Rhesus monkeys (Macaca mulatta) show robust primacy and recency in memory for lists from small, but not large, image sets

The combination of primacy and recency produces a U-shaped serial position curve typical of memory for lists. In humans, primacy is often thought to result from rehearsal, but there is little evidence for rehearsal in nonhumans. To further evaluate the possibility that rehearsal contributes to primacy in monkeys, we compared memory for lists of familiar stimuli (which may be easier to rehearse) to memory for unfamiliar stimuli (which are likely difficult to rehearse). Six rhesus monkeys saw lists of five images drawn from either large, medium, or small image sets. After presentation of each list, memory for one item was assessed using a serial probe recognition test. Across four experiments, we found robust primacy and recency with lists drawn from small and medium, but not large, image sets. This finding is consistent with the idea that familiar items are easier to rehearse and that rehearsal contributes to primacy, warranting further study of the possibility of rehearsal in monkeys. However, alternative interpretations are also viable and are discussed.     

Long-Term Recency in Anterograde Amnesia

Amnesia is usually described as an impairment of a long-term memory (LTM) despite an intact short-term memory (STM). The intact recency effect in amnesia had supported this view. Although dual-store models of memory have been challenged by single-store models based on interference theory, this had relatively little influence on our understanding and treatment of amnesia, perhaps because the debate has centred on experiments in the neurologically intact population. Here we tested a key prediction of single-store models for free recall in amnesia: that people with amnesia will exhibit a memory advantage for the most recent items even when all items are stored in and retrieved from LTM, an effect called long-term recency. People with amnesia and matched controls studied, and then free-recalled, word lists with a distractor task following each word, including the last (continual distractor task, CDFR). This condition was compared to an Immediate Free Recall (IFR, no distractors) and a Delayed Free Recall (DFR, end-of-list distractor only) condition. People with amnesia demonstrated the full long-term recency pattern: the recency effect was attenuated in DFR and returned in CDFR. The advantage of recency over midlist items in CDFR was comparable to that of controls, confirming a key prediction of single-store models. Memory deficits appeared only after the first word recalled in each list, suggesting the impairment in amnesia may emerge only as the participant’s recall sequence develops, perhaps due to increased susceptibility to output interference. Our findings suggest that interference mechanisms are preserved in amnesia despite the overall impairment to LTM, and challenge strict dual-store models of memory and their dominance in explaining amnesia. We discuss the implication of our findings for rehabilitation.     

Primacy and recency effects in extinction and latent inhibition: a selective review with implications for models of learning

In the framework of animal conditioning and human associative learning, primacy and recency effects on acquired stimulus control of behavior refer to the superior influence of first-learned and last-learned associations, respectively. Most contemporary associative models of learning anticipate unwavering recency effects and claim support from numerous published studies. But, for pragmatic reasons, almost all of these studies were conducted under select conditions that favored recency effects. When these conditions are not met, recency effects are far from ubiquitous. We review the literature on primacy and recency effects regarding extinction and latent inhibition (i.e., interference between outcomes), with special emphasis on the impact of certain post-training manipulations and test conditions on conditioned responding. Evidence for recency-to-primacy shifts and for memory integration is examined in light of contemporary models of learning.     

On recency and echoic memory

In short-term memory, the tendency for the last few (recency) items from a verbal sequence to be increasingly well recalled is more pronounced if the items are spoken rather than written. This auditory recency advantage has been quite generally attributed to echoic memory, on the grounds that in the auditory, but not the visual, mode, sensory memory persists just long enough to supplement recall of the most recent items. This view no longer seems tenable. There are now several studies showing that an auditory recency advantage occurs not only in long-term memory, but under conditions in which it cannot possibly be attributed to echoic memory. Also, similar recency phenomena have been discovered in short-term memory when the items are lip-read, or presented in sign-language, rather than heard. This article provides a partial review of these studies, taking a broad theoretical position from which these particular recency phenomena are approached as possible exceptions, to a general theory according to which recency is due to temporal distinctiveness. Much of the fresh evidence reviewed is of a somewhat preliminary nature and it is as yet unexplained by any theory of memory. The need for additional, converging experimental tests is obvious; so too is the need for further theoretical development. Several alternative theoretical resolutions are mentioned, including the possibility that enhanced recency may reflect movement, from sequentially occurring stimulus features, and the suggestion that it may be associated with the primary linguistic mode of the individuals concerned. But special weight is attached to the conjecture that all these recency phenomena might be accounted for in terms of distinctiveness or discriminability. On this view, the enhanced recency effects observed with certain modes, including the auditory mode, are attributed to items possessing greater temporal discriminability in those modes.     

Reactivation in Working Memory: An Attractor Network Model of Free Recall

The dynamic nature of human working memory, the general-purpose system for processing continuous input, while keeping no longer externally available information active in the background, is well captured in immediate free recall of supraspan word-lists. Free recall tasks produce several benchmark memory phenomena, like the U-shaped serial position curve, reflecting enhanced memory for early and late list items. To account for empirical data, including primacy and recency as well as contiguity effects, we propose here a neurobiologically based neural network model that unifies short- and long-term forms of memory and challenges both the standard view of working memory as persistent activity and dual-store accounts of free recall. Rapidly expressed and volatile synaptic plasticity, modulated intrinsic excitability, and spike-frequency adaptation are suggested as key cellular mechanisms underlying working memory encoding, reactivation and recall. Recent findings on the synaptic and molecular mechanisms behind early LTP and on spiking activity during delayed-match-to-sample tasks support this view.     

Dissociable Temporo-Parietal Memory Networks Revealed by Functional Connectivity during Episodic Retrieval

Episodic memory retrieval most often recruits multiple separate processes that are thought to involve different temporal regions. Previous studies suggest dissociable regions in the left lateral parietal cortex that are associated with the retrieval processes. Moreover, studies using resting-state functional connectivity (RSFC) have provided evidence for the temporo-parietal memory networks that may support the retrieval processes. In this functional MRI study, we tested functional significance of the memory networks by examining functional connectivity of brain activity during episodic retrieval in the temporal and parietal regions of the memory networks. Recency judgments, judgments of the temporal order of past events, can be achieved by at least two retrieval processes, relational and item-based. Neuroimaging results revealed several temporal and parietal activations associated with relational/item-based recency judgments. Significant RSFC was observed between one parahippocampal region and one left lateral parietal region associated with relational recency judgments, and between four lateral temporal regions and another left lateral parietal region associated with item-based recency judgments. Functional connectivity during task was found to be significant between the parahippocampal region and the parietal region in the RSFC network associated with relational recency judgments. However, out of the four tempo-parietal RSFC networks associated with item-based recency judgments, only one of them (between the left posterior lateral temporal region and the left lateral parietal region) showed significant functional connectivity during task. These results highlight the contrasting roles of the parahippocampal and the lateral temporal regions in recency judgments, and suggest that only a part of the tempo-parietal RSFC networks are recruited to support particular retrieval processes.     

Non-monotonic Temporal-Weighting Indicates a Dynamically Modulated Evidence-Integration Mechanism

Perceptual decisions are thought to be mediated by a mechanism of sequential sampling and integration of noisy evidence whose temporal weighting profile affects the decision quality. To examine temporal weighting, participants were presented with two brightness-fluctuating disks for 1, 2 or 3 seconds and were requested to choose the overall brighter disk at the end of each trial. By employing a signal-perturbation method, which deploys across trials a set of systematically controlled temporal dispersions of the same overall signal, we were able to quantify the participants’ temporal weighting profile. Results indicate that, for intervals of 1 or 2 sec, participants exhibit a primacy-bias. However, for longer stimuli (3-sec) the temporal weighting profile is non-monotonic, with concurrent primacy and recency, which is inconsistent with the predictions of previously suggested computational models of perceptual decision-making (drift-diffusion and Ornstein-Uhlenbeck processes). We propose a novel, dynamic variant of the leaky-competing accumulator model as a potential account for this finding, and we discuss potential neural mechanisms.     

Decreased Fronto-Temporal Interaction during Fixation after Memory Retrieval

Previous studies have revealed top-down control during memory retrieval from the prefrontal cortex to the temporal cortex. In the present functional MRI study, we investigated whether the fronto-temporal functional interaction occurs even during fixation periods after memory retrieval trials. During recency judgments, subjects judged the temporal order of two items in a study list. The task used in the present study consisted of memory trials of recency judgments and non-memory trials of counting dots, and post-trial fixation periods. By comparing the brain activity during the fixation periods after the memory trials with that during the fixation periods after the non-memory trials, we detected heightened brain activity in the lateral prefrontal cortex, the lateral temporal cortex and the hippocampus. Functional interactions during the fixation periods after the memory vs. non-memory trials as examined using a psychophysiological interaction revealed a decreased interaction from the lateral prefrontal cortex to the lateral temporal cortex, but not to the hippocampus. The functional interaction between the same frontal and temporal regions was also present during the memory trials. A trial-based functional connectivity analysis further revealed that the fronto-temporal interaction was positive and decreased during the fixation periods after the memory trials, relative to the fixation periods after the non-memory trials. These results suggest that the fronto-temporal interaction existed during the post-trial fixation periods, which had been present during the memory trials and temporally extended into the fixation periods.     

Intrusion of stereotyped responding in pigeon spatial memory tasks

Pigeons appear predisposed to respond in stereotyped manners in multi-item spatial memory tasks in which the items are simultaneously presented. We discovered this when we attempted to study primacy and recency serial position effects using a delayed matching of key location task (Experiment 1) and when we attempted to develop a keypack analog of the radial-arm maze task (Experiment 4). In Experiment 1 matching accuracy for first-pecked sample was above chance, approximately chance for the second-pecked sample, and below chance for the third-pecked sample. Experiments 2 and 3 showed that the results of Experiment 1 were due to “superstitious” stereotyped responding: When pigeons were allowed to respond to two or three keys in any order and then respond to the same keys again, they responded in the same order on the second occasion. In Experiment 4, the pigeons successfully avoided pecking previously-pecked keys but did so by pecking the keys in a fixed sequence.     

Synaptic efficacy shapes resource limitations in working memory

Working memory (WM) is limited in its temporal length and capacity. Classic conceptions of WM capacity assume the system possesses a finite number of slots, but recent evidence suggests WM may be a continuous resource. Resource models typically assume there is no hard upper bound on the number of items that can be stored, but WM fidelity decreases with the number of items. We analyze a neural field model of multi-item WM that associates each item with the location of a bump in a finite spatial domain, considering items that span a one-dimensional continuous feature space. Our analysis relates the neural architecture of the network to accumulated errors and capacity limitations arising during the delay period of a multi-item WM task. Networks with stronger synapses support wider bumps that interact more, whereas networks with weaker synapses support narrower bumps that are more susceptible to noise perturbations. There is an optimal synaptic strength that both limits bump interaction events and the effects of noise perturbations. This optimum shifts to weaker synapses as the number of items stored in the network is increased. Our model not only provides a circuit-based explanation for WM capacity, but also speaks to how capacity relates to the arrangement of stored items in a feature space.     

On the Chemical Nature and Origin of Teleonomy

The physico-chemical characterization of a teleonomic event and the nature of the physico-chemical process by which teleonomic systems could emerge from non-teleonomic systems are addressed in this paper. It is proposed that teleonomic events are those whose primary directive is discerned to be non-thermodynamic, while regular (non-teleonomic) events are those whose primary directive is the traditional thermodynamic one. For the archetypal teleonomic event, cell multiplication, the non-thermodynamic directive can be identified as being a kinetic directive. It is concluded, therefore, that the process of emergence, whereby non-teleonomic replicating chemical systems were transformed into teleonomic ones, involved a switch in the primacy of thermodynamic and kinetic directives. It is proposed that the step where that transformation took place was the one in which some pre-metabolic replicating system acquired an energy-gathering capability, thereby becoming metabolic. Such a transformation was itself kinetically directed given that metabolic replicators tend to be kinetically more stable than non-metabolic ones. The analysis builds on our previous work that considers living systems to be a kinetic state of matteras opposed to the traditional thermodynamic states that dominate the inanimate world     

Eye Movements Provide an Index of Veridical Memory for Temporal Order

The present research examined whether eye movements during retrieval capture the relation between an event and its temporal attributes. In two experiments (N=76), we found converging evidence that eye movements reflected the veridicality of memory for temporal order seconds before overt memory judgments, suggesting that these movements captured indirect access to temporal information. These eye movements did not entirely depend on the amount of contextual cueing available (Experiment 1) and reflected the unique ordinal position of an event in a sequence (Experiment 2). Based on our results, we conclude that eye movements reflected the absolute temporal order of past events.     

Fine-Grained,Local Maps and Coarse,Global Representations Support Human Spatial Working Memory

While sensory processes are tuned to particular features, such as an object''s specific location, color or orientation, visual working memory (vWM) is assumed to store information using representations, which generalize over a feature dimension. Additionally, current vWM models presume that different features or objects are stored independently. On the other hand, configurational effects, when observed, are supposed to mainly reflect encoding strategies. We show that the location of the target, relative to the display center and boundaries, and overall memory load influenced recall precision, indicating that, like sensory processes, capacity limited vWM resources are spatially tuned. When recalling one of three memory items the target distance from the display center was overestimated, similar to the error when only one item was memorized, but its distance from the memory items'' average position was underestimated, showing that not only individual memory items'' position, but also the global configuration of the memory array may be stored. Finally, presenting the non-target items at recall, consequently providing landmarks and configurational information, improved precision and accuracy of target recall. Similarly, when the non-target items were translated at recall, relative to their position in the initial display, a parallel displacement of the recalled target was observed. These findings suggest that fine-grained spatial information in vWM is represented in local maps whose resolution varies with distance from landmarks, such as the display center, while coarse representations are used to store the memory array configuration. Both these representations are updated at the time of recall.     

European Neogene rodent communities: explaining family-level replacements through a spatiotemporal approach

We analyse the changes in rodent regional assemblages based on 657 fossil-bearing western European localities distributed ranging from ca. 27 Ma (Late Oligocene) to ca. 3 Ma (Late Pliocene). We compare temporal and spatial patterns in order to identify the factors that drive the evolutions of communities. Regional assemblages are analysed based on the distribution of species richness among families. First, communities are temporally analysed to identify significant changes in their composition. Second, regional communities are spatially compared to indentify diversity gradients. The temporal analysis reveals that communities' evolution is marked by several breaks in their composition, related to either migration or environmental/climatic events. This evolution can be summarised in terms of shifts in the relative abundances of glirids, cricetids and murids within assemblages. In contrast, spatial analysis shows that only environmental changes induce long-lasting changes in diversity gradients. Some observations made on cricetids and murids extant relatives indicate that they have undergone a large dietary diversification enabled by a specific digestive tract (along with the diversification of other life-history traits), whereas glirids are more specialised. The opposite diversity dynamics of these groups emphasises the importance of family-level adaptive potential in diversity conservation issues when facing environmental changes.     

Genotype accounts for intraspecific variation in the timing and duration of multiple,sequential life-cycle events in a willow species

Premise

Phenological variation among individuals within populations is common and has a variety of ecological and evolutionary consequences, including forming the basis for population-level responses to environmental change. Although the timing of life-cycle events has genetic underpinnings, whether intraspecific variation in the duration of life-cycle events reflects genetic differences among individuals is poorly understood.

Methods

We used a common garden experiment with 10 genotypes of Salix hookeriana (coastal willow) from northern California, United States to investigate the extent to which genetic variation explains intraspecific variation in the timing and duration of multiple, sequential life-cycle events: flowering, leaf budbreak, leaf expansion, fruiting, and fall leaf coloration. We used seven clones of each genotype, for a total of 70 individual trees.

Results

Genotype affected each sequential life-cycle event independently and explained on average 62% of the variation in the timing and duration of vegetative and reproductive life-cycle events. All events were significantly heritable. A single genotype tended to be “early” or “late” across life-cycle events, but for event durations, there was no consistent response within genotypes.

Conclusions

This research demonstrates that genetic variation can be a major component underlying intraspecific variation in the timing and duration of life-cycle events. It is often assumed that the environment affects durations, but we show that genetic factors also play a role. Because the timing and duration of events are independent of one another, our results suggest that the effects of environmental change on one event will not necessarily cascade to subsequent events.     

Spontaneous recovery in human instrumental learning: Integration of information and recency to primacy shift

This experiment was conducted to study the effect of changes in the retention interval (RI) on spontaneous recovery within an acquisition-test interference task. College students learned a reversal conditional discrimination to solve a task involving conflicting phases across two training phases. When the test was conducted immediately after training, participants’ performance revealed recency, behaving according to the information received during the last phase. Performance after retention interval averaged the information received across phases, regardless of the length of the RI (1.5, 3, 24 or 48 h). These results are not in agreement with traditonal theories of spontaneous recovery as they predict a recency to primacy shift effect of the RI. An interpretation of spontaneous recovery based on a temporal weighting rule (TWR) is discussed.     

Multiscale representations of community structures in attractor neural networks

Our cognition relies on the ability of the brain to segment hierarchically structured events on multiple scales. Recent evidence suggests that the brain performs this event segmentation based on the structure of state-transition graphs behind sequential experiences. However, the underlying circuit mechanisms are poorly understood. In this paper we propose an extended attractor network model for graph-based hierarchical computation which we call the Laplacian associative memory. This model generates multiscale representations for communities (clusters) of associative links between memory items, and the scale is regulated by the heterogenous modulation of inhibitory circuits. We analytically and numerically show that these representations correspond to graph Laplacian eigenvectors, a popular method for graph segmentation and dimensionality reduction. Finally, we demonstrate that our model exhibits chunked sequential activity patterns resembling hippocampal theta sequences. Our model connects graph theory and attractor dynamics to provide a biologically plausible mechanism for abstraction in the brain.     

Abnormal semantic processing in females with fragile X‐associated tremor/ataxia syndrome

Fragile X‐associated tremor/ataxia syndrome (FXTAS), a neurodegenerative disorder, affects fragile X (FMR1) gene premutation carriers in late life. Studies have shown cognitive impairments in FXTAS including executive dysfunction, working memory and visuospatial deficits. However, less is known about cognition in females with FXTAS. Thus, we examined semantic processing and verbal memory in female FXTAS patients with event‐related potentials (ERPs) and neuropsychological testing. Sixty‐one females (34 FXTAS, M_age = 62.7; 27 controls, M_age = 60.4) were studied with 32‐channel ERPs during a category judgment task in which semantically congruous (50%) and incongruous items were repeated approximately 10–140 seconds later. N400 and P600 amplitude data were submitted to analysis of covariance. Neuropsychological testing demonstrated lower performance in verbal learning and executive function in females with FXTAS. Event‐related potential analyses showed a significant reduction of the N400 congruity effect (incongruous ? congruous) in the FXTAS group. The N400 congruity effect reduction in females with FXTAS was mainly due to increased N400 amplitude to congruous new words. No significant abnormalities of the N400 repetition effect or the P600 repetition effect were found, indicating preserved implicit memory and verbal memory, respectively, in females with FXTAS. The decreased N400 congruity effect suggests abnormal semantic expectancy and/or semantic network disorganization in female FXTAS patients. The enhanced N400 amplitude to congruous new words may reflect decreased cognitive flexibility among FXTAS women, making access to less typical category exemplar words more difficult .     

Modeling daily flowering probabilities: expected impact of climate change on Japanese cherry phenology

Understanding the drivers of phenological events is vital for forecasting species’ responses to climate change. We developed flexible Bayesian survival regression models to assess a 29‐year, individual‐level time series of flowering phenology from four taxa of Japanese cherry trees (Prunus spachiana, Prunus × yedoensis, Prunus jamasakura, and Prunus lannesiana), from the Tama Forest Cherry Preservation Garden in Hachioji, Japan. Our modeling framework used time‐varying (chill and heat units) and time‐invariant (slope, aspect, and elevation) factors. We found limited differences among taxa in sensitivity to chill, but earlier flowering taxa, such as P. spachiana, were more sensitive to heat than later flowering taxa, such as P. lannesiana. Using an ensemble of three downscaled regional climate models under the A1B emissions scenario, we projected shifts in flowering timing by 2100. Projections suggest that each taxa will flower about 30 days earlier on average by 2100 with 2–6 days greater uncertainty around the species mean flowering date. Dramatic shifts in the flowering times of cherry trees may have implications for economically important cultural festivals in Japan and East Asia. The survival models used here provide a mechanistic modeling approach and are broadly applicable to any time‐to‐event phenological data, such as plant leafing, bird arrival time, and insect emergence. The ability to explicitly quantify uncertainty, examine phenological responses on a fine time scale, and incorporate conditions leading up to an event may provide future insight into phenologically driven changes in carbon balance and ecological mismatches of plants and pollinators in natural populations and horticultural crops.     

Effects of DGAVP on verbal memory

Effects of DGAVP (desglycinamide-arginine-vasopressin, a synthetic vasopressin analog) on verbal memory were investigated in 13 healthy male volunteers. Ten word lists, each consisting of 15 words, were presented to the subjects who had to recall them according to a free recall paradigm. The total number of recalled words was not different between DGAVP and placebo treatment; but DGAVP had an effect on memory performance depending on the serial position of the words. It attenuated the primacy effect and enhanced the recency effect of memory performance. The pattern of changes after DGAVP may be consistent with an effect of the peptide on general arousal. Since the experiment was not designed to test influences of DGAVP on arousal, these considerations remain tentative.