Consciousness and Cognition: Hans J. Markowitsch, Angelica Staniloiu

Consciousness and Cognition xxx (2010) xxxxxx
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Consciousness and Cognition

journal homepage: www.elsevier.com/locate/concog
Memory, autonoetic consciousness, and the self q

Hans J. Markowitsch a,b,, Angelica Staniloiu a
a b
Physiological Psychology, University of Bielefeld, Bielefeld, Germany Alfried Krupp Institute for Advanced Study, Greifswald, Germany
a r t i c l e
i n f o
a b s t r a c t
Memory is a general attribute of living species, whose diversication reects both evolutionary and developmental processes. Episodic-autobiographical memory (EAM) is regarded as the highest human ontogenetic achievement and as probably being uniquely human. EAM, autonoetic consciousness and the self are intimately linked, grounding, supporting and enriching each others development and cohesiveness. Their development is inuenced by the socio-culturallinguistic environment in which an individual grows up or lives. On the other hand, through language, textualization and social exchange, all three elements leak into the world and participate to the dynamic shaping and re-shaping of the cultural scaffolding of the self, mental time traveling and EAM formation. Decits in selfrelated processing, autonetic consciousness, emotional processing and mental time traveling can all lead to or co-occur with EAM disturbances, as we illustrate by ndings from EAM impairments associated with neurological or psychiatric disorders. 2010 Elsevier Inc. All rights reserved.
Article history: Available online xxxx Keywords: Dissociative amnesia Self-consciousness Emotion Episodic-autobiographical memory (EAM) Perspective taking Time
1. Introduction Memory is a multi-facetted attribute of all animals and may even be found in rudimentary forms in the ora and in so-called intelligent machines. This speaks for a million year-long process of memory evolution. More recent theories in psychology and the neurosciences have acknowledged this process by proposing different memory systems, especially in phylogenetically advanced species. The endowment with a developed body that has enabled the exploration of the environment over wide distances (embodiment; Pfeifer & Bongard, 2007) has most likely led to the diversication of memories as well as the need to store information long term (cf. Campbell & Garcia, 2009). This is evident in species such as certain birds (Miyata, Gajdon, Huber, & Fujita, 2010; Weir, Chappell, & Kacelnik, 2002), whales and dolphins (Reiss & Marino, 2001), elephants (Plotnik, de Waal, & Reiss, 2006), the great apes (Bard, Todd, Bernier, Love, & Leavens, 2006; Call & Tomasello, 2008; Kitchen, Denton, & Brent, 1996), and New World capuchin monkeys (de Waal, Dindo, Freeman, & Hall, 2005). Furthermore, being a social animal and engaging in cooperative behavior (Brosnan & Bshary, 2010; de Waal & Suchak, 2010; Melis & Semmann, 2010) required and enabled a more exible application of mental capacities (Blakemore, 2010), though it is still debated whether and to what degree animals developed at least rudimentary abilities of foresight, prospection, and theory of mind (e.g., Gilbert & Wilson, 2007; Hare & Tomasello, 2005; Miyata et al., in press; Osvath, 2010; Roberts & Feeney, 2009; Suddendorf, Addis, & Corballis, 2009a; Suddendorf, Corballis, & Collier-Baker, 2009b). Similar to memory, basic selfnon-self distinctions, such as the ones linked to physiological processes of immunity or digestion, are features of all viable species. The main unanswered question is the extent to which different species are capable of higher levels of conscious self-representations and self-awareness. Tulving (2005) has a clear position when stating I argue that only human beings possess
q
Corresponding author. Address: Physiological Psychology, University of Bielefeld, P.O.B. 100131, D-33501 Bielefeld, Germany. Fax: +49 5211066049.
E-mail address: hjmarkowitsch@uni-bielefeld.de (H.J. Markowitsch). 1053-8100/$ - see front matter 2010 Elsevier Inc. All rights reserved. doi:10.1016/j.concog.2010.09.005
This article is part of a special issue of this journal on Brain and Self: Bridging the Gap.
Please cite this article in press as: Markowitsch, H. J., & Staniloiu, A. Memory, autonoetic consciousness, and the self. Consciousness and Cognition (2010), doi:10.1016/j.concog.2010.09.005
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autonoetic episodic memory and the ability to mentally travel into the past and into the future, and that in that sense they are unique. (p. 4). Though debated, his position has been supported by a number of ndings also from human memory research (Botzung, Denkova, & Manning, 2008). Firstly, not all human beings possess the ability for mental time traveling: Patients with severe mental retardation or dementia may lack this capacity and patients with other disorders may show decits in integrating autobiographic memories with autonoetic consciousness and their selfhood. Examples are individuals with Asperger syndrome or autism. Tanweer, Rathbone, and Souchav (2010) found that adult Asperger individuals recalled in comparison to matched controls fewer events from their personal past and rated them much less specically (responding more frequently to them as being just known, but not remembered a nding typical for patients with amnesia; see, e.g., Bengner & Malina, 2008; Hirano, Noguchi, Hosokowa, & Takayama, 2002; Noulhiane et al., 2008; Yonelinas, Kroll, Dobbins, Lazzara, & Knight, 1998). They also made in comparison to the controls fewer social identity statements and provided more abstract, trait-linked identities. Individuals with autism were found to have an atypical neural response pattern to judgments about their self, when their brains were studied with functional neuroimaging methods (Lombardo et al., 2010). The authors also detected that the magnitude of neural self-other distinctions in the ventro medial prefrontal cortex was strongly related to the magnitude of early childhood social impairments (p. 611). Secondly, developmental studies have emphasized socio-culturallinguistic mechanisms that may be unique to the development of EAM. Small children similar to animals live initially in the here and now (Nelson, 2005a, 2005b). Their episodic-autobiographical memory (EAM) develops together with their self, theory of mind capacities, emotional conceptual knowledge and capacity for mental time traveling (Ghetti, DeMaster, Yonelinas, & Bunge, 2010; Rochat, 2010). Social context plays a critical role in the development of all the above neuro-cognitive functions. It was, for example, shown that simply listening to the voice from a tape recorder is not sufcient for learning early aspects of language, but instead infants require the social presence of a person (Adolphs, 2010; Kuhl, Tsao, & Liu, 2003). Theory of mind functions furthermore appear only after children have become experienced verbal communicators (Surian, Caldi, & Sperber, 2007, p. 580). The onset of EAM and the ToM capacities in the offspring (Nelson & Fivush, 2004) depend on the degree of elaboration of the reminiscing style of their mothers. The importance of the social component for the emergence of ToM capacities is also reected by ndings that in institutionalized children ToM capacity correlated with the adultchild ratio (Bedny, Pascual-Leone, & Saxe, 2009). Developmental changes of EAM (e.g. pertaining to autonetic consciousness) extend however beyond childhood into early adolescent years (Picard, Reffuveille, Eustache, & Piolino, 2009) and may include in late childhood (ages 812 years) the ability to suppress memories (Paz-Alonso, Ghetti, Matlen, Anderson, & Bunge, 2009). On the brain level these EAM developmental changes in humans (from infancy to early adulthood) are reected in the extensive structural and functional reorganization of different components of the neural networks supporting EAM, autonoetic consciousness and self-referential processing, ToM capacities and ability for emotional regulation (Shing et al., 2010). From a comparative cognitive-neuroscience perspective, frontopolar cortex (BA10) shows the biggest relative increase between great apes and human beings. BA10 activation recently has been found to be correlated with working memory capacity and general intelligence (Colom, Jung, & Haier, 2007). The basolateral nuclear group of amygdala shows a progressive enlargement from insectivores to prosimians and nally simians (Sarter & Markowitsch, 1985b, p. 348), while vice versa the centromedial nuclear group shows a clear regression along this phylogenetic scale. This is in line with ideas that in more phylogenetically evolved species the basolateral nuclear group expands to encompass higher cognitiveemotional functions such as EAM in the case of human beings (Cahill, Babinsky, Markowitsch, & McGaugh, 1995). One question which remains is concerned with the role of EAM in humans. Does indeed the EAM through its intrinsic feature of mental time traveling play a main function in the survival, as it has lately been emphasized repeatedly? And if the appearance of EAM is indeed adaptive, why did it not occur in other species? Is it in fact possible that certain claims of cognitive differences between humans and other species are the product of an underdeveloped experimental methodology rather than species differences per se? Or may it be the case that the main function of EAM is in fact social a suggestion that was put forth by several authors, though it has not been an explicit focus of extensive experimental investigations yet (Markowitsch & Welzer, 2009; Welzer & Markowitsch, 2005). A hint in favor of this hypothesis comes from the work with patients with Alzheimers dementia: Fargeau et al. (in press) found that the social self was impaired earliest in this patient group. In the current paper, after a presentation of memory systems and their neural correlates, we will provide a review of the relationship between EAM, autonoetic consciousness and self, by preponderantly drawing on the socio-culturallinguistic developmental model advanced by Nelson and Fivush (2004). We will then argue that EAM disturbances can result from deficits in the accurate re-collection of the encoding context, mental time traveling, emotional disturbances or self-related processing. By describing several EAM disturbances associated with both neurological and psychiatric diseases, we will demonstrate that many (especially severe) EAM impairments arise or exist in combination with dysfunctions in the realms of emotion, self, mental time traveling and social functioning. 2. Memory systems Memory is not unitary, but can be deconstructed along a time and content axis, respectively. Along the time axis, memory was traditionally divided into short-term and long-term memory. The short-term memory has a limited capacity of a few bits (47) (Cowan, 2000; Miller, 1956) and encompasses a time range of seconds to minutes. Any information that is not lost and exceeds the limited capacity of short-term memory is assigned to long-term memory stores. The above time-related
dichotomy of memory was later expanded by the addition of working memory by Alan Baddeley (Baddeley, 2000; Baddeley & Hitch, 1974). As captured by its name, working memory refers to working with memory this involves not only time-limited online holding of new information, but also retrieving portions of old, already stored information. Another time-related categorization of memory consists of the distinction between old and new and anterograde and retrograde memories, respectively. The compromised ability to access information that happened before the memory-impairing incident corresponds to retrograde memory impairment, while anterograde memory impairment refers to the compromised capacity to long-term acquire new information after the incident. A different facet of the relation between memory and time is concerned with the phenomenological experience of time in memory disturbances (inner time perception, mental time traveling). The classication of long-term memory systems along the content dimension has undergone several changes, especially since Tulving had proposed a distinction between episodic and semantic memory in 1972. Presently two overlapping classications dominate the memory research literature one was initiated by Larry Squire and the other one was advanced by Endel Tulving. In Squires classication, a main distinction is made between declarative and non-declarative memory. Under declarative memory, episodic and semantic memories that is (biographical) events and general facts are subsumed. Nondeclarative memory contains several other forms of memory, which are considered to be automatically processed. Tulvings content-based classication contains ve long-term memory systems, which are considered to build-up on each other phylo- and ontogenetically. According to the SPI-model (SPI = serial, parallel, independent) that was proposed by Tulving (1995), it is assumed that information is encoded serially into these systems, may be stored in parallel in different systems and can be retrieved independently of the system in which encoding occurred. These ve memory systems distinguish themselves by different levels of consciousness (such as autonoetic, noetic or anoetic) and distinct or partly distinct neural correlates. Wheeler, Stuss, and Tulving (1997, p. 335) dened autonoetic consciousness as the capacity that allows adult humans to mentally represent and to become aware of their protracted existence across subjective time. They differentiated autonoetic from noetic consciousness (knowing) which refers to the awareness of symbolic representations of the world, and from anoetic consciousness that describes the simple awareness of external stimuli. The latter form of consciousness approximates the one evoked by the following citation of Mesulam (2000): the existence of consciousness might be inferred when a living organism responds to environmental events in an adaptive way that is not entirely automatic (p. 93) (cf. Markowitsch, 2003). The hierarchy of the memory systems proposed by Tulving is depicted in Fig. 1. While excluding very basic forms of memories such as habituation, sensitization, classical conditioning, this hierarchy starts with procedural and priming memory systems two simple memory systems that are still devoid of the need for conscious reection upon the environment (anoetic). Procedural memory is mainly motor-based, but includes also sensory and cognitive skills (routines). Examples are riding a bike, skiing, playing piano, or reading words presented in a mirror-image. While procedural memory is largely an action-based memory system, the reverse is true for the priming system: Priming refers to a higher probability to identify stimuli, which were previously perceived in the same (perceptual priming) or a related way (conceptual priming). The perceptual memory system acts consciously (noetically), but on a presemantic level and relies on familiarity judgments. An example is the conscious identication of an apple without hesitance, no matter what color it has or whether it is already half eaten or not. Patients with semantic dementia, who lose the capabilities for language and semantic memory, may still be
Fig. 1. The ve long-term memory systems and their assumed brain bases (for further description see the text).
able to distinguish for example an apple from a peach or pear without the need to access semantic information, by accessing perceptual representations of information via the perceptual memory system. Semantic memory, which was also termed knowledge system or by Tulving and Markowitsch (1998) declarative memory, is context-free and refers to general facts. It is accompanied by noetic awareness. The denition of episodic memory has been submitted to several revisions over the years. While several decades ago the term episodic memory could be applied to laboratory stimuli with a specic embedding in time and place (Tulving, 1972), the eliciting of what and where and when information is no longer regarded as a sufcient condition for fullling the requirement for being an episode memory (episodicity). Nowadays episodic memory is dened as the conjunction of subjective time, autonoetic consciousness and the experiencing self (Tulving, 2005) and subsequently the episodic memory system is currently viewed as being equivalent to the episodic-autobiographical memory system (EAM). Though the term autobiographical is still at times used interchangeably with the term episodic, not all the components of autobiographical memory, however, have an episodic quality. Therefore a distinction is emphasized between autobiographical-episodic memory and autobiographical-semantic memory. The latter refers to knowledge of name, date of birth or self-traits and may be preserved or updated in spite of blocked access to episodic memory for personal events. In comparison to autobiographical-semantic memory that requires noetic awareness only, the EAM presupposes a higher level of consciousness (autonoetic). Autonoetic consciousness entails a sense of self in time and the ability to relive subjective experiences from the encoding context by mentally travelling back in time (Lemogne et al., 2006, p. 260). Given that most EAM reminiscences are affectively-laden, the reliving of the subjective experiences from the encoding context is usually intimately linked to an emotional evaluation of the signicance of these past experiences for oneself and with respect to ones own position in his social and biological environment. This emotional evaluation may in turn shape someones motivation for planning for the future and engaging in future acts. This emphasizes that EAM has not only a retrospective function, such as the conservation of certain conditions, their reproduction, and their localization in the past (Ribot, 1882 p. 10), but also a prospective one. The classications of Squire and Tulving, which were presented above, partly stem from opposing theoretical assumptions on the brains processing of episodic and semantic memory information. Squires theory emphasizes the commonalities between episodic and semantic memory processing, both from a behavioral and an anatomical perspective, while Tulvings theory stresses the dissimilarities. Apart from these two categorizations, we currently witness a search for new models for memory systems, such as a model based on processing modes (Henke, 2010) rather than consciousness. This search may partly be prompted by the grand challenge of consciousness (Seth, 2010), in particular the challenge posed by designing a testing instrument (methodology) that provides an objective estimate (measure) of the subjective phenomenon of autonoetic consciousness or mental time traveling, which can be used in young children and non-human beings (Perner, Kloo, & Rohwer, 2010; Roberts & Feeney, 2009). 3. EAM and the brain The formation of stable long-term EAMs requires several information processing stages (encoding and consolidation). Debate about the process of consolidation still exists, with some authors arguing that the process may extend to years (Haist, Bowden Gore, & Mao, 2001). Once the information is consolidated, it is stored and then usually available for retrieval. Each retrieval is followed by re-encoding of the EAM in the newly present context. According to adherents to the reconsolidation theory, consolidated memories that are recalled by a reminder enter after retrieval a vulnerability (labilization) phase, during which they might become susceptible to disruption or strengthening or incorporation of new information (Forcato et al., 2010); this is followed by a process of stabilization (reconsolidation).The reconsolidation theory constitutes the basis for pharmacological studies in humans that aim to weaken the vivid and disturbing traumatic memories associated with certain forms of post-traumatic stress disorder by interfering with their assumed post-retrieval reconsolidation (Brunet et al., 2008). The incorporation of new information may have as the result a re-contextualization of the retrieved material (Modell, 2006) or the introduction of falsied details that may lead to false memories or confabulations (Loftus, 2000; Loftus & Hoffman, 1989; Loftus & Pickrell, 1995; Borsutzky, Fujiwara, Brand, & Markowitsch, 2008, 2010). Freud wrote about a re-transcription of memories in accordance to fresh circumstances (Masson, 1985, p. 207). And Bartlett (1932) and later Tulving (1995, 2002, 2005) and Schacter (2001) noted that human beings construct and reconstruct their personal memories, perhaps in an attempt to support current aspects of the self and match future goals that are coherent with one individuals goals, self image and system of beliefs (Conway, 2009). Edelman (1998) remarked that every act of memory is to some degree an act of imagination a remark that later on was substantiated by ndings that the hippocampal formation may also be involved in mental construction of complex scenes. Furthermore, Edelman wrote that memory has the properties that allow perception to alter recall and recall to alter perception (1998, cf. Modell, 2006, p. 37). This sentence hints to an important feature of EAM namely its state-dependency. This feature implies that memories are optimally retrieved if the environmental and mood and physical state conditions match those during encoding. Inspired by Semons description (1904), Tulving (1983, 1985, 1995) coined this state-dependency of memory retrieval ecphorizing. Tulving (1983) employed the term ecphory to describe the process by which retrieval cues interact with stored information so that an image or a representation of the information in question appears. A mismatch between encoding and retrieval conditions may lead to a spectrum of memory retrieval disturbances, ranging from common tip-of-the-tongue phenomena to complete pathological retrieval blockades (such as in dissociative amnesic conditions; see below).
The processes associated with the EAMs formation and retrieval engage a widespread network that contains several brain structures and long-distance ber connections (Table 1). Some of these structures show a preferential specialization for mnemonic processing, while others (such as the amygdala) act as hubs that integrate emotive and cognitive functions. This supports the current theoretical approach to memory processing, which blends elements from both the previous localizationist and Gestalt-like theories. Encoding and consolidation of EAM depend on the integrity of brain networks that comprise several structures and their ber connections, including medial temporal lobe, diencephalon and parts of prefrontal cortex. The role of hippocampal formation in the encoding and consolidation of EAM (and semantic memory) has been substantiated by an overwhelming number of ndings from patients with brain damage as well as healthy people who underwent functional brain imaging investigations during test paradigms that tapped on EAM functions. As emotion is according to several authors intrinsic to EAM (see Bluck, Alea, Habermas, & Rubin, 2005), it is not surprising that both the Papez circuit and the basolateral limbic loop (mediodorsal nucleus of the thalamus, subcallosal area, amygdala and interconnecting bers) are engaged in the formation of EAM. The Papez and the basolateral limbic circuits constitute a group of bottleneck structures (Brand & Markowitsch, 2003) that are interconnected and of high relevance for the extraction of the emotional, biological and social signicance of newly incoming information. While prefrontal regions have also been implicated in the formation of EAMs, they are nevertheless more strongly associated with the retrieval of personal events. Recruitment of limbic structures was also evidenced during the retrieval of EAM. Some limbic structures (such as the hippocampus) have been ascribed functions in binding specic details of past events, while others (such as the amygdala) have been opined to charge sensory information with appropriate emotional cues, in order to guide successful memory re-collection of emotionally signicant events. Several studies found that the combined activation of right-hemispheric fronto-temporal regions serves as trigger stations for retrieving stored EAM events (Brand & Markowitsch, 2008; Fink et al., 1996; Kroll, Markowitsch, Knight, & von Cramon, 1997; LaBar & Cabeza, 2006). The corre-
Table 1 Structures involved in EAM processes. Structure Telencephalon, cortical (principally cortical midline structures) Hippocampal formation Entorhinal, perirhinal, parahippocampal cortex Anterior cingulate cortex Posterior cingulate/retrosplenial cortex, supracommissural hippocampus Insula Telencephalon, subcortical Amygdaloid body Basal forebrain (including the septal nuclei, diagonal band of Broca, basal nucleus of Meynert, i.e., cholinergic nuclei) Claustrum Diencephalon (subcortical midline structures) Mediodorsal thalamic nucleus Anterior thalamic nucleus Nonspecic thalamic midline nuclei (e.g., paratenial nucleus) Mammillary nuclei Functional implications (Episodic-autobiographical) memory, spatiotemporal integration, prospection, part of DMN (Semantic) memory Attention, ToM, awareness (VEN in its anterior area), emotion, reward, selfreference EAM, ToM, self-referential processing, part of DMN, imagination, familiarity Sensory-motivational integration, consciousness, reward, empathy, self processing, VEN Emotional-tagging of EAM, self-referential, imagining the future?, emotional awareness, reward and punishment EAM, inhibition, emotional evaluation, inner time perception, reward, social memory Support for conscious processing, sensory integration Encoding/consolidating EAM, consciousness, sleep, emotion, self-referential processing Emotional avoring, attention, support in encoding and consolididating EAM, consciousness? Consciousness, EAM? Encoding/consolidating of EAM, emotion?
Associated regions, especially of the expanded limbic system (paralimbic cortex) Orbitofrontal, medial prefrontal cortex EAM, self, ToM, prospection, part of DMN, time, monitoring veracity or feeling of rightness of a memory, episodic prospection of future rewards, emotional evaluation, social memory, accuracy of self-evaluation, temporal context, cortical midline structure Prefrontal cortex (esp. inferolateral and ventrolateral portions) Support in encoding and retrieving of EAM and semantic memory, ToM, part of mirror neuron system, consciousness, self, part of DMN, EAM retrieval, strategic search of memory, prospection Temporal pole Initiation of recall, memory-related sensory integration, ToM, self Temporo-parietal junction area Involved in EAM retrieval (recollective quality, i.e., vividness, condence), ToM, prospection, part of DMN Precuneus Imagination, ToM, self, EAM Lateral temporal cortex Memory storage (semantic and EAM), part of DMN Inferior parietal lobule Part of DMN, EAM, ToM, self?, mirror neuron system Abbreviations: DMN, default mode network; EAM, episodic-autobiographical memory; ToM, Theory of Mind; VEN, von Economo neurons. For explanations see text.
sponding regional complex in the left hemisphere seems to trigger the retrieval of semantic old memories (Markowitsch, Calabrese, Neufeld, Gehlen, & Durwen, 1999). The preponderant engagement of the right hemisphere in the retrieval of EAM has been linked to several factors, including the ascribed role of the right hemisphere in the processing of emotions and self-awareness (Calabrese et al., 1996; Feinberg & Keenan, 2005; Kaplan, Aziz-Zadeh, Uddin, & Iacobini, 2008; Keenan, Rubin, Racioppi, Johnson, & Barnacz, 2005; Keenan, Wheeler, Gallup, & Pascual-Leone, 2000; LaBar & Cabeza, 2006; Markowitsch, 1999; Schore, 2002). The right hemisphere was hypothesized to be preferentially involved in sympathetic arousal, such as the one that would result from negative feedback, while the left was preferentially linked to parasympathetic quietude (Allman et al., 2010; Wittling, 1997). Several authors have provided evidence for the involvement of the left hemisphere in parasympathetic inuences on cardiovascular functioning, while the right hemisphere was shown to mediate the sympathetic regulation of cardiovascular activity (Foster, Drago, Ferguson, & Harrison, 2008; Wittling, 1997; Wittling, Block, Genzel, & Schweiger, 1998). Anomalies in resting heart rate have been studied in relationship to aggressive behavior and emotional processing in psychopathy and antisocial personality disorder and postulated to arise from a right hemispheric dysfunction or an altered inter-hemispheric connectivity (Raine, 2003; Raine et al., 2003). Alexithymic traits consisting of a difculty with identifying (especially negatively valenced) emotions and feelings, and distinguishing between feelings and the bodily sensations of emotional arousal have also been linked to a dysfunction of the right hemisphere (Moriguchi et al., 2006; Schore, 2002) or an impaired interhemispheric transfer (Romei et al., 2008). They have been connected to a history of early trauma and a heightened susceptibility for dissociative disorders (Staniloiu, Markowitsch, & Brand, 2010). Interestingly, the right uncinate fascicle that links portions of the frontal and temporal lobe was found by a histopathological study to contain 33% more bers and be 27% larger in the right hemisphere than in the contralateral one (Highley, Walker, Esiri, Crow, & Harrison, 2002). The ventral portion of right uncinate fascicle is involved in ecphorizing affect-laden personal events. According to some authors, the uncinate fascicle may also contribute to the formation of memories (Sepulcre et al., 2008). The ventral branch of the uncinate fascicle is additionally a component of an emotional processing circuitry that connects the amygdala with the orbitofrontal cortex and the anterior cingulate cortex. As opposed to other brain ber connections, the uncinate fascicle matures later and more slowly and may continue its development beyond the age of 30 years (Lebel, Walker, Leemans, Phillips, & Beaulieu, 2008). This may enable a higher structural plasticity in relationship to a variety of environmental inuences, including physical or psychological stress-related insults. Microstructural abnormalities of the uncinate fascicle were for example reported in children, who were raised in a neglectful environment (Govindan, Behen, Helder, Makki, & Chugani, 2010). Levine et al. (1998, 2009) described a case of isolated dense retrograde EAM covering the entire life, which occurred after a severe traumatic brain injury and was associated with a focal lesion of the frontal portion of the right uncinate fascicle. Despite normal performance on standard anterograde memory tests, the above-mentioned patient reported a feeling of disconnection from the post-accident autobiographical events. Subsequent rened testing of his anterograde EAM revealed that he assigned signicantly less remember-ratings to his post-injury autobiographical events in comparison to normal subjects, suggesting an impaired rst-person autonoetic connection with his post-accident explicit memories of personal events. Fujie et al. (2008) reported both memory and emotional recognition impairments in amnestic MCI together with abnormalities of the uncinate fascicle. In agreement with the idea that white matter adjacent to cortical associations areas that matures later also undergoes an earlier age-related decline in myelination, Davis et al. (2009) recently found evidence via diffusion tensor tractography of a stronger age-related microstructural white matter change (demyelination) of the uncinate fascicle (especially the right one) in comparison to other ber tracts (such as inferior longitudinal fascicle, cingulum, splenium). These results point to the importance of taking into consideration the integrity of fronto-temporal connections, when performing or analyzing studies that investigate the effects of aging on emotional processing or EAM. We demonstrated in several studies that the retrieval of emotionally-laden autobiographical events relies more on the right temporo-frontal region than on the left (Calabrese et al., 1996; Fink et al., 1996; Kroll et al., 1997; Markowitsch et al., 1993; Driessen et al., 2004). Botzung, Rubin, Miles, Cabeza, and LaBar (2010) also found a right-hemispheric amygdalar activation during the retrieval of highly emotional memories. In patients with dissociative amnesia a condition that is accompanied by a failure of integration of emotive with memory processing functions we found evidence for right hemispheric dysfunction, such as a hypometabolism of the right inferior lateral prefrontal cortex (Brand et al., 2009). Similarly, Tramoni et al. (2009) found subtle structural right-hemispheric prefrontal white matter abnormalities in a case of functional amnesia that occurred on a background of signicant psychological stressors. In a study that investigated the neural correlates of visual retrieval perspective in EAM, Eich et al. (2009) visualized an increase in the right posterior amygdala activity during the recall of eld (rst person perspective) memories, which was interpreted as a higher degree of subjective emotionality, associated with these memories in comparison to those that are retrieved from a third person (theatrical or disembodied) perspective (Northoff et al., 2006). This argument echoes previous suggestions that the boundary between self and not-self is one s emotional attitude about an object or thought (Barresi, 2002, quoted by Northoff et al., 2006, p. 453) and that self-referential processing in subcortical structures may in particular be intimately linked to emotional processing. Along with the emphasized role of the right hemisphere in several self-related tasks (such as self-face processing and selfregulatory control) (Keenan et al., 2000; Schore, 2002; Marsh et al., 2006), a study that compared the retrieval of autobiographical episodes with the retrieval of ctitious episodes visualized a right-hemispheric activation of the amygdala only during the recall of personal authentic events, while the recall of ctitious material activated the retrosplenial/precuneus
area only (Markowitsch, Thiel, et al., 2000). Another study that investigated the neural correlates of deception showed right prefrontal activation associated with rehearsed lies that were part of a coherent story (Ganis, Kosslyn, Stose, Thompson, & Yurgelun-Todd, 2003). Although the latter results could be subjected to different interpretations, one speculation is that through repeated rehearsal, triggered by motivated agendas, self-related deceptive material may become familiar to our narrative self to the point that the deceptive material is not only part of a story we tell others, but also part of a story we tell to ourselves and in some cases a story we may live by. The idea that the right prefrontal cortex might disproportionately contribute to processing the familiarity of information has been advanced by some authors (Dobbins, Foley, Schacter, & Wagner, 2002; Dobbins, Simons, & Schacter, 2004). Recently, Benoit, Gilbert, Volle, and Burgess (2010) showed that there is a relation to closeness and similarity of others to oneself, which leads to more or less activation of the medial rostral prefrontal cortex. In the same study self judgments were also associated with greater activation of right lateral rostral prefrontal cortex and stronger activation of the right insula. In addition to medial rostral prefrontal cortex a cluster in right lateral prefrontal cortex also showed less activity for contrasting self with other judgments to the extent that the other was perceived as more similar. Another pursued explanatory avenue for a preferential right hemispheric lateralization during the retrieval of explicit memories of personal events concerns the relationship between EAM and autonoetic consciousness. Keenan et al. wrote in 2005: The evidence that there is a right-hemsipheric bias in terms of self-awareness is overwhelming. (p. 700) and further, that the right hemisphere is dominant for higher-order consciousness (p. 702). In a recent article, Allman et al. (2010) reported that the right hemisphere of the human postnatal insular-frontal cortex contains signicantly more von Economo neurons (VENs) than the left. The VENs, which have been described in the insula and anterior cingular cortex (ACC), have been ascribed functions in consciousness, intuition, social cognition and regulation of appetite (Allman et al., 2010). Their degeneration was for example found to be associated with loss of emotional awareness, loss of empathy, altered self-consciousness and aberrant eating behaviors in patients with a behavioral variant of fronto-temporal dementia (Seeley, 2008). Especially the right fronto-insular cortex atrophy in these patients was found to correlate with the presence of core clinical symptoms. It was speculated that these symptoms arise from failures to integrate visceral guidance cues with behaviour (Seeley, 2008, p. 704), a speculation that is congruent with other authors idea that self-consciousness is in many ways intertwined to bodily consciousness and that unknowing and knowing consciousness are in fact two poles of a continuum (Vandekerckhove & Panksepp, 2009). A relationship between impairments of conscious access and integrity of large white matter bundles, particularly involving prefrontal cortex, was described by a new study (Reuter et al., 2008). Damage preferentially located to right parieto-occipital or right temporo-occipital cortex was also identied as a neurosubstrate of autoscopic phenomena (Blanke & Metzinger, 2009). While most of the above mentioned results hint to a preferential lateralization for the retrieval of EAM, self-awareness and other self-related tasks, other ndings emphasize the modulation of this lateralization by a wide range of variables. In the case of EAM, these variables comprise gender, valence, nature of stimuli, and remoteness of memories, testing language (in case of bilinguals), visual retrieval perspective and genetic polymorphisms (Buchanan, Tranel, & Adolphs, 2005; Eustache et al., 2004; Markowitsch, 1998/1999; Markowitsch, Vandekerckhove, Lanfermann, & Russ, 2003; Piefke, Weiss, Zilles, Markowitsch, & Fink, 2003). Furthermore, it is increasingly acknowledged nowadays that the networks underpinning mnemonic processing, self-referential processing and consciousness, include structures that are located in both hemispheres. Especially recent studies that aimed to capture the neural correlates of EAM retrieval in settings that approximate the real-world ones (Botzung, LaBar, Kragel, Miles, & Rubin, 2010) have supported the view that EAM retrieval is basically organized bilaterally (Vandekerckhove et al., 2005). Powell, Macrae, Cloutier, Metcalfe, and Mitchell (2009) pointed to the self as a construct of collection of distinct mental operations distributed throughout the brain. They made the important distinction between the self as containing conceptual knowledge of ones own personality traits and the self as an agent (e.g., freely choosing objects or watching passively as one is chosen). This second view of the self is also followed in the work of Vogeley (David et al., 2006; Vogeley et al., 2004). Northoff and Panksepp (2008) proposed that subcortical-cortical midline structures are engaged in self-related processing that supports a primal form of self-representation (a core self), which may exist in other mammalian species as well. Craig (2009) hypothesized that the insula supports a so-called sentient (feeling) self. The fronto-parietal human mirror system areas have been ascribed functions in the experiential understanding of others and were opined to be able to effectively function as bridges between self and other, by co-opting a system for recognizing the actions of others to support self representation functions (Uddin, Iacoboni, Lange, & Keenan, 2007, p. 154). Furthermore it has been pointed out that the human mirror neuron system shares connections with the default mode network (DMN). The DMN comprises several areas of the brain areas (ventral-medial prefrontal cortex, dorso-medial prefrontal cortex, posterior cingulate cortex, retrosplenial cortex, inferior parietal lobule, lateral temporal cortex and hippocampal formation) that display high baseline metabolic activity at rest and are considered to be important for mind-wandering, introspection, prospection and EAM processing. Though not recognized as a core structure of the DMN network, areas of the lateral prefrontal cortex have been reported to be recruited during EAM as well as self-referential processing (Spreng, Mar, & Kim, 2009). As relatively recently Northoff et al. (2006) remarked, distinct concepts of self differ in the class of stimuli and their specic material or content reecting what is called different domains; what remains unclear, however, is what unites these distinct concepts of self allowing us to speak of a self in all cases (p. 440). Feinberg (2005) proposed that we experience a unied and single self, as opposed to multiple selves from multiple hierarchical levels, because lower-order as well as higher-order elements are part of the nested hierarchy of the self (p. 47). Pinker talked about I as being the unity of selfness over time (1997, p. 564). We argue that the experience of the unity of selfness or the personal sameness
over time (Locke, 1689/1964) is closely linked to EAM and autonetic consciousness, whose appearance and maturation enable us to override the awareness of discontinuity or multiplicity and lead us to attain and conserve a feeling of a cohesive personal identity over time and across contexts (Bromberg, 1994). 4. Memory development and the development of a self The development of memory systems starts with systems that involve simple, unconscious (anoetic) processing of information (procedural memory, priming), continues with conscious (noetic) systems and apparently culminates with the EAM system which requires autonoetic consciousness (cf. Piolino, Desgranges, & Eustache, 2009). The EAM system is considered the last human ontogenetic acquisition, but also the highest achievement on the phylogenetic ladder (Tulving, 2005). The latter view is also reected in the cautious employment of the term episodic-like memory to refer to the mnemonic abilities of species (e.g. scrub-jays) that show that they know some scientists even use the expression recall the what and where and how long ago of a salient event (Clayton & Dickinson, 1998; Roberts & Feeney, 2009). Despite increasingly sophisticated studies that have been carried out to examine animals ability for mental time traveling (Roberts & Feeney, 2009), there is at this point in time no conclusive evidential material to counteract the assertion of Tulving (2005, p. 9) that the EAM is probably unique to humans. The present point is that mental time travel always occurs not only in subjective time but also in mental space, and that the mental space of the remembered past and imagined future may be different from the present space. Individuals with (autonoetic) episodic memory can, if the situation calls for it, think here and now about personal happenings in other places and other times. Therefore, if we ask whether other animals have episodic memory, we ask, among other things, whether they can also do so. (Tulving, 2005, p. 7). Tulving (2000, 2002, 2005) assumed the existence of a hierarchy of memory systems which principally follows that of the memory systems listed in Fig. 1 from left to right. It is perhaps on the rst glance counterintuitive that memory for facts (semantic memory) is an earlier ontogenetic acquisition than EAM, given the common observation that the repetition of single similar episodes leads to their generalization and thereby semantization (Fig. 2). However, ndings from human developmental studies emphasize that children rst acquire the semantic memory system and only thereafter their EAM neuro-cognitive capacities emerge (Nelson, 2003, 2005a, 2005b; Nelson & Fivush, 2004; Markowitsch & Welzer, 2009). When infants are asked to talk about what happened yesterday (What happened during lunch yesterday?), they usually make very general semantic statements and do not provide vivid episodes. The emergence of EAM is dependent on or takes place in concert with the development of language and conceptual knowledge, higher levels of self and self-understanding, the ability to understand the feelings and intentions of others, executive functions, working memory, maternal reminiscing style, the capacity for mental time traveling and the maturation of the nervous system (Markowitsch & Welzer, 2009; Morrison & Conway, 2010; Nelson & Fivush, 2004; Picard et al., 2009). The relationship between self and episodic-autobiographical memory is powerful and dynamic. It can be discussed from different angles, such as the phenomenological momentanous experience of self accompanying the act of remembering of the rememberer or the relationship between EAMs and the creation and articulation of narratives that support outlasting self constructs or
Fig. 2. Relations between EAMs and semantic memories. It is assumed that most semantic memories are strengthened by repetition, while the reverse is true when one is repeatedly exposed to similar events.
self-concepts (moral, social, etc.). Similarly to the proposed hierarchical development of the memory systems, several authors put forth models for a hierarchy of self development. This may start with a very basic proto-self that is grounded in the sensory and motor domains (Panksepp, 1998), continue with a pre-linguistic, affective core self (Damasio, 1999; Northoff & Panksepp, 2008), be followed by a cognitive self (Howe & Courage, 1993) and thereafter by an autobiographical (Damasio, 1999) or narrative self (Gallagher, 2000) (Fig. 3). The appearance of a cognitive self late in the second year of life (Howe & Courage, 1993) seems to be a necessary, but not sufcient condition for the EAM development. The EAM establishment entails in addition an enduring sense of self across time. As detected through the delayed self recognition paradigm of Povinelli and his colleagues (Povinelli, Landau, & Perilloux, 1996), an enduring sense of self is typically described in most 4 and 5 years old ones. Around age 4 children also typically succeed on a standard false belief task (Baron-Cohen, Leslie, & Frith, 1985) a task that may require metarepresentational abilities (Perner, 2000). The autonoetic experiencing self or the rememberer features the capacity to exibly travel in mental time and space and a superior awareness of oneself as a person in a social (and biological) environment, with a past and a future. Several studies have concluded that EAM, autonoetic consciousness and mental time traveling appear between the age of 4 to 5 years approximately (Perner, Kloo, & Gornik, 2007; Perner & Ruffmann, 1995; Piolino et al., 2007). Morrison and Conway (2010) suggest that the formation of conceptual knowledge that is abstracted from details in early personal memories needs rst to be established, before specic EAMs can be retrieved. Childhood amnesia would therefore be related to the time until this ability is formed. (This corresponds to Tulvings ideas of state-dependency and encoding specicity; Tulving, 2005; Tulving & Thompson, 1973). The development of EAM and mental time traveling extends however beyond the above mentioned age bracket. Piolino and co-workers investigated the establishment of EAMs and autonoetic consciousness in children up to the age of 11 years (Picard et al., 2009; Piolino et al., 2007). The authors proposed that mental time traveling may be one of the last features of the EAM to become fully operational. Picard et al. (2009) found that the EAM still improves in quality and quantity considerably over the rst ten years of life. Children provide more factlike information when requested to provide old episodes; only the recent episodes of older children are more detailed and authentic, while the remote ones remain mainly semantic. Probably, on the brain level, the establishment of inhibitory processes, acting from the late developing prefrontal cortex (Gibson, 1991; Huttenlocher, 1994) on the posterior (temporo-parietal) association cortex partly accounts for the developmental changes of self-related processing, EAM, and autonoetic consciousness. Findings from lying in a coherent way, where the right prefrontal cortex seems to play a pivotal role, add to this idea (Ganis et al., 2003). The reverse process seems to occur in older age, where autobiographical memories become more semantized (Piolino et al., 2006) (cf. Fig. 2). As children grow up they acquire greater gist memory that may however increase their susceptibility for false recall or recognition of information (Khnel, Woermann, Mertens, & Markowitsch, 2008). The development of EAM may also enhance the ability of children to fantasize (to construct complex mental scenes) as well as to deceive. The capacity for deceiving has not only been linked to EAM (Ganis et al., 2003), but also to theory of mind and empathizing abilities as well as the capacity for emotional regulation (Adolphs, 2010). Profound changes of the sense of self take that take place from adolescence (Sebastian, Burnett, & Blakemore, 2008) to early adulthood are also reected in the quality of EAMs (Oddo et al., 2010). Beginning with early adolescence the individuals start to habitually incorporate the perspectives of others (e.g. peers) during the process of self-evaluation (J. H. Pfeifer,
Fig. 3. Sketch on possible relationships between states of mind and complexity and diversity of representation.
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Dapretto, & Lieberman, 2010). Along this line it was found that the retrieval of recent memories in young women engages the ventro-medial prefrontal cortex, a region known to be recruited not only by self-evaluating tasks, but also by the evaluation of perceived similar others. The complex relationship between the development of self and EAM is captured by the model advanced by Nelson and Fivush (2004), which depicts the stages of EAM development from birth to age ve (Fig. 4). The fact that the model had been conceived before Tulvings last revision of EAM (2005) and therefore reected Tulvings denition of episodic memory from the last century (e.g., Tulving, 1995), in which the embeddedness of episodic memory in time and place was emphasized, does not diminish its powerful argumentation for the important contribution of the socio-cultural milieu to the emergence of EAM (Fig. 4). As Courage and Cowan (2009) pointed out, the concept of context implied by the encoding specicity principle (Tulving & Thompson, 1973) is viewed in the model of Nelson and Fivush (2004) more broadly to encompass family, society and culture (Fivush & Nelson, 2004; Nelson, 2007; Nelson & Fivush, 2004; Pope, Poliakoff, Parker, Boynes, & Hudson, 2007; Zhu, Zhang, Fan, & Han, 2007). The way in which the mothers structure their conversations when they engage in reminiscing with their offspring has a major impact on the onset of earlier EAM and the manner in which the offspring narrate their personal past. The maternal reminiscing style is, however, partly mediated by the nature of the motherchild attachment, with the consequence that insecure attachments may negatively impact on the quality of EAMs. As Valentino, Toth, and Cicchetti (2009) showed, children with a history of abuse or neglect manifest a reduced EAM specicity (overgeneral memory effect). The relationship between the maternal reminiscing style and the development of the EAM of the offspring however extends beyond the familial context to encompass the cultural context (Harbus, 2010). Cultural differences were reported in EAM and were linked to childrearing practices, views of selfhood and past, preferences for high arousal versus low arousal emotional states or degree of attention to social context (Gutchess & Indeck, 2009; Nelson & Fivush, 2004). For example Asians EAMs, in contrast to the ones of Caucasian Americans, were found to emphasize more social interactions and contain more people (Wang & Conway, 2004). Peterson, Wang, and Hou (2009) observed that Canadian children, who talked about re-collections of their EAMs showed a more autonomous self-construal style, while Chinese children showed a more relational one. As Adolphs (2010) pointed out, identifying the source of cultural difference requires the consideration of at least three potential variables: the ethnicity, the culture of origin and the cultural milieu where the testing is performed. Part of the ethnic differences may in fact reect a regional distribution of a certain genetic polymorphism, which may bias the brain connectivity between structures involved in cognitive or emotive functions. The short (S) allele variant of the promoter region of the serotonin transporter gene (5HTTLPR) was for example found to be more prevalent in Japanese populations than in European ones. The S allele has been linked to alterations in microstructure of the uncinate fascicle, decreased gray matter volume in the amygdala and medial prefrontal cortex and altered functional connectivity between the amygdala and the prefrontal cortex (Caspi, Hariri, Holmes, Uher, & Moftt, 2010). In the EAM domains, the 5HTTLPR polymorphism has been hypothesized to modulate the retrieval perspective (Lemogne et al., 2009). The claim that ones individuals representations of self (independent versus interdependent self-construal) is inuenced by the culture of origin has received support from functional neuroimaging studies. Zhu and colleagues (2007) compared subjects from an individualistic (Western) culture and subjects from a collectivistic (Chinese) culture during a task where they had to judge personal trait adjectives with respect to self, mother, or a public person. They found that the medial prefrontal and anterior cingulate areas were activated
Fig. 4. Hypothetical relations in developments from 1 to 5 years of age leading to the emergence of autobiographical memory. Larger arrows indicate more direct inuences; double-headed arrows indicate reciprocal inuences. Years (yr.) in the bottom scale indicate approximate ages when inuences come into play on average in normal development. Areas above the center are presumed to be more endogenous and those below more exogenous as sources of development. Figure and gure legend copied from Fig. 1 of Nelson and Fivush (2004).
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strongest during self-related judgments in subjects of both cultures. In contrast to subjects from Western cultures, in Chinese subjects the medial prefrontal cortex was also activated when making judgments about the mother. The authors concluded that Westerners use the medial prefrontal cortex exclusively for representing their own self. However, Benoit et al. (2010) recently showed that the degree of activation of the medial rostral prefrontal cortex depends on the proximity and similarity of others to oneself. Given the increasing economic globalization and the accompanying frequent migration or relocation, it has been emphasized that one should broaden the concept of culture-sensitivity in human cognition and consider the dynamic shaping and re-shaping, which is not only imposed by culture, but in feedback also by the individual (Han & Northoff, 2008; Vogeley & Roepstorff, 2009). Studies of bilingual or bicultural individuals and migrants suggest that the language and the degree acculturation (that is partly measured by language prociency) can impact on memory performance, cognitive strategies, selfconcept and self-other distinctions. Bilingual Chinese children were found to recruit more elaborate and self-focused EAMs when speaking in English compared to when speaking Chinese (Wang, Shao, & Li, 2010). Kobayashi, Glover, and Temple (2008) assessed the response towards false-belief tasks (in the form of x thinks that y thinks that . . .) in early (children) and late (adult) bilingual Japanese. While they found medial prefrontal cortex activation in the children who had acquired the second language early, adults had more diverse brain activations, suggesting a more automatic processing in the adults and differences in activation dependent on the language used. Perner and Aichhorn (2008) conclude from the results of Kobayashi et al. that the brain physiology . . . might be more liable to cultural and linguistic variation with intriguing links to similar developmental variations (p. 125). Discussions about the role of language in shaping memory (and cognition in general) have a long history. Tulving (2005) asserted that language is not a necessary condition for EAM, but acknowledged that it supports and enriches its development, while others proposed a stronger connection between language and EAM (Nelson & Fivush, 2004; Suddendorf, Addis, et al., 2009). In the last few years it was found that a tribe of Amazon Indians the Pirah lacks many of those attributes which are commonsense for our understanding of the selfhood and existence across time. They do not think of past and future and consequently cannot imagine the life of persons from history or past (Everett, 2005, 2008). The language spoken by the Pirah reportedly only comprises two rudimentary temporal markers (Everett, 2005; Suddendorf, Addis, et al., 2009). These ndings are relevant in the light of the proposal of Suddendorf, Addis, et al. (2009) that language and mental time traveling might have co-evolved. Language might have been preceded by the mimetic gesture. And mental time traveling might have been rooted in a form of embodied cognition, grounded on species-specic propensities for moving in a particular sense. In keeping with the general idea that episodic memory grows out of semantic memory (Tulving, 2005), the ability to mentally travel in time has been viewed as an extension of mental travel in space. The foundation of mental time traveling indeed may include a spatial component. This has been suggested from a comparative biological perspective, from an experimental perspective, and from the results of patients with vestibular nerve damage accompanied by selective hippocampal degeneration. We have argued for a functional shift in hippocampal function from a predominant processing of spatial cues (as in rodents) to a predominant temporal processing in higher primates (Markowitsch & Welzer, 2009; Tulving & Markowitsch, 1997, 1998). Miles, Karpinska, Lumsden, and Macrae (2010) asked subjects to daydream while viewing a display that elicited an illusion of self-motion (vection). Backward vection evoked thinking about the past, while forward vection resulted in preponderantly future-oriented thoughts. And for patients with hippocampal degeneration decits in virtual maze performance and spatial memory were reported, indicating that in addition to the more recent functional attributes of the hippocampal formation, some more phylogenetically ancient ones are still present in human beings (Brandt et al., 2005; Hfner et al., 2007, in press). Another phylogenetically old brain structure, the cerebellum, has been found to play a signicant role in coordinating spacetime relations (Oliveri et al., 2009). The capacity for mental time traveling was opined to offer a survival advantage and perhaps be the main answer for the existence in humans of an EAM system that is fragile and only offers an imperfect repository of the past. Laboratory studies, which investigated the neural correlates of constructive episodic future thinking, conrmed the prospective function of EAM by nding common, but also distinct neural correlates involved in EAM retrieval and episodic future thinking (Addis, Pan, Vu, Laiser, & Schacter, 2009). However some authors questioned the main survival function of EAM, by emphasizing that autonetic self-awareness might bring with it the awareness of ones own nitude a possibly fatal piece of knowledge, full understanding of which would preclude any motivation to survive (Adolphs, 2010, p. 762). So how could these two views be reconciled? Roberts and Feeney depicted mental time traveling like a bicone and emphasized that typically our plans in the forthcoming weeks are more detailed and those for the years to come are more vague and overgeneralized. Consistent with the idea that the healthy individuals generally tend to expect more positive events than is rational to expect, a recent imaging study that related medial prefrontal cortex activity to self-evaluation advanced the hypothesis that the default network activity might be biased towards a positive self-evaluation, instead of an accurate one (Beer, 2007). A positivity memory bias was reported in elderly people, though the neural underpinnings of it are still under debate (Addis, Leclerc, Muscatell, & Kensinger, 2010). Furthermore it has been hypothesized that self-awareness could only have emerged in parallel with mechanisms such as faith and religion (Varki, 2009). Gilbert and Wilson (2007) pointed out to another aspect of future thinking. They remarked that people tend in fact to make several errors during simulation of future events, such as reliance on the most recent memories or the most salient ones. They also offered as a solution for the optimization of future simulation, namely getting advice. Suddendorf, Corballis, et al. (2009) commented on the authors suggestion, stating that it resonates with our proposal that one adaptive function of language may be to allow us to improve our mental time travel by drawing on the descriptions others can offer of what the future may hold (p. 1322). They subsequently proposed that the
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main function of EAM may in fact be a social one. The possible contribution of EAM to social regulation was also implied by a recent study that showed that severe EAM decits interfere with the updating of moral character judgments of others and consequently may inuence the way we perceive and behave towards others (Croft et al., 2010). 5. Disturbances of (autonoetic) consciousness and EAM in neurological diseases Similar to memory and self, consciousness is not regarded as a unitary phenomenon. Seth (2010) distinguishes between conscious level and conscious content. Conscious level is measured by tools such as Glasgow Coma Scale and ranges from brain death over coma to full alertness and awareness of ones environment. Conscious contents differentiate themselves into multimodal sensory contents related to the world; experiences of selfhood, volition and agency, and affective and somatic perceptions (Seth, 2010, p. 1). Damasio (1999) proposed a core consciousness and an extended consciousness, respectively and, as mentioned above, Tulving repeatedly described three forms of consciousness: anoetic, noetic and autonoetic. Block (1995) further made the distinction between phenomenological consciousness and access consciousness. The quest for a neural correlate of level and content consciousness prompted studies of the phenomenal aspects of perception, sleep and wakefulness and vegetative and minimal conscious states as well as investigations of self- awareness (Newen & Vogeley, 2003) in healthy humans or clinical populations with different degrees of cognitive impairments or neurological or psychiatric diseases (Alkire, Hudetz, & Tononi, 2008; Coleman et al., 2009; Langnickel & Markowitsch, 2006; Markowitsch, 1995, 2000, 2003, 2005; Reinhold & Markowitsch, 2007; Staniloiu & Markowitsch, in press). Functional imaging and electrophysiological studies have attempted to disentangle the neural underpinnings of experiences of ownership and agency, selfface recognition or remembering. Theoretical and empirical work has also focused on understanding the interactions between unconscious forms of information processing and the conscious ones. Vandekerckhove and Panksepp proposed that higher forms of consciousness are embedded in the ancient affective soil of anoetic consciousness (2009, p. 1026). Under anesthetic conditions, unconsciousness occurs when the brains ability to integrate information is blocked (Alkire et al., 2008; Mhuircheartaigh et al., 2010); loss or reduction of consciousness was also observed with other various brain changes (e.g., damage, degeneration, hormonal changes). One question in consciousness research is concerned with the necessary and sufcient processes that underpin healthy human consciousness, with some authors favoring the view that the thalamocortical system is the seat of the relevant neural machinery (Seth, 2010, p. 1; Tononi & Edelman, 1998). Autonoetic consciousness is generally regarded to depend on a well-functioning cerebral cortex (Markowitsch, 1995, 2005; Melloni et al., 2007; Uddin, Iacoboni, Lange, & Keenan, 2007). The functions in the spheres of EAM, autonoetic consciousness and self are, to a considerable degree, under the control of portions of the prefrontal cortex, in particular its ventromedial region (Beer, Lombardo, & Bhanji, 2010; Lee et al., 2010; Northoff et al., 2006; Passingham, Bengtsson, & Lau, 2010; Sebastian et al., 2008). However, functional imaging methods revealed that, in fact, the network is broader. There is also a more lateral prefrontal, right-hemispheric engagement (Keenan et al., 2000) and the anterior and posterior cingulate cortex, the precuneus, and the temporo-parietal junction area are engaged in addition to medial prefrontal regions (Buckner & Carroll, 2007; Plateck, Keenan, Gallup, & Mohamed, 2003; Vanhaudenhuyse et al., 2010). A reduction in consciousness together with an inability to form new autobiographical memories long-term may however accompany certain forms of diencephalic brain damage, in particular those affecting thalamic midline structures (Bogen, 1995; Bressler, 1995; Hart, 1995; Markowitsch, Cramon, & Schuri, 1993). We reported the case of a patient with above average intelligence, but severe amnesia due to an infarct that affected the thalamic mediodorsal nucleus bilaterally. After diencephalic damage this patient became totally anterogradely amnesic while his retrograde memory was partially preserved. The patient had a faade of normality when talking to him; he still displayed his premorbid jovial attitude in spite of his memory for new events being in the range of seconds only. On the other hand, the patient was able to acquire the skill of reading words, written in a mirror-image manner and he also demonstrated priming learning, as tested with an incomplete pictures test. He however showed a signicant impairment of his ability to reect on his condition, which was attributed to the severity of his anterograde memory impairment. When he was asked open questions about his memory, he responded that it was normal. And when he was questioned in more focused manner (e.g. Arent there areas where you have problems?), he responded: Yes, I quickly forget my dreams and I quickly forget jokes. When he was asked about politics, he initially replied that he was not interested in politics. When the interviewer persisted with questioning, the patient made general remarks, such as Well, you know, our continuing quarrels with France, you no longer want to hear that. In addition to amnesia and disturbances of insight and autonoetic consciousness (Markowitsch, 2003) the patient showed a tendency to engage in confabulations. Confabulations the production of ctitious narratives might arise from a combination of amnesia, impairments of autonetic consciousness and executive dysfunctions and perhaps impaired inner time perceptions (Borsutzky et al., 2008). They have been described in other diencephalic amnesic conditions (e.g. Korsakoffs syndrome). Damage to the basal forebrain (such as the one associated with aneurysms of anterior communicating arteries) may also lead to an enhanced vulnerability to certain types of false memories (such as provoked confabulations and false recognition in procedural memory) (Borsutzky et al., 2010). The involvement of the thalamus in autonoetic consciousness and the self is also reected by ndings in other patients with bilateral thalamic infarcts, who manifested persisting childish behavior and symptoms suggestive of Ganser syndrome (Fukatsu, Fujii, Yamadori, Nagasawa, & Sakurai, 1997) and a disorientation with respect to time (e.g., time of the day or season of the year) (Kumral, Gulluoglu, & Dramali, 2007; Spiegel, 1982; Spiegel, Wycis, Orchinik, & Freed, 1956).
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The relationship between time and memory and awareness has been emphasized since long and from very different perspectives. Time awareness (Brown, 1990), time orienting (Trivino, Correa, Arnedo, & Lupianez, 2010) and a proper handling of time epochs (Pribram & Tubbs, 1967) and temporal order (St. Jacques, Rubin, LaBar, & Cabeza, 2008) are related especially to the prefrontal cortex, while the proper perception of time and time epochs seems to engage the parietal cortex (Hoff & Ptzl, 1938; Ptzl, 1939, 1942, 1951, 1958) as well as the diencephalic structures as mentioned before (but see also Hfner, 1954). Damage to these areas may disturb the sense of time (including the the ability to successively link events in time) considerably with the consequence that the affected patients also become unable to encode or store new EAMs successfully. On the other hand, the capacity for episodic future thinking (self projection in the future) is an essential feature of EAM and autonoesis (Azry, Collette, Ionta, Fornari, & Blanke, 2009: Addis et al., 2009; Buckner & Carroll, 2007; Levine, Svoboda, Turner, Mandic, & Mackey, 2009; Perner et al., 2010; Schacter, Addis, & Buckner, 2007; Vogeley & Kupke, 2007; Zllig et al., 2007). The ability to re-experience the past events and pre-experience personal future events might also be severely impaired after bilateral medial temporal lobe damage (Levine et al., 2009; Noulhiane et al., 2007; Piolino et al., 2004). The most wellknown and well-studied amnesic (hippocampal) patient, H.M. reportedly made the following statement: Every day is alone, whatever enjoyment Ive had, and whatever sorrow Ive had (Milner, Corkin, & Teuber, 1968, p. 217). This statement suggests that H.M. knew the concept of time and that the life was a continuum. He showed at least partial insight into his memory problems and the ability to reect on them. He furthermore possessed emotional knowledge and likely some capacity to experience feelings (in spite of having bilateral amygdalar damage!). One could speculate that he might have even still felt the lingering feelings associated with the personal experiences (events) from the previous hours, though he was unable to explicitly remember any of those experiences (Feinstein, Duff, & Tranel, 2010). Many other patients with bilateral medial temporal lobe pathology have been found to share similar impairments an inability to encode new EAMs and new facts long-term. The ability of mental time traveling may be impaired both with respect to future and past in patients with major medial temporal lobe damage (e.g., due to encephalitis) and even in some patients with developmental amnesia (Andelman, Hooen, Goldberg, Aizenstein, & Neufeld, 2010; Damasio, Eslinger, Damasio, Van Hoesen, & Cornell, 1985; Kwan, Carson, Addis, & Rosenbaum, 2010). While the above described patients suffered from focal, circumscribed brain damage in so-called bottleneck structures (Brand & Markowitsch, 2003) another group of patients is characterized by pathologies accompanied by more widespread cortical damage, which might result in more global impairments of self-referential processing, autonoetic consciousness and mnemonic abilities. This group includes patients with degenerative forms of dementia (Banks & Weintraub, 2008; Greene, Hodges, & Baddeley, 1995; Gregory, Lough, Stone, Erzinclioglu, & Martin, 2002; OKeeffe et al., 2007; Piolino, Belliard, Desgranges, Perron, & Eustache, 2003). 6. Disturbances of EAM, autonoetic consciousness and self in psychiatric disorders Complete loss of the EAM is rare in psychiatric disorders as well as neurological ones. It has been reported in psychogenic or dissociative amnesic conditions (see below). Less extensive or severe EAM disturbances have, however, been described in several other psychiatric conditions, such as schizophrenia, major depressive disorder, bipolar disorder, post-traumatic stress disorder or personality disorders (e.g. borderline personality disorder). In patients with schizophrenia studies reported impairments of EAM, theory of mind functions, self-referential processing, autonoetic consciousness and emotional processing (Corcoran & Frith, 2003; Satterthwaite et al., 2010). Certain positive symptoms of psychosis, such as paranoid delusions and hallucinations were linked to defects in mnemonic and emotional processing (Satterthwaite et al., 2010). One recent study showed, for example, that auditory verbal hallucinations were preceded by deactivation of the parahippocampus, suggesting that they may arise from spontaneous re-experiencing of memories (Diederen et al., 2010). Patients with active symptoms of major depressive disorder demonstrated abnormalities of self-referential processing, such as excessive self-focus (Grimm et al., 2009) and qualitative changes of the EAMs (reduced specicity of details, increased retrieval from third person-perspective, changes in autonetic consciousness) in comparison to healthy people (Lemogne et al., 2006; Williams & Scott, 1988), which concerned preponderantly the memories of positively-valenced personal events (Lemogne et al., 2006). A third person retrieval perspective for positive EAMs was found to persist in patients with major depressive disorder, even after clinical remission (Bergouignan et al., 2008). Patients with active symptoms of depression often display hopelessness and suicidal thoughts, which were hypothesized to be connected with their difculties with imagining (pre-experiencing) positive personal future events (Sharot, Riccardi, Raio, & Phelps, 2007). These observations are consistent with studies that indicate that memory encoding and retrieval of negative versus positive material engage different brain networks (Markowitsch et al., 2003). In patients with post-traumatic stress disorder, the relationship between a history of trauma and impairments of EAMs in the form of reduced specicity of recalled past events was found to be moderated by the nature of trauma, the age at the onset of trauma (Stokes, Dritschel, & Bekerian, 2004; Valentino et al., 2009) and the cognitive style (Lemogne et al., 2008). In healthy young adults a relationship was observed between a diminution of EAMs and a cognitive style characterized by avoidance of emotional material (Lemogne et al., 2008). Traumatic experiences have also been identied to trigger more global loss of EAM, such as in dissociative or psychogenic amnesic conditions. Psychogenic amnesic conditions have traditionally been viewed as episodes of amnesia, which were
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preponderantly etiologically linked to psychological factors and occurred in the absence of signicant brain damage (as detected by conventional structural imaging techniques). Most cases of psychogenic amnesia were reported to have a retrograde nature. In a few instances anterograde (Markowitsch, Kessler, Kalbe, & Herholz, 1999; Smith et al., 2010; Staniloiu, Markowitsch, et al., 2010) instead of retrograde amnesia (or a combination of the two) has been documented (Markowitsch, Kessler, Van der Ven, Weber-Luxenburger, & Heiss, 1998). The interest in psychogenic amnesia has uctuated over the years, dating back to around 140 years ago (Markowitsch, 1992). The etiological underpinnings of this disorder have from the beginning been a source of debate. This is also reected in the various terms, which have over the years been used to capture this condition, such as epileptic somnolence(Burgl, 1900), hysteria (Breuer & Freud, 1895; Janet, 1907), hysterical state of somnolence, multiple personality (disorder) (Sidis & Goodhart, 1905), dissociative, functional or medically unexplained amnesia or mnestic block syndrome (Markowitsch, 2002). The subsuming of this condition under the concept of hysteria was reinforced by the strong tradition of French psychiatry (Pierre Janet, Jean-Marie Charcot), which was followed by Sigmund Freud, a pupil of Charcot. In the current main ofcial classications of diseases (DSM-IV-TR, 2000 and ICD-10, 1992) earlier diagnostic designations of hysterical or psychogenic amnesia are now preponderantly incorporated by the diagnostic categories of dissociative disorders in DSM-IV-TR and dissociative (conversion) disorders in ICD-10, but also by other diagnostic subcategories such as somatization disorder (in DSM-IV-TR only), post-traumatic stress disorder (PTSD) and acute stress disorder (DSMIV-TR and ICD-10) or certain personality disorders (such as borderline personality disorder). However, the term hysteria is still in use (Stone, Smyth, Carson, Warlow, & Sharpe, 2006; Thomas-Anterion, Guedj, Decousus, & Laurent, 2010). There are authors who still debate the pathogenetic mechanisms or the legitimacy of the diagnostic entity of dissociative amnesia; Pope et al. (2007) consider it to be culture-bound and as having arisen in the 19th century. Though culture denitely plays a role in molding perceptions of trauma, selfhood, memory, future and past, there is nowadays signicant evidence for the contribution of psychological stress or trauma to the genesis of dissociative disorders across a variety of cultures (Staniloiu, Borsutzky, & Markowitsch, 2010). A large body of data has linked psychological stress (in the form of acute massive stress or chronic stress) to a variety of psychiatric and non-psychiatric conditions (Staniloiu, Markowitsch, et al., 2010). Although psychological stress or trauma is necessary for the emergence of dissociative amnesic disorders, it is not a sufcient condition, which in fact has been suggested since Janets time. Genetic dispositions, epigenetic mechanisms, gender, personality traits and the nature, severity, recurrence and the age of onset of trauma all modulate the impact of psychological stress on the structural and functional integrity of the brain network subserving mnemonic processes (see below). In line with Janets view of the mechanism of dissociation as an inability of the personal self to bind together the various mental components in an integrated whole under its control (Janet, 1907, p. 23), dissociative disorders are dened in DSMIV-TR (2000) as disturbances of the integrated organization of memory, perception, consciousness and identity. Among dissociative disorders, amnesia is a prominent symptom of dissociative amnesia, dissociative fugue, dissociative identity disorder (multiple personality disorder) and dissociative trance disorder (possession trance). Apart from dissociation, other psychological mechanisms, such as hyper-suppression or cognitive avoidance (Fujiwara et al., 2008; Lemogne et al., 2008; Tramoni et al., 2009) have been identied to underlie at times concurrently amnesic conditions, which are etiologically linked to psychological factors. These ndings partly explain why some authors still favor the use of the term psychogenic amnesia over the term dissociative (McKay & Kopelman, 2009). Dissociative amnesia has as primary symptom the loss of memory for autobiographical events, especially those of a traumatic nature. This symptom should not be better explained by other somatic diseases (such as closed head injury), normative forgetfulness or other psychiatric disturbances. Psychologically motivated feigning of memory disturbances (factitious disorder) and malingering have to be ruled out (Jenkins, Kapur, & Kopelman, 2009). This is however, not always an easy task as some cases of dissociative amnesia can occur on a forensic background (Markowitsch, 1992, 2010; Markowitsch, Calabrese, et al., 1997). On the other hand the most commonly malingered cognitive impairment by individuals who seek nancial compensation after an accident is memory impairment (Serra, Fadda, Buccione, Caltagirone, & Carlesimo, 2007). A variant of dissociative amnesia is the dissociative fugue. In this condition retrograde amnesia for personal events is accompanied by suddenly leaving the customary environment home, city and workplace and compromised knowledge about personal identity. A failure of integration of self-referential, emotive and cognitive processes can however be observed not only in patients with dissociative amnesic conditions, but also in patients with related symptomatologies, such conversion paralysis, where body parts (most frequently the extremities) become paralyzed in the context of major psychological stress (Burgmer et al., 2006) or body identity integrity disorders (where individuals experience a strong wish for amputation of one of their limbs or for becoming paralyzed) (Oddo et al., 2009). A relevant example of a case of dissociative fugue is the one of a 37-year-old family father, who developed an episode of retrograde amnesia and loss of personal identity, seemingly out the blue (Markowitsch, Fink, Thne, Kessler, & Heiss, 1997). One morning he took his bike, with the plan to go to the nearby bakery to buy roles for breakfast. However, instead of returning for breakfast, he apparently continued to ride his bike for the next couple of days from north to south Germany until he reached the suburbs of a major city. There he met a few people who suggested to him to continue to ride to the central railway station. After doing so, he met a female ofcer from the Salvation Army, who advised him to go the nearby university psychiatric clinic. He followed her advice, went to the hospital and got admitted to the psychiatric ward, where he was diagnosed with psychogenic (dissociative) fugue. In the hospital the patient made friends and appeared not to care about having no access to his past a presentation which has been noted in other patients with dissociative amnesia and was termed la belle indiffrence by Janet (1907). The description of this case points so far to several similarities with other case-reports from literature. It is not uncommon for patients with dissociative fugue to present themselves to the emergency services
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of hospitals or the police stations or wander in railway stations (Kluft, 1988; Kritchevsky, Chang, & Squire, 2004; Loewenstein, 1991) and to be susceptible to inuences (advice) from others (Reinhold & Markowitsch, 2009; Stone et al., 2006). In fact many patients with dissociative disorders are highly hypnotizable on formal testing (Maldonado & Spiegel, 2008). The patients lack of concern about their condition may bear a close link to the impairment of their capacity for mental time traveling, which presupposes an emotional engagement with both past and future (Azry et al., 2009). Later on, the patient described having had a strong need (compulsion) to continue to ride during his fugue episode, which in fact he did for a couple of days. His state of altered (autonetic) consciousness apparently did not interfere with his automatic performance (ambulatory automatism) (Loewenstein, 1991). In fact there are reports that some dissociative states, which are characterized by suspension of critical thought, may even enhance performance of coordinated and complex motor acts, such as in athletes (Maldonado & Spiegel, 2008). The patient also described that during his ride, he at times looked at himself in the windows of shops in the cities, but his face did not look familiar to him. This again suggests an impairment of self-referential processing (self-face processing) and self-awareness, perhaps rooted in a dysfunction of the right hemisphere (see below). We initially tested the patient neuropsychologically. He demonstrated normal anterograde memory; however, he showed no evidence of any retrograde EAM. He later relearned his past, including his autobiographical-semantic knowledge and then could retrieve portions of his past in a neutral, affectless manner. Other cognitive functions were largely spared or regained within weeks. We investigated the patient with functional brain imaging, using a paradigm in which he was confronted with events from his autobiographical past. While the normal subjects activated in the water-PET study predominantly the fronto-temporal region of the right hemisphere (Fink et al., 1996; Kroll, Markowitsch, Knight, & von Cramon, 1997; Markowitsch et al., 1993), the patient activated largely the left hemisphere, which is considered to be relevant for the retrieval of neutral facts or knowledge (Grossi, Trojano, Grasso, & Orsini, 1988; Markowitsch, Calabrese, et al., 1999). As mentioned above, the right temporal-frontal connections are important for ecphorizing the affect-laden personal events. Therefore those imaging ndings suggested that the patients access to his past occurred via his semantic memory system. As it was also transparent from his behavior, the patient lacked that rst-person autonoetic connection and feeling of warmth that is characteristic to EAM retrieval. Interestingly, the patients personality underwent changes as well. While he had been an avid car driver prior to his escape (fugue), afterwards he avoided car driving or riding a bike and said that cars were too fast for human beings. His relinquishing of a previously rewarding behavior (car driving) occurred in the absence of evidence of impairment in his procedural skills. Furthermore, the patient changed his food preferences and gained considerable weight. Interestingly, he ceased having asthma attacks or allergic reactions, conditions which may be viewed as partly having a psychosomatic basis. Alterations in previously rewarding activities (such as eating, smoking, cooking, drinking alcohol) and personality after the onset of psychogenic (dissociative) amnesia have been reported by several other authors (Fujiwara et al., 2008; Tramoni et al., 2009). They may reect the fact that several structures that are involved in EAM processing play also a role in reward and are sensitive to stress (Ulrich-Lai & Herman, 2009). Furthermore, they also converge with a recent view of self as having connections to reward (Enzi, de Greck, Prsch, Tempelmann, & Northoff, 2009). Contrary to old teachers lore, confrontation with his wife did not result in dissolving the patients amnesia. Instead he even thought that the hospital personal wanted to couple him with an unknown woman. He, however, moved back to the family house. Upon arrival, he complained about the existing furniture and tapestry. Nevertheless he pretty much followed all suggestions or directions of his wife instantaneously, as he had done before the onset of the amnesia. A review of the patients developmental history revealed that he had been raised by parents who abused alcohol and fought a lot against each other. The patient was dressed as a girl until starting elementary school because his mother reportedly wanted to have a daughter instead of a son. Later his mother frequently criticized him for being weak and behaving in a girlish manner. She predicted that he would never become successful in life and instead would ruin their business. The patient grew up with very low self-esteem. In his early 20ies, he married a woman who apparently behaved similarly to his mother and possessed the self-condence he was missing. Being not very successful with their business, the couple always suffered from a lack of money. Nevertheless, the patients wife reportedly decided one day that they should leave by car for a longer vacation. Though the patient knew that they would not have enough nancial resources for the suggested trip, he did not object. Three days prior to the date of the intended holidays, he, however started riding his bike along the river Rhine. This case paradigmatically reects the disruption (disintegration) of the cohesiveness between autonoetic consciousness, EAM and the self in the dissociative amnesic condition: The patient had lost rst-person autonoetic access to the personal past of his entire life, while being able to retrieve impersonal semantic facts. He could, however still read, write and calculate. He displayed a blunted affective disposition and did not seem to care about his future life (la belle indifference; Reinhold & Markowitsch, 2009; Stone et al., 2006). Though his knowledge about how to behave in social situations seemed largely preserved, there were several indications that his amnesia might have impacted on his ability to judge the feelings of close others, regulate his social behavior and exibly adapt it to the new circumstances. He for example became closer and friendlier with comrades from the psychiatric ward than with his family, which resulted in signicant interpersonal difculties with his close family members. The patients presentation is similar to the clinical descriptions of other patients with dissociative amnesic conditions, though there are a number of variants of these conditions. One particular variant is the Ganser syndrome, which at times had been described in association with dissociative fugue conditions. The syndrome has been submitted to several diagnostic revisions and debates over the years. In comparison to previous DSM editions, where Ganser syndrome was presented as a Factitious Disorder, Ganser syndrome is currently included under the category of Dissociative Disorders Not Otherwise
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Specied in DSM-IV-TR and it is simply dened by giving approximate answers to questions (vorbeireden). Gansers (Ganser, 1898, 1904, 1965; cf. Feinstein & Hattersley, 1988) original description of the syndrome was, however, much broader than the current DSM-IV-TR one. It included a hysterical semitrance or twilight state, characterized by a tendency to give approximate answers, impairments of consciousness, amnesia and hallucinations, being more consistent with the later views of this disorder as a brief reactive psychosis to stress. In 1904 Ganser wrote on page 34 (our translation): The most prominent phenomenon was that the patients were unable to respond properly to questions of the most simple kind, they were given, though they indicated through their mode of answering that they principally had captured the sense of the questions, and that they revealed in their answers an astonishing lack of knowledge for facts they denitely had possessed or were still possessing. ... After days, and in some cases after repeated remissions and intermissions, a total clarity with recurrence of normal knowledge appeared, whereby for the period of the somnolence a gap of reminiscence remained. Though initially linked to forensic background, Ganser syndrome was also reported in non- forensic contexts. The syndrome was found to affect preponderantly young men with a mean age of 35 years, although there were some case-reports in women and children as well. Both a transient, limited course and a chronic course have been recorded. We investigated two male patients with this syndrome, one with a clear forensic background and one without (Staniloiu et al., 2009). Both patients showed a global deterioration of their intellectual capacities suggestive of a pseudo- dementic picture. The patients became lethargic, lost interest and only reacted when being directly addressed and otherwise relied on the guidance provided by their wives. They lost joy, did not attend to their children and other stimuli of the environment and appeared unable to understand and to react appropriately to even simple commands. The patients appeared to be very insecure. Even very simple questions, for instance, pertaining to their date of birth were not directly answered (I have to look in my passport was, for example, a response to questioning about the date of birth.) Their clinical presentation followed a chronic course, despite treatment. This is congruent with the reports of other authors who evidenced a prolonged course of memory decits in a substantial number of dissociative amnesic conditions (Kritchevsky et al., 2004) and is in contrast with ndings from older studies, that described that memory recovered in most patients within a month from the onset of psychogenic memory loss (Kanzer, 1939). Although in both cases of Ganser syndrome the structural brain imaging investigations were not indicative of organic impairment, in the patient where additional functional imaging was performed, a global signicant reduction in the brain metabolism was visualized. The mentioned functional imaging result and the chronic course of the above mentioned two patients are relevant, in the light of ndings from previous studies. Some of those pointed to a possible organic basis to the psychiatric presentation of the Ganser syndrome that can especially become apparent over time. For example, Ladowsky-Brooks and Fischer (2003) described a patient, who presented with features of Ganser syndrome and severe cognitive decits in other domains. However, the individuals cognitive decline over a period of a year, in combination with ndings from functional imaging, resulted in a diagnosis of fronto-temporal dementia. The connection between stress-related cognitive decline, the potential contribution of glucocorticoids to accelerated aging, fronto-temporal dysfunction, and the later development of dementia was already pointed out by Porter and Landeld (1998). Recent studies have further conrmed a relationship between different forms of stress-related psychopathologies (such as major depressive disorder or post-traumatic stress disorder) and later risk for the development of dementia (e.g. Yaffe et al., 2010). We have studied several dozens of patients with a condition of long-lasting dissociative amnesia. All of them encountered major negative life events that had occurred mostly in early childhood, but in part continued or recurred in their later life (e.g., Markowitsch, Fink, et al., 1997; Markowitsch et al., 1998, 2000; Markowitsch, Kessler, Russ, et al., 1999; Fujiwara et al., 2008; Reinhold & Markowitsch, 2007). In some of these cases there was an additional background of immigration (Fujiwara et al., 2008; Staniloiu, Borsutzky, et al., 2010; Staniloiu, Markowitsch, et al., 2010). Similar cases with a (likely) background of immigration were reported by others (Thomas-Anterion et al., 2010). In our pathogenetic model of dissociative amnesic conditions, we propose that stressful events lead to the release of a variety of stress hormones and initiate a neurotoxicity cascade (Joels & Baram, 2010; Markowitsch, 2000; OBrien, 1997). We assume that the release of stress hormones in particular glucocorticoids is principally responsible for the block in the function of brain structures that are essential for the encoding or retrieval of information. In fact, it is known that several key brain areas involved in memory processing show especially during their periods of development or regression (Lupien, McEwen, Gunnar, & Heim, 2009) an increased susceptibility to the hormonal effects of stress and that enduring alterations in stress hormone responses in response to environment (e.g., early life experiences) might occur via epigenetic mechanisms, which act during windows of heightened vulnerability (McGowan et al., 2009; Szyf, 2007). The effects of stress and environment (Brand & Markowitsch, 2009) on mental health might be clinically observable after a lag period. This may be accounted for by the fact that the early stress effects on synaptic organization and brain structure and function only may become obvious after the completion of synaptic organization (Lupien et al., 2009) We (Markowitsch et al., 1998, 2000; Reinhold, Khnel, Brand, & Markowitsch, 2006) and others (Thomas-Anterion et al., 2010; Tramoni et al., 2009) have found evidence for stress-induced changes of the brain network engaged in mnemonic processes in dissociative amnesic conditions by employing different functional brain imaging methods and structural imaging techniques specialized for estimating white matter integrity. Via these methods we provided evidence for a desynchronization between regions of the brain that are involved in the processing of factual information and regions of the brain that are involved in the processing of emotion, self-awareness and EAM (limbic system, in particular the amygdala and the septal region/basal forebrain). This is consistent with knowledge that receptors for stress hormones exist in the regions of the hippocampal formation and the amygdala structures which synchronize facts and emotions and are therefore crucial for EAM
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(de Kloet, Oiztl, & Joels, 1999; Ellen, Kaut, & Lord, 2009; Joels & Baram, 2010). As both brain imaging results from patients with dissociative amnesic conditions (summarized in Reinhold et al. (2006)) and theoretical accounts point to, the expanded limbic system (Nauta, 1979; Nieuwenhuys, 1996), including the frequently mentioned medial prefrontal/orbitofrontal cortex and the insula (e.g., Ganis, Morris, & Kosslyn, 2009; Modinos, Ormel, & Aleman, 2009) constitutes a structural network for autonoetic consciousness, EAM, and the self. Clinical and imaging ndings from patients with dissociative amnesic conditions re-emphasize the importance of a proper emotional charging for EAMs (Markowitsch & Staniloiu, in preparation-a). There is evidence from many sites that EAM and emotions are strongly interdependent (Sarter & Markowitsch, 1985a). Especially patients with bilateral amygdala damage demonstrate that they are impaired in properly encoding EAM (Cahill et al., 1995; Markowitsch et al., 1994; Siebert, Markowitsch, & Bartel, 2003; see Murty, Ritchey, Adcock, and LaBar (in press) and Markowitsch and Staniloiu (in preparation-b), for reviews). But recent work also shows that there are several interdependencies between different forms of emotions, activation of brain regions and awareness and interpretation of the environmental signals (Amting, Greening, & Mitchell, 2010). 7. Conclusions The self, autonoetic consciousness and EAM are intimately interlocked. The relationship between them evokes the reference to the threefold cord that is not easily broken (Ecclesiastes 4:12). The three cords appear and evolve both ontogenetically and phylogenetically in strong connection with each other, supporting and enriching each others development. Their appearance and development take place in concert with other cognitive and emotive functions the ability of perspective taking, executive functions, language, the feeling of empathy, the ability to reect on oneself, solving abilities with divert thinking, etc. (e.g., Sutin & Robins, 2008). Given the multi-functionality of EAM and the multiple facets of self (Klein & Gangi, 2010; Neisser, 1988) and consciousness, the relationship between EAM, autonoetic consciousness and self can be approached from multiple perspectives. Neisser (1988) distinguished the ecological, the interpersonal, the conceptual, the remembered, and the private self. Klein and Gangi (2010) wrote about the multiplicity of the self and pointed to the composition of the self of EAMs, of semantic representations of ones personality traits, of semantic knowledge of facts about ones life, an experience of continuity through time, a sense of personal agency and ownership, the ability to self-reect, and the physical self. Along the same vein Kaplan et al. (2008) had pointed to the multimodality of the self. Similar to Vandekerckhove and Panksepp (2009), Davis (1996) had described a ow between multiple levels of consciousness: Not one unconscious, not the unconscious, but multiple levels of consciousness and unconsciousness, in an ongoing state of interactive articulation as past experience infuses the present and present experience evokes state-dependent memories of formative interactive representations. Not an onion, which must be carefully peeled, or an archeological site to be meticulously unearthed and reconstructed in its original form, but a childs kaleidoscope in which each glance through the pinhole of a moment in time provides a unique view; a complex organization in which a xed set of colored, shaped, and textured components rearrange themselves in unique crystalline structures determined by way of innite pathways of interconnectedness (p. 197). The overlap between EAM, the self, and autonoetic consciousness is depicted as a Venn-diagram below (Fig. 5).
Fig. 5. EAM, the self, and autonoetic consciousness overlap considerably. They are connected by their embeddedness in time.
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As it is evident from the results of work with patients with neurological or psychiatric diseases, as well as from results on the developing human individual, different components of memory contribute to the formation of the self and make it autonoetic. The dimensions of time as reected in mental time traveling and autonetic consciousness are central to the trans-temporal cohesiveness of self across contexts. As Wheeler and colleagues (1997) put forth (see above), autonoetic consciousness allows the individual to become aware of their protracted existence across subjective time (p. 335) (Fig. 5). Furthermore it gives memory that binding power, which prevents the one-ness from getting lost in the multiplicity (CooperWhite, 2008). This binding (connecting) power of memory that is also metaphorically conveyed by the etymology of the word re-collection (re-collection derives from the word collection and the latter has similar roots as colligation, which means binding) (Casey, 2000) was beautifully captured by Hering. He wrote in 1870 (p. 12): Memory connects innumerable single phenomena into a whole, and just as the body would be scattered like dust in countless atoms if the attraction of matter did not hold it together so consciousness without the connecting power of memory would fall apart in as many fragments as it contains moments. References
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Consciousness and Cognition

Memory, autonoetic consciousness, and the self q

H.J. Markowitsch, A. Staniloiu / Consciousness and Cognition xxx (2010) xxxxxx

H.J. Markowitsch, A. Staniloiu / Consciousness and Cognition xxx (2010) xxxxxx

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