European Journal of Neuroscience

European Journal of Neuroscience, Vol. 28, pp. 829–840, 2008 doi:10.1111/j.1460-9568.2008.06381.x

Autobiographical memory of the recent past following

frontal cortex or temporal lobe excisions

Laila Thaiss* and Michael Petrides

Montreal Neurological Institute and Department of Psychology, McGill University, Montreal, QC, Canada

Keywords: autobiographical memory, event sampling, frontal cortex, patients, temporal lobe, temporal order

Abstract
Previous research has raised questions regarding the necessity of the frontal cortex in autobiographical memory and the role that it
plays in actively retrieving contextual information associated with personally relevant events. Autobiographical memory was studied
in patients with unilateral excisions restricted to the frontal cortex or temporal lobe involving the amygdalo-hippocampal region and in
normal controls using an event-sampling method. We examined accuracy of free recall, use of strategies during retrieval and memory
for specific aspects of the autobiographical events, including temporal order. Patients with temporal lobe excisions were impaired in
autobiographical recall. By contrast, patients with frontal cortical excisions exhibited normal autobiographical recall but were less
likely to use temporal order spontaneously to organize event retrieval. Instruction to organize retrieval by temporal order failed to
improve recall in temporal lobe patients and increased the incidence of plausible intrusion errors in left temporal patients. In contrast,
patients with frontal cortical excisions now surpassed control subjects in recall of autobiographical events. Furthermore, the retrieval
accuracy for the temporal order of diary events was not impaired in these patients. In a subsequent cued recall test, temporal lobe
patients were impaired in their memory for the details of the diary events and their context. In conclusion, a basic impairment in
autobiographical memory (including memory for temporal context) results from damage to the temporal lobe and not the frontal
cortex. Patients with frontal excisions fail to use organizational strategies spontaneously to aid retrieval but can use these effectively if
instructed to do so.

Introduction
Functional neuroimaging has indicated that the prefrontal cortex,
medial and lateral temporal regions, and the posterior cingulate ⁄
retrosplenial cortex play some role in episodic autobiographical
memory (e.g. Gilboa et al., 2004; Svoboda et al., 2006). However,
changes in activity in particular brain regions observed in functional
neuroimaging do not prove that these regions are necessary for
autobiographical memory. The increased activity may reflect general
processes that may be involved during autobiographical memory
retrieval but may not be essential for it (e.g. increased attention in the
case of the frontal cortex). Studies of patients with brain lesions can
establish the necessity and role of a particular brain region but studies in
which the lesions are restricted to the area of interest are rare. For
instance, although some lesion studies have suggested a critical role for
the frontal cortex in autobiographical memory, the patients typically
also had extensive extra-frontal damage due to head injury (Kroll et al.,
1997; Levine et al., 1998), dementia or Korsakoff’s syndrome
(Kopelman, 1989, 1991; Kopelman et al., 1997). It has thus been
difficult to establish the relative importance of the prefrontal cortex in
autobiographical memory from these studies. Nevertheless, it is known
that patients with lesions to the frontal cortex exhibit inefficient use of
semantic organization and search strategies during active recall (Stuss
et al., 1978; Incisa della Rocchetta, 1986; Jetter et al., 1986; Gershberg
Correspondence: Dr L. Thaiss, *present address below.
E-mail: laila.thaiss@mail.mcgill.ca
*Present address: QACCH, 2400 Arbutus Road, Victoria, BC, Canada V8N 1V7.
Received 5 October 2007, revised 20 June 2008, accepted 24 June 2008
& Shimamura, 1995; Petrides, 2000) and hence the influence of such
impairments on autobiographical memory requires examination.
The present study examined patients with excisions restricted to
either the frontal cortex or the anterior part of the temporal lobe,
including significant removal of the amygdalo-hippocampal region,
in order to clarify the relative role of these two brain regions in
autobiographical memory. Previous studies tested autobiographical
memory with structured questionnaires (Dall’Ora et al., 1989;
Kopelman, 1989; Della Sala et al., 1993; Viskontas et al., 2000)
or various adaptations of the cued-word test (Crovitz & Schiffman,
1974; Zola-Morgan et al., 1983; Baddeley & Wilson, 1986;
Moscovitch & Melo, 1997). In these studies, the accuracy of
autobiographical memory recall was verified by asking relatives or
friends about the events or by later retesting the patients. Because
these methods pose difficulties in terms of the interpretation of the
accuracy and quality of the patient’s retrieval, we adopted an
experience-sampling method (Brewer, 1988). This method has the
advantage of providing a detailed and accurate record of the events
that the subjects experienced. In addition, as the personally
experienced events are linked in time and organized in a diary,
their retrieval encourages active search and organization during free
recall, thus permitting examination of these processes in relation to
autobiographical memory. The subject’s spontaneous use of temporal
organizational strategies during free recall was studied separately
from the ability to order events in time. Finally, analysis of the
accuracy and detail of subject’s memories allowed us to explore
whether they relied on generic autobiographical knowledge during
recall, as opposed to having specific episodic memories.

ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
830 L. Thaiss and M. Petrides
Materials and methods
Experimental subjects
Twenty-nine individuals who had undergone unilateral cortical
excisions for the treatment of pharmacologically intractable focal
epilepsy and 16 neurologically normal individuals participated in this
study. All subjects gave written informed consent according to the
Declaration of Helsinki for participation in the study and for release of
magnetic resonance images (MRIs) and neuropsychological assess-
ment results. The study was approved by the ethics committees of the
Montreal Neurological Institute and the Department of Psychology,
McGill University. The subjects were provided with an honorarium for
their participation.
All subjects were fluent speakers of English and ⁄ or French.
Bilingual subjects responded in the language in which they had been
formally educated. Gender, age, language, education and full-scale IQ
information for the subject groups are provided in Table 1. Full-scale
IQ scores were available only for the four patient groups. The groups
did not differ significantly with respect to age (F4,40 = 1.58, P = 0.20),
length of education (F4,40 = 0.45, P = 0.77) or full-scale IQ
(F3,25 = 0.03, P = 0.99).
In order to ensure that the memory performance obtained in this
study could be attributed with confidence to damage of either the
prefrontal cortex or the anterior temporal region, we established very
strict inclusion criteria. Patients were excluded from the study if they
met one or more of the following criteria: (i) seizure disorder arising
from or associated with diffuse cerebral damage; (ii) bilateral
independent electrographic abnormalities; (iii) rapidly growing neo-
plasm; (iv) bilateral or right hemisphere speech representation
as determined by the preoperative intracarotid amobarbital procedure;
(v) damage to the brain that was outside the operated area, such as
damage contralateral to the side of the operation or damage involving
subcortical structures (e.g. basal ganglia and basal forebrain region);
(vi) full-scale IQ below 80 on the Wechsler Adult Intelligence Scale-
Revised; (vii) age above 55 years old and (viii) evidence of impaired
receptive or expressive language abilities.
Frontal cortical excisions
There were seven patients with an excision restricted to the left side
of the prefrontal cortex and four with a right-sided excision. A
postoperative MRI of the brain was available for 10 of the 11 patients
with frontal cortical excisions. For the one patient without a
postoperative MRI, the location and extent of the excision was
determined from a preoperative MRI. The available MRIs were
transformed into the Montreal Neurological Institute standard stereo-
taxic space, which is based on the Talairach standardized proportional
stereotaxic space (Talairach & Tournoux, 1988), in order to compare
the size and location of the excisions across patients. The location and
extent of the surgical resections within the frontal cortex were
Table 1. Subject groups
Gender
Group M F Age (years)*
Left frontal 6 1 29.3 (18–43)
Right frontal 2 2 28.3 (20–35)
Left temporal 5 4 35.2 (22–43)
Right temporal 4 5 36.9 (27–43)
Normal control 7 9 35.1 (23–59)
*For Age, Education and Wechsler full-scale IQ, data are presented as means, with range in parentheses. M, male; F, female; NA, not available.
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
identified and labeled using an interactive three-dimensional drawing
program (DISPLAY) that was developed at the Montreal Neurological
Institute (see Thaiss & Petrides, 2003 for details). The location and
extent of the excisions from the left or right frontal cortex are
illustrated in Fig. 1. The etiology of the seizures included low-grade
tumor (n = 1), arteriovenous malformation (n = 1), cavernous hem-
angioma (n = 2) and idiopathic epilepsy (n = 7). The anterior speech
area (Broca’s region) was spared in all patients with left frontal
excisions. The median time elapsed since surgery for the patients in
the frontal group was 2 years and 1 month, with a range of 3 months
to 25 years.
Temporal lobe excisions
There were 18 patients with unilateral temporal lobe resections in this
study. The unilateral surgical removals in this group of patients
involved either a selective amygdalo-hippocampectomy (four patients
with left- and four with right-sided excisions) or an excision of the
anterior temporal cortex that included the amygdala and hippocampus
(five left- and five right-sided excisions). In the selective amygdalo-
hippocampectomy procedure, a small narrow excision was made
through the first or second temporal gyrus or through the floor of
the superior temporal sulcus, allowing the surgeon to remove the
amygdala (4 ⁄ 5 or total resection) and the anterior 2.5–3.5 cm of the
hippocampal formation together with the surrounding parahippocam-
pal gyrus (2.5–4 cm), while sparing the rest of the temporal neocortex.
In anterior temporal lobe resections, the anterior-most temporal cortex
was removed (ranging from 4 to 6 cm along the Sylvian fissure and
from 4.5 to 6 cm along the base of the temporal lobe), as well as the
amygdala, anterior portion of the hippocampus (between 1.5 and
4 cm) and adjacent parahippocampal cortex (between 1.5 and 4 cm).
Thus, in both the anterior temporal lobe group and the group with
selective amygdalo-hippocampal excisions, a significant proportion of
the amygdala, hippocampus and adjacent parahippocampal cortex was
removed. Note that, in these surgical removals, the entorhinal and
perirhinal cortex is considered to be part of the anterior parahippo-
campal cortex adjacent to the amygdala and the anterior-most part of
the hippocampus. Thus, the excision included the limbic structures on
the medial part of the temporal lobe (amygdala, hippocampus and
adjacent entorhinal ⁄ perirhinal cortex as well as the more posterior
parahippocampal cortex lying next to the removed part of the
hippocampus) in addition to a minor part of the lateral temporal
neocortex in the amygdalo-hippocampectomies and a more sizeable
part of the anterior temporal neocortex in the classic anterior temporal
removals. Figure 2 shows an example MRI (in standardized propor-
tional stereotaxic space) of each of the two types of anterior temporal
lobe excisions.
The etiology of seizures in this group of patients included idiopathic
epilepsy (n = 13), febrile illness (n = 2), low-grade tumor (n = 1),
perinatal illness (n = 1) and meningitis (n = 1). The median time
Language
Education Wechsler
French English (years)* full-scale IQ*
4 3 13.0 (10–17) 98.0 (83–116)
0 4 13.0 (11–17) 96.25 (90–110)
3 6 13.9 (10–17) 96.5 (86–119)
6 3 14.0 (11–17) 100.2 (83–124)
8 8 14.3 (11–19) NA
European Journal of Neuroscience, 28, 829–840
 Autobiographical memory and the frontal cortex 831

Fig. 1. The location and extent of the surgical resections for patients with frontal cortical excisions are illustrated in these brain images. The excisions are shown in
red on the three-dimensional MRI reconstructions and on coronal sections through the site of the excision. Left- or right-sided lesions are represented on the left and
right side of the coronal sections, respectively. The numbers above each coronal section refer to the y (anterior ⁄ posterior) stereotaxic coordinate in standardized
proportional space. The signs of these coordinates refer to sections anterior (+) or posterior ()) to the anterior commissure. The patient’s identifying number and the
volume of each patient’s surgical resection (in cm3) are indicated below the brain images.

ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
European Journal of Neuroscience, 28, 829–840
832 L. Thaiss and M. Petrides
Fig. 2. Examples of the location and extent of the surgical resections in a right-sided selective amygdalo-hippocampectomy and a right anterior temporal lobe
resection are illustrated in these three-dimensional MRI reconstructions and coronal sections. The excision is shown in green. The numbers above each coronal
section refer to the y (anterior ⁄ posterior) stereotaxic coordinate in standardized proportional space. The signs of these coordinates refer to sections anterior (+) or
posterior ()) to the anterior commissure.
elapsed since surgery for the patients with temporal lobe excisions was
1 year and 7 months, with a range of 3 months to 30 years.
Experimental procedures
The subjects filled out a diary form detailing the activity ⁄ activities that
they were engaged in each time they were paged. At 1 week to 10 days
after the diaries had been completed, subjects were contacted and their
autobiographical memory of the activities that they had recorded in their
diaries was tested during a telephone interview. In the first part of the
memory test, we examined the number of activities from the diaries that
the subjects were able to retrieve and their recall accuracy in order to
establish whether the subject groups differed in their free recall of
autobiographical memories. As previous studies had implicated the
prefrontal cortex in the online maintenance and initiation of search for
past events (e.g. Petrides, 2002; Levine, 2004) and perhaps also in the
retrieval of contextual information (e.g. Janowsky et al., 1989;
Shimamura et al., 1990; Schacter et al., 1998), we also examined
whether the subject groups differed in their use of retrieval strategies
during recall (specifically, whether they organized their recall sponta-
neously by the temporal order of events) and in their ability to organize
their memories in the order of occurrence when instructed to do so.
In the second part of the memory test, the quality of the subject’s
autobiographical memories was examined in order to confirm that the
subjects were, in fact, retrieving episodic autobiographical information
(i.e. personally relevant memories from a unique time and place) as
opposed to generic autobiographical knowledge (Brewer, 1988). For
this purpose, a cued recall test was administered in order to examine the
amount of detail that the subjects were able to retrieve about each one
of the events that they had recorded in their diaries (regardless of
whether they had recalled the event in the free recall test). This allowed
us not only to examine the subject’s memories for specific types of
autobiographical information associated with an event (e.g. the day,
who they were with, the preceding activity, etc.) but also whether the
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
details retrieved represented unique information that could not be
inferred and, therefore, true episodic autobiographical memories.
Diaries (event sampling)
Each subject was provided with a pager, 16 diary forms in individual
envelopes, written instructions and a prepaid courier service envelope.
Two consecutive days were chosen by each subject to do the event
sampling and a telephone interview was booked for 1 week to 10 days
following the event sampling. The subjects were told that the study
was about people’s daily activities and that the telephone interview at
the end was ‘to go over how things went’. They were not told that
there would be a memory test during the telephone interview. The
subjects were instructed to fill out a separate diary form each time the
pager beeped, showing the number ‘1111’, and then to seal the form in
its own envelope.
As it was critical that the diaries represent the subject’s current
activities, rather than their memory of an event, the subjects were
instructed to fill out the diary form as soon as they received the paging. If
this was not possible (e.g. a subject was driving) they were required to fill
it out as soon as possible. The subjects wrote down the time that they had
completed each form so that any time delays in recording the event in the
diary could be identified later by the examiner. For events that the
subjects were not willing to disclose, they were told to leave the form
blank. The subjects were asked to list in their diaries the activities that
they were engaged in when paged and to specify: (i) the time; (ii) the day;
(iii) their current location; (iv) the details of the activities and the context
or goal of the activities; (v) their interest level and their feelings at the
time (using a five-point Leikert scale); (vi) the people that they were
with; (vii) the general topic of their conversation; (viii) the activities of
the previous half-hour and (ix) what they planned to do next (see
Supplementary material, Appendix S1).
During the event sampling, the subjects were paged at semi-
random intervals between 1.5 and 4 h apart. Each subject was paged
a total of 12–14 times in the 2-day period. Each subject had to have
European Journal of Neuroscience, 28, 829–840

completed at least 10 diary forms in order to be given the memory
test 1 week later. At the end of the 2-day event-sampling period, the
subjects returned the completed diaries (individually sealed in
envelopes) and the pager to the experimenter by prepaid courier.
Two subjects failed to carry the pager with them during the 2-day
experience-sampling period and did not fill out the diary forms when
paged. For these two subjects, the instructions for the study were
provided again and they repeated the event sampling approximately
5 weeks later. Only their memory of the events during this second
event-sampling period was tested as the first event-sampling expe-
rience was considered invalid.
In this paper, each diary form that was filled out by a subject
over the 2-day event-sampling period is referred to as a diary
‘event’. Each event includes information about the activity (or
activities), context and other details, the persons the subjects were
with, previous activity, etc. Occasionally, a subject’s activity was
the same over several diary forms (e.g. watching TV over several
hours). For the purpose of testing, the different diary forms
containing the same activity were treated as separate events. In
most instances, although the activity was listed as the same at the
top of the diary form, the subjects described different details on
their forms (e.g. ‘watching the news’; ‘now I’m watching Seinfeld,
the cat is sitting on my lap’). When more than one activity was
listed on a diary form (e.g. eating breakfast and reading the
newspaper), these were considered a single diary event.
Telephone interview
The subjects were interviewed over the telephone 1 week to 10 days
after the event-sampling period. They were asked to be alone in a
room during the interview and to be uninterrupted for about 1 h. The
interview was organized in two parts. In the first part, free recall of the
diary events was tested. The subjects verbally listed all of the activities
that they had filled out on the diary forms over the 2-day event-
sampling period. All responses were transcribed verbatim by the
examiner. When free recall was completed, the subjects were told how
many times they had been paged and were asked to recall again as
many events as they could. Finally, in order to examine whether recall
of the diary events improved when a retrieval strategy was employed
during recall, the subjects were asked to recall the events one more
time but this time in the temporal order in which they had occurred
(temporally ordered recall). Those subjects who had recalled their
activities in temporal order during the free recall trial, as verified by
their diary entries (17 subjects), were not asked to recall the events
again.
For the second part of the interview, a cued recall test was given in
order to examine the level of detail of the information that the subjects
could recall for each one of the diary events, including those diary
events that they had failed to retrieve during the free recall attempts.
The examiner had prearranged the diary forms in random order and a
cardboard mask was placed on each diary form so that it covered all of
the responses on the form, except for the brief description of the
activity engaged in. The mask was necessary to avoid biases in the
questioning. The examiner read the activity entered in the diary form
and asked the subject to provide the remaining information by verbally
prompting him ⁄ her with each one of the nine category headings: day,
time, location, details and context, interest level and feelings,
interactions, conversations, previous half-hour and next half-hour. If
more than one activity was listed on the diary form, either both
activities or the activity that provided the most information about the
event was read to the subject by the examiner. If the activity was the
same on two or more of the diary forms, additional identifying
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
European Journal of Neuroscience, 28, 829–840
Autobiographical memory and the frontal cortex 833
information, specifically the time or the day of that particular event,
was provided to the subject.
Scoring
Diaries
The events recorded in the diaries were classified into one of 34 action
categories in order to examine how varied the activities of individual
subjects were over the 2-day period. The relation between the degree
of variability of the activities and the ability to recollect these activities
was examined. It was postulated that more frequently performed
activities (e.g. watching television) would be less salient for subjects
and more difficult to remember. A word count of each diary was also
calculated in order to examine whether the subjects differed in the
amount of information that they had provided and whether this was
correlated with their memory of the autobiographical events.
Two raters who were ‘blind’ to the identity of the subjects and who
were both fluently bilingual (English and French) independently
scored the subject’s memory responses on the free recall and cued
recall tests by comparing these responses to the events recorded in the
subject’s diary forms. They also scored the cued recall responses for
each event according to the quality of the memory trace. The raters
were provided with detailed scoring instructions and were each trained
with five example protocols. The inter-rater reliability was calculated
for all measures and determined to be acceptable (see supplementary
Appendix S1).
Accuracy of free recall
All of the subject’s free recall responses were included in the accuracy
score. Accuracy of free recall was determined on the basis of whether
each activity recalled was consistent with one of the subject’s diary
events (taking into consideration information from diaries that specified
the context, details, what they were doing just before the event, etc). As
the subjects were specifically asked to recall events that they had
recorded on their diary forms, activities that were not consistent with the
diary events were scored as intrusions. The free recall accuracy score
was a proportion calculated as follows: the number of activities correctly
recalled over the total number of diary events recorded.
Organization of free recall
In order to examine the use of organization strategies during free
recall, the free recall responses were scored on a scale from 1 to 3
according to whether the subject had spontaneously organized their
retrieval of the activities by the temporal order of occurrence (score
of 3) or not (score of 1). A score of 2 indicated that there was
some temporal organization apparent in their recall (e.g. the
activities were organized by the day) but it was not used
consistently to aid retrieval of events. The ‘temporal organization’
score was a measure of retrieval strategy and was independent of
whether the subject’s responses corresponded accurately to the
events recorded in the diaries. Furthermore, this score was not
dependent on the accuracy of the temporal order as established by
comparison with the diary events. Instead, the score was determined
on the basis of temporal cues provided by the subject during their recall
(e.g. ‘on the first paging I was still in bed, then it beeped while I was
having breakfast...’ or ‘on Saturday I was paged while I was
shopping...’). The position of activities that had clear temporal
associations (e.g. breakfast, dinner and afternoon nap) was also taken
into consideration in determining this score. For example, breakfast is
one of the first events in the day and therefore if it appeared at the end of
834 L. Thaiss and M. Petrides
a list of activities during recall, it was assumed that at least some events
were misplaced temporally.
Accuracy of event recall when temporally ordered
We also examined whether the instruction to the subjects to organize
their recall by temporal order improved their memory for events. The
subject’s responses after the instruction to recall the diary events in the
correct temporal order was therefore scored for accuracy. The same
scoring criteria were used in this measure as in the free recall test. The
temporally ordered recall score was a proportion calculated as follows:
the number of activities correctly recalled over the total number of
diary events recorded.
Accuracy of temporal order
The accuracy of temporal order was scored separately from the
spontaneous use of temporal organization during recall. This score
was based on whether the subject had recalled the relative sequence of
each activity correctly. The accuracy of temporal order was a
proportion score calculated as follows: the number of recalled
activities that were in the correct sequence over the total number of
activities recalled correctly.
Cued recall
In the cued recall test, the subjects had to retrieve details about each one
of the diary events from nine different categories of information (e.g. the
day and time that the event took place, the persons they were with, etc).
The two independent raters scored responses to each one of the nine
categories of information for each diary event that the subjects had been
questioned on. A three-point scale was used: 1, wrong or no memory of
this information; 2, some correct or inferred information and 3, generally
correct information that contained some non-inferable information. The
data from this test were used to examine whether the subject groups
differed in terms of the type of information that they remembered about
an event. For example, was a particular group of subjects less likely to
remember information about the context of the events?
Quality of memory trace
The question of whether subjects had episodic autobiographical
memories for the events that they recalled, i.e. a memory that
represents a personal experience that happened at a unique time and
location, or whether they relied primarily on autobiographical
knowledge or generic information about what they generally did
was addressed with this qualitative scale. For each subject, the cued
recall responses for each diary event were classified according to
one of seven categories based on a scale defined by Brewer (1988).
The classification scheme represented the quality or type of the
memory trace for the event (e.g. the success with which subjects
recalled specific information about the context and content associ-
ated with the event) and included the following: (i) ‘don’t know’,
i.e. they had no memory of the event; (ii) ‘wrong event’, i.e. they
retrieved a different event (not in the diary) on the basis of the cue; (iii)
‘wrong time slice’, i.e. their memory for the event cued was misplaced in
time compared with the diary entry (e.g. the subject recalled that they
were on their way to the car wash when, in fact, according to the diary,
they were waiting at the car wash for their car); (iv) ‘overt error’, i.e. they
recalled the event but made an overt error in their memory of it (e.g.
remembering that someone was there when, in fact, no-one was there);
(v) ‘inference’, i.e. the retrieved information was probably inferred on
the basis of the cue provided; (vi) ‘correct’ but with little detail; (vii)
‘correct with details’, i.e. the subject included a detailed account of the
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
event that suggested a vivid memory for the event (e.g. specific details of
a conversation or of their thoughts at the time).
Results
The median scores and distribution of the individual scores for the
different tests of memory (specifically, free recall, temporal ordered
recall and temporal organization scores) were similar for the patients
with excisions from the right and left frontal cortex. The data of all
patients with frontal cortex excisions were therefore combined into a
single group of 11 subjects in order to increase statistical power.
Statistical analysis
When the data were based on ordinal scales (e.g. the three-point scales
used for temporal organization and cued recall), non-parametric tests
were used to test the significance of group differences. Post hoc tests
following the Kruskal–Wallis non-parametric anova compared the
differences between ranks to the standard normalized score at
a = 0.05 (unless otherwise indicated), as described by Siegel &
Castellan (1988). All post hoc testing following parametric anovas
used the Neuman–Keuls method with a significance level of 0.05. The
reported significance levels are two-tailed.
Diaries
As described in Experimental procedures, two subjects repeated the
experience-sampling procedure because of problems in the first
attempt and therefore the telephone interview and tests were based on
diary entries from their second event-sampling experience.
Each subject filled out between 10 and 15 diary forms (overall
mean, 13.5, SD 0.92). There was no significant difference between the
four groups in the number of diary forms filled out by subjects
(one-way anova, F3,41 = 2.02, P = 0.12) or in the variability of the
activities (i.e. the number of different categories) reported in the diaries
(one-way anova, F3,41 = 0.94, P = 0.43, n.s.; mean categories, 8.84).
There was no correlation between the variability in the activities over
the 2-day period and the subject’s recall performance (all subjects,
r = )0.07, P = 0.63; just the normal control subjects, r = 0.17,
P = 0.54). See supplementary Appendix S1 for a list of the activities.
A one-way anova on the word count of the diaries for the various
groups indicated a significant difference between the groups
(F3,41 = 3.18, P < 0.05). Patients with left temporal lobe excisions
provided significantly longer (i.e. more words) descriptions of the
events in comparison with all of the other subject groups. There was
no significant correlation between the length of the descriptions in the
diaries and the proportion of diary events correctly recalled during free
recall (all subjects, r = 0.17, P = 0.27; just the normal control
subjects, r = 0.38, P = 0.15).
Free recall
Use of temporal sequence to organize recall of events
Overall, 17 of the 45 subjects spontaneously used temporal order to
structure their retrieval of the diary events. A chi-squared analysis,
used to examine the differences in the frequency of scores across the
groups, was significant [v2(6) = 19.09, P = 0.004]. Further break-
down of this analysis showed significant differences between the
groups in the number of subjects who scored 1 [v2(3) = 7.98,
P < 0.05; v2(3)critical = 7.82, P = 0.05]; specifically, the patients with
European Journal of Neuroscience, 28, 829–840
 Autobiographical memory and the frontal cortex 835

Fig. 3. Frequency with which subjects spontaneously organized their free

recall. The bar graph for each subject group represents the number of subjects
who scored 1, 2 or 3 on a measure of the spontaneous use of organization
strategies during recall. A score of 3 indicates that the subject spontaneously
organized their retrieval by temporal order and 1 means that they did not. A
score of 2 indicates that some organization is apparent. The number of subjects
in each group is indicated in parentheses. Abbreviations: F, frontal cortical
excisions; N, normal control group; LT, left temporal excisions; RT, right
temporal excisions.
Fig. 4. The mean proportion of diary events that were correctly recalled by
each subject group is shown for the free recall condition and for the subsequent
recall condition in which subjects were specifically instructed to temporally
order their recall of the events. Error bars represent SEM. Abbreviations as in
Fig. 3.

frontal excisions were more likely than the other groups to use no
organization in their recall. In addition, patients with frontal cortex
excisions were significantly more likely to score 1 than 2 or 3
and normal control subjects were significantly more likely to score
3 than 2 or 1 [v2(2) = 6.45 and 7.57, P < 0.05, respectively;
v2(2)critical = 5.99, P = 0.05] (see Fig. 3).

Performance in free as opposed to temporally ordered recall

A two-way repeated-measures anova (group · recall type) was used
to examine the proportion of diary events that were correctly recalled
in the initial ‘free recall’ test compared with the subsequent
‘temporally ordered recall’ test in which the subjects were instructed
to retrieve events according to their temporal sequence. This analysis
revealed a significant interaction between the group and the recall type
(F3,41 = 5.51, P = 0.003). Post hoc analyses showed that, during both
the free and temporally ordered recall, patients with left and right
temporal excisions recalled a smaller proportion of the diary events
correctly than normal control subjects and patients with frontal cortical
excisions. In contrast, the patients with frontal cortical excisions did
not differ from normal control subjects in the free recall test and were Fig. 5. The mean proportion of recall responses that were classified as
the only group to show a significant improvement in their recall during intrusions in the free recall condition and in the temporally ordered recall
the subsequent temporally ordered recall test when they were told to condition is shown for each of the subject groups. Error bars represent SEM.
Abbreviations as in Fig. 3.
organize their retrieval by the sequence of occurrence. The perfor-
mance of the frontal group was significantly better than that of the
control group in the temporally ordered recall condition (Fig. 4). between the first free recall and the subsequent temporally ordered
A two-way repeated-measures anova (group · type of recall) was recall trial. Furthermore, in temporally ordered recall, these subjects
used to examine the proportion of activities remembered that were had a significantly higher proportion of responses that were intrusions
intrusion errors (i.e. activities ‘remembered’ but that were not part of compared with any other subject group (Fig. 5).
the diaries) in free recall compared with the later temporally ordered
recall. Results of this analysis revealed a significant interaction Accuracy of the memories for the temporal sequence of events
between the groups and recall type (F3,41 = 3.05, P = 0.039). Post hoc The proportion of correct events that were recalled in the correct
analysis showed that the left temporal lobe group (but not any other temporal sequence was examined using a one-way anova. In this
group) exhibited a significant increase in the proportion of intrusions analysis, the sample size was 44 subjects instead of 45, as one subject

ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
European Journal of Neuroscience, 28, 829–840
836 L. Thaiss and M. Petrides
Fig. 6. The mean proportion of recall responses that were in the correct
temporal sequence in the temporally ordered recall task. Abbreviations as in
Fig. 3. Error bars represent SEM.
from the right temporal group had recalled only one event and
therefore had no temporal accuracy score. There was a significant
difference between the groups (F3,40 = 3.32, P = 0.029) and post hoc
testing showed that the left temporal group was significantly impaired
in recalling the temporal sequence of the events from their diaries
compared with the group with frontal cortical excisions and the normal
control group (Fig. 6).
Cued recall
Quantitative measures
The cued recall test examined the subject’s ability to recall specific
information (from nine categories) about the individual diary events.
The Kruskal–Wallis one-way anova by ranks was used to assess
differences between the groups for each one of the categories of
information. A significance level of a = 0.01 was used to correct for
the multiple comparisons. Results of these analyses revealed
significant group differences for the subject’s memory of the day
on which the event took place [v2(3) = 12.47, P = 0.005], the
amount of detail remembered [v2(3) = 11.35, P = 0.01], memory for
what they had been doing just prior to the event [v2(3) = 11.03,
P = 0.01] and memory for what they said they were going to do next
[v2(3) = 10.11, P = 0.01]. Post hoc tests indicated that the left
temporal group was impaired in their memory of the day on which
events occurred and for the details of the events in comparison with
the normal control group. Patients with right temporal excisions were
impaired in their memory for the day and for what they said they
were going to do next in comparison with the normal control
subjects (Fig. 7).
Qualitative scale
For each subject, the cued recall responses for each diary event were
classified into one of seven categories according to the quality or
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
success of the retrieval. Due to the small number of ‘wrong event’
responses, these were combined with ‘don’t know’ responses, thus
reducing the total number of categories to six. A two-way repeated-
measures anova was used to examine the differences across the
groups in the frequencies of these classifications. There was a
significant interaction between group and the different classifications
(F15,205 = 1.9, P = 0.025). Simple, main effect analyses, using the
pooled error term and degrees of freedom, were used to examine this
interaction. For each one of the subject groups, the pattern of
distribution of responses across the six categories was examined. As
expected, the number of responses classified in each one of the six
categories differed significantly for each one of the groups (frontal
cortex, F5,164 = 9.42, P < 0.01; left temporal, F5,164 = 8.79, P < 0.01;
right temporal, F5,164 = 7.61, P < 0.01; normal control,
F5,164 = 27.82, P < 0.01; Fcritical5,164 = 3.14, P = 0.01). For the
groups with frontal cortical excisions and the normal control subjects,
there were significantly more responses classified as ‘correct’ than any
other category. However, the number of responses classified as
‘correct’ were not significantly different from the number classified as
‘inference’ in the left temporal and right temporal lobe groups. For
each of the two temporal lobe subject groups, responses classified into
these two categories were significantly more common than those
classified in the other categories (Fig. 8).
Discussion
The present study demonstrated that patients with left or right
temporal lobe excisions that included significant involvement of the
limbic medial temporal region (amygdala, hippocampus and
entorhinal ⁄ perirhinal and parahippocampal cortex) are impaired in
autobiographical memory retrieval compared with control subjects and
patients with frontal cortical excisions. Specifically, the patients with
temporal lobe excisions recalled fewer events and fewer details of
these events (Figs 4 and 7) and they were significantly more likely to
produce plausible intrusions drawn from their autobiographical
knowledge when pressed to produce more responses, in comparison
with the other groups of subjects (Fig. 5).
Patients with frontal cortical excisions showed no impairments in
autobiographical memory retrieval compared with the normal control
subjects. These patients, however, differed from the other groups in
that they were less likely to use retrieval strategies spontaneously
during free recall (Fig. 3), a finding consistent with results obtained in
previous studies (e.g. Stuss et al., 1978; Incisa della Rocchetta, 1986;
Jetter et al., 1986; Gershberg & Shimamura, 1995; Petrides, 2000).
Specifically, during free recall the majority of normal control subjects
and many of the temporal lobe patients organized retrieval of their
event memories by the order in which they had occurred. This strategy
was used spontaneously by only three of the 11 patients with frontal
cortical excisions. It is important to note, however, that the patients
with frontal cortical lesions were able to retrieve the temporal order of
the diary events as accurately as normal control subjects when
instructed to do so (Fig. 6). It appears, therefore, that the lack of
strategy use by these patients during free recall is a failure to initiate
the strategy rather than an inability to use it. This finding is again
consistent with results from earlier studies that also showed that
patients with frontal lesions can use strategies when instructed to do so
although they may fail to use them spontaneously (e.g. Jetter et al.,
1986; Hirst & Volpe, 1988; Incisa della Rocchetta & Milner, 1993;
Gershberg & Shimamura, 1995). Interestingly, when the patients with
frontal cortical excisions were provided with a retrieval strategy, they
outperformed the normal control subjects in terms of the retrieval
European Journal of Neuroscience, 28, 829–840
 Autobiographical memory and the frontal cortex 837

Fig. 7. Mean cued recall scores for each of the subject groups for the different aspects of the events (e.g. time, day, current location, etc.). Only the four scales (of
nine) that showed significant differences (at a = 0.01) between the subject groups are presented here. A score of 3 indicates an accurate recollection of that specific
aspect of the event, 2 means that there was incomplete or inferred information and 1 indicates that the information was either not recalled or was incorrect. Error bars
represent SEM. Abbreviations as in Fig. 3.

accuracy of their autobiographical events (Fig. 4). In other words,
even in the face of poor recall organization, the patients with frontal
lesions could remember as well as the normal control subjects who
had the benefit of organized recall and, when instructed to recall their
memories in an organized fashion, the frontal patients recalled more
than the normal subjects. Thus, the present study provides strong and
unambiguous evidence that, when the lesions are restricted to the
frontal cortex, they do not affect the ability to recall autobiographical
memories accurately.
A recent meta-analysis of functional imaging studies of autobio-
graphical memory found that nearly all of the studies reviewed
reported activation of the prefrontal cortex (Svoboda et al., 2006). It
has been suggested that the ventrolateral prefrontal cortex is involved
in the strategic retrieval, verification and controlled retrieval of
information from posterior regions (Fletcher & Henson, 2001;
Petrides, 2002, 2005; Cabeza & St Jacques, 2007) and that the
superior medial and lateral prefrontal cortex and dorsal component of
the anterior cingulate and frontopolar cortex are involved in memory
reconstruction (Cabeza & Nyberg, 2000; Duncan & Owen, 2000;
Fletcher & Henson, 2001). How is it possible then that the patients
with frontal cortex excisions performed so well in the autobiographical
memory tests in the present study?
It has been argued that lateral frontal cortical lesions are not likely
to affect memory retrieval when retrieval can be based on the clear
unambiguous recognition of familiar as opposed to unfamiliar stimuli
or when strong and unambiguous stimulus-to-context relations can
trigger the retrieval of specific information (Petrides, 2002, 2005). The
involvement of the lateral prefrontal cortex and, in particular, the
ventrolateral prefrontal cortex increases as retrieval requires active
top-down control processing when memory information is embedded
in complex and ambiguous relations (Petrides, 2002, 2005). The
present autobiographical memory experiences were embedded in their
own unique contexts. Thus, during free recall, memories of specific
events were triggered by their unambiguous associations with
particular contexts. We did not examine here autobiographical
memory retrieval under conditions in which the same stimuli enter
into multiple and ambiguous relations with various contexts that
would tax prefrontal control processing because the question that we
aimed to address was whether autobiographical memory per se (and
not resolution of ambiguity in memory) depends on prefrontal cortex.

ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
European Journal of Neuroscience, 28, 829–840
838 L. Thaiss and M. Petrides
Fig. 8. Cued recall responses were classified according to one of six qualitative categories. The mean number of cued recall responses in each category is shown
here for each of the subject groups. Error bars represent SEM. Abbreviations as in Fig. 3.
The findings provided a clear answer, i.e. autobiographical memory
depends more on the classical medial temporal (i.e. hippocampal)
declarative memory circuit rather than the prefrontal cortex. It remains
to be tested whether the prefrontal cortex would become increasingly
necessary with increasing ambiguity in the relations of autobiograph-
ical information in memory.
The present study also addressed another major issue with regard
to the role of prefrontal cortex in memory. It has often been
suggested that patients with frontal cortical excisions have specific
deficits in retrieving the temporal context of long-term memories
(Janowsky et al., 1989; Kopelman et al., 1997; Schacter et al.,
1998). This suggestion is largely an inference based on the well-
known failure of patients with frontal lesions on the recency task
(Milner, 1971; Milner et al., 1991). In this task, subjects observe a
long list of items (words, concrete pictures or abstract designs) and,
occasionally, they are presented with either pairs of stimuli, both of
which were seen in the sequence (e.g. the 25th stimulus vs. the 39th
stimulus), or pairs in which one of the stimuli was seen but the other
is novel. When presented with the pairs, subjects have to decide
whether the stimuli were previously seen or not and, if both were
seen, which of the two items appeared earlier in the long sequence.
Patients with frontal cortical excisions are not impaired in recogniz-
ing whether particular stimuli are familiar or novel but are severely
impaired in judging the relative order of two familiar stimuli (Milner,
1971; Milner et al., 1991). As the long sequence exceeds by far the
subject’s immediate memory span and there is no external structure
(e.g. no contextual information) in the arbitrary long sequence of
unrelated items that are presented, the judgment of the relative
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
recency ⁄ order of any two test items is by necessity a judgment of
subtle differences in their relative saliency ⁄ familiarity (e.g. Milner,
1971; Shimamura et al., 1990; Milner et al., 1991; Mangels, 1997).
A clear distinction must be made between the cognitive requirements
for the retrieval of the temporal order of everyday experiences from
memory that are embedded in a rich fabric of contextual information,
as in the autobiographical memory studied here, and the relative
order judgments on the recency task in which stimuli are presented
in a long list with non-existent or impoverished contextual informa-
tion and there is no basis on which to infer or reconstruct order
information other than by making judgments of subtle differences in
the relative saliency ⁄ familiarity of stimuli.
As pointed out above, in autobiographical memory, events are
embedded in a variety of rich contextual cues, including the time of
day when something happened, and are linked by multiple other
associations that can be used to retrieve the order of events. In the
latter case, order is largely an inference based on a series of strong
contextually related experiences. For instance, the memory of riding
a bicycle in the park triggers the preceding memory of having
stopped at the grocery store to buy my favorite soft drink and the
following memory of stopping to make a phone call from the pay-
phone on my way back home from the park. Temporal order in these
types of serially related memories is unlikely to depend on the same
neural circuit as judgments of the relative order between stimuli
presented in a long, arbitrary and unstructured sequence, which has
been shown to depend on the prefrontal cortex. Temporal order in
autobiographical memory is likely to be another aspect of the
relations between stimuli coded by the hippocampal memory system.
European Journal of Neuroscience, 28, 829–840

The present results are consistent with this view. The patients with
temporal lobe excisions that exhibited impairments in autobiograph-
ical temporal order retrieval also had a basic deficit in autobio-
graphical event retrieval. By contrast, memory for the temporal
context of autobiographical events (which is different from serial
order in long lists of arbitrary unstructured stimuli) was not impaired
in patients with frontal cortical excisions. Although we have not
tested the present group of patients with frontal cortical excisions on
the recency test, we would expect them to be impaired because they
are the same type of frontal patient population tested in the classic
studies of recency by Milner and her colleagues (Milner, 1971;
Milner et al., 1991).
Interestingly, in comparison with normal control subjects and
patients with frontal cortical excisions, patients with left temporal lobe
excisions recalled fewer autobiographical events correctly and,
furthermore, were not helped by instructions to structure their recall
according to the temporal context of the experiences (Fig. 4). They
were also less accurate during recall in temporally ordering the events
that they had recorded in their diaries (Fig. 6) and in recalling the
specific details, including contextual information, about the diary
events (Fig. 7). These results are consistent with previous evidence
that these patients have a primary, generalized episodic (declarative)
memory impairment (Milner, 1971; Barr et al., 1990; Squire & Zola-
Morgan, 1991; Viskontas et al., 2000) and their present impairment on
autobiographical memory can be attributed to their generally poorer
memory trace for the events from the 2-day sampling period. Thus, the
evidence does not support the assumption that temporal context in
autobiographical memory is a distinct dimension that is dependent on
the prefrontal cortex. Instead, the present evidence suggests that
temporal contextual information in long-term memory is just one of
the many attributes of contextual ⁄ associative information that is
encoded by posterior cortical association areas in interaction with the
limbic medial temporal lobe region (Petrides, 2000; Addis et al.,
2004). This conclusion is also supported by previous studies in which
patients with left temporal lesions were impaired in their memory for
contextual aspects of events (Spiers et al., 2001; Steinvorth et al.,
2005).
The fact that further prompting (i.e. subjects were told how many
times they were paged and asked if they could recall any more
events) resulted in increased intrusion rates in patients with left
temporal lobe lesions, but not in those with frontal excisions (Fig. 4),
is of considerable interest. These intrusions would not have been
obvious without the diaries. The use of the event-sampling
methodology (Brewer, 1988), therefore, allowed us to show that
ordinary and plausible memory distortions do occur in the autobio-
graphical recall of patients with temporal lobe lesions. An earlier
study by Moscovitch & Melo (1997) did not find a significant
increase in such memory distortions in the autobiographical recall of
non-confabulating amnesic patients compared with normal control
subjects. The method used by those investigators to verify memories,
however, may not have been sensitive to higher rates of plausible
intrusions, such as those found among left temporal lobe patients in
the present study.
What exactly do we mean by plausible intrusions? The subject’s
memories for the events during the event-sampling procedure had
presumably been practiced or ‘highlighted’ by the fact that they had
recorded them in detail in the diaries. For the recall test, the subjects
were specifically asked to retrieve the events from the 2-day period
that they had recorded. Although the raters considered whether the
responses were consistent with the diary when they scored for the
accuracy of the subject’s event memories (e.g. a response that was
consistent with the detail of an event was scored as correct even if it
ª The Authors (2008). Journal Compilation ª Federation of European Neuroscience Societies and Blackwell Publishing Ltd
European Journal of Neuroscience, 28, 829–840
Autobiographical memory and the frontal cortex 839
was not the main activity listed), it nevertheless remains a possibility
that those items scored as ‘intrusions’ included true memories that
had not been recorded (or referred to) in the diaries. For example,
these ‘intrusions’ could have been the result of memories retrieved
from the wrong time frame, thus representing an error in the
retrieval of the temporal context of an event. Although one can
question whether ‘intrusion’ is an accurate description for this type
of recall error, the fact remains that only subjects with left temporal
lobe lesions showed elevations in this type of error compared with
control subjects. Furthermore, the analysis of the cued recall
responses, both in terms of the ability to retrieve detailed
information and the quality of their responses, supported the
interpretation that these subjects had weaker memory traces for
the diary events and were more likely to rely on generic semantic
autobiographical memories during recall than the other subject
groups. Specifically, patients with left temporal lobe excisions
appeared to resort to making educated guesses when retrieving
autobiographical information, as often as they actually remembered
an autobiographical event (Fig. 8). In contrast, the cued recall of
events by patients with frontal cortical excisions and normal control
subjects were significantly more likely to be accurate autobiograph-
ical episodic memories (i.e. the subjects retrieved unique information
that could not have been inferred) than inferences based on their
autobiographical knowledge.
In conclusion, the present study design permitted the uncovering of
new information about the neural basis of autobiographical memory.
The use of an experience-sampling method allowed us to verify
personally experienced events. Although some of the events may have
been mundane, everyday events (e.g. having breakfast), asking
subjects to reflect and write about specific details of the event
highlighted those aspects of the event that were personally relevant
and salient to the subject. This information was not only important in
verifying the accuracy of the subject’s memories but also allowed us to
explore the level of detail of their subsequent memories and to
distinguish between generic ‘knowledge-based’ retrieval about them-
selves and true episodic recall of the events. As we were able to
demonstrate, patients with temporal lobe lesions were in fact more
reliant on this generic autobiographical knowledge to fill in the ‘gaps’
of their memories for the events.
Finally, the results of this patient study provided evidence that basic
autobiographical memory is critically dependent on the medial
temporal lobe region, just as with other forms of declarative memory.
The findings also suggested that the lateral frontal cortex may aid in
the efficient retrieval of events through its initiation of strategic
processes but do not provide any support for claims that the frontal
cortex plays a necessary or critical role in autobiographical memory.
Furthermore, the widespread assumption that temporal order in long-
term episodic memory is a distinct dimension that relies on the lateral
prefrontal cortex did not receive support from the present data. On the
basis of these findings, we argue that temporal order in normal,
everyday, long-term declarative memory is just another type of
contextual information that is encoded and retrieved like other
contextual cues (e.g. spatial context). Temporal order information in
this sense relies on the limbic medial temporal lobe region like other
types of contextual information.
Supplementary material
The following supplementary material may be found on http://
www.blackwell-synergy.com
Appendix S1. Supplementary information. Method.
840 L. Thaiss and M. Petrides
Please note: Blackwell Publishing are not responsible for the content
or functionality of any supplementary materials supplied by the
authors. Any queries (other than missing material) should be directed
to the correspondence author for the article.
Acknowledgements
The present study was supported by grant 7466 to M.P. from the Natural
Sciences and Engineering Research Council of Canada and by a Canadian
Institute of Health Research Doctoral Research Award to L.T. We thank
Jennifer Novek for help with figure preparation.
Abbreviations
F, frontal cortical excisions; LT, left temporal excisions; MRI, magnetic
resonance image; N, normal control group; RT, right temporal excisions.
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