Arc length coding by interference of theta frequency oscillations may underlie context-dependent hippocampal unit data and episodic memory function

Many memory models focus on encoding of sequences by excitatory recurrent synapses in region CA3 of the hippocampus. However, data and modeling suggest an alternate mechanism for encoding of sequences in which interference between theta frequency oscillations encodes the position within a sequence based on spatial arc length or time. Arc length can be coded by an oscillatory interference model that accounts for many features of the context-dependent firing properties of hippocampal neurons observed during performance of spatial memory tasks. In continuous spatial alternation, many neurons fire selectively depending on the direction of prior or future response (left or right). In contrast, in delayed non-match to position, most neurons fire selectively for task phase (sample vs. choice), with less selectivity for left versus right. These seemingly disparate results are effectively simulated by the same model, based on mechanisms similar to a model of grid cell firing in entorhinal cortex. The model also simulates forward shifting of firing over trials. Adding effects of persistent firing with reset at reward locations addresses changes in context-dependent firing with different task designs. Arc length coding could contribute to episodic encoding of trajectories as sequences of states and actions.     

Visual learning of statistical relations among nonadjacent features: Evidence for structural encoding

Recent results suggest that observers can learn, unsupervised, the co-occurrence of independent shape features in viewed patterns (e.g., Fiser & Aslin, 2001). A critical question with regard to these findings is whether learning is driven by a structural, rule-based encoding of spatial relations between distinct features or by a pictorial, template-like encoding, in which spatial configurations of features are embedded in a “holistic” fashion. In two experiments, we test whether observers can learn combinations of features when the paired features are separated by an intervening spatial “gap”, in which other, unrelated features can appear. This manipulation both increases task difficulty and makes it less likely that the feature combinations are encoded simply as larger unitary features. Observers exhibited learning consistent with earlier studies, suggesting that unsupervised learning of compositional structure is based on the explicit encoding of spatial relations between separable visual features. More generally, these results provide support for compositional structure in visual representation.     

Multidimensional encoding of visual form

The encoding of stimulus dimensions of visual form in the human S was investigated under conditions of threshold exposure durations. Three stimulus dhnensions defined on a spatial grating were investigated: spatial line frequency, grating orientation, and orientation of a transverse break in the lines of grating. Results support the conclusion that, within the general category of visual form, different primary stimulus dimensions such as spatial frequency and orientation may be encoded simultaneously, whereas, when the two stimulus components for report are defined on the same dimension (orientation), overall performance is consistent with the predictions of a single channel model.     

Differentiating between verbal and spatial encoding using eye-movement recordings

Visual information processing is guided by an active mechanism generating saccadic eye movements to salient stimuli. Here we investigate the specific contribution of saccades to memory encoding of verbal and spatial properties in a serial recall task. In the first experiment, participants moved their eyes freely without specific instruction. We demonstrate the existence of qualitative differences in eye-movement strategies during verbal and spatial memory encoding. While verbal memory encoding was characterized by shifting the gaze to the to-be-encoded stimuli, saccadic activity was suppressed during spatial encoding. In the second experiment, participants were required to suppress saccades by fixating centrally during encoding or to make precise saccades onto the memory items. Active suppression of saccades had no effect on memory performance, but tracking the upcoming stimuli decreased memory performance dramatically in both tasks, indicating a resource bottleneck between display-controlled saccadic control and memory encoding. We conclude that optimized encoding strategies for verbal and spatial features are underlying memory performance in serial recall, but such strategies work on an involuntary level only and do not support memory encoding when they are explicitly required by the task.     

Space and movement in working memory

Encoding seen movement of another human body requires visuo-spatial processing, and recall involves motor activity. However, encoding whole body movement patterns is affected differently by patterned and spatial secondary tasks, and this difference is reversed for encoding of spatial targets for movement (Smyth, Pearson, & Pendleton, 1988). The experiments reported here investigate the rehearsal of such movement patterns and their recall over unfilled and filled intervals. Performing, watching, or encoding a sequence of spatial positions while carrying a memory load of movement patterns did not affect recall of those movements, whereas performing, watching, or encoding a further set of patterned movements reduced the number recalled from the original set. However, memory for a series of locations in space was not affected by watching patterned movements during the interval, and only order information was affected by watching movement to a series of spatial locations during the interval. The results are discussed in terms of the independence of rehearsal mechanisms for spatial sequencing and movement patterns.     

Are spatial frequencies integrated from coarse to fine?

The existence of a temporal anisotropy in the integration of spatial frequencies, such that spatial frequencies are integrated more effectively if they are available from low to high through time, has been examined in a series of experiments. In the first experiment, the first three harmonics of a square wave were presented in a low-to-high or a high-to-low sequence in a temporal two-interval forced-choice experiment. Subjects were asked to indicate which sequence appeared to resemble a square wave more. A high-to-low sequence of spatial frequencies was judged to more resemble the target than the low-to-high sequence. These results support a temporal anisotropy in the integration of spatial frequencies of exactly the opposite form to that suggested from previous results. Further experiments established that this was not due to task differences or to subjects basing their decision on the final spatial frequency shown. An interpretation is offered in which an isotropic mechanism for spatial-frequency integration is combined with a recency bias.     

The role of location indexes in spatial perception: a sketch of the FINST spatial-index model

This paper hypothesizes a resource-limited mechanism, called a FINST, for individuating or indexing visual features, as distinct from encoding their type or location. FINSTs have the property that they index features in a way that is transparent to their retinal location, and hence under certain conditions succeed in "pointing to" scene locations. The basic assumption is that no operation upon sets of features can occur unless all the features to which the operation applies are first FINSTed. A number of applications of this hypothesis are explored in this paper, including applications to phenomena such as the spatial stability of visual percepts, the ability to track several independently moving targets in parallel, the ability to detect a class of spatial relations requiring the use of "visual routines", various mental imagery phenomena, and the ability to encode complex shapes for recognition. In addition, the possibility is examined that such indexes might be used to bind perceived locations to arguments in motor commands, thereby allowing some forms of perceptual-motor coordination. Several additional assumptions are introduced for this purpose, including the postulation of other indexes (called ANCHORS) for non-visually sensed locations. It is assumed that ANCHORS can be bound to FINSTs, thus allowing cross-referencing of visually detected locations to locations given within a proprioceptive or motor-command frame of reference.     

Visual persistence of spatially filtered images

Periodic stimuli such as sine-wave gratings and checkerboard patterns have been used in many studies of visual perception. It is well known that with such stimuli, visual persistence increases as spatial frequency increases and as contrast decreases. It is not clear, however, that similar relationships obtain for aperiodic stimuli such as natural images. Digitized images of objects (a face and a vase) were submitted to two-dimensional Fourier analysis. Four pairs of spatial frequency band-limited images were created for each image. Each pair consisted of a normal (NP) and a scrambled (SP) phase version, with the magnitude spectrum and space-averaged luminance the same within each pair. Filter bandwidths were one octave wide. Threshold persistence was measured for each spatially filtered image. Visual persistence for SP images increased significantly as spatial frequency increased, whereas no significant differences were found for NP images. This suggests that the temporal processing of complex, aperiodic visual images is influenced by the spatial frequency and contrast of local features within the image and cannot be predicted by space-averaged estimates of contrast and spatial frequency.     

Integrating visual images and visual percepts across time and space

Recent studies suggest that visual information can be integrated over a relatively long delay (> 1500 ms) to form a more complete representation (image-percept integration). The current studies investigated whether this process can occur between stimuli that differ in their spatial properties. Participants viewed two dot arrays that filled all but one space in a square or rectangular grid when combined, and reported the missing space. The arrays differed either in size or orientation. Performance reached a comparable level as when spatial properties were matched. However, such performance depends on at least two processes. We suggest an early encoding process and a later image formation/spatial attention reallocation process are required. The flexibility of the image-percept integration process suggests a strong mechanism to form more complete or detailed representations over time, even when the retinal size and orientation of the scene may change between successive views.     

The effect of eccentricity and the adapting level on the café wall illusion

The café wall pattern is composed of rows of alternating light and dark tiles, and alternate rows are shifted by one fourth of a cycle. The rows of tiles are separated by narrow horizontal mortar lines whose luminance is between those of the dark and the light tiles. Although the mortar lines are physically parallel, they are perceived to be tilted, which is known as the café wall illusion. In this study, an energy-based model for encoding orientation is implemented in order to estimate the strength of the café wall illusion, and it is shown that the estimated orientation depends on the spatial frequency to which each orientation-encoding unit is tuned. The estimation of mortar line orientation from an orientation-encoding unit tuned to a lower spatial frequency was greater than that from a unit tuned to a higher spatial frequency. It is assumed that the perceived mortar line orientation is the result of an integration of responses from the orientation-encoding units tuned to various spatial frequencies. This leads to the prediction that under viewing conditions in which responses from orientation-encoding units tuned to a higher spatial frequency are presumably weakened, the strength of the café wall illusion increases. In agreement with this prediction, it is shown that the café wall illusion is stronger when the café wall image is presented at the periphery or is observed under low luminance levels. On the other hand, the weighted averaging of the estimated mortar orientations across spatial frequencies overestimates the perceived orientation of the mortar lines. This suggests that the final percept of the café wall illusion could be determined by some kind of nonlinear interaction, such as an inhibitory interaction, between orientation-encoding units.     

A configural effect in visual short-term memory for features from different parts of an object

Previous studies have shown that change detection performance is improved when the visual display holds features (e.g., a colour and an orientation) that are grouped into different parts of the same object compared to when they are all spatially separated (Xu, 2002a, 2002b). These findings indicate that visual short-term memory (VSTM) encoding can be "object based". Recently, however, it has been demonstrated that changing the orientation of an item could affect the spatial configuration of the display (Jiang, Chun, & Olson, 2004), which may have an important influence on change detection. The perceptual grouping of features into an object obviously reduces the amount of distinct spatial relations in a display and hence the complexity of the spatial configuration. In the present study, we ask whether the object-based encoding benefit observed in previous studies may reflect the use of configural coding rather than the outcome of a true object-based effect. The results show that when configural cues are removed, the object-based encoding benefit remains for features (i.e., colour and orientation) from different parts of an object, but is significantly reduced. These findings support the view that memory for features from different parts of an object can benefit from object-based encoding, but the use of configural coding significantly helps enlarge this effect.     

Cue-dependent effects in recall of categorized lists

Noncued and cued recall of categorized lists are similar in a number of ways: They show the same rate of gain in item recall with increasing category size; they both produce serial position effects within categories; and they both show a “some-or-none” pattern of with-category recall frequency. Cued and noncued recall differ in other respects: There is a great improvement in item and in category recall attendant on the provision of cues, and only noncued recall displays a list position effect. The points of similarity are taken to indicate that both cued and noncued recall measure the same underlying processes. The points of differences suggest the utility of cueing procedures in identifying the origin of characteristics of recall, according to the principle that any feature of noncued recall that is altered by provision of cues must originate in the retrieval process; features resistant to such modification represent encoding or storage effects.     

Learning to discriminate simple sinusoidal gratings is task specific

Learning transfer effects of a modified spatial frequency discrimination task with simple sinusoidal gratings on various untrained test tasks, testing performance in relative position, local width and global size discrimination, have been investigated. Six subjects were exposed to grating stimuli of varying spatial frequency and size but constant relative position while they had to respond only to spatial frequency. In the course of 7 consecutive days all subjects showed significant reduction of spatial frequency discrimination thresholds. Comparison of discrimination thresholds for the untrained test tasks, taken before and after the learning epoch, reveals complete learning transfer to spatial frequency discrimination with gratings of constant size and variable relative position, but complete lack of transfer when grating spatial frequency is shifted about one octave lower. Further, there is improvement of relative position discrimination and width discrimination of single luminance bars, which is not very specific for the learnt spatial frequency. Although size variation was part of the learning procedure, discrimination of global stimulus size did not improve. Generally, the observed scheme of learning transfer reveals that there is learning only for stimulus attributes that are behaviorally relevant in the learning task. The differential scheme of improvement and code usage in the test tasks strongly indicates involvement of higher stages capable of independent access to different coding domains, as well as attentionally guided attribute selection and suppression. Supported by other recent findings, it is suggested that discrimination learning can be understood as a higher level process of gradual refinement of code selection out of a rich code base provided by lower level stages.     

Face processing in different brain areas,and critical band masking

Neurophysiological evidence is described showing that some neurons in the macaque inferior temporal visual cortex have responses that are invariant with respect to the position, size, view, and spatial frequency of faces and objects, and that these neurons show rapid processing and rapid learning. Critical band spatial frequency masking is shown to be a property of these face‐selective neurons and of the human visual perception of faces. Which face or object is present is encoded using a distributed representation in which each neuron conveys independent information in its firing rate, with little information evident in the relative time of firing of different neurons. This ensemble encoding has the advantages of maximizing the information in the representation useful for discrimination between stimuli using a simple weighted sum of the neuronal firing by the receiving neurons, generalization, and graceful degradation. These invariant representations are ideally suited to provide the inputs to brain regions such as the orbitofrontal cortex and amygdala that learn the reinforcement associations of an individual's face, for then the learning, and the appropriate social and emotional responses generalize to other views of the same face. A theory is described of how such invariant representations may be produced by self‐organizing learning in a hierarchically organized set of visual cortical areas with convergent connectivity. The theory utilizes either temporal or spatial continuity with an associative synaptic modification rule. Another population of neurons in the cortex in the superior temporal sulcus encodes other aspects of faces such as face expression, eye‐gaze, face view, and whether the head is moving. These neurons thus provide important additional inputs to parts of the brain such as the orbitofrontal cortex and amygdala that are involved in social communication and emotional behaviour. Outputs of these systems reach the amygdala, in which face‐selective neurons are found, and also the orbitofrontal cortex, in which some neurons are tuned to face identity and others to face expression. In humans, activation of the orbitofrontal cortex is found when a change of face expression acts as a social signal that behaviour should change; and damage to the human orbitofrontal and pregenual cingulate cortex can impair face and voice expression identification, and also the reversal of emotional behaviour that normally occurs when reinforcers are reversed.     

Effect on numerosity judgment of grouping of tones by auditory channels

Previous research suggests that numerosity judgments for sequences of tones improve when the sequence is structured such that equal and small groups of tones alternate between the ears. The present. study systematically investigates the effects of the structure of tone sequences on number judgment. Tone frequency is chosen as the grouping principle instead of spatial location. In the first experiment, sequences with equal groups of 2, 3, 4, 5, 6, or 7 tones (groups alternating between 800 and 1, 250 Hz) were compared with monotonous sequences. At a slow repetition rate of the tones (280 msec onset to onset), grouping the sequence deteriorated the numerosity judgment. At II fast repetition rate of the tones (100 msec onset to onset) grouping improved numerosity judgment, but only if the group size did not exceed 4 to 5 tones. In the second experiment, the equality of group size in the sequence as a necessary condition for the improvement was investigated. It was found that tone sequences comprised of equal groups were judged more accurately in number than sequences comprised of unequal groups. These results seem to give support for the existence of an auditory subitizing process. The results of this study are also compared with the results of studies in which sequences of tones, alternating one by om! between locations or frequencies, had to be judged in number.     

Grapheme–color synesthesia can enhance immediate memory without disrupting the encoding of relational cues

Previous evidence has suggested that grapheme–color synesthesia can enhance memory for words, but little is known about how these photisms cue retrieval. Often, the encoding of specific features of individual words can disrupt the encoding of ordered relations between words, resulting in an overall decrease in recall accuracy. Here we show that the photisms arising from grapheme–color synesthesia do not function like these item-specific cues. The influences of high and low word frequency on the encoding of ordered relations and the accuracy of immediate free recall were compared across a group of 10 synesthetes and 48 nonsynesthetes. The main findings of Experiment 1 showed that the experience of synesthesia had no adverse effect on the encoding of ordered relations (as measured by input–output correspondence); furthermore, it enhanced recall accuracy in a strictly additive fashion across the two word frequency conditions. Experiment 2 corroborated these findings by showing that the synesthetes only outperformed the nonsynesthetes when the materials involved words and letters, not when they involved digits and spatial locations. Altogether, the present findings suggest that synesthesia can boost immediate memory performance without disrupting the encoding of ordered relations.     

The posterior parietal cortex and long-term memory representation of spatial information

The hypothesis to be explored in this chapter is based on the assumption that the posterior parietal cortex (PPC) is directly involved in representing a subset of the spatial features associated with spatial information processing and plays an important role in perceptual memory as well as long-term memory encoding, consolidation, and retrieval of spatial information. After presentation of the anatomical location of the PPC in rats, the nature of PPC representation based on single spatial features, binding of visual features associated with visual spatial attention, binding of object-place associations associated with acquisition and storage of associations where one of the elements is a spatial component, and binding of ideothetic and allothetic information in long-term memory is discussed. Additional evidence for a PPC role in mediation of spatial information in long-term storage is offered. Finally, the relationship between the PPC and the hippocampus from a systems and dynamic point view is presented.     

Distraction shrinks space

Research investigating how people remember the distance of paths they walk has shown two apparently conflicting effects of experience during encoding on subsequent distance judgments. By the feature accumulation effect, discrete path features such as turns, houses, or other landmarks cause an increase in remembered distance. By the distractor effect, performance of a concurrent task during path encoding causes a decrease in remembered distance. In this study, we ask the following: What are the conditions that determine whether the feature accumulation or the distractor effect dominates distortions of space? In two experiments, blindfolded participants were guided along two legs of a right triangle while reciting nonsense syllables. On some trials, one of the two legs contained features: horizontally mounted car antennas (gates) that bent out of the way as participants walked past. At the end of the second leg, participants either indicated the remembered path leg lengths using their hands in a ratio estimation task or attempted to walk, unguided, straight back to the beginning. In addition to response mode, visual access to the paths and time between encoding and response were manipulated to determine whether these factors would affect feature accumulation or distractor effects. Path legs with added features were remembered as shorter than those without, but this result was significant only in the haptic response mode data. This finding suggests that when people form spatial memory representations with the intention of navigating in room-scale spaces, interfering with information accumulation substantially distorts spatial memory.     

Recognition memory of spatial location information: Another failure to support automaticity

Hasher and Zacks (1979) claimed that spatial location information is automatically encoded. Evaluation of the empirical basis for this claim, however, casts doubt on some of the evidence for the automaticity position. This evaluation led to four experiments in which five criteria for testing the automaticity of cognitive processes were examined using a recognition task. Results of these experiments clearly show thatrecognition memory for spatial location information is influenced by intention, age of subjects, competing task loads, practice, and individual differences. The reported results, which extend those reported by Naveh-Benjamin (1987) for spatial locationrecall memory, are at odds with the claim that memory for spatial location information is exclusively mediated by automatic encoding processes. The concept of automaticity and the appropriateness of the criteria suggested for testing the automaticity of cognitive processes are discussed in light of the current results and recent findings on other features of the environment (e.g., frequency of occurrence) previously claimed to be automatically encoded.     

Geometric modules in animals' spatial representations: a test with chicks (Gallus gallus domesticus)

Recent work has shown that in place-finding tasks rats rely on the geometric relations between the goal object and the shape of the environment. We tested young chickens (Gallus gallus domesticus) on similar tasks in a reference memory paradigm to determine whether differences exist between species in the ability to use geometric and nongeometric spatial information. The main findings were that chicks: (a) encoded and used both geometric and nongeometric (featural) information; (b) did not use the overall spatial arrangement of the features; (c) relied primarily on nongeometric cues when faced with contradictory information. Two mechanisms are evident in chicks' spatial representations: a metric frame for encoding the spatial arrangement of surfaces as surfaces and a cue-guidance system for encoding conspicuous landmarks near the target. The possibility of hierarchical organization and species differences in these two mechanisms are discussed.