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The perception of temporal order or simultaneity of stimuli is almost always explained in terms of independent-channels models, such as perceptual-moment, triggered-moment, and attention-switching models. Independent-channels models generally posit that stimuli are processed in separate peripheral channels and that their arrival-time difference at a central location is translated into an internal state of order (simultaneity) if it reaches (misses) a certain threshold. Non-monotonic and non-parallel psychometric functions in a ternary-response task provided critical evidence against a wide range of independent-channels models. However, two independent-channels models have been introduced in the last decades that can account for such shapes by considering misreports of internal states (response-error model) or by assuming that simultaneity and order judgments rely on distinct sensory and decisional processes (two-stage model). Based on previous ideas, we also consider a two-threshold model, according to which the same arrival-time difference may need to reach a higher threshold for order detection than for successiveness detection. All three models were fitted to various data sets collected over a period of more than a century. The two-threshold model provided the best balance between goodness of fit and parsimony. This preference for the two-threshold model over the two-stage model and the response-error model aligns well with several lines of evidence from cognitive modeling, psychophysics, mental chronometry, and psychophysiology. We conclude that the seemingly deviant shapes of psychometric functions can be explained within the framework of independent-channels models in a simpler way than previously assumed.

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Although the hippocampus has been implicated in both the temporal organization of memories and association of scene elements, some theoretical accounts posit that the role of the hippocampus in episodic memory is largely atemporal. In this study, we set out to explore this discrepancy by identifying hippocampal activity patterns related to scene construction while participants performed a temporal order memory task. Participants in the fMRI scanner were shown a sequence of photographs, each consisting of a central object and a contextual background scene. On each retrieval trial, participants were shown a pair of the original photographs (FULL), objects from the scenes without the background (OBJ), or background contexts without the main foreground object (BACK). In the temporal order judgment (TOJ) task, participants judged the temporal order of the pair of scenes; in the Viewing trials, two identical scenes were shown without any task. First, we found that the anterior hippocampus-particularly the CA1 and subiculum-showed similar patterns of activation between the BACK and OBJ conditions, suggesting that scene construction occurred spontaneously during both TOJ and Viewing. Furthermore, neural markers of scene construction in the anterior hippocampus did not apply to incorrect trials, showing that successful temporal memory retrieval was functionally linked to scene construction. In the cortex, time-processing areas, such as the supplementary motor area and the precuneus, and scene-processing areas, such as the parahippocampal cortex, were activated and functionally connected with the hippocampus. Together, these results support the view that the hippocampus is concurrently involved in scene construction and temporal organization of memory and propose a model of hippocampal episodic memory that takes both processes into account.

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Studies have shown that enactment improves memory; however, in daily life, our memories of motor events often exhibit a relative temporal order. Therefore, this study examined whether enactment promotes relative temporal order memory. In Experiment 1, a sequential recall task and a subject-performed task were used to explore whether enactment encoding improved relative temporal order memory. The results showed that the relative temporal order memory of the enactment-encoding group was significantly better than that of the verbal-encoding group, indicating that enactment promoted relative temporal order memory. Since temporal order memory is often affected by spatial cues, in Experiment 2, we further controlled spatial cues and used a 2 (spatial cues: consistent with temporal order, vs. no cues) × 2 (encoding type: verbal vs. enactment) design to explore whether spatial cues influence the effect of enactment encoding on temporal order memory. The results showed that compared with verbal encoding, enactment encoding significantly improved relative temporal order memory. However, no effect of spatial cues on relative temporal order memory was found. Our study confirmed that enactment encoding promotes relative temporal order memory performance independent of spatial cues.

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Our actions shape our everyday experience: what we experience, how we perceive, and remember it are deeply affected by how we interact with the world. Performing an action to deliver a stimulus engages neurophysiological processes which are reflected in the modulation of sensory and pupil responses. We hypothesized that these processes shape memory encoding, parsing the experience by grouping self- and externally generated stimuli into differentiated events. Participants encoded sound sequences, in which either the first or last few sounds were self-generated and the rest externally generated. We tested recall of the sequential order of sounds that had originated from the same (within event) or different sources (across events). Memory performance was not higher for within-event sounds, suggesting that actions did not structure the memory representation. However, during encoding, we observed the expected electrophysiological response attenuation for self-generated sounds, together with increased pupil dilation triggered by actions. Moreover, at the boundary between events, physiological responses to the first sound from the new source were influenced by the direction of the source switch. Our results suggest that introducing actions creates a stronger contextual shift than removing them, even though actions do not directly contribute to memory performance. This study contributes to our understanding of how interacting with sensory input shapes experiences by exploring the relationships between action effects on sensory responses, pupil dilation, and memory encoding. Importantly, it challenges the notion of a meaningful contribution from low-level neurophysiological mechanisms associated with action execution in the modulation of the self-generation effect.

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Numerous studies have demonstrated the influence of musical expertise on spoken language processing; however, its effects on reading comprehension remain largely unexplored. This study aims to investigate the role of musical expertise in sentence comprehension, particularly concerning the processing of temporal order. Using two self-paced reading experiments, we examined individuals' responses to two-clause sentences connected by the temporal connectives "before" or "after". "After" sentences consistently presented events in their actual order of temporal occurrence, while "before" sentences described events in reverse temporal order. In both experiments, our analyses of reading times consistently uncovered a significant temporal order effect, with words immediately following the temporal connectives being processed slower in "before" sentences compared to "after" sentences. This suggests the presence of immediate online processing costs associated with "before" sentences. Notably, these processing costs were found to be attenuated in individuals with musical expertise compared to those without. However, analyses of comprehension accuracy showed no advantage of musicians over non-musicians. Specifically, in Experiment 1, the two groups showed no difference in comprehension accuracy, while in Experiment 2, musicians exhibited lower accuracy rates compared to non-musicians in both "before" and "after" sentences. These results suggest that musical expertise may attenuate online processing costs associated with complex linguistic constructs, but could not promote reading accuracy. We concluded that music training is associated with a restricted effect on written sentence comprehension.

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Dopamine D1 receptor agonists improve spatial working memory, but their effects on temporal order memory, particularly prone to the effects of aging, have not been studied. Two D1 agonists, PF6256142 (PF) and 2-methyldihydrexidine (2MDHX), were examined for their effects in a rodent temporal order recognition task. Our results are consistent with the hypothesis that there is an age-related decline in rodent temporal order memory. The data also show that either agonist rescues the poor memory performance with a large effective size. Interestingly, the optimal effective dose varied among individual rats of different age groups. PF showed greater potency for older rats, whereas 2MDHX showed better overall population effectiveness. Both PF and 2MDHX have high intrinsic activity at rodent D1-mediated cAMP synthesis. Conversely, at D1-mediated ß-arrestin recruitment, PF has essentially no intrinsic activity, whereas 2MDHX is a super-agonist. These findings suggest that D1 agonists have potential to treat age-related cognitive decline, and the pattern of functional selectivity may be useful for developing drugs with an improved therapeutic index.

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The temporal component of episodic memory has been recognized as a sensitive behavioral marker in early stage of Alzheimer's disease (AD) patients. However, parallel studies in AD animals are currently lacking, and the underlying neural circuit mechanisms remain poorly understood. Using a novel AppNL-G-F knock-in (APP-KI) rat model, the developmental changes of temporal order memory (TOM) and the relationship with medial prefrontal cortex and perirhinal cortex (mPFC-PRH) circuit were determined through in vivo electrophysiology and microimaging technique. We observed a deficit in TOM performance during the object temporal order memory task (OTOMT) in APP-KI rats at 6 month old, which was not evident at 3 or 4 months of age. Alongside behavioral changes, we identified a gradually extensive and aggravated regional activation and functional alterations in the mPFC and PRH during the performance of OTOMT, which occurred prior to the onset of TOM deficits. Moreover, coherence analysis showed that the functional connectivity between the mPFC and PRH could predict the extent of future behavioral performance. Further analysis revealed that the aberrant mPFC-PRH interaction mainly attributed to the progressive deterioration of synaptic transmission, information flow and network coordination from mPFC to PRH, suggesting the mPFC dysfunction maybe the key area of origin underlying the early changes of TOM. These findings identify a pivotal role of the mPFC-PRH circuit in mediating the TOM deficits in the early stage of AD, which holds promising clinical translational value and offers potential early biological markers for predicting AD memory progression.

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Our brain constantly integrates afferent information, such as visual and tactile information, to perceive the world around us. According to the maximum-likelihood estimation (MLE) model, imprecise information will be weighted less than precise, making the multisensory percept as precise as possible. Individuals with fibromyalgia (FM), a chronic pain syndrome, show alterations in the integration of tactile information. This could lead to a decrease in their weight in a multisensory percept or a general disruption of multisensory integration, making it less beneficial. To assess multisensory integration, 15 participants with FM and 18 pain-free controls performed a temporal-order judgment task in which they received pairs of sequential visual, tactile (unisensory conditions), or visuotactile (multisensory condition) stimulations on the index and the thumb of the non-dominant hand and had to determine which finger was stimulated first. The task enabled us to measure the precision and accuracy of the percept in each condition. Results indicate an increase in precision in the visuotactile condition compared to the unimodal conditions in controls only, although we found no intergroup differences. The observed visuotactile precision was correlated to the precision predicted by the MLE model in both groups, suggesting an optimal integration. Finally, the weights of the sensory information were not different between the groups; however, in the group with FM, higher pain intensity was associated with smaller tactile weight. This study shows no alterations of the visuotactile integration in individuals with FM, though pain may influence tactile weight in these participants.

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The objective of this study was to investigate the impact of music training on the processing of temporal order in Mandarin sentence reading using event-related potentials (ERPs). Two-clause sentences with temporal connectives ("before" or "after") were presented to both musicians and non-musicians. Additionally, a verbal N-back task was utilized to evaluate the participants' working memory capacities. The findings revealed that musicians, but not nonmusicians, demonstrated a more negative amplitude in the second clauses of "before" sentences compared with "after" sentences. In the N-back task, musicians exhibited faster reaction times than nonmusicians in the two-back condition. Furthermore, a correlation was observed between the ERP amplitude differences (before vs. after) and reaction time differences in the N-back task (0-back vs. 2-back) among musicians. These findings suggested that music training enhances the depth of temporal order processing, potentially mediated by improvements in working memory capacity.

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We review the evidence for the conceptual association between arithmetic and space and quantify the effect size in meta-analyses. We focus on three effects: (a) the operational momentum effect (OME), which has been defined as participants' tendency to overestimate results of addition problems and underestimate results of subtraction problems; (b) the arithmetic cueing effect, in which arithmetic problems serve as spatial cues in target detection or temporal order judgment tasks; and (c) the associations between arithmetic and space observed with eye- and hand-tracking studies. The OME was consistently found in paradigms that provided the participants with numerical response alternatives. The OME shows a large effect size, driven by an underestimation during subtraction while addition was unbiased. In contrast, paradigms in which participants indicated their estimate by transcoding their final estimate to a spatial reference frame revealed no consistent OME. Arithmetic cueing studies show a reliable small to medium effect size, driven by a rightward bias for addition. Finally, eye- and hand-tracking studies point to replicable associations between arithmetic and eye or hand movements. To account for the complexity of the observed pattern, we introduce the Adaptive Pathways in Mental Arithmetic (APiMA) framework. The model accommodates central notions of numerical and arithmetic processing and helps identifying which pathway a given paradigm operates on. It proposes that the divergence between OME and arithmetic cueing studies comes from the predominant use of non-symbolic versus symbolic stimuli, respectively. Overall, our review and findings clearly support an association between arithmetic and spatial processing.

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Emotional stimuli (e.g. words, images) are often remembered better than neutral stimuli. However, little is known about how memory is affected by an environmentally induced emotional state (without any overtly emotional occurrences) - the focus of this study. Participants were randomly assigned to discovery (n = 305) and replication (n = 306) subsamples and viewed a desktop virtual environment before rating their emotions and completing objective (i.e. item, temporal-order, duration) and subjective (e.g. vividness, sensory detail, coherence) memory measures. In both samples, a Partial Least Squares Correlation analysis showed that an emotional state characterised by high negative emotion (i.e. threat, fear, anxiety) and arousal was reliably associated with better memory in both objective (i.e. item) and subjective (i.e. vividness and sensory detail) domains. No reliable associations were observed for any temporal memory measures (objective or subjective). Thus, an environmentally induced state of negative emotion corresponds with enhanced memory for indices of episodic memory pertaining to "what" happened, but not necessarily "when" it happened.

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Here's an all-too-familiar scenario: Person A is staring at person B, and then B turns toward A, and A immediately looks away (a phenomenon we call 'gaze deflection'). Beyond perceiving lower-level properties here - such as the timing of the eye/head turns - you can also readily perceive seemingly higher-level social dynamics: A got caught staring, and frantically looked away in embarrassment! It seems natural to assume that such social impressions are based on more fundamental representations of what happened when - but here we show that social gaze dynamics are unexpectedly powerful in that they can actually alter (and even reverse) the perceived temporal order of the underlying events. Across eight experiments, observers misperceived B as turning before A, when in fact they turned simultaneously - and even when B was turning after A. Additional controls confirmed that this illusion depends on visual processing (vs. being driven solely by higher-level interpretations), and that it is specific to the perception of social agents (vs. non-social objects). This demonstrates how social perception is tightly integrated into our perceptual experience of the world, and can have powerful consequences for one of the most basic properties that we can perceive: what happens when.

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Temporal order memory is impaired in autism spectrum disorder (ASD) and schizophrenia (SCZ). These disorders, more prevalent in males, result in abnormal dendritic spine pruning during adolescence in layer 3 (L3) medial prefrontal cortex (mPFC), yielding either too many (ASD) or too few (SCZ) spines. Here we tested whether altering spine density in neural circuits including the mPFC could be associated with impaired temporal order memory in male mice. We have shown that α4ßδ GABAA receptors (GABARs) emerge at puberty on spines of L5 prelimbic mPFC (PL) where they trigger pruning. We show here that α4ßδ receptors also increase at puberty in L3 PL (P < 0.0001) and used these receptors as a target to manipulate spine density here. Pubertal injection (14 d) of the GABA agonist gaboxadol, at a dose (3 mg/kg) selective for α4ßδ, reduced L3 spine density by half (P < 0.0001), while α4 knock-out increased spine density âˆ¼ 40 % (P < 0.0001), mimicking spine densities in SCZ and ASD, respectively. In both cases, performance on the mPFC-dependent temporal order recognition task was impaired, resulting in decreases in the discrimination ratio which assesses preference for the novel object: -0.39 ± 0.15, gaboxadol versus 0.52 ± 0.09, vehicle; P = 0.0002; -0.048 ± 0.10, α4 KO versus 0.49 ± 0.04, wild-type; P < 0.0001. In contrast, the number of approaches was unaltered, reflecting unchanged locomotion. These data suggest that altering α4ßδ GABAR expression/activity alters spine density in L3 mPFC and impairs temporal order memory to mimic changes in ASD and SCZ. These findings may provide insight into these disorders.

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Purpose: Alterations in spatial attention have been reported in people with chronic pain and may be relevant to understanding its cortical mechanisms and developing novel treatments. There is conflicting evidence as to whether people with Complex Regional Pain Syndrome (CRPS) have reduced visuospatial attention to their affected limb and/or its surrounding space, with some evidence that these deficits may be greater in personal (bodily) space. We aimed to test the competing hypotheses of whether the visuospatial attentional bias is specific to the personal space of the affected limb or generalizes to the personal space of other parts of the affected side of the body. Patients and Methods: Using visual Temporal Order Judgement tasks, we measured spatial attention in the personal space of the hands and feet of patients with upper (n=14) or lower (n=14) limb CRPS and pain-free controls (n=17). Participants judged the order of two light flashes presented at different temporal offsets on each of their hands or feet. Slower processing of the flash on one side relative to the other reflects reduced attention to that side of space. Results: Controls prioritized stimuli on the non-dominant (left) relative to dominant side, consistent with the well-documented normal leftward bias of attention (ie "pseudoneglect"). Regardless of the location (upper or lower limb) of the pain or visual stimuli, people with CRPS showed no such asymmetry, representing reduced attention to the affected side (compared to the greater attention of controls to their non-dominant side). More severe CRPS symptoms were associated with a greater tendency to deprioritize stimuli on the affected side. Conclusion: Our findings suggest that relative visuospatial bias in CRPS is generalized to the personal space of the affected side of the body, rather than being specific to the personal space of the CRPS-affected limb.

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Research has shown that children on the autism spectrum and adults with high levels of autistic traits are less sensitive to audiovisual asynchrony compared to their neurotypical peers. However, this evidence has been limited to simultaneity judgments (SJ) which require participants to consider the timing of two cues together. Given evidence of partly divergent perceptual and neural mechanisms involved in making temporal order judgments (TOJ) and SJ, and given that SJ require a more global type of processing which may be impaired in autistic individuals, here we ask whether the observed differences in audiovisual temporal processing are task and stimulus specific. We examined the ability to detect audiovisual asynchrony in a group of 26 autistic adult males and a group of age and IQ-matched neurotypical males. Participants were presented with beep-flash, point-light drumming, and face-voice displays with varying degrees of asynchrony and asked to make SJ and TOJ. The results indicated that autistic participants were less able to detect audiovisual asynchrony compared to the control group, but this effect was specific to SJ and more complex social stimuli (e.g., face-voice) with stronger semantic correspondence between the cues, requiring a more global type of processing. This indicates that audiovisual temporal processing is not generally different in autistic individuals and that a similar level of performance could be achieved by using a more local type of processing, thus informing multisensory integration theory as well as multisensory training aimed to aid perceptual abilities in this population.

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Introduction: Our brain continuously maps our body in space. It has been suggested that at least two main frames of reference are used to process somatosensory stimuli presented on our own body: the anatomical frame of reference (based on the somatotopic representation of our body in the somatosensory cortex) and the spatial frame of reference (where body parts are mapped in external space). Interestingly, a mismatch between somatotopic and spatial information significantly affects the processing of bodily information, as demonstrated by the "crossing hand" effect. However, it is not clear if this impairment occurs not only when the conflict between these frames of reference is determined by a static change in the body position (e.g., by crossing the hands) but also when new associations between motor and sensory responses are artificially created (e.g., by presenting feedback stimuli on a side of the body that is not involved in the movement). Methods: In the present study, 16 participants performed a temporal order judgment task before and after a congruent or incongruent visual-tactile-motor- task in virtual reality. During the VR task, participants had to move a cube using a virtual stick. In the congruent condition, the haptic feedback during the interaction with the cube was provided on the right hand (the one used to control the stick). In the incongruent condition, the haptic feedback was provided to the contralateral hand, simulating a sort of 'active' crossed feedback during the interaction. Using a psychophysical approach, the point of subjective equality (or PSE, i.e., the probability of responding left or right to the first stimulus in the sequence in 50% of the cases) and the JND (accuracy) were calculated for both conditions, before and after the VR-task. Results: After the VR task, compared to the baseline condition, the PSE shifted toward the hand that received the haptic feedback during the interaction (toward the right hand for the congruent condition and toward the left hand for the incongruent condition). Dicussion: This study demonstrated the possibility of inducing spatial biases in the processing of bodily information by modulating the sensory-motor interaction between stimuli in virtual environments (while keeping constant the actual position of the body in space).

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Remembering personal past events and their order is important. These capacities are essential to episodic and autobiographical memory theories, are needed in the creation of life stories and vital in forensic settings. As important as memory for events and their order are, relatively little is known about their development and the underlying neural processes that support them. Further, there is a paucity of studies that have examined memory and its development for autobiographical, yet controlled, events. The objective of this study was to examine memory for the temporal order of naturalistic "real world" events by directly comparing 7-11-year-olds and adults using both behavioral and event-related potential (ERP) measures. Participants photographed events at a local museum and after a delay, we used their photographs to test their memory for the temporal order of pairs of the events. We experimentally manipulated the temporal distance between the event pairs (whether the two events photographed in the pair had a short or long temporal distance between them). A memory asymmetry manipulation was also included such that at retrieval, photographs were either presented in forward direction (photograph on the top configuration was taken before photograph shown on the bottom) or vice versa. Children and adults showed sensitivity to temporal distance between events based on behavior (in some instances accuracy was higher for long compared to short temporal distance) and ERP (differential neural processing for short and long temporal distance conditions). Only adults showed sensitivity to the memory asymmetry manipulation, and only when the events occurred within a short temporal distance. A larger study is needed to confirm the present "proof of concept" study results. There is strong potential of this photo paradigm approach, combining naturalistic events with ERP, in future developmental studies, and would further our understanding of how memory behavior and the neural processes underlying memory operate in the "real world."

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Cross-modal correspondences refer to associations between stimulus features across sensory modalities. Previous studies have shown that cross-modal correspondences modulate reaction times for detecting and identifying stimuli in one modality when uninformative stimuli from another modality are present. However, it is unclear whether such modulation reflects changes in modality-specific perceptual processing. We used two psychophysical timing judgment tasks to examine the effects of audiovisual correspondences on visual perceptual processing. In Experiment 1, we conducted a temporal order judgment (TOJ) task that asked participants to judge which of two visual stimuli presented with various stimulus onset asynchronies (SOAs) appeared first. In Experiment 2, we conducted a simultaneous judgment (SJ) task that asked participants to report whether the two visual stimuli were simultaneous or successive. We also presented an unrelated auditory stimulus, simultaneously or preceding the first visual stimulus, and manipulated the congruency between audiovisual stimuli. Experiment 1 indicated that the points of subjective simultaneity (PSSs) between the two visual stimuli estimated in the TOJ task shifted according to the audiovisual correspondence between the auditory pitch and visual features of vertical location and size. However, these audiovisual correspondences did not affect PSS estimated using the SJ task in Experiment 2. The different results of the two tasks can be explained through the response bias triggered by audiovisual correspondence that only the TOJ task included. We concluded that audiovisual correspondence would not modulate visual perceptual timing and that changes in modality-specific perceptual processing might not trigger the congruency effects reported in previous studies.

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Our brains have a propensity to integrate closely-timed auditory and visual stimuli into a unified percept; a phenomenon that is highly malleable based on prior sensory experiences, and is known to be altered in clinical populations. While the neural correlates of audiovisual temporal perception have been investigated using neuroimaging and electroencephalography techniques in humans, animal research will be required to uncover the underlying cellular and molecular mechanisms. Prior to conducting such mechanistic studies, it is important to first confirm the translational potential of any prospective animal model. Thus, in the present study, we conducted a series of experiments to determine if rats show the hallmarks of audiovisual temporal perception observed in neurotypical humans, and whether the rat behavioral paradigms could reveal when they experienced perceptual disruptions akin to those observed in neurodevelopmental disorders. After training rats to perform a temporal order judgment (TOJ) or synchrony judgment (SJ) task, we found that the rats' perception was malleable based on their past and present sensory experiences. More specifically, passive exposure to asynchronous audiovisual stimulation in the minutes prior to behavioral testing caused the rats' perception to predictably shift in the direction of the leading stimulus; findings which represent the first time that this form of audiovisual perceptual malleability has been reported in non-human subjects. Furthermore, rats performing the TOJ task also showed evidence of rapid recalibration, in which their audiovisual temporal perception on the current trial was predictably influenced by the timing lag between the auditory and visual stimuli in the preceding trial. Finally, by manipulating either experimental testing parameters or altering the rats' neurochemistry with a systemic injection of MK-801, we showed that the TOJ and SJ tasks could identify when the rats had difficulty judging the timing of audiovisual stimuli. These findings confirm that the behavioral paradigms are indeed suitable for future testing of rats with perceptual disruptions in audiovisual processing. Overall, our collective results highlight that rats represent an excellent animal model to study the cellular and molecular mechanisms underlying the acuity and malleability of audiovisual temporal perception, as they showcase the perceptual hallmarks commonly observed in humans.

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(1) Background: Since the emergence of severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), the evolutionary traits of its variants have been revealed. However, the temporal order of the majority of mutations harbored by variants after the closest ancestors (or precursors), as "missing links", remains unclear. In this study, we aimed to unveil such missing links based on analyses of S protein homology by focusing on specimens with incomplete sets of S protein mutations in a variant. (2) Methods: Prevariant and postvariant mutations were defined as those before and after the variant's development, respectively. A total of 6,758,926 and 14,519,521 genomes were obtained from the National Center for Biotechnology Information and the GISAID initiative, respectively, and S protein mutations were detected based on BLASTN analyses. (3) Results: The temporal order of prevariant mutations harbored by 12 variants was deduced. In particular, the D950N mutation in the Mu variant shows V-shaped mutation transitions, in which multiple routes of evolution were combined and resulted in the formation of a V-shaped transition, indicating recombination. (4) Conclusions: Many genome data for SARS-CoV-2 unveiled the candidate precursors of Mu variant based on a data-driven approach to its prevariant mutations in each nation.

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