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Autobiographical Memory Transformation Across Consolidation The Journal of Neuroscience, March 27, 2013 • 33(13):5435–5436 • 5435 Journal Club Editor’s Note: These short reviews of a recent paper in the Journal, written exclusively by graduate students or postdoctoral fellows, are intended to mimic the journal clubs that exist in your own departments or institutions. For more information on the format and purpose of the Journal Club, please see http://www.jneurosci.org/misc/ifa_features.shtml. Autobiographical Memory Transformation across Consolidation Ruud M. W. J. Berkers1,2 and Marlieke T. R. van Kesteren1,3 1Donders Institute for Brain, Cognition, and Behaviour, Radboud University Nijmegen, 6500 HB Nijmegen, The Netherlands, and Departments of 2Cognitive Neuroscience and 3Anatomy, Radboud University Nijmegen Medical Centre, 6500 HB Nijmegen, The Netherlands Review of Bonnici et al. One of the greatest mysteries of the brain the MTL, and specifically the hippocampus, separable systems using complementary is how it can recall particular events viv- is important for the initial encoding of epi- mechanisms to learn and store informa- idly, sometimes after many decades, sodic memories, but memories later be- tion (McClelland et al., 1995). The hip- whereas the bulk of day-to-day experi- come embedded in a neocortical network pocampus is thought to be a fast-learning ences are forgotten. This question has (Squire and Alvarez, 1995) and therefore are system that rapidly stores complete sparked a vigorous search for the memory not affected by MTL lesions. events and reinstates patterns of activity engram, i.e., the mnemonic representa- The dawn of functional magnetic res- in the neocortex when events are re- tions in the brain. Seminal work by Lash- onance imaging (fMRI) provided neuro- experienced, whereas the slow-learning ley (1950) suggested that the engram is scientists with a tool that appeared ideally neocortical system gradually stores accu- widely distributed in the brain. His exper- equipped to investigate the engram mulating statistical regularities and gener- iments demonstrated that progressive throughout the brain. Overall, fMRI stud- alities across ensembles of experiences. lesions of a rat’s brain induced similar ies examining remote autobiographical The main nodes in this slower-learning progressive memory impairment on spatial memory retrieval demonstrated activity neocortical network have been shown to memory tasks, but the exact location of tis- in a widely distributed brain network, include the ventromedial prefrontal cor- sue resection did not determine the extent of including the hippocampus (Maguire, tex (vmPFC), the retrosplenial cortex the deficit. Later, specific memory impair- 2001). This unexpected hippocampal ac- (RSC), and temporal cortex (Maguire, ments were reported in patients with se- tivation during remote retrieval of vivid, 2001). lective lesions of the hippocampus, a episodically detailed memories led to the Recent research has started to investi- substructure of the medial temporal lobe hypothesis that hippocampal activation gate the functional integration of large- (MTL). Famous studies of patients with might not necessarily be related to the age scale networks during memory encoding, such lesions, such as H.M. (Scoville and Mil- of the memory but rather to its vividness consolidation, and retrieval. A study by ner, 1957), typically report both antero- (Gilboa et al., 2004; Winocur et al., 2010). Bonnici et al. (2012) adds to these devel- grade (deficits in forming new memories) This account suggests that the hippocam- opments by using multivariate pattern and retrograde (deficits in retrieving old pus has a persistent role in vivid retrieval analysis (MVPA) techniques to decode memories) amnesia specifically for episodic of episodic autobiographical memories, distributed representations of both recent memories. Nonetheless, retrograde amnesia whereas semantic, abstract concepts can and remote autobiographical memories was often found to be graded: events that also be retrieved independently from the within previously defined regions of inter- occurred years before the injury were still MTL after consolidation. If this hypothe- ests. Specifically, decoding was used to remembered, whereas more recent events sis is correct, the finding that features of establish whether certain brain regions were forgotten. Hence, it was proposed that remote memories are preserved after contained memory-specific information MTL damage can be explained if one as- during retrieval of remote and recent Received Jan. 18, 2013; revised Feb. 14, 2013; accepted Feb. 20, 2013. sumes that autobiographical memories events. Individual accounts of life events Correspondence should be addressed to Ruud Berkers, Donders Insti- can be transformed over time, thereby los- were obtained through an autobiographi- tute for Brain, Cognition, and Behaviour, Radboud University Nijmegen, ing episodic detail and becoming abstract cal memory interview, of which three re- P.O. Box 9101, 6500 HB Nijmegen, The Netherlands. E-mail: [email protected]. and semanticized (Winocur et al., 2010). mote (10 years old) and three recent (2–3 DOI:10.1523/JNEUROSCI.0246-13.2013 Computational models conceptualize weeks old) events were selected and Copyright©2013theauthors 0270-6474/13/335435-02$15.00/0 the hippocampus and the neocortex as matched in terms of rated vividness. 5436 • J. Neurosci., March 27, 2013 • 33(13):5435–5436 Berkers and van Kesteren • Autobiographical Memory Transformation While in the scanner, participants were added episodic features together with few formation. A specific memory, in particu- cued to vividly recall these events. Subse- remaining original event-specific details lar an autobiographical one, is suggested quently, a multivariate classifier was and previously extracted overlapping (se- to consist of multiple features that are rep- trained to distinguish multivoxel activity mantic) features of earlier events. Statistical resented by different brain regions at the patterns corresponding to either remote regularities and generalities can in turn be- same time. Along with functional connec- or recent autobiographical memories. come contextualized into a more abstract tivity analyses, MVPA methods can be The rationale here is that, if the retrieved (self-referential) schema, mediated by the used to determine what information these memory can be decoded based on activity neocortex in general and the vmPFC in par- different regions contribute and how they patterns measured in a certain brain re- ticular (van Kesteren et al., 2012). In con- are functionally integrated. A next step gion, then that region must represent in- trast, recent memories are more likely to would be to analyze feature dimensions of formation for at least one of the remote or retain a more specific episodic quality, with autobiographical memories, such as regu- recent memories. The results show that fewer semantic features extracted from re- larity across ensembles of events and viv- representations of both recent and remote peated experiences. idness, and how events differing in terms autobiographical memories were widely In the study by Bonnici et al. (2012), of these dimensions can be decoded from distributed and could be decoded with remote memory traces might have been different brain regions. This approach above-chance accuracy from activity pat- influenced by the vivid retrieval during might ultimately disentangle and decon- terns in the MTL, vmPFC, RSC, and tem- the pre-study interview 1 week earlier, and struct the computational roles of multiple poral cortex, respectively. Differences this may have biased the reconstruction of brain regions in autobiographical mem- between remote and recent memories the event on the basis of stored specific ory retrieval. were observed in the vmPFC, in which de- episodic and general semantic features. coding accuracy was significantly better Therefore, although the authors were References Bonnici HM, Chadwick MJ, Lutti A, Hassabis D, for remote than for recent events. These careful to control for vividness, it is prob- Weiskopf N, Maguire EA (2012) Detecting results support a distributed model in able that, when participants were in- representations of recent and remote autobi- which functional integration across hip- structed to retrieve the event as vividly as ographical memories in vmPFC and hip- pocampus and neocortex underlie autobi- possible, they might have (perhaps even pocampus. J Neurosci 32:16982–16991. ographical memory retrieval. However, unconsciously) inserted new details or re- CrossRef Medline these results raise important questions for constructed an experience based on se- Gilboa A, Winocur G, Grady CL, Hevenor SJ, Moscovitch M (2004) Remembering our additional study, because it is unclear why mantic features extracted from multiple past: functional neuroanatomy of recollection remote and recent memories can be de- previous events. For example, a person of recent and very remote personal events. coded equally well from the hippocampus who was at many parties you attended, Cereb Cortex 14:1214–1225. CrossRef and why remote memories can be de- but was not at this particular party, may be Medline coded better from the vmPFC. inserted into the memory. Lashley KS (1950) In search of the engram. In: Possible explanations for the findings The central point here is that, by using Symposium of the society for experimental bi- ology, Vol 4. New York: Cambridge UP. of Bonnici et al. (2012) can emerge by this particular procedure, previously se-
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