CLINICAL EEG and NEUROSCIENCE
Memory Encoding and Retrieval in the Aging Brain
David Friedman, Doreen Nessler and Ray Johnson, Jr.
Key Words
Cognitive Aging of Memory
Encoding and Retrieval
Episodic Memory (EM) Effect
Event-Related Potentials
Left Inferior Prefrontal Negativity
Semantic Selection
ABSTRACT
Decline in episodic memory, the encoding and retrieval
of autobiographical events, is a hallmark of normal cognitive
aging. Although the primary causes of this decline remain
elusive, event-related brain potential (ERP) studies have
contributed to an understanding of age-related episodic
memory failure. These data reveal that, although the
retrieval-based episodic memory (EM) effect does not differ
dramatically between young and older adults, the acquisi-
tion-related data suggest a decline in episodic encoding (i.e.,
semantic elaboration) with increasing age. We conclude
that, at the current state of knowledge, encoding deficiencies
are more important than retrieval deficits in understanding
the causes of episodic memory decline in the older adult.
INTRODUCTION
The study of how humans encode, store and retrieve
mnemonic information has had a long scientific history,
beginning formally with the classical studies of Ebbinghaus
in the late 19th century.1 The relatively recent development
of techniques allowing the investigation of how these
processes are instantiated in the brain has provided an
unprecedented in vivo window on mnemonic function in the
actively performing human. Although much has been
learned about how (and by which brain networks) memo-
ries are encoded and retrieved in the normal young adult,
relatively less is known about how aging affects the func-
tioning of these different processes. Normal aging is
accompanied by well-documented failures in episodic
memory, although their root causes are unclear.2
Understanding the sources of encoding and/or retrieval dif-
ficulties is, therefore, critical to a more comprehensive
knowledge of episodic memory function in old age and,
eventually, for designing remedial programs aimed at ame-
liorating mnemonic deficiencies in the elderly.
Two brain imaging technologies have been used to chart
changes in episodic memory in normally aging older adults,
event-related brain potentials (ERPs) and event-related
functional magnetic resonance imaging (efMRI). Although
2
©2007 VOL. 38 NO.
the efMRI technique is highly accurate in localizing areas of
activation within the brain, the sluggishness of the hemody-
namic response makes it impossible to determine precisely
the temporal sequence and duration of brain region activa-
tions. By contrast, the ERP method enables the collection of
very precise temporal information, at the cost of less accu-
rate spatial resolution than efMRI. The ERP technique,
therefore, enables the viewing of brain activity related to the
formation and retrieval of memories in temporal units that
are consistent with the speed of cognitive processes, i.e.,
milliseconds. Hence, ERP studies of cognitive aging can
provide detailed information on age-related differences in
the timing and duration of specific mnemonic processes that
contribute to encoding and retrieval.
Episodic memory is comprised of personal memories
that include specific spatio-temporal information about the
context in which the event occurred.3 Episodic memory is
investigated most often using paradigms that assess
recognition memory. In these paradigms, participants are
tested by asking them to judge whether each item was
encountered previously during an encoding (i.e., study)
phase or is new. Recognition memory is generally thought
to rely on two processes, one relatively automatic and fast-
acting, labeled familiarity, and the other, more effortful,
slower and requiring conscious deliberation, labeled recol-
lection.4 For example, one can recognize with certainty a
person’s face as having been seen before, but not be able
to recall the contextual details that accompanied the initial
episode (i.e., where and when the person was initially
encountered). This is an instance of a recognition judg-
ment based on familiarity in the absence of recollection.
Given that the contextual details were encoded during the
original event and an efficient retrieval strategy, the miss-
ing features can often be recalled and reintegrated with the
information provided by the face (i.e., recollected), result-
ing in a reconstructed episodic memory.
Because the retrieval of episodes depends upon how
the information was initially encoded, we first review stud-
David Friedman, Professor and Research Scientist VII, Doreen Nessler,
Associate Research Scientist and Research Scientist II, Cognitive
Electrophysiology Laboratory, New York State Psychiatric Institute, New
York, New York, and Ray Johnson, Jr., Professor, Department of
Psychology, Queens College, CUNY, Flushing, New York, New York.
Address requests for reprints to D. Friedman, PhD, Cognitive
Electrophysiology Laboratory, NY Psychiatric Institute, Unit 6, 1051
Riverside Drive, New York, N.Y. 10032, USA.
Email: df12@columbia.edu
CLINICAL EEG and NEUROSCIENCE
ies of ERP activity recorded during encoding phases for
subsequent recognition testing. We then review the results
of ERP investigations of retrieval.
Encoding-related studies
Relative to non-semantic, or shallow encoding (for
example, processing orthographic features of words),
semantic or deep encoding (as when making living/non-liv-
ing judgments about words) leads to a greater likelihood
that those words will be recognized on a subsequent mem-
ory test.5 By now, a large number of studies have shown
that the degree to which an item’s memory trace is
enriched during encoding is determined by the amount one
elaborates upon the information retrieved from semantic
memory,6 a phenomenon that can be labeled episodic
encoding. Hence, semantic retrieval and episodic encoding
appear to work in tandem, perhaps with semantic retrieval
occurring prior to episodic encoding.
ERP investigators have assessed encoding-related
processing by quantifying the differences in neural activity
associated with individual items during an encoding phase
based on whether or not those items are subsequently
remembered. A subtraction of the ERPs elicited by study
items that are subsequently forgotten from those that are
subsequently remembered yields the subsequent memory
effect (SME), a measure of encoding.7,8 SMEs usually, but
not always, have a positive polarity that is widely distrib-
uted across the scalp. In their investigation, Paller and col-
leagues9 had subjects encode words under deep (e.g., is it
living?) and shallow (e.g., does the word contain 2 vow-
els?) conditions. In accord with Craik and Lockhart,5
semantic encoding engendered higher rates of recognition
success compared to non-semantic encoding. Importantly,
SMEs, which onset at roughly 400 ms and lasted about
600 ms, were reliable under semantic but not non-seman-
tic conditions. Hence, SMEs have been associated with
semantic encoding activity because their magnitude is pos-
itively related to the extent of deep encoding processes
during acquisition.9 The question arises, however, as to
whether SMEs are related to subsequent recollection-
compared to familiarity-based retrievals.
One method by which the distinction between recollec-
tion and familiarity has been demonstrated is with the
“Remember/Know” paradigm.10 In this paradigm, for items
judged old, subjects indicate whether their recognition deci-
sion was accompanied by the retrieval of contextual detail, a
“remember” (R) judgment (i.e., a recollection-based retriev-
al), or was based on a feeling of familiarity with the item, a
“know” (K) judgment (i.e., a familiarity-based retrieval).
Moreover, items that are deeply encoded are associated
subsequently with a greater proportion of R judgments,
whereas items that are shallowly encoded are associated
subsequently with a greater proportion of K responses.
The R/K paradigm was used by Friedman and Trott11 to
assess whether recollection- and familiarity-based
3
©2007 VOL. 38 NO. 1
retrievals had their origin during encoding. During the study
phase, participants were asked to memorize two nouns
embedded in sentences for a subsequent recognition test.
During the recognition test, subjects decided whether
nouns were new or old and, for old nouns, made an addi-
tional R/K judgment. The study phase ERPs were aver-
aged into three classes on the basis of subsequent recog-
nition performance: 1) subsequently correctly recognized
and given an R judgment, 2) subsequently correctly recog-
nized and given a K judgment, and 3) subsequently unrec-
ognized (i.e., a miss). SMEs were computed by subtracting
the ERPs associated with subsequently missed items from
those associated with subsequent R and K judgments. For
young adults, reliable SMEs were observed in the interval
from 400 - 900 ms for study items associated with subse-
quent R judgments but not subsequent K judgments.
These SME data suggest that recollection- and familiarity-
based recognition judgments at retrieval had their origin in
the type of encoding activity engaged in during study.
In addition to young adult participants, Friedman and
Trott11 recruited older adults. There was some basis for
expecting age-related differences in ERP activity because
the elderly had been shown to produce a smaller percentage
of remember judgments than the young,12 and to rely more
on familiarity than recollection when these two processes
are placed in opposition.13,14 Both findings suggest an age-
related imbalance in the recollective component of recogni-
tion memory. Hence, Friedman and Trott11 determined
whether aging interacted with the familiarity/recollection dis-
tinction in modulating encoding-related ERP activity. Com-
pared with the data of the young described above, for the
elderly, reliable SMEs were observed in association with
both subsequent R and K judgments. Friedman and Trott11
concluded that, during the study phase, older subjects did
not differentially encode those items that would be subse-
quently associated with retrieval of contextual details
(remember) from those that would not (know). Furthermore,
these data implicated age-related deficits in encoding as
contributing to episodic memory deficits in the elderly.
These conclusions received some support from the dis-
tribution of SME activity across the scalp. For the young, in
addition to positive electrical activity, the SME for R judg-
ments was associated with negative activity over left inferi-
or prefrontal scalp (LIPFS); there was no evidence of neg-
ative activity over LIPFS in the data of the elderly.13 The
negative activity could have reflected retrieval of informa-
tion from semantic memory, as R. Johnson and cowork-
ers15 observed negative-going ERP activity over LIPFS in
association with semantic but not orthographic decisions.
Such results fit well with those of hemodynamic studies
showing that the retrieval of semantic memories and the
encoding of episodic memories depend upon activity in
regions of the LIPF Cortex (LIPFC16). Assuming that the
blood flow and ERP data share at least some common
CLINICAL EEG and NEUROSCIENCE ©2007 VOL. 38 NO. 1

Figure 1
Left panel. Grand mean difference waveforms (High-selection minus Low-selection) recorded over left prefrontal scalp electrodes. The
data for the young (solid lines) and elderly (dashed lines) are superimposed. Early (light gray shading; 400-800 ms) and late (dark gray
shading; 1200-1400 ms) regions of the waveforms are indicated. Additionally, the black shading in the early interval indicates the region
where the ERPs of the young were more negative than those of the elderly. Time lines every 600 ms with arrows indicating stimulus
onset. Right panel. Surface potential, spline-interpolated scalp maps (based on the averaged reference computed on a 62-channel,
extended 10-20 montage41) are depicted for the early and late measured intervals computed on the young and elderly waveforms depict-
ed in the left panel. Dots represent the 62 scalp locations.

brain generators, together they suggest that the older
adults of the Friedman and Trott investigation11 may have
been impaired in semantic encoding, which could have
been responsible for their lower recognition success.
Supporting evidence for this conclusion has come from
behavioral investigations documenting that older adults, in
the absence of a specific encoding strategy, do not use
semantic elaboration spontaneously.17 Hence, because
volunteers in the Friedman and Trott11 investigation were
asked simply to memorize the nouns without any specific
encoding instructions, it is difficult to determine from those
data whether the putative encoding deficiency in the elder-
ly was due to a failure of semantic retrieval, a failure of
episodic encoding or both.
To investigate further the nature of encoding processes
in the elderly, Nessler and colleagues18,19 exerted greater
control over the use of encoding strategies with a seman-
tic selection paradigm adapted from Thompson-Schill and
coworkers.20 In our modified version of the object classifi-
cation task described by Thompson-Schill et al.,20 young
and older participants were asked to study words under
Low- and High-selection conditions. The Low-selection
task required a decision as to whether a previously pre-
sented picture (e.g., lion) matched the meaning of a to-be-
remembered word (e.g., lion). In the High-selection condi-
tion, by contrast, a decision was made concerning whether
a previously presented adjective (e.g., heavy) described a
feature of the to-be-remembered word (e.g., feather).
While both decisions necessitated the retrieval of the
semantic concept, the High-selection decision required, in
addition, the selection of specific semantic features from
several alternatives. This design enabled us to assess
age-related changes in ERP activity under conditions in
which young and old participants engaged in qualitatively
similar types of encoding activity (see Rugg and Morcom21
for other methodological issues in ERP research on aging).
Following the encoding phase, a recognition test on the
items studied under Low- and High-selection conditions
enabled us to determine whether the age-related decline in
recognition memory was due to failures in semantic
retrieval/selection and/or episodic encoding.
During the encoding phase, older adults performed as
well as the young adults, indicating that they were well able
to retrieve semantic information and select among compet-
ing alternatives in the High-selection condition. However,
during the subsequent recognition test, older adults per-
formed reliably more poorly than the young adults. Figure 1
depicts the encoding-related ERP activity elicited over
LIPFS in the young and elderly adults obtained by subtract-
ing the activity elicited in the Low-selection condition from
that elicited in the High-selection condition. These differ-
ence waveforms show for both young and old adults signif-
icant negative activity in the early period (400-800 ms) over
LIPFS. This selection-related effect is sustained in the
young adult data into the late time interval (1200-1400 ms),
but dramatically attenuated in the older adult waveforms.
Additionally, the young show greater negativity than the eld-
erly in the early interval. Hence, there appear to be three

4
CLINICAL EEG and NEUROSCIENCE
negativities: an early, selection-related activity present in
both young and old ERPs; an additional, early negativity
present only in the waveforms of the young; and a late,
selection-related negativity, again present only in the ERPs
of the young. Consistent with their equivalent behavioral
performance, the magnitude of the early, selection-related
negativity was the same in the young and elderly, suggest-
ing that this brain activity reflects semantic retrieval/selec-
tion. As the young outperformed the elderly during recogni-
tion testing, the additional early negativity and the late neg-
ativity presumably reflect episodic encoding, i.e., additional
semantic elaboration performed on the products of seman-
tic retrieval/selection. On this view, our data support the
conclusion that the episodic memory deficit in the elderly in
the current paradigm is a result of a failure to engage in
additional episodic encoding which would have resulted in
more richly encoded memory traces, thereby ensuring a
higher rate of successful retrieval. The topographic maps
illustrated in Figure 1 (right panel) demonstrate dramatical-
ly attenuated negative activity in the late interval over LIPFS
for the elderly. Hence, the hemodynamic data reviewed ear-
lier and these topographic data suggest that the age-relat-
ed encoding difficulties may have resulted from a failure to
activate LIPFC in the late interval.
The Friedman and Trott11 and Nessler et al.19 data
suggest that encoding deficiencies may underlie the
well-documented episodic memory decline in elderly
samples. We turn now to a consideration of ERP data
recorded at retrieval.
Retrieval-related studies
Recognition processes can be studied by separately
averaging the ERP activity elicited by old items that are cor-
rectly recognized (i.e., hits) and new items that are correct-
ly rejected. A subtraction of the latter from the former
reveals the episodic memory (EM) effect, an index of
retrieval.22 In young adults, a series of EM effects with dis-
tinct temporal and topographic patterns have been
observed fairly consistently in many investigations of recog-
nition memory, suggesting that they reflect unique stages of
mnemonic processing.22 For example, relatively early in the
retrieval process, between approximately 300 and 500 ms,
a medial frontal, positive-going, EM effect has been associ-
ated with the familiarity component of recognition memory.
This association with familiarity is based upon observations
that its amplitude is similar to 1) correctly recognized old
items regardless of whether they are endorsed with remem-
ber or know judgments,23 2) correctly recognized old items
regardless of whether the contextual details from the origi-
nal episode are correctly identified,24 and 3) correctly recog-
nized old items and falsely recognized items that are highly
similar to the previously studied old items, i.e., “lures.”25,26
On the other hand, later in the retrieval sequence, a subse-
quent left parietal, positive-going, EM effect (500-800 ms)
has been associated with recollection, as its amplitude is
5
©2007 VOL. 38 NO. 1
larger to 1) correctly recognized old items associated with
remember compared to know judgments,23 2) correctly rec-
ognized old items associated with correct compared to
incorrect source judgments,27 and 3) correctly recognized
old items compared to falsely recognized, highly similar lure
items.25 Consistent with the medial prefrontal and left pari-
etal EM effects reflecting distinct mnemonic processes,
recent data have indicated that they are associated with
reliably different scalp distributions, suggesting different
intracranial generators.24,28,29
Later-onsetting, longer duration EM effects have also
been reported. For example, an asymmetric, positive, right
prefrontal EM effect has been observed, although a con-
sensus regarding its functional significance has yet to be
arrived at. It onsets roughly with the diminution of the left
parietal EM effect and reaches peak amplitude typically fol-
lowing mean reaction time for the old/new recognition deci-
sion. The post-response timing of this component has led
some authors to suggest that it may reflect a cognitive con-
trol function, such as evaluating the products of retrieval to
support ongoing memory performance.30
Based on the finding that the elderly rely more on famil-
iarity than recollection, and assuming that the medial pre-
frontal EM effect reflects familiarity-based processing, one
expectation is that this EM effect would be of similar mag-
nitude in the ERPs of the young and elderly. Unfortunately,
few studies have directly assessed age-related changes in
the medial prefrontal EM effect. In the Remember/Know
study of Trott et al23 described above, the medial prefrontal
EM effect (300-500 ms) was of similar magnitude and
topography in the young and old adult waveforms in asso-
ciation with correctly recognized old items given either R or
K judgments (see also Wegesin et al31). Along with a simi-
lar percentage of old items attracting know judgments in
the young and elderly, these limited ERP data suggest an
intact familiarity mechanism in the elderly.
By contrast with the paucity of data on age-related
changes in the medial prefrontal EM effect, somewhat more
is known about the left parietal EM effect. Based on the
behavioral literature emphasizing age-related deficits in rec-
ollection and the ERP data reviewed earlier suggesting that
the left parietal EM effect indexes contextual retrieval, one
might expect smaller-magnitude left parietal EM effects in
the ERPs of the elderly compared to those of the young. To
the contrary, this has generally not proven to be the case.
Initial investigations by Friedman et al,32 Rugg et al33 and
Swick and Knight34 did not use behavioral indices presumed
to reflect familiarity and recollection. In later studies, howev-
er, measures considered proxies for recollection and famil-
iarity were collected and, hence, the results of these inves-
tigations provide a firmer basis for conclusions about aging
e ffects on the left parietal EM effect. Both Mark and Rugg35
and Trott et al23 solicited R and K as well as source judg-
ments from young and older adults. In both investigations,
CLINICAL EEG and NEUROSCIENCE
the left parietal EM effect in response to correctly recog-
nized old items attracting R judgments (or correct source
judgments) did not differ in magnitude or scalp topography
between young and older adults. Similar phenomena were
observed by Wegesin and coworkers.31 Despite this similar-
ity, in all of these investigations older adults showed greater
decrements in source compared to item memory relative to
young adults.36 Moreover, the temporal precision of the ERP
method enabled the investigators in all three studies to
determine that the EM effects were prolonged in older
adults by about 100 ms relative to those of the young. This
finding appears to be consistent with the cognitive slowing
that is typically observed in older adults.37 Although limited,
these data collectively suggest that, to the extent that the
left parietal EM effect reflects the retrieval of source-speci-
fying information, older adults, albeit with prolonged pro-
cessing time, do not appear to differ from young adults in
this aspect of memory retrieval.
The results of age-related investigations of the right-
prefrontal EM effect are not at all in agreement. This may
not be surprising given the somewhat imprecise definition
of “post-retrieval processing” that has been ascribed to this
EM effect. In two studies of recognition memory with
source judgments, Trott and colleagues23 and Wegesin et
al31 observed robust right prefrontal EM effects in their
young adult participants but failed to find similar activity in
the waveforms of their older adults. On the other hand,
Mark and Rugg35 and Li et al,38 also in source memory par-
adigms, did observe reliable right prefrontal EM effects in
their older adult waveforms. Given the elusive nature of the
behavioral correlates of this EM effect and the contradicto-
ry age-related findings, it is difficult to comment with any
confidence on the import of this EM effect in understanding
age-associated changes in episodic memory.
CONCLUSIONS
Clearly, there are too few age-related ERP studies of
episodic memory to permit firm conclusions. Nevertheless,
some tentative implications of the studies reviewed here
can be advanced. The available data suggest that encod-
ing deficiencies may play a larger role in influencing
episodic memory deficits in older adults. This interpretation
is consistent with the results of divided attention studies,
REFERENCES
1. Ebbinghaus H. Memory: a contribution to experimental psy-
chology. New York: Dover Publications; 1964.
2. Light LL. Memory and aging: four hypotheses in search of
data. Ann Rev Psychol 1991; 42: 333-376.
3. Tulving E. What is episodic memory? Curr Direct Psychol
Sci 1993; 2: 67-70.
6
©2007 VOL. 38 NO. 1
which demonstrate greater episodic memory deficits in
both young and elderly adults when attention is divided at
encoding, with a concomitant decrease in LIPFC activity,
compared to when it is divided at retrieval.39 The encoding-
related data of Friedman and Trott11 and Nessler and col-
leagues18,19 suggest that an underlying mechanism may be
a lack of spontaneous elaboration of the products retrieved
during semantic processing (i.e., episodic encoding) per-
haps resulting from a failure to recruit LIPFC. The limited,
retrieval-based ERP data suggest relatively intact familiar-
ity- and recollection-based processing in the elderly.
However, this does not account for the consistently poorer
source memory performance of older relative to younger
adults despite similar-magnitude, left parietal EM effects in
young and elderly samples. This disparity suggests that, in
addition to encoding deficiencies, other, as yet under- or
un-explored, retrieval mechanisms might contribute to this
episodic memory deficit. For example, age-related differ-
ences in retrieval orientation, the differential processing of
test items according to how the items were studied, have
recently been demonstrated.40 Further study of this type of
phenomenon and its impact on age-related episodic mem-
ory decline is clearly warranted. Additionally, future work
needs to address current gaps in knowledge by recording
ERP activity during both encoding and retrieval phases.
Although older adults may not spontaneously elaborate
upon the products of semantic retrieval, they do benefit
from increased processing demands when given an appro-
priate encoding strategy.18,19 Hence, future studies should
include this type of levels of processing manipulation.6
These kinds of investigations will undoubtedly bring us
closer to a better understanding of the root causes of age-
related decline in episodic memory.
ACKNOWLEDGMENTS
The writing of this manuscript was supported by NIA
Grants AG05213 and AG09988 and by the New York State
Department of Mental Hygiene. We are grateful to Charles
L. Brown III for computer programming and technical
assistance, Ms. Efrat Schori and Ms. Letecia Latif for their
assistance in the recruiting and screening of volunteers
and Mr. Michael Bersick for various aspects of data collec-
tion and analysis.
4. Yonelinas AP. The nature of recollection and familiarity: a
review of 30 years of research. J Mem Lang 2002; 46:
441-517.
5. Craik FIM, Lockhart S. Levels of processing: a framework
for memory research. J Verb Learn Verb Behav 1972; 11:
671-684.
CLINICAL EEG and NEUROSCIENCE
6. Craik FI. Levels of processing: past, present and future?
Memory 2002; 10: 305-318.
7. Paller KA, Wagner AD. Transforming experience into memo-
ry: observations of mind and brain. Trends Cogn Sci 2002; 6:
93-102.
8. Sanquist TF, Rohrbaugh JW, Syndulko K, Lindsley DB.
Electrocortical signs of levels of processing: perceptual
analysis and recognition memory. Psychophysiology 1980;
17: 568-576.
9. Paller KA, Kutas M, Mayes AR. Neural correlates of encod-
ing in an incidental learning paradigm. Electroencephalogr
Clin Neurophysiol 1987; 67: 360-371.
10. Tulving E. Memory and consciousness. Can Psychologist
1985; 26: 1-12.
11. Friedman D, Trott C. An event-related potential study of
encoding in young and older adults. Neuropsychologia 2000;
38: 542-557.
12. Parkin AJ, Walter BM. Recollective experience, normal
aging, and frontal dysfunction. Psychol Aging 1992; 7: 290-
298.
13. Friedman D. Age-associated changes in episodic memory:
event-related potential (ERP) investigations of recollection
and familiarity. In: Zimmer HD, Mecklinger A, Lindenberger U,
(eds). Binding in Human Memory: A Neurocognitive
Approach. New York: Oxford. In press.
14. Jennings JM, Jacoby LL. An opposition procedure for detect-
ing age-related deficits in recollection: telling effects of repe-
tition. Psychol Aging 1997; 12: 352-61.
15. Johnson R Jr, Barnhardt J, Grossman S, Adler N, Schindler
D. Levels of processing effects on memory encoding and
retrieval: an ERP mapping study. Psychophysiology 2001;
38: S53.
16. Wagner AD, Schacter DL, Koutstaal MRW, Maril A, Dale AM,
Rosen BR, et al. Building memories: remembering and for-
getting of verbal experiences as predicted by brain activity.
Science 1998; 281: 1188-1191.
17. Hashtroudi S, Parker ES, Luis JD, Reisen CA. Generation
and elaboration in older adults. Exper Aging Res 1989; 15:
73-78.
18. Nessler D, Johnson R, Bersick M, Friedman D. On why the
elderly have normal semantic retrieval but deficient episodic
encoding: a study of left inferior frontal ERP activity.
Neuroimage 2006; 30: 299-312.
19. Nessler DR, Johnson J, Bersick M, Friedman D. Age-related
episodic encoding deficits are associated with early attenua-
tion of a left inferior prefrontal negativity. J Cogn Neurosc
2005; 17: 136.
20. Thompson-Schill SL, D’Esposito M, Aguirre GK, Farah MJ.
Role of left inferior prefrontal cortex in retrieval of semantic
knowledge: a reevaluation. Proc Natl Acad Sci USA 1997; 94:
14792-14797.
21. Rugg MD, Morcom AM. The relationship between brain activ-
ity, cognitive performance and aging: the case of memory. In:
Cabeza R, Nyberg L, Park D, (eds). Cognitive Neuroscience
of Aging: Linking Cognitive and Cerebral Aging. New York:
Oxford University Press; 2005: 132-154.
22. Friedman D, Johnson R. Event-related potential (ERP) stud-
ies of memory encoding and retrieval: a selective review.
Microsc Res Tech 2000; 51: 6-28.
©2007 VOL. 38 NO. 1
23. Trott CT, Friedman D, Ritter W, Fabiani M, Snodgrass JG.
Episodic priming and memory for temporal source: event-
related potentials reveal age-related differences in prefrontal
functioning. Psychol Aging 1999; 14: 390-413.
24. Friedman D. ERP studies of recognition memory: differential
effects of familiarity, recollection and episodic priming. Cogn
Sci 2004; 1: 81-121
25. Curran T. Brain potentials of recollection and familiarity. Mem
Cogn 2000; 28: 923-938.
26. Nessler D, Mecklinger A, Penney TB. Event related brain
potentials and illusory memories: the effects of differential
encoding. Cogn Brain Res 2001; 10: 283-301.
27. Wilding EL, Rugg MD. An event-related potential study of
recognition memory with and without retrieval of source.
Brain 1996; 119: 889-905.
28. Johnson R Jr, Kreiter K, Russo B, Zhu J. A spatio-temporal
analysis of recognition-related event-related brain potentials.
Intl J Psychophysiol 1998; 29: 83-104.
29. Rugg MD, Yonelinas AP. Human recognition memory: a cog-
nitive neuroscience perspective. Trends Cogn Sci 2003; 7:
313-319.
30. Wilding EL, Sharpe H. Episodic memory encoding and
retrieval: recent insights from event-related potentials. In: Zani
A, Proverbio AM, (eds). The Cognitive Electrophysiology of
Mind and Brain. New York: Academic Press; 2002: 169-196.
31. Wegesin DJ, Friedman D, Varughese N, Stern Y. Age-related
changes in source memory retrieval: an ERP replication and
extension. Cogn Brain Res 2002; 13: 323-338.
32. Friedman D, Berman S, Hamberger M. Recognition memory
and ERPs: age-related changes in young, middle-aged and
elderly adults. J Psychophysiol 1993; 7: 181-201.
33. Rugg MD, Mark RE, Gilchrist J, Roberts RC. ERP repetition
effects in indirect and direct tasks: effects of age and
interitem lag. Psychophysiology 1997; 34: 572-586.
34. Swick D, Knight RT. Event-related potentials differentiate the
effects of aging on word and nonword repetition in explicit
and implicit memory tasks. J Exp Psychol Learn Mem Cogn
1997; 23: 123-142.
35. Mark RE, Rugg MD. Age effects on brain activity associated
with episodic memory retrieval: an electrophysiological study.
Brain 1998; 121: 861-873.
36. Spencer WD, Raz N. Differential effects of aging on memory
for content and context: a meta-analysis. Psychol Aging
1995; 10: 527-539.
37. Salthouse TA. The processing-speed theory of adult age dif-
ferences in cognition. Psychol Rev 1996; 103: 403-428.
38. Li J, Morcom AM, Rugg MD. The effects of age on the neural
correlates of successful episodic retrieval: an ERP study.
Cogn Affect Behav Neurosci 2004; 4: 279-293.
39. Anderson ND, Iidaka T, Cabeza R, Kapur S, McIntosh AR,
Craik FI. The effects of divided attention on encoding- and
retrieval-related brain activity: a PET study of younger and
older adults. J Cogn Neurosci 2000; 12: 775-792.
40. Morcom AM, Rugg MD. Effects of age on retrieval cue pro-
cessing as revealed by ERPs. Neuropsychologia 2004; 42:
1525-1542.
41. Sharbrough F, Chatrian GE, Lsser RP, Luders H, Nuwer M,
Picton TW. Guidelines for Standard Electrode Position
Nomenclature Bloomfield: American EEG Society; 1990.