Time-dependent transformations of memory representations differ along the long axis of the hippocampus

Research has shown that sleep is beneficial for the long-term retention of memories. According to theories of memory consolidation, memories are gradually reorganized, becoming supported by widespread, distributed cortical networks, particularly during postencoding periods of sleep. However, the effects of sleep on the organization of memories in the hippocampus itself remains less clear. In a 3-d study, participants encoded separate lists of word–image pairs differing in their opportunity for sleep-dependent consolidation. Pairs were initially studied either before or after an overnight sleep period, and were then restudied in a functional magnetic resonance imaging (fMRI) scan session. We used multivariate pattern similarity analyses to examine fine-grained effects of consolidation on memory representations in the hippocampus. We provide evidence for a dissociation along the long axis of the hippocampus that emerges with consolidation, such that representational patterns for object–word memories initially formed prior to sleep become differentiated in anterior hippocampus and more similar, or overlapping, in posterior hippocampus. Differentiation in anterior hippocampal representations correlated with subsequent behavioral performance. Furthermore, representational overlap in posterior hippocampus correlated with the duration of intervening slow wave sleep. Together, these results demonstrate that sleep-dependent consolidation promotes the reorganization of memory traces along the long axis of the hippocampus.

The hippocampus has long been considered critical for encoding new memories; however, the effects of consolidation on hippocampal memory traces has remained an area of active research. Memories are thought to be stabilized for the long term as they become distributed across neocortical networks (Buzsáki 1989; Alvarez and Squire 1994; McClelland et al. 1995), a process supported by mechanisms during sleep (Diekelmann and Born 2010; Rasch and Born 2013). Whereas much research has been devoted to understanding the hippocampal contributions to the long-term retention of memories, open questions remain in considering how sleep-dependent consolidation affects the organization of hippocampal traces.The hippocampus has previously been shown to be critical for binding disparate elements of an experience together (Cohen and Eichenbaum 1993; Davachi 2006). Theories suggest that the hippocampus quickly encodes new experiences, while the cortex, with a slower learning rate, gradually comes to represent the central features from this hippocampal trace, resulting in abstracted memories that can be integrated into long-term cortical stores (McClelland et al. 1995). Prior research has demonstrated evidence for a coordinated hippocampal–cortical dialogue during sleep (Andrade et al. 2011; Bergmann et al. 2012; Ngo et al. 2020) as well as enhanced hippocampal–cortical functional connectivity after learning, facilitating the retention of memories (Tambini et al. 2010; Tompary et al. 2015; Murty et al. 2017; Cowan et al. 2021). Reports suggest consolidation results in more integrated cortical memory traces in the cortex (Richards et al. 2014; Tompary and Davachi 2017; Cowan et al. 2020); however, it remains an open question whether the active consolidation processes that support memory reorganization across hippocampal–cortical networks also transform hippocampal memory traces.Research on the fate of the hippocampal trace with consolidation has often focused on questions about the permanence of memories in the hippocampus. Theories of systems consolidation have classically debated whether the hippocampal trace is time-limited (Alvarez and Squire 1994), or, rather, whether the hippocampus continues to represent memories in perpetuity (Nadel and Moscovitch 1997; Winocur and Moscovitch 2011; Moscovitch et al. 2016; Sekeres et al. 2018a). Another theory posits that while the original hippocampal trace is transient, during retrieval the hippocampus reconstructs details of an experience from cortical traces (Barry and Maguire 2019). Much research in this vein has focused on investigating changes in hippocampal blood-oxygenation level-dependent (BOLD) univariate activation with time (Bosshardt et al. 2005a,b; Takashima et al. 2006, 2009; Gais et al. 2007; Sterpenich et al. 2007, 2009; Yamashita et al. 2009; Milton et al. 2011; Watanabe et al. 2012; Ritchey et al. 2015; Baran et al. 2016; Dandolo and Schwabe 2018) and the effects of hippocampal lesions in animals and humans (Winocur et al. 2001; Frankland and Bontempi 2005; Winocur and Moscovitch 2011; Moscovitch et al. 2016) with mixed results. Interestingly, pinpointing these effects along the long axis of the hippocampus has also proven unclear. Some reports have found that only the anterior hippocampus exhibits time-dependent changes in retrieval-related univariate activation, with evidence of decreases with delay (Takashima et al. 2006; Milton et al. 2011; Dandolo and Schwabe 2018), but also evidence of greater activation for more remote, compared with recent, memories (Bosshardt et al. 2005a,b). At the same time, other studies have found decreases in univariate activation only in the posterior hippocampus (Bosshardt et al. 2005b; Takashima et al. 2009; Yamashita et al. 2009; Milton et al. 2011; Watanabe et al. 2012; Ritchey et al. 2015; Sekeres et al. 2018b).Because of these conflicting findings, instead of asking just about dependence or overall changes in activation in the hippocampus, theories and empirical research have instead increasingly considered the organization of memory representations in the hippocampus (Robin and Moscovitch 2017; Sekeres et al. 2018a). Broadly, using representational similarity analyses, several studies have shown that hippocampal memory representations tend to become differentiated over learning, particularly for memories with overlapping content (LaRocque et al. 2013; Schlichting et al. 2015; Chanales et al. 2017; Brunec et al. 2020). Furthermore, it has been suggested that information is represented at different scales or “granularity” along the long axis of the hippocampus, in line with place field size differences (Kjelstrup et al. 2008; Komorowski et al. 2013), with anterior hippocampus representing more similar, coarse-grained, or gist-like information, while the posterior hippocampus represents fine-grained, detail-oriented representations (Evensmoen et al. 2013; Poppenk et al. 2013; Robin and Moscovitch 2017; Brunec et al. 2018, 2020). However, limited work has investigated whether this representational organization is altered with consolidation. Reports have shown that memory representations sharing overlapping content become more similar over a delay (Tompary and Davachi 2017; Audrain and McAndrews 2020), yet other work has found that hippocampal representations were not modulated by time (Ritchey et al. 2015; Ezzyat et al. 2018). Intriguingly, reports indicating greater differentiation in memories in anterior compared with posterior hippocampus with consolidation (Tompary and Davachi 2017; Dandolo and Schwabe 2018; Ezzyat et al. 2018) raise the possibility that the representational granularity along the anteroposterior axis may be transformed with consolidation. Thus, more work is needed to understand how consolidation influences the representational structure of memories in the hippocampus. In particular, despite much research connecting sleep to consolidation (Diekelmann and Born 2010; Rasch and Born 2013), it remains unknown whether sleep-dependent processes facilitate such delay-dependent transformations to the hippocampus.Active processes in the sleeping brain seem to be optimized for systems consolidation. Currently, the best mechanistic evidence for sleep-dependent consolidation comes from studies on hippocampal replay showing the repeated reactivation of encoding-related patterns of hippocampal activity (Buzsáki 1989; Wilson and McNaughton 1994; Girardeau and Zugaro 2011), which seems to be coordinated with replay in areas of the cortex (Ji and Wilson 2007; Peyrache et al. 2009; Wierzynski et al. 2009). It is thought that the coupling between oscillations during non-REM sleep stages (particularly slow wave sleep [SWS])—including sharp wave ripples that support replay, thalamocortical spindles, and slow oscillations—facilitates the hippocampal–cortical dialogue and information transfer to the cortex (Buzsáki 1996; Sirota et al. 2003; Steriade 2006; Clemens et al. 2011; Mölle and Born 2011; Staresina et al. 2015). Indeed, our previously published work from the present study provided supporting evidence that the density of thalamocortical sleep spindles (11–16 Hz) during overnight sleep is related to enhanced hippocampal–cortical functional connectivity measures, and increased similarity, or greater representational overlap, among memories in the ventromedial prefrontal cortex (vmPFC) (Cowan et al. 2020). Yet, while some prior work has shown that features of sleep, including spindle density and the duration of non-REM SWS, are related to decreased retrieval-related hippocampal activation for memoranda learned prior to sleep (Takashima et al. 2006; Baran et al. 2016; Hennies et al. 2016), it remains unclear how the reactivation of hippocampal traces during replay may impact the way memories are organized along the long axis of the hippocampus.To examine the effects of sleep-dependent consolidation on the neural representation of memories in the hippocampus, we designed a within-participant 3-d study using overnight polysomnography (PSG), functional magnetic resonance imaging (fMRI), and behavioral measures of memory (Fig. 1). In this study, aspects of which have been previously published (Cowan et al. 2020), participants first studied a list of word–image pairs before sleeping overnight (Sleep List), during which PSG was recorded. Upon waking in the morning, participants studied a new list of pairs (Morning List). The word–image pairs from these two lists were then restudied while undergoing an fMRI scan, intermixed with a third, novel list of pairs (Single Study List). Associative memory was tested immediately after the scan and again 24 h later. We compared measures of multivariate pattern similarity and univariate BOLD signal for the lists learned prior to, or after, sleep to probe how modulating the opportunity for sleep-dependent consolidation impacts the way memories are organized across the long axis of the hippocampus. Furthermore, our design allowed us to examine how features of overnight sleep are related to the representational organization of memories learned prior to the sleep period, as well as the behavioral benefit of changes to the organization of these memories. Thus, our study provides a novel examination of the effects of sleep-dependent consolidation on the representation of memories along the long axis of the hippocampus.Open in a separate windowFigure 1.Study design. For all encoding and restudy sessions, participants were asked to form an association between a word and an image. Participants first encoded the Sleep List (blue) before sleeping overnight while polysomnography was recorded. The next morning (day 2), participants encoded a second set of novel word–image pairs (Morning List). After a short delay (∼2 h), participants restudied these two sets of pairs, intermixed with novel pairs (Single Study List) in the functional magnetic resonance imaging (fMRI) scanner. Source memory was tested immediately after the scan and after a 24-h delay (day 3).     

Fichte's striving subject

In this paper I argue that Fichte's attempt to reconcile the dualism of concept and intuition requires the overcoming of any idea of a thing‐in‐itself. At the same time he preserves the idea of an external constraint on the I's self‐positing. This central role for the realist constraint of the check conflicts with recent interpretations of Fichte that see his project as advocating the exclusivity of the space of reasons. The striving subject confronts and unifies the opposition between the realistic and idealistic elements in the Wissenschaftslehre. It is argued that as striving, reason's drive for self‐determination is a process of self‐transformation, as consciousness confronts the limitations of its inadequate explanations of the objects of experience.     

Reasons and practical possibility

According to a common thesis about normative reasons for action, you have a reason to perform a given action only if you can act for that reason. This thesis has long had broad appeal and is intended to capture the practical character of practical reasons. I’ll call it the ‘Practicality Thesis’. Recently, however, various writers have developed subtly different objections to it, each designed to show that there can be actions you have a reason to perform even though you could not act for that reason – because, were you aware of the reason-giving facts in the ways needed to act for the reason, it would no longer be a reason for you to so act. This article defends the Practicality Thesis against such objections. It considers some extant defences but shows that these are inadequate. It then advances an alternative approach intended to counter any structurally similar objection.     

Election predictions: Reply

Contrary to Aubert's claim, my paper on election predictions does not seek to draw empirical conclusions from mathematical premisses alone. The empirical premiss, approximated by the continuity assumption, is that sufficiently small changes in the predicted vote will cause only small changes in the actual vote. The technical criticisms by Øfsti and Østerberg of the reaction function are answered by specifying the function's domain. Other criticisms are also answered, and the reply concludes by placing the election prediction theorem in the context of other theorizing about human expectations and outguessing phenomena.     

Monaural Loudness Adaptation for Middle-Intensity Middle-Frequency Signals: The Importance of Measurement Technique

Using the Simple Adaptation technique (SA) and the Ipsilateral Comparison Paradigm (ICP), the authors studied monaural loudness adaptation to a middle-intensity [60 dB(A)] tone at signal frequencies of 250, 1000, and 4000 Hz in the left and right ears. Adaptation effects were absent when the SA procedure was used. However, they were observed uniformly across all frequency values with the ICP, a result that challenges the assertion in the literature, on the basis of SA measures, that loudness adaptation for middle-intensity signals occurs only at frequencies above 4000 Hz. The ICP features periodic intensity modulations (±10 dB relative to the base signal) to accommodate listeners' needs for referents by which they can gauge subtle changes in the loudness of the adapting tone, a key component that is missing in the SA method. Adaptation effects in this investigation were similar in both ears, supporting the equal susceptibility assumption common in loudness adaptation studies.     

Specificity of Task Constraints and Effects of Visual Demonstrations and Verbal Instructions in Directing Learners' Search During Skill Acquisition

In the present study, the efficacy of visual demonstrations and verbal instructions as instructional constraints on the acquisition of movement coordination was investigated. Fifteen participants performed an aiming task on 100 acquisition and 20 retention trials, under 1 of 3 conditions: a modeling group (MG), a verbally directed group (VDG), and a control group (CG). The MG observed a model intermittently throughout acquisition, whereas the VDG was verbally instructed to use the model's movement pattern. Participants in the CG received neither form of instruction. Kinematic analysis revealed that compared with verbal instructions or no instructions, visual demonstrations significantly improved participants' approximation of the model's coordination pattern. No differences were found in movement outcomes. Coordination data supported the visual perception perspective on observational learning, whereas outcome data suggested that the modeling effect is mainly a function of task constraints, that is, the novelty of a movement pattern.     

Latent self-oriented and socially prescribed perfectionism: Tests of factorial,convergent, and discriminant validity using self- and informant reports

Using latent variables for self-oriented perfectionism (i.e., demanding perfection of oneself) and socially prescribed perfectionism (i.e., perceiving others are demanding perfection of oneself) has advantages. However, few studies have specifically examined the psychometric properties of these latent variables. The present study addresses this shortcoming by testing the factorial, convergent, and discriminant validity of self-oriented and socially prescribed perfectionism latent variables using self- and informant reports. It was hypothesized that (a) the factor structure of the self-oriented and socially prescribed perfectionism latent variables would be supported, (b) self- and informant reports would moderately converge, and (c) the self-oriented and socially prescribed perfectionism latent variables would be discriminable. A sample of 242 undergraduate women and 378 informants (218 mothers, 160 fathers) was recruited. Undergraduate women completed self-reports, and mothers and fathers completed informant reports. Results were consistent with hypotheses, thereby supporting (a) the factorial validity of the self-oriented and socially prescribed latent variables using self- and informant reports, (b) the convergent validity of the self-oriented and socially prescribed latent variables via moderately to strongly correlated self- and informant reports, and (c) the discriminant validity of the self-oriented and socially prescribed perfectionism latent variables. Findings suggest research using these latent variables is warranted.     

Taking a Turn Toward the Masculine: The Impact of Mortality Salience on Implicit Leadership Theories

The present research investigates the influence of subtle death-related thoughts (i.e., mortality salience) on people's images of effective leaders (i.e., their implicit leadership theories [ILTs]). We test the prediction that mortality salience will change the content of these implicit theories to be more gender stereotypical such that individuals will conceive of effective leaders in a significantly more masculine, or agentic, manner. To test this prediction, we assessed the communal and agentic components of participants' ILTs after they were presented with a mortality salience or control manipulation. Results show that priming individuals to think about their mortality with two open-ended questions resulted in a significant shift in their ILTs such that an effective leader is described in significantly more agentic terms compared to the control condition. This masculine shift in people's ILTs was demonstrated in both women and men, and mortality salience did not influence perceptions of effective leaders' communal traits. This work contributes to research on gender bias in leadership, ILTs, and terror management theory and has implications for female leaders.     

The Effects of Warnings and Speeding on Scale Scores and Convergent Validity of Conscientiousness

We examined the effects of warnings and speeding on scale scores and convergent validity of a measure of Conscientiousness in a faking context (N = 329). A completely crossed 2 × 2 experimental design was used in which instructions (no warning or a warning) and speeding (with or without a time limit) were manipulated. No statistically significant effects on scale scores or convergent validity were evidenced for speeding. Warning participants did decrease Conscientiousness scores by almost 1 standard deviation (d = .91). Warnings also moderated the relationship between self- and observer-ratings of Conscientiousness such that the relationship between self- and observer-ratings was statistically significant and positive in the warned conditions (r _partial = .29, p < .01, n = 136), but this same relationship was statistically nonsignificant (r _partial = ?.14, p > .05, n = 148) in the unwarned conditions.