects of stress in the CSD task.     

Coherence and recurrency: maintenance,control and integration in working memory

Working memory (WM), including a ‘central executive’, is used to guide behavior by internal goals or intentions. We suggest that WM is best described as a set of three interdependent functions which are implemented in the prefrontal cortex (PFC). These functions are maintenance, control of attention and integration. A model for the maintenance function is presented, and we will argue that this model can be extended to incorporate the other functions as well. Maintenance is the capacity to briefly maintain information in the absence of corresponding input, and even in the face of distracting information. We will argue that maintenance is based on recurrent loops between PFC and posterior parts of the brain, and probably within PFC as well. In these loops information can be held temporarily in an active form. We show that a model based on these structural ideas is capable of maintaining a limited number of neural patterns. Not the size, but the coherence of patterns (i.e., a chunking principle based on synchronous firing of interconnected cell assemblies) determines the maintenance capacity. A mechanism that optimizes coherent pattern segregation, also poses a limit to the number of assemblies (about four) that can concurrently reverberate. Top-down attentional control (in perception, action and memory retrieval) can be modelled by the modulation and re-entry of top-down information to posterior parts of the brain. Hierarchically organized modules in PFC create the possibility for information integration. We argue that large-scale multimodal integration of information creates an ‘episodic buffer’, and may even suffice for implementing a central executive.     

Discrimination of what,when, and where: Implications for episodic-like memory in rats

We investigated the discrimination of what, when, and where in rats (n = 5) using an eight-arm radial maze. Rats received daily training consisting of forced-choice visits to four baited arms, one of which was randomly chosen each day to contain chocolate (Phase 1). In Phase 2, all eight arms were available. After a short (30 min) retention interval (RI), the four arms that were not available in Phase 1 provided food. After a long (4 h) RI, the four remaining arms plus the arm containing chocolate provided food (i.e., the chocolate arm replenished). The rats visited the chocolate location after the long RI more than after the short RI. Next, chocolate was paired with lithium chloride, and subsequent testing used the long RI. The rats visited the chocolate location less after the taste-aversion manipulation than in previous training, demonstrating knowledge of what, when, and where. Implications for episodic-like memory are discussed.     

Effects of association and repetition in memory for temporal order

Temporal order recognition memory has been examined previously with tasks involving a recency judgment between a pair of items in a preceding string. Recency judgments are impaired when the earlier item is repeated. The present study employed the comparative recency judgment paradigm, with the lists composed of words. The effect of the inclusion in the list of a high associate of the earlier test item was examined and compared to the effect of repetition. Associative interference was observed, but not in all conditions. Direction of association was a significant factor. The results were interpreted in terms of a model of word recognition proposed by Morton.     

Selection of objects and tasks in working memory

When people hold several objects (such as digits or words) in working memory and select one for processing, switching to a new object takes longer than selecting the same object as that on the preceding processing step. Similarly, selecting a new task incurs task- switching costs. This work investigates the selection of objects and of tasks in working memory using a combination of object-switching and task-switching paradigms. Participants used spatial cues to select one digit held in working memory and colour cues to select one task (addition or subtraction) to apply to it. Across four experiments the mapping between objects and their cues and the mapping between tasks and their cues were varied orthogonally. When mappings varied from trial to trial for both objects and tasks, switch costs for objects and tasks were additive, as predicted by sequential selection or resource sharing. When at least one mapping was constant across trials, allowing learning of long-term associations, switch costs were underadditive, as predicted by partially parallel selection. The number of objects in working memory affected object-switch costs but not task-switch costs, counter to the notion of a general resource of executive attention.     

Visual working memory capacity for objects from different categories: a face-specific maintenance effect

The capacity of visual working memory was examined when complex objects from different categories were remembered. Previous studies have not examined how visual similarity affects object memory, though it has long been known that similar-sounding phonological information interferes with rehearsal in auditory working memory. Here, experiments required memory for two or four objects. Memory capacity was compared between remembering four objects from a single object category to remembering four objects from two different categories. Two-category sets led to increased memory capacity only when upright faces were included. Capacity for face-only sets never exceeded their nonface counterparts, and the advantage for two-category sets when faces were one of the categories disappeared when inverted faces were used. These results suggest that two-category sets which include faces are advantaged in working memory but that faces alone do not lead to a memory capacity advantage.     

Contribution of the dorsal subiculum to memory for temporal order and novelty detection using objects,odors, or spatial locations in the rat

The contribution of the dorsal subiculum (DS) to memory for temporal order and novelty detection was assessed using a spontaneous exploration paradigm with objects (visual/tactile stimuli), odors, or spatial locations (Hunsaker, Fieldsted, Rosenberg, & Kesner, 2008). Rats with selective excitotoxic lesions of the DS were compared to sham-operated rats (SHAM) in the two exploration tests. In temporal order tests, two previously explored stimuli were presented and normal rats typically show a preference for exploring the stimulus that was first explored compared to the other stimulus. In novelty detection tests, a familiar and a new stimulus were presented and normal rats typically have a preference for exploring new stimuli. In temporal order tests, results indicated that Group SHAM explored significantly more the first than the last stimulus they met when the stimuli were odors or objects. In addition, SHAM rats predictably displayed a significant preference for the new stimulus in the novelty detection tests with objects, odors, and spatial locations. Group DS did not differ from controls on the temporal order and the novelty detection tests with objects or odors. However, on the novelty detection test with spatial locations, Group DS differed from Group SHAM. These results suggest that the DS is necessary for the memory of spatial locations but not of objects and odors.     

Full scenes produce more activation than close-up scenes and scene-diagnostic objects in parahippocampal and retrosplenial cortex: an fMRI study

We used fMRI to directly compare activation in two cortical regions previously identified as relevant to real-world scene processing: retrosplenial cortex and a region of posterior parahippocampal cortex functionally defined as the parahippocampal place area (PPA). We compared activation in these regions to full views of scenes from a global perspective, close-up views of sub-regions from the same scene category, and single objects highly diagnostic of that scene category. Faces were included as a control condition. Activation in parahippocampal place area was greatest for full scene views that explicitly included the 3D spatial structure of the environment, with progressively less activation for close-up views of local scene regions containing diagnostic objects but less explicitly depicting 3D scene geometry, followed by single scene-diagnostic objects. Faces did not activate parahippocampal place area. In contrast, activation in retrosplenial cortex was greatest for full scene views, and did not differ among close-up views, diagnostic objects, and faces. The results showed that parahippocampal place area responds in a graded fashion as images become more completely scene-like and include more explicit 3D structure, whereas retrosplenial cortex responds in a step-wise manner to the presence of a complete scene. These results suggest scene processing areas are particularly sensitive to the 3D geometric structure that distinguishes scenes from other types of complex and meaningful visual stimuli.     

Neuroanatomical correlates of processing in visual and visuospatial working memory

Working memory is traditionally seen as being organised in a modular way with a central executive orchestrating at least two slave systems (phonological loop and visuospatial sketch pad). Neuroanatomical correlates of the visual and visuospatial subsystems and the central executive are discussed in this article. A series of experiments are presented yielding evidence for a differentiation into active and passive processing in working memory as well as their neuroanatomical correlates in the prefrontal cortex. Data, yielding evidence for an interaction and separation of visual and visuospatial working memory are presented and discussed. Further results are presented which suggest a convergence of these two systems with increasing working memory demands. The discussed findings will give new insight in the organisation of visual and visuospatial working memory on the anatomical level.     

Encoding location and serial order in auditory working memory: evidence for separable processes

In this study, we investigated the interactions between temporal and spatial information in auditory working memory. In two experiments, participants were presented with sequences of sounds originating from different locations in space and were then asked to recall either their position or their serial order. In Experiment 1, attention during encoding was manipulated by contrasting 'pure' blocks (i.e., location-only or serial-order-only trials) to 'mixed' blocks (i.e., different percentages of spatial and serial-order trials). In Experiment 2, 'pure' blocks were contrasted to blocks in which spatial and serial-order trials were intermixed with a third task requiring a semantic categorization of sounds. Results from both experiments showed that, whereas serial-order recall is linearly affected by the simultaneous encoding of a concurrent feature, the recall of position is mostly unaffected by concurrent feature encoding. Contrastingly, overall performance level was lower for spatial recall than serial recall. We concluded that serial order and location of items appear to be independently encoded in auditory working memory. Serial order is easier to recall, but strongly affected by the processing of concurrent item dimensions, while item location is more difficult to recall, but relatively automatic, as shown by its strong resistance to interfering dimensions in encoding.     

No sex differences in spatial location memory for abstract designs

Previous research has demonstrated a female advantage, albeit imperfectly, on tests of object location memory where object identity information is readily available. However, spatial and visual elements are often confounded in the experimental tasks used. Here spatial and visual memory performance was compared in 30 men and 30 women by presenting 12 abstract designs in a spatial array for recall and recognition (visual memory) and spatial location ("object" location memory). Object location memory was measured via a sensitive absolute displacement score defined as the distance in mms between the position assigned to the object during recall and the actual position it originally occupied. There were no sex differences in either the visual or spatial location tests. Controlling for age and estimated IQ scores made no impact on the results. These data suggest an absence of a sex difference in purely visual and spatial aspects of object location memory.     

Electrophysiological evidence for colour effects on the naming of colour diagnostic and noncolour diagnostic objects

In this study, we investigated the level of visual processing at which surface colour information improves the naming of colour diagnostic and noncolour diagnostic objects. Continuous electroencephalograms were recorded while participants performed a visual object naming task in which coloured and black-and-white versions of both types of objects were presented. The black-and-white and the colour presentations were compared in two groups of event-related potentials (ERPs): (1) The P1 and N1 components, indexing early visual processing; and (2) the N300 and N400 components, which index late visual processing. A colour effect was observed in the P1 and N1 components, for both colour and noncolour diagnostic objects. In addition, for colour diagnostic objects, a colour effect was observed in the N400 component. These results suggest that colour information is important for the naming of colour and noncolour diagnostic objects at different levels of visual processing. It thus appears that the visual system uses colour information, during naming of both object types, at early visual stages; however, for the colour diagnostic objects naming, colour information is also recruited during the late visual processing stages.     

No evidence for object alternation impairment in obsessive-compulsive disorder (OCD)

Recent neuroimaging studies have consistently ascribed the orbito-frontal cortex (OFC) a pivotal role in the pathogenesis of obsessive-compulsive disorder (OCD). Cognitive tests presumed sensitive to this region, such as the Object Alternation Task (OAT), are considered important tools to verify this assumption and to investigate the impact of cortical dysfunction on behavior. The aim of the present study was to assess if patients with OCD show enhanced perseveration errors on the OAT relative to healthy controls taking into account several potential moderators, especially comorbid depression and OCD subtype. Thirty-five OCD patients and 18 healthy controls underwent the OAT as well as the Trail-Making Tests (TMT) A and B. In line with prior studies, OCD patients were slowed on both TMT tasks. In contrast, samples performed similarly on the OAT. While the latter finding does not invalidate the assumption that the OFC is affected in OCD, dysfunctions involving this region may be more subtle than often claimed and likely encompass only a small subset of functional domains hosted in the OFC.     

Electrophysiological indices of memory for temporal order in early childhood: implications for the development of recollection

The ability to recall contextual details associated with an event begins to develop in the first year of life, yet adult levels of recall are not reached until early adolescence. Dual‐process models of memory suggest that the distinct retrieval process that supports the recall of such contextual information is recollection. In the present investigation, we used both behavioral and electrophysiological measures to assess the development of memory for contextual details, as indexed by memory for temporal order, in early childhood. Results revealed age‐related improvements in memory for temporal order despite similar levels of memory for the individual items themselves. Furthermore, this pattern of recall was associated with specific components in the electrophysiological response. Consistent with electrophysiological research in adults, distributed, positive‐going activity late in the waveform was associated with increases in recall of contextual details and the development of recollective processes.