Forgetting dynamics for items of different categories.

How the dynamic evolution of forgetting changes for different material types is unexplored. By using a common experimental paradigm with stimuli of different types, we were able to directly cross-examine the emerging dynamics and found that even though the presentation sets differ minimally by design, the obtained curves appear to fall on a discrete spectrum. We also show that the resulting curves do not depend on physical time but rather on the number of items shown. All measured curves were compatible with our previously developed mathematical model, hinting to a potential common underlying mechanism of forgetting.

Changes in Cortical Coherence Supporting Complex Visual and Social Processing in Adolescence.

Despite our differences, there is much about the natural visual world that most observers perceive in common. Across adults, approximately 30% of the brain is activated in a consistent fashion while viewing naturalistic input. At what stage of development is this consistency of neural profile across individuals present? Here, we focused specifically on whether this mature profile is present in adolescence, a key developmental period that bridges childhood and adulthood, and in which new cognitive and social challenges are at play. We acquired fMRI data evoked by a movie shown twice to younger (9-14 years old) and older adolescents (15-19 years old) and to adults, and conducted three key analyses. First, we characterized the consistency of the neural response within individuals (across separate runs of the movie), then within individuals of the same age group, and, last, between age groups. The neural consistency within individuals was similar across age groups with reliable activation in largely overlapping but slightly different cortical regions. In contrast, somewhat differing regions exhibited higher within-age correlations in both groups of adolescents than in the adults. Last, across the whole cortex, we identified regions evincing different patterns of maturation across age. Together, these findings provide a fine-grained characterization of functional neural development in adolescence and uncover signatures of widespread change in cortical coherence that supports the emerging mature stereotypical responses to naturalistic stimuli. These results also offer a more nuanced account of development that obeys neither a rigid linear progression nor a large qualitative change over time.

Fundamental Law of Memory Recall.

Human memory appears to be fragile and unpredictable. Free recall of random lists of words is a standard paradigm used to probe episodic memory. We proposed an associative search process that can be reduced to a deterministic walk on random graphs defined by the structure of memory representations. The corresponding graph model can be solved analytically, resulting in a novel parameter-free prediction for the average number of memory items recalled (R) out of M items in memory: R=sqrt[3πM/2]. This prediction was verified with a specially designed experimental protocol combining large-scale crowd-sourced free recall and recognition experiments with randomly assembled lists of words or common facts. Our results show that human memory can be described by universal laws derived from first principles.

Memory States and Transitions between Them in Attractor Neural Networks.

Human memory is capable of retrieving similar memories to a just retrieved one. This associative ability is at the base of our everyday processing of information. Current models of memory have not been able to underpin the mechanism that the brain could use in order to actively exploit similarities between memories. The current idea is that to induce transitions in attractor neural networks, it is necessary to extinguish the current memory. We introduce a novel mechanism capable of inducing transitions between memories where similarities between memories are actively exploited by the neural dynamics to retrieve a new memory. Populations of neurons that are selective for multiple memories play a crucial role in this mechanism by becoming attractors on their own. The mechanism is based on the ability of the neural network to control the excitation-inhibition balance.

Practice makes perfect in memory recall.

A large variability in performance is observed when participants recall briefly presented lists of words. The sources of such variability are not known. Our analysis of a large data set of free recall revealed a small fraction of participants that reached an extremely high performance, including many trials with the recall of complete lists. Moreover, some of them developed a number of consistent input-position-dependent recall strategies, in particular recalling words consecutively ("chaining") or in groups of consecutively presented words ("chunking"). The time course of acquisition and particular choice of positional grouping were variable among participants. Our results show that acquiring positional strategies plays a crucial role in improvement of recall performance.

Effects of long-term representations on free recall of unrelated words.

Human memory stores vast amounts of information. Yet recalling this information is often challenging when specific cues are lacking. Here we consider an associative model of retrieval where each recalled item triggers the recall of the next item based on the similarity between their long-term neuronal representations. The model predicts that different items stored in memory have different probability to be recalled depending on the size of their representation. Moreover, items with high recall probability tend to be recalled earlier and suppress other items. We performed an analysis of a large data set on free recall and found a highly specific pattern of statistical dependencies predicted by the model, in particular negative correlations between the number of words recalled and their average recall probability. Taken together, experimental and modeling results presented here reveal complex interactions between memory items during recall that severely constrain recall capacity.

Motion-induced blindness as a noisy excitable system.

Perceptual disappearance of a salient target induced by a moving texture mask (MIB: Motion-Induced Blindness) is a striking effect, currently poorly understood. Here, we investigated whether the dynamics of MIB qualify as an excitable system. Excitable systems exhibit fast switches from one state to another (e.g., visible/invisible) induced by an above-threshold perturbation and stimulus-independent dynamics, followed by a refractory period. In the experiments, disappearance was induced by masks consisting of slowly rotating radial bars with a gap at the target location, leading to periodic perturbation of the visual field around the target (a bright parafoveal spot). When passed around the target location, masks frequently induced an abrupt target disappearance, pointing to locality. As expected from excitable systems, the disappearance time was not affected by additional bars crossing the target during invisibility, and there was little dependence on the mask configuration. After the target reappeared, it stayed for at least 0.5-2 s (the refractory period). Therefore, the dynamics governing MIB represent an example of an excitable system, where the transition to the invisible state is induced by the mask. The dynamics that follow were determined mostly by the internal network properties.

Retroactive interference model of forgetting.

Memory and forgetting constitute two sides of the same coin, and although the first has been extensively investigated, the latter is often overlooked. A possible approach to better understand forgetting is to develop phenomenological models that implement its putative mechanisms in the most elementary way possible, and then experimentally test the theoretical predictions of these models. One such mechanism proposed in previous studies is retrograde interference, stating that a memory can be erased due to subsequently acquired memories. In the current contribution, we hypothesize that retrograde erasure is controlled by the relevant "importance" measures such that more important memories eliminate less important ones acquired earlier. We show that some versions of the resulting mathematical model are broadly compatible with the previously reported power-law forgetting time course and match well the results of our recognition experiments with long, randomly assembled streams of words.

Effects of order on memory of event times.

Memorizing time of an event may employ two processes (1) encoding of the absolute time of events within an episode, (2) encoding of its relative order. Here we study interaction between these two processes. We performed experiments in which one or several items were presented, after which participants were asked to report the time of occurrence of items. When a single item was presented, the distribution of reported times was quite wide. When two or three items were presented, the relative order among them strongly affected the reported time of each of them. Bayesian theory that takes into account the memory for the events order is compatible with the experimental data, in particular in terms of the effect of order on absolute time reports. Our results suggest that people do not deduce order from memorized time, instead people's memory for absolute time of events relies critically on memorized order of the events.

Emergence of hierarchical organization in memory for random material.

Structured information is easier to remember and recall than random one. In real life, information exhibits multi-level hierarchical organization, such as clauses, sentences, episodes and narratives in language. Here we show that multi-level grouping emerges even when participants perform memory recall experiments with random sets of words. To quantitatively probe brain mechanisms involved in memory structuring, we consider an experimental protocol where participants perform 'final free recall' (FFR) of several random lists of words each of which was first presented and recalled individually. We observe a hierarchy of grouping organizations of FFR, most notably many participants sequentially recalled relatively long chunks of words from each list before recalling words from another list. Moreover, participants who exhibited strongest organization during FFR achieved highest levels of performance. Based on these results, we develop a hierarchical model of memory recall that is broadly compatible with our findings. Our study shows how highly controlled memory experiments with random and meaningless material, when combined with simple models, can be used to quantitatively probe the way meaningful information can efficiently be organized and processed in the brain.

Inverse modeling of human contrast response.

Mathematical singularities found in the Signal Detection Theory (SDT) based analysis of the 2-Alternative-Forced-Choice (2AFC) method [Katkov, M., Tsodyks, M., & Sagi, D. (2006a). Analysis of two-alternative force-choice Signal Detection Theory model. Journal of Mathematical Psychology, 50, 411-420; Katkov, M., Tsodyks, M., & Sagi, D. (2006b). Singularities in the inverse modeling of 2AFC contrast discrimination data. Vision Research, 46, 256-266; Katkov, M., Tsodyks, M., & Sagi, D. (2007). Singularities explained: Response to Klein. Vision Research, doi:10.1016/j.visres.2006.10.030] imply that contrast discrimination data obtained with the 2AFC method cannot always be used to reliably estimate the parameters of the underlying model (internal response and noise functions) with a reasonable number of trials. Here we bypass this problem with the Identification Task (IT) where observers identify one of N contrasts. We have found that identification data varies significantly between experimental sessions. Stable estimates using individual session data showed Contrast Response Functions (CRF) with high gain in the low contrast regime and low gain in the high contrast regime. Noise Amplitudes (NA) followed a decreasing function of contrast at low contrast levels, and were practically constant above some contrast level. The transition between these two regimes corresponded approximately to the position of the dipper in the Threshold versus Contrast (TvC) curves that were computed using the estimated parameters and independently measured using 2AFC.

Synaptic Correlates of Working Memory Capacity.

Psychological studies indicate that human ability to keep information in readily accessible working memory is limited to four items for most people. This extremely low capacity severely limits execution of many cognitive tasks, but its neuronal underpinnings remain unclear. Here we show that in the framework of synaptic theory of working memory, capacity can be analytically estimated to scale with characteristic time of short-term synaptic depression relative to synaptic current time constant. The number of items in working memory can be regulated by external excitation, enabling the system to be tuned to the desired load and to clear the working memory of currently held items to make room for new ones.

Singularities in the inverse modeling of 2AFC contrast discrimination data.

Analytical calculations show that two-alternative force-choice data are not always suitable for specifying the parameters of the underlying discrimination model. Experimentally, we show here that in the case of contrast discrimination in humans, a variety of models spanning a large range of parameters can explain the data within an experimental error. Monte-Carlo simulations indicate that the number of trials in psychophysical experiments is not the limiting factor in estimating the parameters in contrast discrimination. These results can therefore explain the contradictory conclusions made by different groups about the relationship between the response to contrast and the noise amplitude.

Visual perception of order-disorder transition.

Our experience with the natural world, as composed of ordered entities, implies that perception captures relationships between image parts. For instance, regularities in the visual scene are rapidly identified by our visual system. Defining the regularities that govern perception is a basic, unresolved issue in neuroscience. Mathematically, perfect regularities are represented by symmetry (perfect order). The transition from ordered configurations to completely random ones has been extensively studied in statistical physics, where the amount of order is characterized by a symmetry-specific order parameter. Here we applied tools from statistical physics to study order detection in humans. Different sets of visual textures, parameterized by the thermodynamic temperature in the Boltzmann distribution, were designed. We investigated how much order is required in a visual texture for it to be discriminated from random noise. The performance of human observers was compared to Ideal and Order observers (based on the order parameter). The results indicated a high consistency in performance across human observers, much below that of the Ideal observer, but well-approximated by the Order observer. Overall, we provide a novel quantitative paradigm to address order perception. Our findings, based on this paradigm, suggest that the statistical physics formalism of order captures regularities to which the human visual system is sensitive. An additional analysis revealed that some order perception properties are captured by traditional texture discrimination models according to which discrimination is based on integrated energy within maps of oriented linear filters.

Neural Network Model of Memory Retrieval.

Human memory can store large amount of information. Nevertheless, recalling is often a challenging task. In a classical free recall paradigm, where participants are asked to repeat a briefly presented list of words, people make mistakes for lists as short as 5 words. We present a model for memory retrieval based on a Hopfield neural network where transition between items are determined by similarities in their long-term memory representations. Meanfield analysis of the model reveals stable states of the network corresponding (1) to single memory representations and (2) intersection between memory representations. We show that oscillating feedback inhibition in the presence of noise induces transitions between these states triggering the retrieval of different memories. The network dynamics qualitatively predicts the distribution of time intervals required to recall new memory items observed in experiments. It shows that items having larger number of neurons in their representation are statistically easier to recall and reveals possible bottlenecks in our ability of retrieving memories. Overall, we propose a neural network model of information retrieval broadly compatible with experimental observations and is consistent with our recent graphical model (Romani et al., 2013).

Word length effect in free recall of randomly assembled word lists.

In serial recall experiments, human subjects are requested to retrieve a list of words in the same order as they were presented. In a classical study, participants were reported to recall more words from study lists composed of short words compared to lists of long words, the word length effect. The world length effect was also observed in free recall experiments, where subjects can retrieve the words in any order. Here we analyzed a large dataset from free recall experiments of unrelated words, where short and long words were randomly mixed, and found a seemingly opposite effect: long words are recalled better than the short ones. We show that our recently proposed mechanism of associative retrieval can explain both these observations. Moreover, the direction of the effect depends solely on the way study lists are composed.

Decision criteria in dual discrimination tasks estimated using external-noise methods.

According to classical signal detection theory (SDT), in simple detection or discrimination tasks, observers use a decision parameter based on their noisy internal response to set a boundary between "yes" and "no" responses. Experimental paradigms where performance is limited by internal noise cannot be used to provide an unambiguous measure of the decision criterion and its variability. Here, unidimensional external noise is used to estimate a criterion and its variability in stimulus space. Within this paradigm, the criterion is defined as the stimulus value separating the two response alternatives. This paradigm allows the assessment of interactions between criteria assigned to different targets in dual tasks. Previous studies suggested that observers' criteria interacted or even collapsed to one (hence, nonoptimal) criterion. An alternative interpretation of those results is that observers equated their false alarm (FA) rates. The external-noise method enables the confrontation of the two hypotheses. It is shown that the variability of observers' criterion in stimulus space is about 1.6 times their measured sensory threshold, suggesting that the presence of external noise increases decision uncertainty. Observers' stimulus criterion settings are close to SDT predictions in single tasks, but not in dual tasks where the two criteria tend to "attract" each other. Observers maintain distinct FA rates even when SDT predicts equal rates. Observers trained in psychophysics or provided with basic notions of SDT exemplified with the present experimental design manage to better separate their criteria in some conditions.

Lateral facilitation--no effect on the target noise level.

The detection threshold of a centrally placed Gabor target is reduced in the presence of aligned high-contrast Gabor patches that are optimally spaced from the target (Polat & Sagi, 1993). Here we determined whether threshold reduction is due to signal enhancement or to decreased signal response variability (internal noise), using a recently developed analysis for a Signal Detection Theory (SDT)-based contrast-identification paradigm (Katkov, Tsodyks, & Sagi, 2007a). We found that flankers did not affect internal noise, but instead caused increased target response when collinear with it, in agreement with the lateral facilitation effect. Based on these results, we concluded that lateral facilitation can be explained by signal enhancement only, and that uncertainty-based models do not provide a satisfactory description of the data.