Recollection and familiarity: Examining controversial assumptions and new directions

“…and Brown, 2006; Diana et al., 2007), and have led to the development of a number of dual-process theories of recognition (e.g., Atkinson and Juola, 1974; Mandler, 1980; Jacoby, 1991; Yonelinas, 1994; Aggleton and Brown, 1999; Reder et al., 2000; Kelley and Wixted, 2001; Rotello et al., 2004).”

“For example, in a review of remember/know, source memory and ROC studies that included recollection and familiarity contrasts, 16 of 19 studies showed that the hippocampus was involved in recollection, whereas only two showed hippocampal involvement in familiarity (Eichenbaum et al., 2007).”

“…1998; Kopelman et al., 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests of item recognition (e.g., Milner, 1971; Milner et al., 1985; Janowsky et al., 1989; Shimamura et al., 1990; Johnson et al., 1997; but see Swick et al., 2006).”

“This idea is supported by evidence that PFC lesions disrupt recall more than recognition (for reviews, see Wheeler et al., 1995, 1997; but see Kopelman and Stanhope, 1998; Kopelman et al., 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests…”

“In studies of patients with relatively selective hippocampal damage, recall is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al.,…”

“Evidence from a broad range of paradigms has provided support for these claims (for earlier reviews, see Aggleton and Brown, 1999, 2001; Yonelinas, 2002; Eichenbaum et al., 2007).”

“The signal detection and threshold assumptions are consistent with recent neurocomputational models of recognition (e.g., Norman and O’Reilly, 2003; Elfman et al., 2008 also see Murdock, 1974).”

“In collaboration with Howard Eichenbaum, Charan Ranganath, and Rachel Diana, we have taken preliminary steps toward integrating these results with the DPSD model (Eichenbaum et al., 2007; Diana et al., 2007).”

“However, the neuroimaging literature suggests that the PFC is involved in both recollection and familiarity during encoding and retrieval (Henson et al., 2000; Dobbins et al., 2004; Yonelinas et al., 2005; Blumenfeld and Ranganath, 2007).”

“In addition to perception and short-term memory, the MTL also appears to play a role in the processing and detection of novelty (for reviews, see Ranganath and Rainer, 2003; Nyberg, 2005; Kumaran and Maguire, 2007, 2009).”

“, 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests of item recognition (e.g., Milner, 1971; Milner et al., 1985; Janowsky et al., 1989; Shimamura et al., 1990; Johnson et al., 1997; but see Swick et al., 2006).”

“For example, Mayes et al. (2007) have argued that individuals are able to use familiarity to link aspects of an event that come from the same processing domain (e.g., a face–face pair), whereas across-domain associations (e.g., a face–name pair) require recollection.”

“However, the neuroimaging literature suggests that the PFC is involved in both recollection and familiarity during encoding and retrieval (Henson et al., 2000; Dobbins et al., 2004; Yonelinas et al., 2005; Blumenfeld and Ranganath, 2007).”

“The latter finding is interesting in light of prior work suggesting that the PRc and PHc seem to be particularly important for processing of object and spatial information, respectively (Bar and Aminoff, 2003; Pihlajamaki et al., 2004; Buffalo et al., 2006; Litman et al., 2009).”

“VC 2010 WILEY-LISS, INC. model, which is often treated as the classical single-process model (e.g., Egan, 1958; Donaldson, 1996; Dunn, 2004), does not provide an adequate account for results from associative or source memory tests (e.g., Kelley and Wixted, 2001; Decarlo, 2002; Rotello et al., 2004;…”

“Second, the PHc is often associated with recollection, even in cases in which there are little or no spatial retrieval demands (e.g., Ranganath et al., 2003; Dolcos et al., 2005; Fenker et al., 2005; Yonelinas et al., 2005).”

“The finding that MTL amnesia leads to symmetrical item recognition ROCs has now been replicated numerous times (e.g., Yonelinas et al., 2002; Aggleton et al., 2005; Cipolotti et al., 2006; Wais et al., 2006; Kirwan et al., 2010).”

“A large number of such studies have now been published and they provide converging evidence that the hippocampus is involved primarily in recollection (for reviews, see Eichenbaum et al., 2007; Skinner and Fernandes, 2007; Wais, 2008).”

“This pattern of results has now been replicated in well over 50 published experiments (for a review, see Yonelinas and Parks, 2007; for similar results in studies of rats see Sauvage et al., 2008).”

“Hippocampus (e.g., Manns et al., 2003, Stark and Squire, 2003; Wais et al., 2006; Gold et al., 2006b, Kirwan et al., 2010).”

“ROCs have been examined in odor recognition studies in rats, and have indicated that selective hippocampal lesions impair recollection but spare familiarity-based recognition (Fortin et al., 2004).”

“This prediction was initially tested by examining recognition ROCs in patients with amnesia with extensive medial temporal lobe (MTL) damage (Yonelinas et al., 1998).”

“…found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed and Squire, 1997; Manns and Squire, 1999; Kopelman et al., 2007).”

“…that hippocampal damage is associated with deficits in perceptual or short-term memory tasks involving complex spatial scenes or relational stimuli (Lee et al., 2005a,b; Hannula et al., 2006; Olson et al., 2006; Piekema et al., 2006; for reviews, see Bussey and Saksida, 2007; Graham et al., 2010).”

“…that hippocampal damage is associated with deficits in perceptual or short-term memory tasks involving complex spatial scenes or relational stimuli (Lee et al., 2005a,b; Hannula et al., 2006; Olson et al., 2006; Piekema et al., 2006; for reviews, see Bussey and Saksida, 2007; Graham et al., 2010).”

“…that hippocampal damage is associated with deficits in perceptual or short-term memory tasks involving complex spatial scenes or relational stimuli (Lee et al., 2005a,b; Hannula et al., 2006; Olson et al., 2006; Piekema et al., 2006; for reviews, see Bussey and Saksida, 2007; Graham et al., 2010).”

“This idea is supported by evidence that PFC lesions disrupt recall more than recognition (for reviews, see Wheeler et al., 1995, 1997; but see Kopelman and Stanhope, 1998; Kopelman et al., 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests…”

“In fact, there is now evidence that the PRc is important in complex visual discrimination tasks that make little explicit demands on long-term memory (Bussey et al., 2002; Lee et al., 2005a,b, 2007; for a review, see Bussey and Saksida, 2007).”

“However, the neuroimaging literature suggests that the PFC is involved in both recollection and familiarity during encoding and retrieval (Henson et al., 2000; Dobbins et al., 2004; Yonelinas et al., 2005; Blumenfeld and Ranganath, 2007).”

“6B; Bowles et al., 2007).”

“In fact, there is now evidence that the PRc is important in complex visual discrimination tasks that make little explicit demands on long-term memory (Bussey et al., 2002; Lee et al., 2005a,b, 2007; for a review, see Bussey and Saksida, 2007).”

“In fact, there is now evidence that the PRc is important in complex visual discrimination tasks that make little explicit demands on long-term memory (Bussey et al., 2002; Lee et al., 2005a,b, 2007; for a review, see Bussey and Saksida, 2007).”

“, 2008), and hippocampal damage selectively disrupts recollection (Aggleton et al., 2005).”

“…in studies of source recognition (Diana et al., 2008, 2010), associative recognition of word pairs (e.g., Quamme et al., 2007; Haskins et al., 2008; for related ERP results see Rhodes and Donaldson, 2007, 2008), and associative recognition for intact and rearranged faces (Yonelinas et al., 1999).”

“Familiarity can contribute to accurate associative and source recognition judgments when pairs of items, or items and sources, are ‘‘unitized’’ (i.e., treated as single items rather than as arbitrary pairings; Yonelinas et al., 1999; Yonelinas, 1999; Quamme et al., 2007).”

“The relationship between novelty detection and recognition memory processes is not yet fully understood, but one possibility is that the efficacy of encoding is directly related to the novelty of the study item (Tulving et al., 1994; Tulving and Kroll, 1995).”

“…and Brown, 2006; Diana et al., 2007), and have led to the development of a number of dual-process theories of recognition (e.g., Atkinson and Juola, 1974; Mandler, 1980; Jacoby, 1991; Yonelinas, 1994; Aggleton and Brown, 1999; Reder et al., 2000; Kelley and Wixted, 2001; Rotello et al., 2004).”

“However, in a recent fMRI study (Haskins et al., 2008), word pairs encoded in a unitized fashion were associated with increased activity in the PRc, and activation in this region during encoding predicted subsequent familiarity-based associative recognition responses (for similar results at…”

“…whereas damage that includes both the hippocampus and the surrounding MTL leads to deficits in both recollection and familiarity (e.g., Verfaellie and Treadwell, 1993; Knowlton and Squire, 1995; Blaxton and Theodore, 1997; Schacter et al., 1996, Schacter et al., 1997; Yonelinas et al., 1997).”

“Hippocampus (e.g., Manns et al., 2003, Stark and Squire, 2003; Wais et al., 2006; Gold et al., 2006b, Kirwan et al., 2010).”

“…exhibit more pronounced deficits on associative and source memory tests than on item recognition tests (e.g., Gold et al., 2006a, experiment 1; Holdstock et al., 2005, Mayes et al., 2002; Turriziani et al., 2004; but see Gold et al., 2006a, experiment 2; Stark et al., 2002; Gold et al., 2006b).”

“In studies of patients with relatively selective hippocampal damage, recall is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed a…”

“In studies of patients with relatively selective hippocampal damage, recall is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed a…”

“…or the fact that items that are associated with correct source recollection can lead to different levels of recognition confidence, have sometimes been interpreted as evidence against threshold-based models (Slotnick et al., 2000; Slotnick and Dodson, 2005; Wixted, 2007; Mickes et al., 2009).”

“…that hippocampal damage is associated with deficits in perceptual or short-term memory tasks involving complex spatial scenes or relational stimuli (Lee et al., 2005a,b; Hannula et al., 2006; Olson et al., 2006; Piekema et al., 2006; for reviews, see Bussey and Saksida, 2007; Graham et al., 2010).”

“(A) Source recognition ROCs for items encoded under high and low unitization conditions (Diana et al., 2008).”

“…or the fact that items that are associated with correct source recollection can lead to different levels of recognition confidence, have sometimes been interpreted as evidence against threshold-based models (Slotnick et al., 2000; Slotnick and Dodson, 2005; Wixted, 2007; Mickes et al., 2009).”

“In addition to perception and short-term memory, the MTL also appears to play a role in the processing and detection of novelty (for reviews, see Ranganath and Rainer, 2003; Nyberg, 2005; Kumaran and Maguire, 2007, 2009).”

“, 2007), and it can play a role in associative and source memory under conditions that promote unitization of item–item or item–source information (Staresina and Davachi, 2006; Haskins et al., 2008; Diana et al., 2010).”

“, 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests of item recognition (e.g., Milner, 1971; Milner et al., 1985; Janowsky et al., 1989; Shimamura et al., 1990; Johnson et al., 1997; but see Swick et al., 2006).”

“Several studies examining patients with fornix lesions have indicated that these patients exhibit selective deficits in recollection (e.g., Carlesimo et al., 2007; Vann et al., 2009).”

“…whereas damage that includes both the hippocampus and the surrounding MTL leads to deficits in both recollection and familiarity (e.g., Verfaellie and Treadwell, 1993; Knowlton and Squire, 1995; Blaxton and Theodore, 1997; Schacter et al., 1996, Schacter et al., 1997; Yonelinas et al., 1997).”

“…exhibit more pronounced deficits on associative and source memory tests than on item recognition tests (e.g., Gold et al., 2006a, experiment 1; Holdstock et al., 2005, Mayes et al., 2002; Turriziani et al., 2004; but see Gold et al., 2006a, experiment 2; Stark et al., 2002; Gold et al.,…”

“For example, Sherman et al. (2003) extended the DPSD model by proposing a ‘‘variable recollection dual-process model’’ in which recollection is assumed to be a threshold process, but the items that are above the recollective threshold form a Gaussian distribution (for related ideas see Macho, 2002;…”

“Hippocampus (e.g., Manns et al., 2003, Stark and Squire, 2003; Wais et al., 2006; Gold et al., 2006b, Kirwan et al., 2010).”

“The latter finding is interesting in light of prior work suggesting that the PRc and PHc seem to be particularly important for processing of object and spatial information, respectively (Bar and Aminoff, 2003; Pihlajamaki et al., 2004; Buffalo et al., 2006; Litman et al., 2009).”

“…example, studies using a variety of process estimation methods, such as the remember/ know, ROC, and source recognition paradigms have indicated that PFC damage can impair both recollection and familiarity (Duarte et al., 2005; Farovik et al., 2008; MacPherson et al., 2008; Kishiyama et al., 2009).”

“…severe deficits in their ability to recollect details about prior events, but often show remarkably preserved familiarity-based discrimination (Huppert and Piercy, 1978), the DPSD model assumes that the hippocampus is particularly important in forming and retrieving the arbitrary associations…”

“…al., 2009), whereas damage that includes both the hippocampus and the surrounding MTL leads to deficits in both recollection and familiarity (e.g., Verfaellie and Treadwell, 1993; Knowlton and Squire, 1995; Blaxton and Theodore, 1997; Schacter et al., 1996, Schacter et al., 1997; Yonelinas et…”

“…exhibit more pronounced deficits on associative and source memory tests than on item recognition tests (e.g., Gold et al., 2006a, experiment 1; Holdstock et al., 2005, Mayes et al., 2002; Turriziani et al., 2004; but see Gold et al., 2006a, experiment 2; Stark et al., 2002; Gold et al., 2006b).”

“…is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed and Squire, 1997; Manns and Squire, 1999; Kopelman…”

“This has been verified in studies of associative recognition (Giovanello et al., 2006; Quamme et al., 2007) and source recognition (Diana et al.”

“Volumetric data linking hypoxia to hippocampal atrophy were not available in the previous study, but this was directly addressed in a similar study of healthy aging (Yonelinas et al., 2007), which showed that reductions in hippocampal volume were associated with declines in recollection, but not familiarity.”

“In fact, fornix damage appears to lead to selective recollection impairments in both anterograde and retrograde amnesia (Gilboa et al., 2006).”

“…in studies of source recognition (Diana et al., 2008, 2010), associative recognition of word pairs (e.g., Quamme et al., 2007; Haskins et al., 2008; for related ERP results see Rhodes and Donaldson, 2007, 2008), and associative recognition for intact and rearranged faces (Yonelinas et al., 1999).”

“A number of studies have suggested that the PFC might be particularly important for recollection (Janowsky et al., 1989; Wheeler et al., 1997; Davidson and Glisky, 2002; Wheeler and Stuss, 2003).”

“In addition to perception and short-term memory, the MTL also appears to play a role in the processing and detection of novelty (for reviews, see Ranganath and Rainer, 2003; Nyberg, 2005; Kumaran and Maguire, 2007, 2009).”

“Although this issue has yet to be systematically examined, evidence from several single case studies demonstrates that right hemisphere hippocampal damage is sufficient to impair both recollection and familiarity for scenes (Carlesimo et al., 2001, Cipolotti et al., 2006; Bird et al., 2007, 2008).”

“…ROC, and process dissociation procedures have now verified that selective hippocampal damage disrupts recollection but not familiarity (e.g., Bastin et al., 2004; Aggleton et al., 2005; Bird et al., 2008; Brandt et al., 2008; Peters et al., 2008; Turriziani et al., 2008b; Jager et al.,…”

“However, the neuroimaging literature suggests that the PFC is involved in both recollection and familiarity during encoding and retrieval (Henson et al., 2000; Dobbins et al., 2004; Yonelinas et al., 2005; Blumenfeld and Ranganath, 2007).”

“…example, studies using a variety of process estimation methods, such as the remember/ know, ROC, and source recognition paradigms have indicated that PFC damage can impair both recollection and familiarity (Duarte et al., 2005; Farovik et al., 2008; MacPherson et al., 2008; Kishiyama et al., 2009).”

“Second, the PHc is often associated with recollection, even in cases in which there are little or no spatial retrieval demands (e.g., Ranganath et al., 2003; Dolcos et al., 2005; Fenker et al., 2005; Yonelinas et al., 2005).”

“The latter finding is interesting in light of prior work suggesting that the PRc and PHc seem to be particularly important for processing of object and spatial information, respectively (Bar and Aminoff, 2003; Pihlajamaki et al., 2004; Buffalo et al., 2006; Litman et al., 2009).”

“…familiarity-based item recognition responses (for reviews, see Eichenbaum et al., 2007), and it can play a role in associative and source memory under conditions that promote unitization of item–item or item–source information (Staresina and Davachi, 2006; Haskins et al., 2008; Diana et al., 2010).”

“…recollection’’ (Yonelinas and Jacoby, 1996; Parks, 2007), and it can bias estimates of recollection and familiarity, and can play havoc with imaging or patient studies that use very restrictive measures of source memory to assess recollection and familiarity (e.g., Wais et al., 2010).”

“…1998; Kopelman et al., 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests of item recognition (e.g., Milner, 1971; Milner et al., 1985; Janowsky et al., 1989; Shimamura et al., 1990; Johnson et al., 1997; but see Swick et al., 2006).”

“For example, Yonelinas et al. (2002; also see Quamme et al., 2004) obtained recall and recognition data from a large group of mild hypoxic patients expected to have selective hippocampal damage, and used structural equation modeling to examine the effects of hypoxia on the latent variables…”

“This idea is supported by evidence that PFC lesions disrupt recall more than recognition (for reviews, see Wheeler et al., 1995, 1997; but see Kopelman and Stanhope, 1998; Kopelman et al., 2007), and damage to the PFC is associated with more pronounced deficits in tests of source memory than tests…”

“Behavioral evidence indicates that both recollection and familiarity are increased when items are processed or presented in a novel manner at study (Dobbins et al., 1998; Kishiyama and Yonelinas, 2003).”

“Several studies examining patients with fornix lesions have indicated that these patients exhibit selective deficits in recollection (e.g., Carlesimo et al., 2007; Vann et al., 2009).”

“Although this issue has yet to be systematically examined, evidence from several single case studies demonstrates that right hemisphere hippocampal damage is sufficient to impair both recollection and familiarity for scenes (Carlesimo et al., 2001, Cipolotti et al., 2006; Bird et al., 2007, 2008).”

“For example, Sherman et al. (2003) extended the DPSD model by proposing a ‘‘variable recollection dual-process model’’ in which recollection is assumed to be a threshold process, but the items that are above the recollective threshold form a Gaussian distribution (for related ideas see Macho, 2002;…”

“As illustrated in Figure 6B, additional studies using process estimation methods such as remember/know, ROC, and process dissociation procedures have now verified that selective hippocampal damage disrupts recollection but not familiarity (e.g., Bastin et al., 2004; Aggleton et al., 2005; Bird et al., 2008; Brandt et…”

“…1996; Dunn, 2004), does not provide an adequate account for results from associative or source memory tests (e.g., Kelley and Wixted, 2001; Decarlo, 2002; Rotello et al., 2004; for a review see Yonelinas and Parks, 2007), although it can account for results from some item recognition tests.”

“Hippocampus (e.g., Manns et al., 2003, Stark and Squire, 2003; Wais et al., 2006; Gold et al., 2006b, Kirwan et al., 2010).”

“Moreover, the PRc has been shown to be sensitive to stimulus repetition in semantic judgment tasks (O’Kane et al., 2005; Voss et al., 2009), and it appears to be necessary for conceptual implicit memory (Wang et al., in press).”

“In studies of patients with relatively selective hippocampal damage, recall is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed a…”

“…made highly similar (e.g., a study list consisting of 200 photos of very similar suburban houses), source recognition ROCs (i.e., was the picture initially on the left or right side of the screen?) become much more curved than is typically the case in tests of source memory (Elfman et al., 2008).”

“In the absence of histological evidence, however, it is possible that the familiarity deficits observed in this group were due to damage outside the hippocampus (Yonelinas et al., 2004).”

“…example, studies using a variety of process estimation methods, such as the remember/ know, ROC, and source recognition paradigms have indicated that PFC damage can impair both recollection and familiarity (Duarte et al., 2005; Farovik et al., 2008; MacPherson et al., 2008; Kishiyama et al., 2009).”

“…have now verified that selective hippocampal damage disrupts recollection but not familiarity (e.g., Bastin et al., 2004; Aggleton et al., 2005; Bird et al., 2008; Brandt et al., 2008; Peters et al., 2008; Turriziani et al., 2008b; Jager et al., 2009), whereas damage that includes both the…”

“Hippocampus (e.g., Manns et al., 2003, Stark and Squire, 2003; Wais et al., 2006; Gold et al., 2006b, Kirwan et al., 2010).”

“Moreover, patients with lesions to the hippocampus do not benefit from stimulus novelty, as indexed using the von Restorff paradigm, in either recollection or familiarity (Kishiyama et al., 2004).”

“In studies of patients with relatively selective hippocampal damage, recall is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed a…”

“In addition, different types of recollection may be differentially diagnostic and so may support different levels of confidence (cf. Yonelinas, 1999).”

“…example, studies using a variety of process estimation methods, such as the remember/ know, ROC, and source recognition paradigms have indicated that PFC damage can impair both recollection and familiarity (Duarte et al., 2005; Farovik et al., 2008; MacPherson et al., 2008; Kishiyama et al., 2009).”

“Although procedures like free recall and remember/know tests do not easily lend themselves to animal studies, the ROC procedures can be readily adapted (for a review, see Eichenbaum et al., 2010).”

“Although this issue has yet to be systematically examined, evidence from several single case studies demonstrates that right hemisphere hippocampal damage is sufficient to impair both recollection and familiarity for scenes (Carlesimo et al., 2001, Cipolotti et al., 2006; Bird et al., 2007, 2008).”

“As illustrated in Figure 6B, additional studies using process estimation methods such as remember/know, ROC, and process dissociation procedures have now verified that selective hippocampal damage disrupts recollection but not familiarity (e.g., Bastin et al., 2004; Aggleton et al., 2005; Bird et al., 2008; Brandt et…”

“…recollection but not familiarity (e.g., Bastin et al., 2004; Aggleton et al., 2005; Bird et al., 2008; Brandt et al., 2008; Peters et al., 2008; Turriziani et al., 2008b; Jager et al., 2009), whereas damage that includes both the hippocampus and the surrounding MTL leads to deficits in both…”

“In studies of patients with relatively selective hippocampal damage, recall is often found to be disrupted to a greater extent than recognition (Vargha-Khadem et al., 1997; Holdstock et al., 2000; Baddeley et al., 2001; Mayes et al., 2001; Bastin et al., 2004; Holdstock et al., 2008; Adlam et al., 2009; but see Reed a…”

“…the cortical medial temporal lobe (MTL) regions appear to support episodic memory function in conjunction with the hippocampus, a number of memory models suggest dissociable representations and processes linked to these subregions [Norman, 2010; Eichenbaum et al., 2007; Yonelinas et al., 2010].”