Retrieval Intention Modulates the Effects of Directed Forgetting Instructions on Recollection

The neurocognitive basis of memory retrieval is often examined by investigating brain potential old/new effects, which are differences in brain activity between successfully remembered repeated stimuli and correctly rejected new stimuli in a recognition test. In this study, we combined analyses of old/new effects for words with an item-method directed-forgetting manipulation in order to isolate differences between the retrieval processes elicited by words that participants were initially instructed to commit to memory and those that participants were initially instructed to forget. We compared old/new effects elicited by to-be-forgotten (TBF) words with those elicited by to-be-remembered (TBR) words in both an explicit-memory test (a recognition test) and an implicit-memory test (a lexical-decision test). Behavioral results showed clear directed forgetting effects in the recognition test, but not in the lexical decision test. Mirroring the behavioral findings, analyses of brain potentials showed evidence of directed forgetting only in the recognition test. In this test, potentials from 450–650 ms (P600 old/new effects) were more positive for TBR relative to TBF words. By contrast, P600 effects evident during the lexical-decision test did not differ in magnitude between TBR and TBF items. When taken in the context of prior studies that have linked similar parietal old/new effects to the recollection of episodic information, these data suggest that directed-forgetting effects manifest primarily in greater episodic retrieval by TBR than TBF items, and that retrieval intention may be important for these directed-forgetting effects to occur.     

Stimulus dependency of object-evoked responses in human visual cortex: an inverse problem for category specificity

Many studies have linked the processing of different object categories to specific event-related potentials (ERPs) such as the face-specific N170. Despite reports showing that object-related ERPs are influenced by visual stimulus features, there is consensus that these components primarily reflect categorical aspects of the stimuli. Here, we re-investigated this idea by systematically measuring the effects of visual feature manipulations on ERP responses elicited by both structure-from-motion (SFM)-defined and luminance-defined object stimuli. SFM objects elicited a novel component at 200-250 ms (N250) over parietal and posterior temporal sites. We found, however, that the N250 amplitude was unaffected by restructuring SFM stimuli into meaningless objects based on identical visual cues. This suggests that this N250 peak was not uniquely linked to categorical aspects of the objects, but is strongly determined by visual stimulus features. We provide strong support for this hypothesis by parametrically manipulating the depth range of both SFM- and luminance-defined object stimuli and showing that the N250 evoked by SFM stimuli as well as the well-known N170 to static faces were sensitive to this manipulation. Importantly, this effect could not be attributed to compromised object categorization in low depth stimuli, confirming a strong impact of visual stimulus features on object-related ERP signals. As ERP components linked with visual categorical object perception are likely determined by multiple stimulus features, this creates an interesting inverse problem when deriving specific perceptual processes from variations in ERP components.     

Unconscious context control of visual perception of simple stimuli: A study using evoked potentials

The effect of nonsemantic context on the perception of simple nonverbal visual stimuli has been studied in ten healthy volunteers by the event-related potential (ERP) method. The nonsemantic context was specified by the formation of a memory trace of a test visual stimulus via its repeated presentation without any instruction except gaze fixation. Then, this stimulus randomly alternated with control stimuli that did not form memory traces before their presentation. It has been found that an ERP in the interval 260–340 ms after presentation of a simple nonverbal stimulus significantly differs from the control ERPs. The results suggest that some stages of the processing of visual stimuli may be modified by nonsemantic context.     

Attribute Pair-Based Visual Recognition and Memory

Background

In the human visual system, different attributes of an object, such as shape, color, and motion, are processed separately in different areas of the brain. This raises a fundamental question of how are these attributes integrated to produce a unified perception and a specific response. This “binding problem” is computationally difficult because all attributes are assumed to be bound together to form a single object representation. However, there is no firm evidence to confirm that such representations exist for general objects.

Methodology/Principal Findings

Here we propose a paired-attribute model in which cognitive processes are based on multiple representations of paired attributes. In line with the model''s prediction, we found that multiattribute stimuli can produce an illusory perception of a multiattribute object arising from erroneous integration of attribute pairs, implying that object recognition is based on parallel perception of paired attributes. Moreover, in a change-detection task, a feature change in a single attribute frequently caused an illusory perception of change in another attribute, suggesting that multiple pairs of attributes are stored in memory.

Conclusions/Significance

The paired-attribute model can account for some novel illusions and controversial findings on binocular rivalry and short-term memory. Our results suggest that many cognitive processes are performed at the level of paired attributes rather than integrated objects, which greatly facilitates the binding problem and provides simpler solutions for it.     

Electrophysiological correlates of object location and object identity processing in spatial scenes

The ability to quickly detect changes in our surroundings has been crucial to human adaption and survival. In everyday life we often need to identify whether an object is new and if an object has changed its location. In the current event-related potential (ERP) study we investigated the electrophysiological correlates and the time course in detecting different types of changes of an objecṫs location and identity. In a delayed match-to-sample task participants had to indicate whether two consecutive scenes containing a road, a house, and two objects, were either the same or different. In six randomly intermixed conditions the second scene was identical, one of the objects had changed its identity, one of the objects had changed its location, or the objects had switched locations. The results reveal different time courses for the processing of identity and location changes in spatial scenes. Whereas location changes elicited a posterior N2 effect, indicating early mismatch detection, followed by a P3 effect reflecting post-perceptual processing, identity changes elicited an anterior N3 effect, which was delayed and functionally distinct from the N2 effect found for the location changes. The condition in which two objects switched position elicited a late ERP effect, reflected by a P3 effect similar to that obtained for the location changes. In sum, this study is the first to cohesively show different time courses for the processing of location changes, identity changes, and object switches in spatial scenes, which manifest themselves in different electrophysiological correlates.     

Comparison of preferences for object properties in the rat using paired- and free-choice paradigms

A common method for testing preference for objects is to determine which of a pair of objects is approached first in a paired-choice paradigm. In comparison, many studies of preference for environmental enrichment (EE) devices have used paradigms in which total time spent with each of a pair of objects is used to determine preference. While each of these paradigms gives a specific measure of the preference for one object in comparison to another, neither method allows comparisons between multiple objects simultaneously. Since it is possible that several EE objects would be placed in a cage together to improve animal welfare, it is important to determine measures for rats’ preferences in conditions that mimic this potential home cage environment. While it would be predicted that each type of measure would produce similar rankings of objects, this has never been tested empirically. In this study, we compared two paradigms: EE objects were either presented in pairs (paired-choice comparison) or four objects were presented simultaneously (simultaneous presentation comparison). We used frequency of first interaction and time spent with each object to rank the objects in the paired-choice experiment, and time spent with each object to rank the objects in the simultaneous presentation experiment. We also considered the behaviours elicited by the objects to determine if these might be contributing to object preference. We demonstrated that object ranking based on time spent with objects from the paired-choice experiment predicted object ranking in the simultaneous presentation experiment. Additionally, we confirmed that behaviours elicited were an important determinant of time spent with an object. This provides convergent evidence that both paired choice and simultaneous comparisons provide valid measures of preference for EE objects in rats.     

On the interactivity in the human visual cortex caused by specific and nonspecific inflows

Summary An interactivity (Sato, 1969) with respect not only to the interstimulus intervenal between the conditioned and test stimuli but also with respect to the time after the test stimulus was defined as a new generalized recovery function.The contour maps of the above interactivities of the cerebral evoked potentials elicited by the paired flash stimuli and by the conditioned cutaneous and test flash stimuli, respectively, were obtained in the adult human visual cortex.It was suggested to a certain degree by the autocorrelograms of background occipital EEGs and average VEPs that the more prominent VEPs are elicited, the more predominant are the alpha waves in the occipital background EEG, and vice versa. In the contour maps of the interactivity by the paired flash stimuli, cyclic changes of facilitory mountains and occlusive valleys were regular and conspicuous in the subject who exhibited predominant alpha waves in his background EEG, whereas the cyclic changes were less regular and less marked in the subject with few alpha waves in the background EEG.The cyclic changes in the former subject appeared at an interval of about 100 msec at an interstimulus interval of around 100 msec. This acts favorably in the contribution of the cyclic change in interactivity via natural stimulations to the generation of alpha waves in spontaneous EEG. The main facilitory mountains and occlusive valleys in the maps were arranged in a line parallel to the ordinate (axis of the interstimulus interval), which suggest that the conditioned flash will affect in a specific fashion each response compoment elicited via specific afferent inflows in the VEP timelocked to the test flash stimulus. On the contrary, those in the maps obtained by the conditioned cutaneous and test flash stimuli were diagonal to the abscissa (time after the test stimulus) and ordinate, and/or the peaks of facilitory mountains and bottoms of occlusive valleys tended to form a line parallel to the abscissa, which indicates that as the cutaneous somatosensory afferent inflow converges to the visual cortex nonspecific interactivities are exhibited in each component in the VEP.     

The Processing of Object Identity Information by Women and Men

The study examined whether women excel at tasks which require processing the identity of objects information as has been suggested in the context of the well-known object location memory task. In a computer-simulated task, university students were shown simulated indoor and outdoor house scenes. After studying a scene the students were presented with two images. One was the original image and the other a modified version in which one object was either rotated by ninety degrees or substituted with a similar looking object. The participants were asked to indicate the original image. The main finding was that no sex effect was obtained in this task. The female and male students did not differ on a verbal ability test, and their 2D:4D ratios were found to be comparable.     

Evolution and the expression of biases: situational value changes the endowment effect in chimpanzees

Cognitive and behavioral biases, which are widespread among humans, have recently been demonstrated in other primates, suggesting a common origin. Here we examine whether the expression of one shared bias, the endowment effect, varies as a function of context. We tested whether objects lacking inherent value elicited a stronger endowment effect (or preference for keeping the object) in a context in which the objects had immediate instrumental value for obtaining valuable resources (food). Chimpanzee subjects had opportunities to trade tools when food was not present, visible but unobtainable, and obtainable using the tools. We found that the endowment effect for these tools existed only when they were immediately useful, showing that the effect varies as a function of context-specific utility. Such context-specific variation suggests that the variation seen in some human biases may trace predictably to behaviors that evolved to maximize gains in specific circumstances.     

The Positional-Specificity Effect Reveals a Passive-Trace Contribution to Visual Short-Term Memory

The positional-specificity effect refers to enhanced performance in visual short-term memory (VSTM) when the recognition probe is presented at the same location as had been the sample, even though location is irrelevant to the match/nonmatch decision. We investigated the mechanisms underlying this effect with behavioral and fMRI studies of object change-detection performance. To test whether the positional-specificity effect is a direct consequence of active storage in VSTM, we varied memory load, reasoning that it should be observed for all objects presented in a sub-span array of items. The results, however, indicated that although robust with a memory load of 1, the positional-specificity effect was restricted to the second of two sequentially presented sample stimuli in a load-of-2 experiment. An additional behavioral experiment showed that this disruption wasn’t due to the increased load per se, because actively processing a second object – in the absence of a storage requirement – also eliminated the effect. These behavioral findings suggest that, during tests of object memory, position-related information is not actively stored in VSTM, but may be retained in a passive tag that marks the most recent site of selection. The fMRI data were consistent with this interpretation, failing to find location-specific bias in sustained delay-period activity, but revealing an enhanced response to recognition probes that matched the location of that trial’s sample stimulus.     

The functional neuroanatomy of implicit-motion perception or representational momentum

BACKGROUND: When we view static scenes that imply motion - such as an object dropping off a shelf - recognition memory for the position of the object is extrapolated forward. It is as if the object in our mind's eye comes alive and continues on its course. This phenomenon is known as representational momentum and results in a distortion of recognition memory in the implied direction of motion. Representational momentum is modifiable; simply labelling a drawing of a pointed object as 'rocket' will facilitate the effect, whereas the label 'steeple' will impede it. We used functional magnetic resonance imaging (fMRI) to explore the neural substrate for representational momentum. RESULTS: Subjects participated in two experiments. In the first, they were presented with video excerpts of objects in motion (versus the same objects in a resting position). This identified brain areas responsible for motion perception. In the second experiment, they were presented with still photographs of the same target items, only some of which implied motion (representational momentum stimuli). When viewing still photographs of scenes implying motion, activity was revealed in secondary visual cortical regions that overlap with areas responsible for the perception of actual motion. Additional bilateral activity was revealed within a posterior satellite of V5 for the representational momentum stimuli. Activation was also engendered in the anterior cingulate cortex. CONCLUSIONS: Considering the implicit nature of representational momentum and its modifiability, the findings suggest that higher-order semantic information can act on secondary visual cortex to alter perception without explicit awareness.     

Short-Term Memory Affects Color Perception in Context

Color-based object selection — for instance, looking for ripe tomatoes in the market — places demands on both perceptual and memory processes: it is necessary to form a stable perceptual estimate of surface color from a variable visual signal, as well as to retain multiple perceptual estimates in memory while comparing objects. Nevertheless, perceptual and memory processes in the color domain are generally studied in separate research programs with the assumption that they are independent. Here, we demonstrate a strong failure of independence between color perception and memory: the effect of context on color appearance is substantially weakened by a short retention interval between a reference and test stimulus. This somewhat counterintuitive result is consistent with Bayesian estimation: as the precision of the representation of the reference surface and its context decays in memory, prior information gains more weight, causing the retained percepts to be drawn toward prior information about surface and context color. This interaction implies that to fully understand information processing in real-world color tasks, perception and memory need to be considered jointly.     

Effects of Acute Cortisol Administration on Perceptual Priming of Trauma-Related Material

Intrusive memories are a hallmark symptom of posttraumatic stress disorder (PTSD). They reflect excessive and uncontrolled retrieval of the traumatic memory. Acute elevations of cortisol are known to impair the retrieval of already stored memory information. Thus, continuous cortisol administration might help in reducing intrusive memories in PTSD. Strong perceptual priming for neutral stimuli associated with a “traumatic” context has been shown to be one important learning mechanism that leads to intrusive memories. However, the memory modulating effects of cortisol have only been shown for explicit declarative memory processes. Thus, in our double blind, placebo controlled study we aimed to investigate whether cortisol influences perceptual priming of neutral stimuli that appeared in a “traumatic” context. Two groups of healthy volunteers (N = 160) watched either neutral or “traumatic” picture stories on a computer screen. Neutral objects were presented in between the pictures. Memory for these neutral objects was tested after 24 hours with a perceptual priming task and an explicit memory task. Prior to memory testing half of the participants in each group received 25 mg of cortisol, the other half received placebo. In the placebo group participants in the “traumatic” stories condition showed more perceptual priming for the neutral objects than participants in the neutral stories condition, indicating a strong perceptual priming effect for neutral stimuli presented in a “traumatic” context. In the cortisol group this effect was not present: Participants in the neutral stories and participants in the “traumatic” stories condition in the cortisol group showed comparable priming effects for the neutral objects. Our findings show that cortisol inhibits perceptual priming for neutral stimuli that appeared in a “traumatic” context. These findings indicate that cortisol influences PTSD-relevant memory processes and thus further support the idea that administration of cortisol might be an effective treatment strategy in reducing intrusive reexperiencing.     

Novel object exploration in mice: not all objects are created equal

Object exploration is an increasingly popular experimental paradigm in behavioral sciences. We have begun a series of studies with mice specifically looking at the parameters that influence behaviors in this test. The aim of the present study was to examine the effect of object type on performance in the object exploration test. More specifically, adult male C57BL/6J mice were trained and tested using objects that could be climbed (CLIMB) or with those that could only be touched (TOUCH). The results show that activity is affected by the presentation of objects, with object type interacting with some of these changes. C57 mice explored objects that can be climbed over significantly longer than objects that can only be touched and a more rapid habituation was observed using objects that could only be touched. Robust recognition memory was observed in both groups of mice, however mice in the CLIMB group exhibited a significantly greater discrimination index compared to mice in the TOUCH group. Taken together, these findings demonstrate that the selection of objects is of critical importance and it is recommended that special attention be given to the functional (affordant) properties of the objects to-be-used in future studies.     

Landmark recognition in Alzheimer's dementia: spared implicit memory for objects relevant for navigation

Background

In spatial navigation, landmark recognition is crucial. Specifically, memory for objects placed at decision points on a route is relevant. Previous fMRI research in healthy adults showed higher medial-temporal lobe (MTL) activation for objects placed at decision points compared to non-decision points, even at an implicit level. Since there is evidence that implicit learning is intact in amnesic patients, the current study examined memory for objects relevant for navigation in patients with Alzheimer’s dementia (AD).

Methodology/Principal Findings

21 AD patients participated with MTL atrophy assessed on MRI (mean MMSE = 21.2, SD = 4.0), as well as 20 age- and education-matched non-demented controls. All participants watched a 5-min video showing a route through a virtual museum with 20 objects placed at intersections (decision points) and 20 at simple turns (non-decision points). The instruction was to pay attention to the toys (half of the objects) for which they were supposedly tested later. Subsequently, a recognition test followed with the 40 previously presented objects among 40 distracter items (both toys and non-toys). Results showed a better performance for the non-toy objects placed at decision points than non-decision points, both for AD patients and controls.

Conclusion/Significance

Our findings indicate that AD patients with MTL damage have implicit memory for object information relevant for navigation. No decision point effect was found for the attended items. Possibly, focusing attention on the items occurred at the cost of the context information in AD, whereas the controls performed at an optimal level due to intact memory function.     

Modulation by dopamine of population spikes in area CA1 hippocampal neurons elicited by paired stimulus pulses

Extracellular recording techniques were used to study the effects of dopamine on postactivation excitability of rat area CA1 hippocampal neurons maintained in vitro. Population spikes were elicited by delivery of conditioning and test stimulus pulses to afferent fibers. The interval between the conditioning and test volley was set to separate delivery of stimuli by 10 to 80 msec. The effect of superfusion or microtopical application of dopamine (DA) on population responses to test stimulus pulses was studied. When paired stimulus volleys, separated by brief intervals (up to 40 msec), were delivered to afferent fibers, paired-pulse suppression (PPS) was indicated by the amplitude of the population spike elicited by the test volley being smaller than that elicited by the conditioning volley. When paired volleys were separated by longer intervals (40 to 80 msec), the response elicited by the test volley was larger in amplitude than that elicited by the conditioning volley, indicating paired-pulse facilitation (PPF). Following exposure to DA, the amplitude of the population response elicited by the conditioning volley was larger than the amplitude before exposure to DA. This effect was long-lasting, enduring for tens of minutes. However, when the amplitude of the conditioning population response was held constant, the PPS was decreased, indicating disinhibition. It is suggested that dopamine produces a long-lasting attenuation of an intervening inhibitory influence onto CA1 pyramidal neurons.     

A reduction in hippocampal GABAA receptor α5 subunits disrupts the memory for location of objects in mice

The memory for location of objects, which binds information about objects to discrete positions or spatial contexts of occurrence, is a form of episodic memory particularly sensitive to hippocampal damage. Its early decline is symptomatic for elderly dementia. Substances that selectively reduce α5‐GABA_A receptor function are currently developed as potential cognition enhancers for Alzheimer's syndrome and other dementia, consistent with genetic studies implicating these receptors that are highly expressed in hippocampus in learning performance. Here we explored the consequences of reduced GABA_Aα5‐subunit contents, as occurring in α5(H105R) knock‐in mice, on the memory for location of objects. This required the behavioral characterization of α5(H105R) and wild‐type animals in various tasks examining learning and memory retrieval strategies for objects, locations, contexts and their combinations. In mutants, decreased amounts of α5‐subunits and retained long‐term potentiation in hippocampus were confirmed. They exhibited hyperactivity with conserved circadian rhythm in familiar actimeters, and normal exploration and emotional reactivity in novel places, allocentric spatial guidance, and motor pattern learning acquisition, inhibition and flexibility in T‐ and eight‐arm mazes. Processing of object, position and context memories and object‐guided response learning were spared. Genotype difference in object‐in‐place memory retrieval and in encoding and response learning strategies for object–location combinations manifested as a bias favoring object‐based recognition and guidance strategies over spatial processing of objects in the mutants. These findings identify in α5(H105R) mice a behavioral–cognitive phenotype affecting basal locomotion and the memory for location of objects indicative of hippocampal dysfunction resulting from moderately decreased α5‐subunit contents.     

Nerve Growth Factor Differentially Affects Spatial and Recognition Memory in Aged Rats

In rats, object discrimination depends on the integrity of the cholinergic system, thus it could be expected that nerve growth factor (NGF) can improve the behavior in aged subjects. The interactive effect of age and cholinergic improvement was assessed behaviorally in young and aged rats. Animals were injected by infusion of NGF into the lateral ventricles and they were tested in two behavioral tasks: an object-location and an object-recognition task. Spatial and recognition memory were assessed in an open field containing five different objects. Rats were submitted to six consecutive sessions. Both age-groups showed comparable habituation of exploratory response in Session 1–4. Discrimination index (DI) was calculated to assess responses to spatial change in Session 5 and object change in Session 6. Control young and aged rats were able to discriminate between familiar and novel object, however DI was lower in aged rats. Treatment with NGF induced decline of object discrimination in both age-groups. Different results were obtained in spatial displacement test. NGF was able to improve spatial memory in aged rats, but had no effect in young controls. These data confer on NGF potential role in improving spatial but not episodic memory in aged rats.