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Volume 16, Number 13,
Issue of July 1, 1996
pp. 4275-4282
Copyright ©1996 Society for Neuroscience
Cerebral Representation of One's Own Past: Neural Networks
Involved in Autobiographical Memory
Gereon R. Fink1, 2,
Hans
J. Markowitsch3,
Mechthild Reinkemeier3,
Thomas Bruckbauer1,
Josef Kessler1, and
Wolf-Dieter Heiss1, 2
1 Max-Planck-Institut für Neurologische
Forschung, D-50931 Köln, Germany,
2 Universitätsklinik für Neurologie der
Universität zu Köln, D-50924 Köln, Germany, and
3 Physiologische Psychologie, Universität
Bielefeld, D-33501 Bielefeld, Germany
 ABSTRACT
- INTRODUCTION
- MATERIALS AND METHODS
- RESULTS
- DISCUSSION
- FOOTNOTES
- REFERENCES
ABSTRACT 
We studied the functional anatomy of affect-laden autobiographical
memory in normal volunteers. Using H215O
positron emission tomography (PET), we measured changes in relative
regional cerebral blood flow (rCBF). Four rCBF measurements were
obtained during three conditions: REST, i.e., subjects lay at rest (for
control); IMPERSONAL, i.e., subjects listened to sentences containing
episodic information taken from an autobiography of a person they did
not know, but which had been presented to them before PET scanning
(nonautobiographical episodic memory ecphory); and PERSONAL, i.e.,
subjects listened to sentences containing information taken from their
own past (autobiographical episodic memory ecphory).
Comparing IMPERSONAL with REST (nonautobiographical episodic memory
ecphory) resulted in relative rCBF increases symmetrically in both
temporal lobes including the temporal poles and medial and superior
temporal gyri. The same loci, however, with a stronger lateralization
to the right hemisphere were activated in the comparison PERSONAL to
REST (autobiographical episodic memory ecphory). In addition, the right
temporomesial, right dorsal prefrontal, right posterior cingulate
areas, and the left cerebellum were activated. A comparison of PERSONAL
and IMPERSONAL (autobiographical vs nonautobiographical episodic memory
ecphory) demonstrated a preponderantly right hemispheric activation
including primarily right temporomesial and temporolateral cortex,
right posterior cingulate areas, right insula, and right prefrontal
areas. The right temporomesial activation included hippocampus,
parahippocampus, and amygdala.
These results suggest that a right hemispheric network of temporal,
together with posterior, cingulate, and prefrontal, areas is engaged in
the ecphory of affect-laden autobiographical information.
Key words:
PET;
autobiographical memory;
episodic memory;
retrieval;
cingulate;
limbic system
INTRODUCTION
Memory commonly is subdivided into a number of
forms (Squire and Knowlton, 1995 ; Tulving, 1995), and it is assumed
that different brain regions contribute differentially to the various
forms of memory processing (Squire et al., 1993; Markowitsch, 1995a).
Two of the main forms of memory are episodic memory and semantic memory
(Tulving, 1995), also subsumed under declarative memory (Squire, 1987).
Episodic memory is supposed to deal with individual
(``autobiographical'') episodes that are definable with respect to
time and locus. Semantic memory contains impersonal facts (e.g.,
knowledge about the world), which we need for verbal and nonverbal
interaction with our environment.
The usefulness of the distinction between episodic and semantic memory
sometimes has been argued (Cermak and Craik, 1979), given the fact that
semantic memory frequently constitutes a generalization from initially
episodically encoded information. However, there are experiments
demonstrating both acquisition of semantic in the absence of the
ability to acquire episodic information (Tulving et al., 1991) and
impairments in acquiring both forms (Gabrieli et al., 1988; Verfaellie
and Cermak, 1994). Although both forms most likely engage overlapping
limbic system structures for information acquisition (Buckner and
Tulving, 1995), memory retrieval seems to depend on different brain
regions and, furthermore, seems to rely on different systems for the
retrieval of episodic and semantic information.
Results of studies performed with positron emission tomography (PET)
suggest a left hemispheric preponderance for the retrieval of semantic
information and a right hemispheric preponderance for the retrieval of
episodic information (Fletcher et al., 1995a; Tulving et al.,
1994a,b,c, 1996).
In most clinical studies on amnesics, it has been shown that the
retrieval of episodic (autobiographical) memory is much more affected
than that of semantic memory (general knowledge) (Markowitsch,
1995a,b). Reasons for this may be sought in the uniqueness of episodic
information (Damasio, 1990), in its higher emotional character (Sarter
and Markowitsch, 1985a,b; Markowitsch et al., 1994), or in the brain
regions in which damage affects ecphory of episodic versus general
knowledge material (Markowitsch, 1995a,b). Tulving (1983) used 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. It is of interest to investigate
whether in the intact human brain, the ecphory of presented
autobiographical material will affect different brain regions compared
with the retrieval of semantically closely similar material, which is,
however, not associated by the subject with his or her own past.
In this study, we wanted to demonstrate the functional anatomy of
ecphory of affect-laden autobiographical material. We used PET to image
significant changes in relative regional cerebral blood flow (rCBF) as
an index of changes in local neuronal activity in the brains of normal
human subjects when they listened to sentences containing affect-laden
episodic autobiographical information. We then intended to relate those
relative rCBF changes to changes obtained when the same subjects
listened to sentences containing semantically similar episodic but
nonautobiographical material. Based on evidence available from human
and nonhuman lesion data (LeDoux, 1995; Markowitsch, 1995a,b; Sarter
and Markowitsch, 1985a,b), we hypothesized that the anterolateral and
temporomesial parts of the temporal lobes (including the amygdala) as
well as the inferolateral prefrontal cortex were involved in the
ecphory of past personal experiences.
MATERIALS AND METHODS
Subjects. Seven normal, healthy male volunteers (age
21-37 years) were recruited. All subjects were right-handed and had no
history or evidence of any medical, neurological, or psychiatric
disease. Informed consent was obtained before participation from all
subjects. The study was approved by the local ethics committee of the
University of Cologne, and permission to administer radioactivity was
obtained from the federal administration authorities.
Paradigm design. The experiment involved 12 sequential
measurements of relative rCBF and consisted of one control condition
(REST) and two testing conditions (IMPERSONAL, PERSONAL), which were
presented in a counterbalanced order (ABCCBAACBBCA). The order of scans
was counterbalanced rather than randomized to avoid variable order
artifacts attributable to the small number of subjects studied. The
conditions were as follows: REST, i.e., subjects were studies in a
resting condition with eyes closed and no auditory or visual
stimulation (baseline condition, for control); IMPERSONAL, i.e.,
subjects were studied during auditory presentation of sentences that
contained episodic information taken from an autobiography of a person
they did not know, but which had been presented to them about 1 hr
before PET scanning; PERSONAL, i.e., subjects were studied while they
listened to sentences containing episodic information taken from their
own past (autobiographical information). The sentences contained brief
episodes from significant events of each subject's past or someone
else's past, respectively. Examples for sentences are: ``When you
were 15 you took part in a swimming marathon and succeeded to swim 10 miles,'' and ``He tore off his shirt to demonstrate his scars to the
nurse.'' Subjects were instructed to imagine what happened to the
person in the described situations (IMPERSONAL) or to imagine what
happened to themselves in the described situations (PERSONAL). Ten
sentences were presented during IMPERSONAL and PERSONAL situations. The
information presented during PERSONAL had been obtained in
semistandardized interviews weeks before PET scanning. Information was
collected from childhood, adolescence, and early adulthood; the
subjects were kept unaware of the purpose of the interviews.
PET-scanning techniques. Relative rCBF was measured by
recording the regional distribution of cerebral radioactivity after the
intravenous injection of 15O-labeled water
(15O is a positron emitter with a half-life of
2.1 min) (Mazziotta et al., 1985; Fox and Mintun, 1989). The PET
measurements were carried out using a CTI ECAT Exact HR PET scanner
(CTI, Knoxville, TN) with a total axial field of view of 15 cm covering
the whole brain (Wienhard et al., 1994). Data were acquired in
three-dimensional mode (Townsend et al., 1991) with interdetector
collimating septa removed, allowing the collection of reliable data of
regional brain activation from a single subject (Watson et al.,
1993).
For each measurement of relative rCBF, 10 mCi of
H215O were given intravenously. Emission data
were collected sequentially over 90 sec after tracer arrival in the
brain. This process was repeated for each emission scan, with 10 min
between scans to allow for adequate decay of radioactivity. This gave
an estimate of rCBF. Because no arterial blood samples were taken
(Mazziotta et al., 1985), no calibration was possible and the term
``relative'' rCBF used in this paper implies that there was no
absolute quantification of rCBF. The emission scan data then were
corrected for effects of radiation attenuation by the skull (by means
of a transmission scan taken before the first relative rCBF
measurement). After attenuation correction, the data were reconstructed
to 47 transverse planes by three-dimension filtered-back projection
using a Hanning filter of cutoff frequency 0.4 cycles per pixel. The
resolution of the resulting images was 5.5 mm (at full-width
half-maximum).
Magnetic resonance imaging. On a separate occasion, a
magnetic resonance (MR) image of each subject's brain was obtained to
exclude morphological-pathological abnormalities. This was performed
with a 1 Tesla system (Magnetom, Siemens, Germany) using a FLASH
sequence (flip angle, 40°; repetition time, 40 msec; echo time, 15 msec) producing 64 transaxial T1-weighted
tomograms.
Image processing. All calculations and image manipulations
were performed on a Sparc workstation (Sun Computers). ANALYZE and
PROMATLAB software (MathWorks) was used to calculate and display
images. Statistical parametric mapping (SPM) software (Wellcome
Department of Cognitive Neurology, London, UK) was used for image
realignment and smoothing and to create statistical maps of significant
relative rCBF changes (Friston et al., 1995a,b).
Realignment, transformation, and smoothing of PET images.
Using SPM software (Friston et al., 1995a), all PET scans were
realigned to the first emission scan to correct for any head movement.
The PET images then were transformed into a standard stereotactic
anatomical space (Talairach and Tournoux, 1988; Friston et al., 1995a)
using linear proportions and a nonlinear sampling algorithm. This
allowed accounting for differences in brain size and shape. PET images
then were filtered using a low-pass Gaussian filter to reduce the
variance attributable to individual anatomical variability and to
improve signal-to-noise ratio (Friston et al., 1995a). The resulting
pixel size in stereotactic space was 2 × 2 mm with an interplane
distance of 4 mm. The group data were expressed in terms of standard
stereotactic coordinates in the x-, y-, and
z-axes (defined in Table
1).
Statistical analysis. After stereotactic normalization, the
main effects of the two testing conditions (IMPERSONAL, PERSONAL) were
estimated on pixel-by-pixel basis using SPM (Friston et al., 1995b).
Global differences in CBF were first covaried out (Friston et al.,
1995b), treating global activity as the covariate. This controlled for
systematic state-dependent differences in global blood flow associated
with the different conditions. Then for each pixel across all subjects
and all scans, the mean values were calculated for the control (REST)
and the two activation tasks separately, and comparisons of the means
thereafter were made using t statistics (Friston et al.,
1995b) subsequently transformed into normally distributed Z
statistics. The resulting set of Z values constituted a
statistical parametric map (SPM{t} map)
(Friston et al., 1995b). For the group, data significance was set to
p < 0.05, corrected for multiple nonindependent
comparisons. In addition, PET data were analyzed for each individual
subject in an identical way to the group data to allow single-subject
data analysis.
Localization of activations. The stereotactic coordinates of
the pixels of local maximum significant changes in relative rCBF within
areas of significant relative rCBF change associated with the different
tasks were determined. The anatomical localization of these local
maxima was assessed by referring to the standard stereotactic atlas of
Talairach and Tournoux (1988) and validated additionally after
superimposition of the SPM{t} maps on an
arbitrary MR image that had undergone stereotactic transformation into
the same standard stereotactic space (Friston et al., 1995a).
Planned comparisons of differences. Three planned
comparisons were made. First, a comparison was made of the condition
associated with neutral nonautobiographical episodic memory ecphory
(IMPERSONAL) and the control (REST). Second, a comparison of
autobiographical episodic memory ecphory (PERSONAL) with the control
(REST) was performed. Third, autobiographical memory ecphory (PERSONAL)
was compared with nonautobiographical memory ecphory (IMPERSONAL).
These comparisons were intended to identify those cortical areas
concerned with the modalities in question (i.e., autobiographical
memory ecphory and episodic memory ecphory in general). For
completeness of data analysis and presentation, the respective reverse
comparisons also were calculated.
RESULTS
Nonautobiographical episodic memory ecphory versus
baseline (IMPERSONAL-REST)
Table 1, IMPERSONAL-REST summarizes the principal
areas with increases in relative rCBF associated with the difference
between IMPERSONAL and REST. Figure 1 provides a
pictorial demonstration of the areas with relative rCBF increase. There
are highly symmetrical bilateral increases in the temporal lobes,
namely the superior and medial temporal gyri including both temporal
poles and Brodmann area 21 (BA 21). Decreases in relative
rCBF associated with the task were seen in the medial (BA 7)
and the lateral inferior parietal cortex (BA 40)
bilaterally, the right fusiform gyrus (BA 19/37),
the left caudate nucleus, and the midbrain (Table 2a,
IMPERSONAL-REST).
Fig. 1.
Relative rCBF increases during nonautobiographical
and autobiographical episodic memory retrieval. Relative rCBF increases
for the group of seven subjects associated with nonautobiographical
episodic memory retrieval (IMPERSONAL-REST),
autobiographical episodic memory retrieval (PERSONAL-REST),
and autobiographical memory (PERSONAL-IMPERSONAL). Areas of
significant (p < 0.05, corrected for multiple
nonindependent comparisons) relative rCBF increases associated with the
different tasks are shown as through projections onto representations
of the standard stereotactic space as defined by Talairach and Tournoux
(1988). The sagittal images (left) view the brain from the
side, the coronal images (middle) view the brain from the
back, and the transverse images (right) view the brain from
the top. R, Right; VAC, vertical plane through
the anterior commissure; VPC, vertical plane through the
posterior commissure. Numbers at axes refer to coordinates
of stereotactic space. The exact coordinates of the local maxima within
areas of activation and their Z statistics are given in
Table 1.
[View Larger Version of this Image (80K GIF file)]
Autobiographical episodic memory ecphory versus
baseline (PERSONAL-REST)
Comparing the presentation of autobiographical sentences with the
resting state (PERSONAL-REST) similarly resulted in relative rCBF
increases in both medial and superior temporal gyri, however, with a
lateralization toward the right hemisphere (Table 1,
PERSONAL-REST, Fig. 1). Again, the temporopolar cortex was
activated bilaterally, although more pronounced on the right than on
the left, and right dorsal frontal and right posterior cingulate areas
were activated together with the cerebellum (primarily left). Relative
rCBF decreases associated with the task were seen in the inferior
lateral parietal cortex (BA 40) bilaterally, the inferior
occipital cortex (BA 19), and the right dorsolateral
prefrontal cortex (BA 9) (Table 2
PERSONAL-REST).
Autobiographical versus nonautobiographical episodic memory
ecphory (PERSONAL-IMPERSONAL)
The comparison of the autobiographical sentences versus the
neutral sentences condition (PERSONAL-IMPERSONAL) demonstrated
preponderantly right hemispheric relative rCBF increases. These were
seen in the lateral and medial aspects of the right temporal lobe
including hippocampal, parahippocampal, and amygdaloid areas, the right
anterior insula, the right posterior cingulate area, the right
temporoparietal junction, and right prefrontal cortex (Table 1
PERSONAL-IMPERSONAL, Figs. 1, 2). Relative
rCBF decreases associated with this task again were seen in the lateral
inferior parietal cortex (BA 40/39) bilaterally, the left
inferior occipital cortex (BA 19), the right fusiform gyrus
(BA 18), and the left inferior frontal cortex (BA
10) (Table 2
PERSONAL-IMPERSONAL).
Fig. 2.
Functional anatomy of temporal activations during
affect-laden autobiographical memory. Same data as in Figure 1, but
here the group SPM{t} map has been sectioned
in sagittal, coronal, and transverse planes and is displayed on top of
an arbitrary MR image that has been normalized spatially to the same
anatomical space. The red cross-hair indicates the local
maximum within the area of activation. The color bar
indicates the Z statistics achieved (Z value).
This figure details the functional anatomy of temporal activations
associated with autobiographical memory (PERSONAL-IMPERSONAL) and
their relationship to underlying anatomy. Note that the activations are
predominantly on the right (left image corresponds to
subjects's left) and include temporomedial, temporolateral, and
insular areas.
[View Larger Version of this Image (103K GIF file)]
Single-subject data analysis
In five subjects, the pattern of activations was congruent to the
group one. One subject failed to show any significant activation; the
remaining showed primarily temporo-occipital activations.
DISCUSSION
The present study demonstrates functional neuronal activity
associated with the ecphory of autobiographical memory in a network of
primarily right hemispheric regions including temporomedial and
temporolateral cortex, amygdala and hippocampus-parahippocampus,
insula, posterior cingulate cortex, temporoparietal cortex, and
prefrontal cortex. Some of these areas are known to be part of or are
at least closely associated with retrieval of episodic long-term
memory. Furthermore, they are part of the expanded limbic system
network (Nauta, 1979) and are involved in affect-based information
coding.
Experimental design
A number of recent PET-studies investigated the brain regions
engaged in memory processing (see also Heiss et al., 1992; Frackowiak,
1994; Buckner and Tulving, 1995; McCarthy, 1995). Most closely related
to our experiment are studies focusing on an anatomical basis for the
retention of long-term verbal memory (Andreasen et al., 1995) or on
encoding or retrieval processes in verbal and visual memory (Grasby et
al., 1994; Kapur et al., 1994; Shallice et al., 1994; Tulving et al.,
1994a,b,c, 1996; Buckner et al., 1995; Fletcher et al., 1995a; Kapur et
al., 1995; Petrides et al., 1995) .
Whereas in the above studies, subjects actually acquired new
information before being scanned, in the present experiment, subjects
had to ecphorize (``re-evoke'') own past (autobiographical) events
taken from their childhood, adolescence, and early adulthood. While
being scanned, they listened to sentences, each describing an important
episode of their past. Neuronal activation during this condition was
compared with activation during a second condition in which subjects
listened to sentences containing semantically very similar material
taken, however, from another person that the subjects did not know. For
both conditions, subjects were instructed to imagine what had happened
during the described situation. By necessity, studying autobiographical
memory ecphory (PERSONAL) involves retrieval of episodic memory.
Because we were interested in the ecphory of autobiographical old
memory rather than of episodic memory per se, the latter was subtracted
from the first (IMPERSONAL-PERSONAL).
Areas activated
Medial temporal lobes
In our study, there was no temporomedial activation associated
with the ecphory of nonautobiographical memory and, indeed, only few
PET studies actually have reported activation of medial temporal lobes
with memory tasks (Squire et al., 1992; Grasby et al., 1993; Kapur et
al., 1995). By contrast, autobiographical memory ecphory led to strong
activation of left and right temporomesial areas including hippocampus
and amygdala. Interestingly, this activation was much more pronounced
on the right. Our data suggest that autobiographical memory ecphory
engages amygdaloid and hippocampal regions. This is in good accordance
with the idea that limbic structures of the temporal cortex are of
crucial importance for affect-sensitive mnestic processing.
Lateral temporal lobes
In the present study, both the middle and superior temporal gyri
were involved in nonautobiographical and autobiographical memory
ecphory. Areas activated included BA 21 and previously have been
associated with word-sentence recognition and understanding (Mazoyer
et al., 1993). However, when comparing PERSONAL with IMPERSONAL,
significant activations were seen in all three right temporal gyri, in
particular, the middle and superior temporal gyri. This suggests that
autobiographical memory ecphory, in addition to sentence recognition
and understanding required in the studied task, engages these brain
areas. This is concordant with experimental findings stressing the
importance of neocortical temporal areas in episodic memory
representation (Merzenich and Sameshima, 1993; Sakai and Miyashita,
1994) and supports the notion of a multimodal representation of
episodic memory.
Temporal poles
The temporal poles have been shown to be part of the cortical
representation of speech. This was demonstrated when subjects listened
to stories (Mazoyer et al., 1993) or read stories in silence (Fletcher
et al., 1995b). Both activation conditions (IMPERSONAL, PERSONAL)
involved the listening and understanding of sentences with narrative
character. These regions, anatomically distinct from the medially
situated allocortical temporal regions necessary for memory encoding
and consolidation (Markowitsch, 1995a), also seem to be involved in the
ecphory of autobiographical memory as a comparison of IMPERSONAL and
PERSONAL revealed activation of the right temporal pole.
Insula
Insular activation has been reported in many PET experiments. We
hypothesize that the insular activation during autobiographical memory
ecphory may be attributed to a strong activation of the limbic system
caused by the high emotional impact of the task.
Cingulate cortex
Clinical studies have established a posterior cingulate cortex
contribution to episodic memory (Rudge and Warrington, 1993).
Activation of this region has been described during encoding of verbal
memory (Grasby et al., 1993, 1994; Kapur et al., 1994; Fletcher et al.,
1995a). In the present study, the posterior cingulate cortex was
activated during the ecphory of autobiographical, but not of
nonautobiographical, memory. Reasons for this activation remain
speculative, although an activation of this important neural link
between the prefrontal cortex and the hippocampus (Goldman-Rakic et
al., 1984) causes little surprise. Furthermore, PERSONAL versus
IMPERSONAL ecphory may require differential attentional mechanisms
during the two activation tasks.
Prefrontal cortex
Activation of prefrontal cortex during memory-associated tasks has
been observed previously, and it has been proposed that such activation
may be related to the degree of monitoring and verification involved in
a recall task (Fletcher et al., 1995a). Additional support to this
hypothesis is provided by clinical observations of retrograde amnesia
in patients with combined damage of anterolateral temporal areas and
prefrontal areas (Markowitsch, 1995b), suggesting that the prefrontal
cortex acts as a kind of control center for effortful initiation of
recall (Jetter et al., 1986) and the sequencing and organizing of
information (Milner et al., 1985; Goldman-Rakic and Friedman, 1991;
Stuss et al., 1994; Goldman-Rakic, 1995). This, together with the
anterolateral temporal cortex which provides a connecting link to
posterior cortical centers of integration as the major storage places
of engrams (Markowitsch et al., 1985), implies an important role of the
prefrontal cortex in the initiation of directed ecphory. The results on
prefrontal activation obtained in the present study are in good
agreement with the above ideas, because the prefrontal cortex is
activated during the ecphory of autobiographical memory retrieval.
Cerebellum
A contralateral cerebellar activation has been observed
during numerous PET studies on memory tasks (Barker et al., 1991;
Grasby et al., 1994; Molchan et al., 1994; Tulving et al., 1994b;
Andreasen et al., 1995; Buckner and Tulving, 1995; Buckner et al.,
1995) and was found in the present study during autobiographical memory
ecphory. Andreasen et al. (1995) speculated recently that the
cerebellum may participate in memory processing apart from procedural
and automatic aspects of memory. Evidence for such a role comes from
both animal studies and neuropsychological experiments in patients with
acquired brain damage (Schmahmann, 1992; Daum et al., 1993; Grafman et
al., 1993; Leiner et al., 1993) .
Areas deactivated
The decreases associated with the differences among the conditions
seem to reflect reduced visual and spatial imagery (occipital and
parietal regions) (Ogden, 1993; Kosslyn et al., 1993), reduced effort
in language comprehension (BA 40, Wernicke area) (Démonet et al.,
1993), and less effort in general attentive functions (prefrontal
regions) (Tulving et al., 1994c).
Laterality
Based on PET activation studies (Shallice et al., 1994; Tulving et
al., 1994a,b,c, 1996; Nyberg et al., 1996) and clinical data from
neurological patients with selective retrograde amnesia in the episodic
(Kapur et al., 1992; Markowitsch et al., 1993a,b) or semantic memory
domain (De Renzi et al., 1987; Grossi et al., 1988), it has been
hypothesized that the right anterior temporal and right inferolateral
prefrontal cortex, structures interconnected via the ventral branch of
the uncinate fascicle, are involved in episodic memory retrieval,
whereas the same structural combination in the left hemisphere may be
responsible for semantic memory retrieval. The present findings,
indicating a huge preponderance of activation in the right hemisphere,
support this hypothesis for the episodic memory system in general and
for autobiographical memory retrieval in particular.
Further observed predominantly right-sided activation in the
periamygdaloid, hippocampal-parahippocampal, insular, and posterior
cingulate regions also may be explained as a correlate for the
frequently found emotional character of autobiographical memories
(Sarter and Markowitsch, 1985a,b; Adolphs et al., 1994, 1995;
Barbas, 1995; Cahill et al., 1995; Devinsky et al., 1995; LeDoux,
1995). Alternatively, one might speculate that at least in part the
right-sided preponderance of activations might be linked to the
self-representation involved in autobiographic memory ecphory.
Possible confounds
Sentences from the nonautobiographical and the autobiographical
conditions by necessity differed in that during the autobiographical
condition, material was presented that was (1) familiar to the subjects
for many years, (2) affectively more relevant, and (3) more personal.
Any or all of these factors could contribute to the described
activations. Furthermore, because subjects had to imagine what had
happened to another person during the nonautobiographical condition as
opposed to imagining what had happened to themselves during the
autobiographical condition, comparing the two conditions also could
reflect blood flow changes associated with imagining oneself versus
imagining others. These potential confounds will have to be addressed
by additional functional imaging studies.
PET (Martin et al., 1995, 1996) and neuropsychological evidence
(Shallice and Kartsounis, 1993) argue for the existence of separable
cortical regions involved in the storage or activation of different
semantic properties. One therefore might speculate that knowledge about
oneself could be another class of semantic knowledge. However, if such
a restricted kind of ``autobiographical knowledge module'' existed,
it most likely would recruit additional information such as
emotional-affective flavor from other regions (e.g., the amygdala),
making it altogether a wider network than those suggested presently for
singular categories of semantic information.
Clinical implications
The process of autobiographical memory ecphory is of particular
importance in patients with retrograde amnesia. It has been assumed
that their ``lost'' episodic information still exists in the brain as
engrams that cannot be retrieved consciously (Markowitsch, 1995b). Both
in patients with so-called focal retrograde amnesia and with
psychogenic amnesia, stored information most likely remains available
(Kapur, 1993; Markowitsch, 1995b), although it seems to be inaccessible
(Tulving and Pearlstone, 1966), i.e., it cannot be ecphorized. Although
it can be only speculated which mechanisms may be responsible for such
an inaccessibility, the present results provide some evidence for the
brain regions involved in autobiographical memory ecphory under normal
conditions.
Our study shows that ecphory of autobiographical memories is dependent
on a restricted network of mostly right hemispheric brain areas that
are in part different from those activated by retrieval of episodic
nonautobiographical memory. Nevertheless, the circuits for the ecphory
of active autobiographical and nonautobiographical memory are related.
Together with the results from neuropsychological investigations on
single cases with selective retrograde amnesia, this study provides
strong evidence that right hemispheric temporal cortical areas are the
key regions for autobiographical memory ecphory. They are supported by
surrounding right hemispheric ``satellite'' regions such as the
amygdala, hippocampus-parahippocampus, posterior cingulate, and
insular and prefrontal cortex.
FOOTNOTES
Received Dec. 19, 1995; revised April 16, 1996; accepted April 18, 1996.
We are grateful to our volunteers who made this study possible. We
thank both anonymous reviewers for helpful comments.
Correspondence should be addressed to Dr. Gereon Fink, Wellcome
Department of Cognitive Neurology, Institute of Neurology, 12 Queen
Square, London WC1N 3BG, UK.
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52
Memory
