Accepted Manuscript

Title: Dissociating Memory Traces and Scenario Construction

in Mental Time Travel
Author: Sen Cheng Markus Werning Thomas Suddendorf
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S0149-7634(15)30160-3
http://dx.doi.org/doi:10.1016/j.neubiorev.2015.11.011
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3-9-2015
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23-11-2015

Please cite this article as: Cheng, S., Werning, M., Suddendorf, T.,Dissociating
Memory Traces and Scenario Construction in Mental Time Travel, Neuroscience and
Biobehavioral Reviews (2015), http://dx.doi.org/10.1016/j.neubiorev.2015.11.011
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Title: Dissociating Memory Traces and Scenario Construction in Mental Time Travel
Authors and author addresses:
Prof. Dr. Sen Cheng
Mercator Research Group "Structure of
Memory"
Department of Psychology
Ruhr University Bochum
Universittsstr. 150
44801 Bochum
Germany
Office Phone: +49 (234) 32 27136
Fax:
+49 (234) 32 14463
Email: sen.cheng@rub.de

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Corresponding author: Sen Cheng

Prof. Dr. Thomas Suddendorf

Early Cognitive Development Centre
School of Psychology
University of Queensland
Brisbane, Queensland 4072
Australia
Office Phone: +61 7 3365 8341
Fax:
+61 7 3365 4466
Email: t.suddendorf@psy.uq.edu.au

Prof. Dr. Markus Werning

Department of Philosophy II
Mercator Research Group "Structure of
Memory"
Ruhr University Bochum
Universittsstr. 150
44801 Bochum
Germany
Office Phone: +49 (234) 32 24734
Fax:
+49 (234) 32 14463
Email: markus.werning@rub.de

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Dissociating Memory Traces and Scenario Construction in

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Mental Time Travel

Sen Chenga,b,*, Markus Werninga,c, and Thomas Suddendorf d

Mercator Research Group Structure of Memory, Ruhr-University Bochum, Bochum,

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Germany
Faculty of Psychology, Ruhr-University Bochum, Bochum, Germany

Department of Philosophy II, Ruhr-University Bochum, Bochum, Germany

Early Cognitive Development Centre, School of Psychology, University of Queensland,

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corresponding author: sen.cheng@rub.de

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Abstract

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St Lucia, Queensland, Australia

There has been a persistent debate about how to define episodic memory and whether it
is a uniquely human capacity. On the one hand, many animal cognition studies employ
content-based criteria, such as the what-where-when criterion, and argue that nonhuman
animals possess episodic memory. On the other hand, many human cognition studies
emphasize the subjective experience during retrieval as an essential property of episodic
memory and the distinctly human foresight it purportedly enables. We propose that both
perspectives may examine distinct but complementary aspects of episodic memory by
drawing a conceptual distinction between episodic memory traces and mental time
travel. Episodic memory traces are sequential mnemonic representations of particular,
personally experienced episodes. Mental time travel draws on these traces, but requires
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other components to construct scenarios and embed them into larger narratives. Various
nonhuman animals may store episodic memory traces, and yet it is possible that only
humans are able to construct and reflect on narratives of their lives and flexibly

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compare alternative scenarios of the remote future.

Keywords: mental time travel; episodic memory; scenario; episodic memory trace;

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animal cognition; episodic-like memory

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1 Introduction

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When Tulving (1972) first conceptualized episodic memory as memory of personallyexperienced events, he hypothesized that episodic memories could be distinguished on

the basis of the information that they contain (Fig. 1). He suggested that [e]pisodic
memory receives and stores information about temporally dated episodes or events, and

temporal-spatial relations among these events. (Tulving, 1972, p. 385) This criterion

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later came to be known as the what-where-when (Griffiths et al., 1999) or WWW

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criterion (Suddendorf and Busby, 2003). Tulving (1972) thought that, on this basis,
episodic memories could be distinguished from other memory systems such as semantic
or procedural memory. However, Tulving (1985) later realized that the WWW criterion
was neither necessary nor sufficient to define episodic memory. It is possible, for
instance, to know what happened where and when (e.g., such as details about the battle
of Hastings) without the events having been personally experienced and recalled. He
therefore changed his definition to one that emphasized the distinct subjective
experience during the retrieval of episodic memories. Tulving (1985) suggested that the
recall of the three different types of memory: procedural, semantic and episodic is
characterized by varying levels of conscious awareness: anoetic, noetic, and autonoetic,
respectively. Roughly speaking, anoetic consciousness lacks awareness of the content of
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the memory, noetic consciousness includes awareness of the content, and auto-noetic
consciousness includes the awareness that one self has been part of the recalled event.
The experience during retrieval of episodic memories was likened to mental time travel
(MTT) into the past (Suddendorf and Corballis, 1997; Tulving, 1985), a virtual reliving

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of the past (Fig. 1).

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Although it took some time to develop behavioral paradigms to test whether nonhuman
animals store memories of combined WWW information (Clayton and Dickinson,

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1998), once published, the what, where, when approach to studying episodic memory

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became very popular in animal cognition studies. The WWW criterion has been tested
in a variety of species and purportedly passed by scrub jays (Clayton and Dickinson,

1998; Dally et al., 2006); pigeons (Zentall et al., 2001); black-capped chickadee
(Feeney et al., 2009); magpies (Zinkivskay et al., 2009); mice (Dere et al., 2005); rats

(Babb and Crystal, 2005; Eacott et al., 2005); meadow voles (Ferkin et al., 2008);

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Yucatan minipigs (Kouwenberg et al., 2009); dogs (Kaminski et al., 2008); rhesus
monkey (Hoffman et al., 2009, but see Hampton et al., 2005); and chimpanzees and

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bonobos (Martin-Ordas et al. 2010, but see Dekleva et al. 2011).

Clayton and Dickinson (1998) introduced the term episodic-like memory to
acknowledge that memory of what, where, when might not be the same as human
episodic memory and, at the same time, to suggest that there might be a close link
between the two. While a number of authors maintain this cautious middle ground,
others swing more towards the extremes. On the one hand, some students of human
episodic memory insist that, no matter what information a nonhuman animal might be
able to retrieve about its past, without evidence for autonoetic consciousness we should
not attribute episodic memory to them (Klein, 2013; Tulving, 2001). On the other hand,
some researchers of memory in nonhuman animals argue that if a memory in nonhuman
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animals can be operationalized in a way similar to episodic memory in humans, then the
subjective experience is immaterial and nonhuman animals should be considered to
have episodic memory (Eichenbaum et al., 2005).

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Suddendorf and colleagues suggest a third perspective. The subjective experience, in

itself, is nothing that evolution can directly select for (Suddendorf and Busby, 2003;

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Suddendorf and Corballis, 2010, 2008, 2007, 1997). It is consequences for fitness that
matter, and these can only be found in how certain characteristics effect survival and

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reproduction in the present and the future. Given that mental time travel into the past

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and into the future appear to draw on many of the same neurocognitive mechanisms,
they may be regarded as two sides of the same coin (Dudai and Carruthers 2005; Byrne

et al. 2007; Schacter et al. 2007; Suddendorf & Corballis, 1997). Rather than being a
system designed to capture the past, Suddendorf and Corballis (e.g., 2007) suggest that

MTT evolved primarily to enable foresight. By considering potential future events, one

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can plan and prepare in ways that convey distinct fitness benefits. Species that are
capable of episodic memory should hence be able to demonstrate flexible foresight in

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their behavior, yet there is little indication that nonhuman animals do (Suddendorf and
Corballis 2010, see also Raby and Clayton 2009).
What complicates the debate further is that the neural mechanisms of episodic memory
are still not well understood. The hippocampus is the brain region most closely
associated with episodic memory in humans (Harand et al., 2012; Nadel et al., 2000;
Scoville and Milner, 1957; Zola-Morgan et al., 1986). In rodents (OKeefe and
Dostrovsky, 1971) and bats (Ulanovsky and Moss, 2007), numerous studies have found
neurons in the hippocampus that are selectively active in certain spatial locations. A
study reported so-called place cells even in humans (Ekstrom et al., 2003). Based on the
early physiological results, OKeefe and Nadel (1978) suggested that the hippocampus
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forms a mental representation of space and that space plays such a central role in
episodic memory that the hippocampus is essential for episodic memory. However, this
view does not easily account for observations that the hippocampus is required for
temporal learning that does not involve learning spatial locations (Fortin et al., 2002;

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McEchron et al., 1998). Recently, Michael Corballis (2013), once one of the most

ardent supporters of the uniquely human hypothesis, suggested that neural sequences

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of place cells that are generated in the rat hippocampus while the animal is resting (Diba

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and Buzski, 2007; Foster and Wilson, 2006; Lee and Wilson, 2002; Pfeiffer and Foster,
2013; for a review, see Buhry et al., 2011) might be evidence of MTT. His collaborator

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Thomas Suddendorf disagrees and maintains their original position (Suddendorf, 2013a).
This controversy exemplifies the difficulty of relating neurophysiological data directly

to cognitive processes (Balter, 2013).

To move beyond the current impasse, we suggest that the three perspectives (focussing

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on WWW information, autonoetic consciousness, or foresight) may examine distinct but

complementary aspects of episodic memory and mental time travel. We propose a

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conceptual model in which episodic memory traces are separate from the capacity to
travel mentally in time. This simple distinction creates a new perspective that is
different from each of the three previous perspective, but, at the same time, integrates
key aspects of each of them. Based on our model we postulate that some nonhuman
animals have the capacity to encode, store and retrieve episodic memory traces, while
potentially lacking other critical components of mental time travel. Tulving (2001)
suggested that nonhuman animals may have memories of specific past events, but that
this was not the same as episodic memory since they lacked autonoetic consciousness.
By contrast, we suggest that humans and nonhuman animals share the same type of
episodic memory traces, as many researchers in nonhuman animal memory would have
it. At the same time, we suggest that mental time travel involves a range of additional
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components and thus, that it remains possible that there is something distinctly human
about the capacity. In particular, there is no compelling evidence to suggest that other
species have the ability to flexibly construct and reflect on alternative scenarios of the
remote future. The first-person feel associated with episodic memory (i.e., autonoetic

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consciousness) is not essential for our distinction but can be accommodated in the

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model.

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2 Components of Mental Time Travel

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It has been argued that mental time travel is not an encapsulated module but the product
of the workings of a number of components (Suddendorf and Busby, 2003). Suddendorf

and Corballis (2007) suggested that a theater production metaphor could help identify
the components involved in successful MTT. They pointed out that there must be mental

analogs to a stage to hold the scenario, a playwright to construct the narrative, actors

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that behave according to our notions of folk psychology, the set that is constructed
according to folk physics, a director to orchestrate rehearsals, an executive producer to

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put the play into action, and a broadcaster to communicate these scenarios. These
components of the theater metaphor can be associated with the development of distinct
cognitive capacities, such as working memory, recursive embedding, theory of mind,
executive functions and language (Suddendorf and Redshaw, 2013). However, for these
components to come together and allow us to construct events displaced in time, one
has to have some memory traces to work with. To these we therefore turn first.

2.1 Episodic Memory Traces

Mental time travel into past and future draws on memories of past experiences (e.g.,
Szpunar and McDermott, 2008). In order to provide us with some useful information
about how to act in the world that we live in and thereby increase our chances of
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survival and reproduction, MTT must be grounded in our general knowledge of the
world and in our personally experienced episodes. We refer to these two sources as
semantic information1 and episodic memory traces, respectively.

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Semantic information are general facts that were extracted from multiple experiences. It
is mostly categorical and refers to prototypical properties of objects and relationships

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between them (Collins and Quillian, 1969; Quillian, 1966; Tulving, 1972). It can also

include idiosyncratic properties of and relationships between particular objects, as long

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as they are consistent across different episodes, such as the name of a particular person

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(Quian Quiroga et al., 2005) or information of the kind The Eiffel Tower is in Paris.
Semantic information can be highly detailed and essential for mental scenario

construction. To use the theater metaphor, semantic information, for instance, provides
the folk physics necessary to build the set: the objects of a scenario typically behave in

accordance with folk physical laws. Or it supplies the folk psychology to direct the

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actors: psychological knowledge constrains how participants of a scenario typically act.

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Episodic memory traces are fairly faithful representations of personally experienced

episodes (Tulving, 1972). They are highly specific in that each trace represents one
particular experienced episode. They are stored incidentally without a special trigger
(Tulving, 1972). In the vast majority of cases, the potential importance of a particular
episodic memory trace is not discernable at the time of storage and the usefulness of the

It is commonly assumed that semantic information is stored in semantic memory and

that semantic memory is a distinct memory system (Squire and Zola-Morgan, 1988).
Even though semantic information is an important ingredient for mental time travel, as
we conceive it here, our model remains agnostic about the source of semantic
information as well as how and where it is stored in the brain.
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stored information might only become apparent in the future. To make it more explicit,
we use the term episodic memory trace to refer to the information that is stored in the
brain about a particular episode and episodic memory to refer to the outcome of scenario

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construction when the information about the episode is retrieved.

A distinction between episodic memory and episodic memory traces has also been

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suggested by Klein (2013), albeit for a very different reason. Klein does not regard the

memory trace as special to episodic memory. In fact, he suggests that the same memory

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trace could be retrieved as semantic memory if the trace was associated with noetic

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consciousness during retrieval, implying that autonoetic consciousness is an add-on to

the memory trace rather than a property of it. By contrast, Cheng and Werning (2015)

recently suggested that episodic memory traces can be distinguished from other memory
traces based on their sequential nature and their link to a specific, experienced episode.

We acknowledge that there might be no clear line that divides episodic memory traces

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from semantic information (Klein, 2013; McKoon and Ratcliff, 1986; Toth and Hunt,
1999), since some cases cannot be easily categorized as either one or the other, leaving

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the possibility that the two are different by degree rather than by kind. Nevertheless, this
controversy is orthogonal to our objective here since our model does not rely on a strict
distinction between episodic memory traces and semantic information. What our model
requires is that some information is linked to one specific experienced episode (episodic
memory trace), whereas other information more generally applies to multiple,
personally experienced or not, episodes (semantic information).
A common view of memory traces is that they store most, if not all, aspect of the
experienced episode for later retrieval (Bernecker, 2010; Martin and Deutscher, 1966;
Patihis et al., 2014). This storage view contrasts with the constructive or generative
view of memory (Michaelian, 2011a, 2011b), according to which information is
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constructed during remembering and this information may bear little resemblance to the
information in the original experience. These two views do not necessarily have to be
mutually exclusive. Experimental evidence suggests that episodic memory traces store
only the gist of an episode (Bartlett, 1932; Deese, 1959; Gernsbacher, 1985; Koutstaal

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and Schacter, 1997; Roediger and McDermott, 1995; Sachs, 1967). For instance,

subjects who saw a cat crossing the road in front of their car are unlikely to remember

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later the exact coloration of the cats fur, what music was playing in the radio, whether

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there were pedestrians in the distance, or other peripheral features of the episode. We

suggest that the gist reflects the information content from the most abstract level that is

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relevant in that one episode (Koutstaal and Schacter, 1997; Oliva, 2005). The relevant
level is established dynamically for each episode depending on the context and attention.

For instance, most subjects are unlikely to remember the color of a cats nose, after
briefly seeing the cat, but they probably would if asked to pay attention to this feature

beforehand. In some cases, such as the preceding example, the most abstract level could

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in fact be quite detailed, but in most cases it is quite coarse. Frequently, the gist includes

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only highly abstract information and details, if required, are constructed from semantic
information during retrieval. For instance, subjects might report that the cat ran onto the
sidewalk after crossing the road, because in their experience most roads have sidewalks,
even if in that particular episode the road had none. Our model thus incorporates aspects
of both the storage and constructive views.
The proposed distinction between episodic memory traces and episodic memory allows
us to separate the content of episodic memory from other components necessary for
mental scenario construction. There is now considerable evidence that various species
store episodic memory traces, as they are conceptualized here. We discuss some of this
evidence in section 3.1. Their mental time travel capacities may, however, be limited
because of shortcomings in other components involved in scenario construction.
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2.2 Scenario Construction

The construction of mental scenarios, as illustrated by Suddendorf and Corballis (2007)
theater metaphor, depends on a range of sophisticated component capacities that enable

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dynamic representations of situations. This scenario construction, or what Cheng and

Werning call mnemonic simulation when applied to the retrieval of episodic memories

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(Cheng and Werning, 2015; Werning and Cheng, 2014), is different from the concept of
scene construction, which Hassabis and Maguire (2007) introduced. The latter refers

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to scenes that are static spatial arrangements while we view scenarios as extending in

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time as well as space. Another difference is that scenes are isolated entities, whereas
scenarios can be embedded to allow constructions of larger narratives and reflection

(Suddendorf, 2013b). Scenarios can contain information about participants, settings,

interactions and sequences of events. We are able to explore a scenario by, for instance,

focusing on different participants, taking different perspectives, and examining the

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spatial layout and the sequence of events. For instance, when we put down our keys, we
frequently do not make a conscious effort to remember where we placed them. Later

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when we are looking for our keys, we might be able to reconstruct our actions around
that time to infer where the keys are. Thus, the construction of mental scenarios allows
us to make new inferences that we did not make at the time when the episode occurred.
MTT into the future and the past use largely overlapping neural mechanisms (Byrne et
al., 2007; Dudai and Carruthers, 2005; Schacter et al., 2007; Suddendorf and Corballis,
2007) but differ in the objective. During MTT into the past, scenarios are constructed
that more or less accurately represent a past episode. However, since episodic memory
traces only contain the gist of the episode, we hypothesize that semantic information is
typically used to aid scenario construction. This process occurs even if subjects are
explicitly instructed only to retrieve a past experience and might account for the
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unreliable nature of human episodic memory (Loftus and Pickrell, 1995; Loftus et al.,
1978; Roediger and McDermott, 1995; Schacter, 2002; Zaragoza and Lane, 1994).
During MTT into the future, the motivation is to construct and compare scenarios about
potential future episodes that can guide behavior. One way to imagine future episodes is

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simply to think that past events will reoccur. But humans can do much more. Humans

can mentally construct scenarios they have never experienced before by combining and

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recombing basic elements into novel constellations and comparing them in terms of

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likelihood, desirablity and so forth (e.g. Gilbert, 2007; Suddendorf and Corballis, 2007).
After all, the future is uncertain and often involves entirely new situations for an

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individual. Nonetheless, scenarios of future events are often based in part on episodic
memory traces. For instance, episodic memory of having packed too much luggage for

the last vacation, might lead us to try to avoid that error during preparation for our
upcoming vacation. Through repetition of this process, we form semantic information

about what to pack for vacations, such that we may no longer need to construct specific

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scenarios when packing for future vacations. The relative contributions of episodic

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memory traces and semantic information during scenario construction may vary
depending on availability, the task demand, and other factors (c.f. Szpunar et al., 2014)
It is important to note that the mechanisms involved in mental scenario construction
enable more than just access to what happened and what might happen..For instance,
they allow us to consider counterfactuals that could have occurred in the past but did not
(De Brigard et al., 2013). They also allow us to imagine the perspectives of other people
(Suddendorf and Corballis, 1997), and even to entertain entirely fictional scenarios that
do not relate to actual past, present nor future situations (Hassabis et al., 2007;
Suddendorf et al., 2009).
Given the complexity of the MTT system, it may not be a fruitful approach to ask
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whether a species does or does not have episodic memory (Beran, 2014; c.f., Templer
and Hampton, 2013). Rather, we propose to focus on studying the extent of their
scenario building capacities and on how these abilities correlate with the extent to which
the components underlying MTT are developed. This kind of analyses has been

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conducted in some detail on the development of mental time travel in children.

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(Suddendorf and Redshaw, 2013). Different limits have different consequences.

For instance, to construct and explore mental scenarios, one needs a sufficient working

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memory capacity (Baddeley, 2000; Baddeley and Hitch, 1974). Ones working memory

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capacity limits the number of information chunks that can be used while constructing
scenarios and therefore the detailedness and spatio-temporal extent of the scenario. It

has been estimated that adult human working memory capacity is as small as 4 chunks
(Cowan, 2001), and lower in other great apes (Balter, 2010; Read, 2008), although the

measurement of the capacity remains contentious. To combine offline representations of

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objects, actions and actors requires generativity, i.e., recursive capacity, which is limited
by the embedding depth. The embedding depth in turn depends on the working memory

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capacity. Constructing a realistic scenario depends on considerable understanding of

how objects and actors behave, i.e., sufficient folk physics and folk psychology, or in
other words, on the complexity and extent of relevant semantic information. Evaluation,
comparison and selection of scenarios requires significant metacognitive and executive
capacities. Here, too, the embedding depth might be a limiting factor. Furthermore, the
properties of the episodic memory traces that are available as bases for scenario
construction may play a role in the outcome. Future research may examine the influence
of the temporal span (i.e., the time interval between the first and last event in an episode)
or of the overall number of traces stored on the range of scenarios individuals construct.
An individuals capacity for mental time travel may be determined by the combination
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of all of the parameters discussed above, and likely by additional factors that we have
not considered here.

Our current model does not require a priori that episodic memory retrieval is

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2.3 How autonoetic consciousness might fit into the model

accompanied by autonoetic consciousness. However, it can accommodate conscious

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awareness during retrieval in humans in a number of potential ways. Consider a simple

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example. An eye witness who is recounting the details of a crime that she experienced
first-hand and a judge who is recounting the same crime after reading the eyewitness

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account. According to Tulving, the eye witness remembering the crime is accompanied
by autonoetic consciousness, whereas the judges remembering the crime is

accompanied by noetic consciousness (even though the judges remembering reading

about the crime is associated with autonoetic consciousness). Klein (2014) suggested

that autonoetic consciousness in humans has the function to indicate the presence of

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retrieved episodic memory as opposed to imaginations and predictions. However, where

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might the difference in conscious awareness come from?

Meta-representation account: One possibility is that noetic and autonoetic
consciousness are related to the different informational contents of meta-representations
(Redshaw, 2014; Suddendorf, 1999). When a person constructs a scenario during
memory retrieval, they also have information about the nature and source of the
memory. This information is represented in the meta-representation. Autonoetic
consciousness might simply consist in the presence of a meta-representation that one
recalls the information about an episode as a result of ones own experience of that
episode. In our example above, the eyewitness meta-represents that her memory is
grounded in having witnessed the crime, while the judge meta-represents that her
memory is based on having read the written account of another person.
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Viewpoint-dependence account: Autonoetic consciousness might arise through the point

of view a person assumes in the constructed scenario that includes the episode (Russell
and Hanna, 2012; Russell et al., 2011). The scenario of an eyewitness has a first-person
perspective: The person is at the center of the scenario, what happens in the scenario is

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related to the person, and the scenario includes her own reactions. The involvement may

be minimal, as in the case of a witness who is merely present in the episode but does not

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influence its course. By contrast, the judge in our example might be able to construct a

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scenario of the crime, but that scenario does not have a first-person perspective, her

body is not at the center of the scenario, and the events in the scenario are not related to

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herself nor does it involve her reactions.

Transparent simulation account: Several authors have suggested that conscious

experience is characteristically linked to what is called phenomenal transparency

(Harman, 1990; Metzinger, 2003; Moore, 1903; Werning, 2010). An occurring mental

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state is phenomenally transparent if only its content-properties are accessible to the

subject at that moment. In other words: A subjects experience of a scenario is

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phenomenally transparent if, and only if, her having that experience is for her just as if
the scenario were present (Werning, 2010). Having the experience is in particular not
for her as if, in addition to the scenario, a representation of the scenario were present.
When the judge reads about the crime, she has access to the crime and its properties, i.e.
the properties of the content of the court record. However, there is something else she
has access to while reading: namely the words of the court record, which is the
representation: the bearer of content. The situation is very different for the eye-witness:
Unless one wants to appeal to a sense-datum theory of perception, she has direct,
unmediated access to the crime. Proponents of the phenomenal-transparency view, hold
that most perceptions, but also certain hallucinations (as in the phenomenon of hearing
voices, in which subjects take these voices for real), are phenomenally transparent,
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whereas many other mental states, e.g., imaginations are typically not (Werning, 2004).2
When you imagine a gun, you not only access the properties of the gun, but also have
access to the mental image, i.e., the representation or content bearer.
Even though experiencing a scenario during remembering is not as transparent as a

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perception of the scenario would be, characterizations of remembering as re-

experiencing or reliving the past suggest that autonoetic consciousness in certain

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respects resembles phenomenally transparent mental states. Viewing remembering as a

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quasi phenomenally transparent simulation of a scenario might, therefore, account for

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autonoetic consciousness.

In conclusion, our model suggests that autonoetic consciousness is not an essential

property of episodic memory, but it nonetheless is or can be associated with human

episodic memory.

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3 Experimental support for our theoretical ideas

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Our central hypothesis that the process underlying MTT can be dissociated into episodic
memory traces and scenario building makes a number of testable predictions, which we
compare to experimental results in the following. The proposed dissociation is
consistent with evidence suggesting that episodic memory emerges gradually during
human development (Suddendorf and Redshaw, 2013). Even though children appear to

In an fMRI study with schizophrenic patients, Dierks et al. (1999) compared auditory

hallucinations to auditory imaginations. In the case of auditory hallucination differential

activity in Heschls gyrus, a part of the primary auditory cortex, was measured. The
location of activity was similar, however, only unilaterally so, to the actual perception
of auditory stimuli.
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possess adult-like episodic memory only by the age of about four (Hayne and Imuta,
2011; Nelson, 1993; Perner and Ruffman, 1995; Scarf et al., 2013), children as young as
2 years old show evidence for storing and retrieving episodic memory traces (Bauer et
al., 1998; Wenner and Bauer, 1999). In these latter studies, the children were able to re-

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enact a complex sequence of actions after observing it only once. Importantly, the

sequence was arbitrary, and therefore the children could not use semantic information

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about the natural sequential order of events to reproduce it. If, however, there are so-

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called enabling relations between the elements of the sequence, even younger toddlers
(in these studies, 16 months olds) could remember the sequence. Our interpretation of

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these results is that children are first able reproduce a sequence based on semantic
information, then develop the ability to store and retrieve arbitrary sequences as

episodic memory traces, and later develop the capacity to construct nested scenarios and
to destill more complex semantic information. The final stages of the development of

episodic memory continue well after age 4, when increases in working memory capacity,

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among others factors, lead to the construction of more complex episodic memories and

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larger narratives (Suddendorf and Redshaw, 2013).

The distinction between episodic memory and episodic memory traces also allows us to
relate findings from nonhuman animal studies to those in humans, which we turn to
next.

3.1 Episodic memory traces in the brain

While several brain regions are certainly involved in storing and retrieving episodic
memory traces, current evidence suggests that, in mammals, the associative binding of
the items involved in the episode and the temporal sequence of events in the episode
critically depends on the hippocampus (Cheng and Werning, 2015; Cheng, 2013; Fortin
et al., 2002; Levy, 1996; Wood et al., 1999). Binding has been suggested before to be a
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central property of episodic memory (Clayton et al., 2003; Crystal, 2010). Our model
leads to, at least, three major implications about hippocampal function that are well
supported by observations in humans and other mammals, in particular rodents.

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First, the responses of hippocampal neurons are invariant with respect to the details of
the presented stimuli. In rodents (OKeefe and Dostrovsky, 1971) and bats (Ulanovsky

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and Moss, 2007), place cell activity is independent of the view that the subject sees at

that location and specific details of the environment (Muller et al., 1994). Furthermore,

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Quian Quiroga et al. (2005) found that hippocampal neurons in humans are activated

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when subjects see photos of certain people, irrespective of their hair style, clothes, and
pose; and even when only their written name was presented. These results suggest that

hippocampal neurons responds to the gist in the stimuli, rather than to their visual
details (Cheng and Werning, 2013; Werning and Maye, 2007).

Second, hippocampal neurons respond to different types of information and form

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arbitrary associations between different pieces of information. Cells in human

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hippocampus integrate information across different sensory modalities, responding to

the auditory presentation of a persons name as well as visual cues related to that person
(Quian Quiroga et al., 2009; Quiroga, 2012). In rodents, studies have found that
hippocampal cells respond to complex conjunctions of different types of stimuli. For
instance, in a classical conditioning paradigm, hippocampal cells responded to the
conditioned stimulus, a tone, but not to the same tone if it was not paired with an
unconditioned stimulus (Moita et al., 2003). In addition, the cells responses were
modulated by the location of the animal. Their conjunctive and multimodal responses
make it plausible that hippocampal neurons are involved in storing episodic memory
traces that represent unique and highly diverse episodes.
Third, the hippocampus is important for learning temporal sequences and bridging
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Page 18 of 33

temporal gaps. Experimental studies suggest that the hippocampus is required for
sequence learning (Fortin et al., 2002), trace eye blink conditioning in humans
(McGlinchey-Berroth et al., 1997) and in rodents (Weiss et al., 1999), and trace fear
conditioning in humans (Clark and Squire, 1998) and in rodents (McEchron et al., 1998).

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In addition, functional magnetic resonance imaging revealed increased hippocampal

activation during trace as compared to delay conditioning (Cheng et al., 2008). For a

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representation of temporal sequences, see Cheng and Werning (2015).

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more detailed discussion of experimental evidence linking the hippocampus to the

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We conclude that there is strong evidence that the hippocampus plays an important role
in the storage of episodic memory traces in both human and nonhuman animals.

3.2 Scenario construction in episodic memory

Scenario construction of past episodes during the retrieval of episodic memories has

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two important implications for human episodic memory.

First, as described above during episodic memory retrieval many details are

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reconstructed based on semantic information rather than solely on the basis of episodic
memory traces. Early evidence for this effect was reported by Bartlett (1932) who
observed that English subjects over time did not remember details of the Canadian
Indian Folklore "War of the Ghosts" as they were in the original story, but adjusted them
to their own cultural expectations. Better controlled laboratory studies into the
reconstructive nature of memories are provided by the DeeseRoedigerMcDermott
(DRM) paradigm (Roediger and McDermott, 1995). After studying a list of
semantically related words, subjects will name other semantically related words that
were not on the study list when trying recalling the study words.
Second, information from different episodic memory traces are pooled together during
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episodic memory retrieval. Confabulations often do result, but need not, when distinct
episodic memory traces are combined fallaciously. Imaging experiments suggest that
episodes indeed form units of human cognition in that brain activity patterns are more
similar within an episode than patterns from distinct episodes (Ezzyat and Davachi,

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2011). It is noteworthy that this occurs even when subjects are not consciously aware of
the fact that episodes were defined by hidden statistical rules (Schapiro et al., 2013).

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Nevertheless, as time passes it becomes difficult for subjects to keep separate

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information that they were exposed to during two distinct episodes. This was mostly

studied using the misattribution paradigm, where information that was provided in post-

Marsh et al., 2008; Zaragoza and Lane, 1994).

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session questions was fallaciously attributed to the original session (Loftus et al., 1978;

The results of psychological experiments thus support our model, in which, during

episodic memory retrieval, scenarios are constructed based on episodic memory traces

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enriched with semantic information. The ultimate reason why episodic memory may be
vulnerable to integration errors and biases may be that scenario construction evolved

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primarily to enable flexible simulation of future situations, rather than the faithful
recreation of the past (Schacter and Addis 2007; Suddendorf and Corballis 2007).
We acknowledge that it is far simpler to obtain evidence for scenario construction in
humans since they can use language to communicate the scenarios they produce. Indeed
humans capacity for flexible communication may have evolved for that very purpose
(Suddendorf et al., 2009). However, behavioral evidence for scenario construction could
be obtained by other means in principle. It is just that previous attempts at obtaining
such evidence have not been compelling (Suddendorf and Corballis, 2010). Recent
eletrophysiological findings in rodents are also inconclusive. While hippocampal place
cells are activated in sequences that were never experienced before (Dragoi and
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Tonegawa, 2011; Gupta et al., 2010), there is no indication that these serve a function. It
remains possible that these neural sequences are generated randomly by a neural
network (Azizi et al., 2013) . More relevant, perhaps, is the observation that, in familiar
environments, place cell sequences predict upcoming movement trajectories (Pfeiffer

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and Foster, 2013). However, these sequences appear to correlate with planning a

trajectory to a current goal rather than reflect the flexible construction of scenarios. In

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summary, there is no compelling evidence for scenario construction in non-human

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animals.

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4 Conclusion

We have suggested that disparate findings from human and nonhuman animal studies in
episodic-like and episodic memory can be integrated into a fruitful model. We propose

that it may be useful to conceive of episodic memory traces as one of several

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components that are important for MTT, but that episodic memory traces can exist
without these other components and may be present early in human development as

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well as in nonhuman animals. It might therefore be possible to directly apply insights

into the neural mechanisms of episodic-like memory in nonhuman animals to
understanding human episodic memory traces. At the same time, we are skeptical that
nonhuman animals have developed all the other components of MTT and suggest that it
remains quite possible that only humans are able to construct nested scenarios and
reflect on them with a view to flexibly secure their future needs.

5 Acknowledgements
This work was supported by a grant from the Stiftung Mercator to S.C. and M.W., grant
SFB 874, project B2 from the German Research Foundation (Deutsche
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Forschungsgemeinschaft, DFG) to S.C. and by an Australian Research Council grant

DP140101341 to T.S. The sponsors had no role in study design, data collection, analysis,
and interpretation, decision to publish, or preparation of the manuscript.

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6 References

cr

Azizi, A.H., Wiskott, L., Cheng, S., 2013. A computational model for preplay in the
hippocampus. Front. Comput. Neurosci. 7, 161. doi:10.3389/fncom.2013.00161

us

Babb, S.J., Crystal, J.D., 2005. Discrimination of what, when, and where: Implications
for episodic-like memory in rats. Learn. Motiv. 36, 177189.
doi:10.1016/j.lmot.2005.02.009

an

Baddeley, A., 2000. The episodic buffer: a new component of working memory? Trends
Cogn. Sci. 4, 417423.

Baddeley, A.D., Hitch, G., 1974. Working Memory. Psychol. Learn. Motiv. 8, 4789.
doi:10.1016/S0079-7421(08)60452-1

Balter, M., 2013. Animal cognition. Can animals envision the future? Scientists spar
over new data. Science 340, 909. doi:10.1126/science.340.6135.909

te

Balter, M., 2010. Evolution of behavior. Did working memory spark creative culture?
Science 328, 1603. doi:10.1126/science.328.5975.160

Ac
ce
p

Bartlett, F.C., 1932. Remembering: a study in experimental and social psychology.

Cambridge University Press.
Bauer, P.J., Hertsgaard, L. a, Dropik, P., Daly, B.P., 1998. When even arbitrary order
becomes important: developments in reliable temporal sequencing of arbitrarily
ordered events. Memory 6, 16598. doi:10.1080/741942074
Beran, M.J., 2014. Animal Memory: Rats Bind Event Details into Episodic Memories.
Curr. Biol. 24, R1159R1160. doi:10.1016/j.cub.2014.11.019
Bernecker, S., 2010. Memory: a philosophical study. Oxford University Press.
Buhry, L., Azizi, A.H., Cheng, S., 2011. Reactivation, replay, and preplay: How it
might all fit together. Neural Plast. 2011, 111. doi:10.1155/2011/203462
Byrne, P., Becker, S., Burgess, N., 2007. Remembering the past and imagining the
future: a neural model of spatial memory and imagery. Psychol. Rev. 114, 340
375. doi:10.1037/0033-295X.114.2.340
Cheng, D.T., Disterhoft, J.F., Power, J.M., Ellis, D.A., Desmond, J.E., 2008. Neural
substrates underlying human delay and trace eyeblink conditioning. Proc. Natl.
Acad. Sci. U. S. A. 105, 810813. doi:10.1073/pnas.0800374105
22
Page 22 of 33

Cheng, S., 2013. The CRISP theory of hippocampal function in episodic memory.
Front. Neural Circuits 7, 88. doi:10.3389/fncir.2013.00088
Cheng, S., Werning, M., 2015. What is Episodic Memory if it is a Natural Kind?
Synthese. doi:10.1007/s11229-014-0628-6

ip
t

Cheng, S., Werning, M., 2013. Composition and replay of mnemonic sequences: The
contributions of REM and slow-wave sleep to episodic memory. Behav. Brain Sci.
36, 610611. doi:10.1017/S0140525X13001234

cr

Clark, R.E., Squire, L.R., 1998. Classical conditioning and brain systems: the role of
awareness. Science 280, 7781. doi:10.1126/science.280.5360.77

us

Clayton, N.S., Bussey, T.J., Dickinson, A., 2003. Can animals recall the past and plan
for the future? Nat. Rev. Neurosci. 4, 68591. doi:10.1038/nrn1180
Clayton, N.S., Dickinson, A., 1998. Episodic-like memory during cache recovery by
scrub jays. Nature 395, 2724. doi:10.1038/26216

an

Collins, A.M., Quillian, M.R., 1969. Retrieval time from semantic memory. J. Verbal
Learning Verbal Behav. 8, 240247. doi:10.1016/S0022-5371(69)80069-1

Corballis, M.C., 2013. Mental time travel: a case for evolutionary continuity. Trends
Cogn. Sci. 17, 56. doi:10.1016/j.tics.2012.10.009

Cowan, N., 2001. The magical number 4 in short-term memory: a reconsideration of

mental storage capacity. Behav. Brain Sci. 24, 87114; discussion 114185.
doi:10.1017/S0140525X01003922

te

Crystal, J.D., 2010. Episodic-like memory in animals. Behav. Brain Res. 215, 23543.
doi:10.1016/j.bbr.2010.03.005

Ac
ce
p

Dally, J.M., Emery, N.J., Clayton, N.S., 2006. Food-caching western scrub-jays keep
track of who was watching when. Science 312, 16625.
doi:10.1126/science.1126539
De Brigard, F., Addis, D.R., Ford, J.H., Schacter, D.L., Giovanello, K.S., 2013.
Remembering what could have happened: Neural correlates of episodic
counterfactual thinking. Neuropsychologia 51, 24012414.
doi:10.1016/j.neuropsychologia.2013.01.015
Deese, J., 1959. On the prediction of occurrence of particular verbal intrusions in
immediate recall. J. Exp. Psychol. 58, 1722. doi:10.1037/h0046671
Dekleva, M., Dufour, V., de Vries, H., Spruijt, B.M., Sterck, E.H.M., 2011.
Chimpanzees (Pan troglodytes) fail a what-where-when task but find rewards by
using a location-based association strategy. PLoS One 6, e16593.
doi:10.1371/journal.pone.0016593
Dere, E., Huston, J.P., De Souza Silva, M. a, 2005. Episodic-like memory in mice:
simultaneous assessment of object, place and temporal order memory. Brain Res.
23
Page 23 of 33

Brain Res. Protoc. 16, 109. doi:10.1016/j.brainresprot.2005.08.001

Diba, K., Buzski, G., 2007. Forward and reverse hippocampal place-cell sequences
during ripples. Nat. Neurosci. 10, 12411242. doi:10.1038/nn1961

ip
t

Dierks, T., Linden, D.E., Jandl, M., Formisano, E., Goebel, R., Lanfermann, H., Singer,
W., 1999. Activation of Heschls gyrus during auditory hallucinations., Neuron.
doi:10.1016/S0896-6273(00)80715-1
Dragoi, G., Tonegawa, S., 2011. Preplay of future place cell sequences by hippocampal
cellular assemblies. Nature 469, 397401. doi:10.1038/nature09633

cr

Dudai, Y., Carruthers, M., 2005. The Janus face of Mnemosyne. Nature 434, 567567.
doi:10.1038/434567a

us

Eacott, M.J., Easton, A., Zinkivskay, A., 2005. Recollection in an episodic-like memory
task in the rat. Learn. Mem. 12, 2213. doi:10.1101/lm.92505

an

Eichenbaum, H., Fortin, N.J., Ergorul, C., Wright, S.P., Agster, K.L., 2005. Episodic
recollection in animals: If it walks like a duck and quacks like a duck... Learn.
Motiv. 36, 190207. doi:10.1016/j.lmot.2005.02.006

Ekstrom, A.D., Kahana, M.J., Caplan, J.B., Fields, T.A., Isham, E.A., Newman, E.L.,
Fried, I., Dale, J., Kandil, F., Nishida, S., 2003. Cellular networks underlying
human spatial navigation. Nature 425, 184188. doi:10.1038/nature01964

Ezzyat, Y., Davachi, L., 2011. What constitutes an episode in episodic memory?
Psychol. Sci. 22, 24352. doi:10.1177/0956797610393742

Ac
ce
p

te

Feeney, M.C., Roberts, W. a, Sherry, D.F., 2009. Memory for what, where, and when in
the black-capped chickadee (Poecile atricapillus). Anim. Cogn. 12, 76777.
doi:10.1007/s10071-009-0236-x
Ferkin, M.H., Combs, A., delBarco-Trillo, J., Pierce, A.A., Franklin, S., 2008. Meadow
voles, Microtus pennsylvanicus, have the capacity to recall the what, where,
and when of a single past event. Anim. Cogn. 11, 14759. doi:10.1007/s10071007-0101-8
Fortin, N.J., Agster, K.L., Eichenbaum, H.B., 2002. Critical role of the hippocampus in
memory for sequences of events. Nat. Neurosci. 5, 45862. doi:10.1038/nn834
Foster, D.J., Wilson, M.A., 2006. Reverse replay of behavioural sequences in
hippocampal place cells during the awake state. Nature 440, 680683.
doi:10.1038/nature04587
Gernsbacher, M.A., 1985. Surface information loss in comprehension. Cogn. Psychol.
17, 324363. doi:10.1016/0010-0285(85)90012-X
Gilbert, D.T., 2007. Prospection: Experiencing the Future 317, 13511354.
doi:10.1126/science.1144161
Griffiths, D., Dickinson, A., Clayton, N., 1999. Episodic memory: What can animals
24
Page 24 of 33

remember about their past? Trends Cogn. Sci. 3, 7480. doi:10.1016/S13646613(98)01272-8

Gupta, A.S., van der Meer, M.A.A., Touretzky, D.S., Redish, A.D., 2010. Hippocampal
replay is not a simple function of experience. Neuron 65, 695705.
doi:10.1016/j.neuron.2010.01.034

cr

ip
t

Hampton, R.R., Hampstead, B.M., Murray, E. a., 2005. Rhesus monkeys (Macaca
mulatta) demonstrate robust memory for what and where, but not when, in an
open-field test of memory. Learn. Motiv. 36, 245259.
doi:10.1016/j.lmot.2005.02.004

us

Harand, C., Bertran, F., La Joie, R., Landeau, B., Mzenge, F., Desgranges, B.,
Peigneux, P., Eustache, F., Rauchs, G., 2012. The hippocampus remains activated
over the long term for the retrieval of truly episodic memories. PLoS One 7,
e43495. doi:10.1371/journal.pone.0043495

an

Harman, G., 1990. The Intrinsic Quality of Experience. Philos. Perspect. 4, 3151.

Hassabis, D., Kumaran, D., Vann, S.D., Maguire, E.A., 2007. Patients with
hippocampal amnesia cannot imagine new experiences. Proc. Natl. Acad. Sci. U.
S. A. 104, 17261731. doi:10.1073/pnas.0610561104
Hassabis, D., Maguire, E.A., 2007. Deconstructing episodic memory with construction.
Trends Cogn. Sci. 11, 299306. doi:10.1016/j.tics.2007.05.001

te

Hayne, H., Imuta, K., 2011. Episodic memory in 3- and 4-year-old children. Dev.
Psychobiol. 53, 317322. doi:10.1002/dev.20527

Ac
ce
p

Hoffman, M.L., Beran, M.J., Washburn, D. a, 2009. Memory for what, where, and
when information in rhesus monkeys (Macaca mulatta). J. Exp. Psychol. Anim.
Behav. Process. 35, 14352. doi:10.1037/a0013295
Kaminski, J., Fischer, J., Call, J., 2008. Prospective object search in dogs: mixed
evidence for knowledge of What and Where. Anim. Cogn. 11, 36771.
doi:10.1007/s10071-007-0124-1
Klein, S.B., 2014. Autonoesis and Belief in a Personal Past: An Evolutionary Theory of
Episodic Memory Indices. Rev. Philos. Psychol. 427447. doi:10.1007/s13164014-0181-8
Klein, S.B., 2013. Making the case that episodic recollection is attributable to
operations occurring at retrieval rather than to content stored in a dedicated
subsystem of long-term memory. Front. Behav. Neurosci. 7, 3.
doi:10.3389/fnbeh.2013.00003
Koutstaal, W., Schacter, D.L., 1997. Gist-Based False Recognition of Pictures in Older
and Younger Adults. J. Mem. Lang. 37, 555583. doi:10.1006/jmla.1997.2529
Kouwenberg, A.-L., Walsh, C.J., Morgan, B.E., Martin, G.M., 2009. Episodic-like
memory in crossbred Yucatan minipigs (Sus scrofa). Appl. Anim. Behav. Sci. 117,
25
Page 25 of 33

165172. doi:10.1016/j.applanim.2009.01.005
Lee, A.K., Wilson, M.A., 2002. Memory of sequential experience in the hippocampus
during slow wave sleep. Neuron 36, 11831194.

ip
t

Levy, W.B.W., 1996. A sequence predicting CA3 is a flexible associator that learns and
uses context to solve hippocampal-like tasks. Hippocampus 6, 579590.
doi:10.1002/(SICI)1098-1063(1996)6:6<579::AID-HIPO3>3.0.CO;2-C

cr

Loftus, E.F., Miller, D.G., Burns, H.J., 1978. Semantic integration of verbal information
into a visual memory. J. Exp. Psychol. Hum. Learn. Mem. 4, 1931.
doi:10.1037//0278-7393.4.1.19

us

Loftus, E.F., Pickrell, J.E., 1995. The formation of false memories. Psychiatr. Ann. 25,
720725.

an

Marsh, E.J., Eslick, A.N., Fazio, L.K., 2008. False Memories, in: Byrne, J.H. (Ed.),
Learning and Memory: A Comprehensive Reference. Academic Press, Oxford, pp.
221238.
Martin, C.B., Deutscher, M., 1966. Remembering. Philos. Rev. 75, 161.
doi:10.2307/2183082

Martin-Ordas, G., Haun, D., Colmenares, F., Call, J., 2010. Keeping track of time:
evidence for episodic-like memory in great apes. Anim. Cogn. 13, 33140.
doi:10.1007/s10071-009-0282-4

Ac
ce
p

te

McEchron, M.D., Bouwmeester, H., Tseng, W., Weiss, C., Disterhoft, J.F., 1998.
Hippocampectomy disrupts auditory trace fear conditioning and contextual fear
conditioning in the rat. Hippocampus 8, 63846. doi:10.1002/(SICI)10981063(1998)8:6&lt;638::AID-HIPO6&gt;3.0.CO;2-Q
McGlinchey-Berroth, R., Carrillo, M.C., Gabrieli, J.D., Brawn, C.M., Disterhoft, J.F.,
1997. Impaired trace eyeblink conditioning in bilateral, medial-temporal lobe
amnesia. Behav. Neurosci. 111, 87382.
McKoon, G., Ratcliff, R., 1986. Automatic activation of episodic information in a
semantic memory task. J. Exp. Psychol. Learn. Mem. Cogn. 12, 10815.
Metzinger, T., 2003. Phenomenal transparency and cognitive self-reference.
Phenomenol. Cogn. Sci. 2, 353393. doi:10.1023/B:PHEN.0000007366.42918.eb
Michaelian, K., 2011a. Generative memory. Philos. Psychol. 24, 323342.
doi:10.1080/09515089.2011.559623
Michaelian, K., 2011b. The information effect: constructive memory, testimony, and
epistemic luck. Synthese 190, 24292456. doi:10.1007/s11229-011-9992-7
Moita, M.A., Rosis, S., Zhou, Y., LeDoux, J.E., Blair, H.T., 2003. Hippocampal place
cells acquire location-specific responses to the conditioned stimulus during
auditory fear conditioning. Neuron 37, 485497.
26
Page 26 of 33

Moore, G.E., 1903. The Refutation of Idealism. Mind 12, 433453.

Muller, R.U., Bostock, E., Taube, J.S., Kubie, J.L., 1994. On the directional firing
properties of hippocampal place cells. J. Neurosci. 14, 723551.

ip
t

Nadel, L., Samsonovich, A., Ryan, L., Moscovitch, M., 2000. Multiple trace theory of
human memory: computational, neuroimaging, and neuropsychological results.
Hippocampus 10, 35268. doi:10.1002/1098-1063(2000)10:4<352::AIDHIPO2>3.0.CO;2-D

cr

Nelson, K., 1993. The Psychological and Social Origins of Autobiographical Memory.
Psychol. Sci. 4, 714. doi:10.1111/j.1467-9280.1993.tb00548.x

us

OKeefe, J., Dostrovsky, J., 1971. The hippocampus as a spatial map. Preliminary
evidence from unit activity in the freely-moving rat. Brain Res. 34, 171175.
doi:10.1016/0006-8993(71)90358-1

an

OKeefe, J., Nadel, L., 1978. The Hippocampus as a Cognitive Map. Clarendon Press,
Oxford.

Oliva, A., 2005. Gist of the scene, in: Itti, L., Rees, G., Tsotsos, J.K. (Eds.),
Neurobiology of Attention. Elsevier Academic Press, London, pp. 251256.
doi:10.1016/B978-012375731-9/50045-8

Patihis, L., Ho, L.Y., Tingen, I.W., Lilienfeld, S.O., Loftus, E.F., 2014. Are the
memory wars over? A scientist-practitioner gap in beliefs about repressed
memory. Psychol. Sci. 25, 51930. doi:10.1177/0956797613510718

Ac
ce
p

te

Perner, J., Ruffman, T., 1995. Episodic memory and autonoetic consciousness:
developmental evidence and a theory of childhood amnesia. J. Exp. Child Psychol.
doi:10.1006/jecp.1995.1024
Pfeiffer, B.E., Foster, D.J., 2013. Hippocampal place-cell sequences depict future paths
to remembered goals. Nature 497, 749. doi:10.1038/nature12112
Quian Quiroga, R., Kraskov, A., Koch, C., Fried, I., 2009. Explicit encoding of
multimodal percepts by single neurons in the human brain. Curr. Biol. 19, 1308
13. doi:10.1016/j.cub.2009.06.060
Quian Quiroga, R., Reddy, L., Kreiman, G., Koch, C., Fried, I., Quian Quiroga, R.,
2005. Invariant visual representation by single neurons in the human brain. Nature
435, 11021107. doi:10.1038/nature03687
Quillian, M.R., 1966. Semantic Memory. Bedford, MA.
Quiroga, R.Q., 2012. Concept cells: the building blocks of declarative memory
functions. Nat. Rev. Neurosci. 13, 58797. doi:10.1038/nrn3251
Raby, C.R., Clayton, N.S., 2009. Prospective cognition in animals. Behav. Processes
80, 314324. doi:10.1016/j.beproc.2008.12.005
Read, D.W., 2008. Working Memory: A Cognitive Limit to Non-Human Primate
27
Page 27 of 33

Recursive Thinking Prior to Hominid Evolution. Evol. Psychol. 6, 676714.

Redshaw, J., 2014. Does metarepresentation make human mental time travel unique?
Wiley Interdiscip. Rev. Cogn. Sci. 5, 519531. doi:10.1002/wcs.1308

ip
t

Roediger, H.L., McDermott, K.B., 1995. Creating false memories: Remembering words
not presented in lists. J. Exp. Psychol. Learn. Mem. Cogn. 21, 803814.
doi:10.1037/0278-7393.21.4.803

cr

Russell, J., Cheke, L.G., Clayton, N.S., Meltzoff, A.N., 2011. What can What-WhenWhere (WWW) binding tasks tell us about young childrens episodic foresight?
Theory and two experiments. Cogn. Dev. 26, 356370.
doi:10.1016/j.cogdev.2011.09.002

us

Russell, J., Hanna, R., 2012. A minimalist approach to the development of episodic
memory. Mind Lang. 27, 2954. doi:10.1111/j.1468-0017.2011.01434.x

an

Sachs, J.S., 1967. Recopition memory for syntactic and semantic aspects of connected
discourse. Percept. Psychophys. 2, 437442. doi:10.3758/BF03208784

Scarf, D., Gross, J., Colombo, M., Hayne, H., 2013. To have and to hold: Episodic
memory in 3- and 4-year-old children. Dev. Psychobiol. 55, 125132.
doi:10.1002/dev.21004

Schacter, D.L., 2002. The Seven Sins of Memory: How the Mind Forgets and
Remembers. Houghton Mifflin.

te

Schacter, D.L., Addis, D.R., 2007. The cognitive neuroscience of constructive memory:
remembering the past and imagining the future. Philos. Trans. R. Soc. B Biol. Sci.
362, 773786. doi:10.1098/rstb.2007.2087

Ac
ce
p

Schacter, D.L., Addis, D.R.R., Buckner, R.L., 2007. Remembering the past to imagine
the future: the prospective brain. Nat. Rev. Neurosci. 8, 657661.
doi:10.1038/nrn2213
Schacter, D.L., Guerin, S.A., St Jacques, P.L., 2011. Memory distortion: an adaptive
perspective. Trends Cogn. Sci. 15, 467474. doi:10.1016/j.tics.2011.08.004
Schapiro, A.C., Rogers, T.T., Cordova, N.I., Turk-Browne, N.B., Botvinick, M.M.,
2013. Neural representations of events arise from temporal community structure.
Nat. Neurosci. 16, 48692. doi:10.1038/nn.3331
Scoville, W.B., Milner, B., 1957. Loss of recent memory after bilateral hippocampal
lesions. J. Neurol. Neurosurg. Psychiatry 20, 1121.
Squire, L.R., Zola-Morgan, S., 1988. Memory: brain systems and behavior. Trends
Neurosci. 11, 1705.
Suddendorf, T., 2013a. Mental time travel: continuities and discontinuities. Trends
Cogn. Sci. 17, 1512. doi:10.1016/j.tics.2013.01.011
Suddendorf, T., 2013b. The Gap: The Science of What Separates Us from Other
28
Page 28 of 33

Animals. Basic Books, New York, NY.

Suddendorf, T., 1999. The rise of the metamind, in: Corballis, M.C., Lea, S.E.G. (Eds.),
The Descent of Mind: Psychological Perspectives on Hominid Evolution. Oxford
University Press, London, pp. 218260.
doi:10.1093/acprof:oso/9780192632593.003.0012

ip
t

Suddendorf, T., Addis, D.R., Corballis, M.C., 2009. Mental time travel and the shaping
of the human mind. Philos. Trans. R. Soc. B Biol. Sci. 364, 13171324.
doi:10.1098/rstb.2008.0301

cr

Suddendorf, T., Busby, J., 2003. Mental time travel in animals? Trends Cogn. Sci. 7,
391396. doi:10.1016/S1364-6613(03)00187-6

us

Suddendorf, T., Corballis, M.C., 2010. Behavioural evidence for mental time travel in
nonhuman animals. Behav. Brain Res. 215, 292298.
doi:10.1016/j.bbr.2009.11.044

an

Suddendorf, T., Corballis, M.C., 2008. Episodic memory and mental time travel, in:
Dere, E., Huston, J.P., Easton, A. (Eds.), Handbook of Episodic Memory Research.
Elsevier, Amsterdam, pp. 3142.

Suddendorf, T., Corballis, M.C., 2007. The evolution of foresight: What is mental time
travel, and is it unique to humans? Behav. Brain Sci. 30, 299313; discussion 313
51. doi:10.1017/S0140525X07001975

te

Suddendorf, T., Corballis, M.C., 1997. Mental time travel and the evolution of the
human mind. Genet. Soc. Gen. Psychol. Monogr. 123, 13367.

Ac
ce
p

Suddendorf, T., Redshaw, J., 2013. The development of mental scenario building and
episodic foresight. Ann. N. Y. Acad. Sci. 1296, 13553. doi:10.1111/nyas.12189
Szpunar, K.K., McDermott, K.B., 2008. Episodic future thought and its relation to
remembering: evidence from ratings of subjective experience. Conscious. Cogn.
17, 3304. doi:10.1016/j.concog.2007.04.006
Szpunar, K.K., Spreng, R.N., Schacter, D.L., 2014. A taxonomy of prospection:
Introducing an organizational framework for future-oriented cognition: Fig. 1.
Proc. Natl. Acad. Sci. 111, 1841418421. doi:10.1073/pnas.1417144111
Templer, V.L., Hampton, R.R., 2013. Episodic memory in nonhuman animals. Curr.
Biol. 23, R801R806. doi:10.1016/j.cub.2013.07.016
Toth, J.P., Hunt, R.R., 1999. Not one versus many, but zero versus any: structure and
function in the context of the multiple memory systems debate, in: Foster, J.K.,
Jelicic, M. (Eds.), Memory: Systems, Process, or Function? Oxford University
Press, Oxford, pp. 232272.
Tulving, E., 2001. Episodic memory and common sense: how far apart? Philos. Trans.
R. Soc. Lond. B. Biol. Sci. 356, 150515. doi:10.1098/rstb.2001.0937
29
Page 29 of 33

Tulving, E., 1985. Memory and Consciousness. Can. J. Psychol. 26, 126.
Tulving, E., 1972. Episodic and Semantic Memory, in: Tulving, E., Donaldson, W.
(Eds.), Organization of Memory. Academic Press, Inc., New York, NY, USA, pp.
381402.

ip
t

Ulanovsky, N., Moss, C.F., 2007. Hippocampal cellular and network activity in freely
moving echolocating bats. Nat. Neurosci. 10, 224233. doi:10.1038/nn1829

cr

Weiss, C., Bouwmeester, H., Power, J.M., Disterhoft, J.F., 1999. Hippocampal lesions
prevent trace eyeblink conditioning in the freely moving rat. Behav. Brain Res. 99,
12332.

us

Wenner, J. a., Bauer, P.J., 1999. Bringing order to the arbitrary: One- to two-year-olds
recall of event sequences. Infant Behav. Dev. 22, 585590. doi:10.1016/S01636383(00)00013-8

an

Werning, M., 2010. Descartes discarded? Introspective self-awareness and the problems
of transparency and compositionality. Conscious. Cogn. 19, 751761.
doi:10.1016/j.concog.2010.07.003

Werning, M., 2004. Self-awareness and imagination, in: Sgua, J. (Ed.), The
Explanation of Human Interpretation. Colibri, Lisbon, pp. 369377.

te

Werning, M., Cheng, S., 2014. Is Episodic Memory a Natural Kind?-A Defense of the
Sequence Analysis, in: Bello, P., Guarini, M., McShane, M., Scassellati, B. (Eds.),
Proceedings of the 36th Annual Conference of the Cognitive Science Society.
Cognitive Science Society, Austin, TX, pp. 964969.

Ac
ce
p

Werning, M., Maye, A., 2007. The cortical implementation of complex attribute and
substance concepts: Synchrony, frames, and hierarchical binding. Chaos Complex.
Lett. 2, 435452.
Wood, E.R., Dudchenko, P.A., Eichenbaum, H., 1999. The global record of memory in
hippocampal neuronal activity. Nature 397, 613616. doi:10.1038/17605
Zaragoza, M.S., Lane, S.M., 1994. Source misattributions and the suggestibility of
eyewitness memory. J. Exp. Psychol. Learn. Mem. Cogn. 20, 934945.
doi:10.1037/0278-7393.20.4.934
Zentall, T.R., Clement, T.S., Bhatt, R.S., Allen, J., 2001. Episodic-like memory in
pigeons. Psychon. Bull. Rev. 8, 68590.
Zinkivskay, A., Nazir, F., Smulders, T. V, 2009. What-where-when memory in magpies
(Pica pica). Anim. Cogn. 12, 11925. doi:10.1007/s10071-008-0176-x
Zola-Morgan, S., Squire, L.R., Amaral, D.G., 1986. Human amnesia and the medial
temporal region: enduring memory impairment following a bilateral lesion limited
to field CA1 of the hippocampus. J. Neurosci. 6, 295067.

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7 Figure Captions
Figure 1: Schematic of three different models of episodic memory. See the main text for a

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with Mental Time Travel (MTT). S&C 1997: Suddendorf and Corballis, 1997

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more detailed description. The dashed box indicates the components most closely associated

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Highlights

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Mental time travel (MTT) involves the construction of scenarios

Scenarios contain information about participants, settings, interactions and
events
Scenarios are constructed based on episodic memory traces and semantic
information
Some nonhuman animals appear to encode, store and retrieve episodic
memory traces
Only humans seem to construct, reflect and flexibly compare alternative
scenarios

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