Developmental Science 10:2 (2007), pp 183– 189 DOI: 10.1111/j.1467-7687.2007.00548.

REPORT
Blackwell Publishing Ltd

Crawling is associated with more flexible memory retrieval

by 9-month-old infants
Jane Herbert,1 Julien Gross2 and Harlene Hayne2
1. Department of Psychology, University of Sheffield, UK
2. Department of Psychology, University of Otago, New Zealand

Abstract
In the present experiment, we used a deferred imitation paradigm to explore the effect of crawling on memory retrieval by
9-month-old human infants. Infants observed an experimenter demonstrate a single target action with a novel object and their
ability to reproduce that action was assessed after a 24-hr delay. Some infants were tested with the demonstration stimulus in
the demonstration context and some infants were tested with a different stimulus in a different context. Half of the infants in
each test condition were crawling at the time of participation and half were not. Both crawling and non-crawling infants exhibited
retention when tested with the demonstration stimulus in the demonstration context, but only infants who were crawling by
9 months of age exhibited retention when tested with a different stimulus in a different context. These findings demonstrate
that the onset of independent locomotion is associated with more flexible memory retrieval during the first year of life.

Introduction the mobile conjugate reinforcement paradigm (Hayne,

During the course of daily life, we typically encounter
stimuli that cue retrieval of prior memories. For most
normal adults, the range of effective retrieval cues is
remarkably broad. A scent, a word, or a picture can
activate highly detailed memories of experiences that
took place hours, days, or even years earlier. In some
instances, the cue that initiates retrieval may bear little
resemblance to the original experience. Furthermore, the
complex way in which our memories are organized
means that retrieving one memory may cue retrieval of
other, related memories as well. The flexible nature of
adult memory retrieval is a key ingredient in many
highly sophisticated cognitive skills, allowing us to
access and use our memories in circumstances that are
very different from those in which the memories were
originally established.
Although memory retrieval by human adults is highly
flexible, memory retrieval by human infants is not (for a
review, see Hayne, 2004). A large body of research has
shown that, for infants, memory retrieval occurs if and
only if the cues encountered at the time of retrieval
match the cues present at the time of encoding almost
exactly. The requisite relation between encoding and
retrieval during infancy was originally demonstrated in
Greco, Earley, Griesler & Rovee-Collier, 1986; Sullivan,
Rovee-Collier & Tynes, 1979). For example, Hayne et al.
trained 3-month-old infants to kick their feet to produce
movement in a five-object mobile. When infants were
tested 24 hrs later, those who were tested with the origi-
nal training mobile exhibited perfect retention, but those
who were tested with a mobile that contained more than
one novel object exhibited no retention at all. The highly
specific nature of infant memory retrieval has been
replicated under a number of different conditions. For
example, following training in the mobile conjugate
reinforcement paradigm, the absence of a distinctive
odor that was present during training has been shown to
impair memory retrieval, and substituting the training
odor with another distinctive odor eliminates memory
retrieval altogether (Rubin, Fagen & Carroll, 1998).
Further research has shown that the specificity of
infant memory retrieval originally documented in the
mobile conjugate reinforcement paradigm is not simply
a reflection of the operant training procedure that is
used to establish the memory in the first place. For
example, deferred imitation involves an observation-only
learning procedure whereby infants watch an experi-
menter perform a series of actions with objects and the
infants’ ability to reproduce those actions is assessed

Address for correspondence: Jane Herbert, Department of Psychology, University of Sheffield, Sheffield S10 2TN, United Kingdom; e-mail:
J.S.Herbert@sheffield.ac.uk

© 2007 The Authors. Journal compilation © 2007 Blackwell Publishing Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UK and
350 Main Street, Malden, MA 02148, USA.
184 Jane Herbert et al.
after a delay. Deferred imitation tasks are widely used in
studies of infant memory development and are considered
to yield a nonverbal measure of declarative memory
(McDonough, Mandler, McKee & Squire, 1995; for a
review, see Hayne, 2004). Memory performance by young
infants in the deferred imitation task, like memory per-
formance by young infants tested in operant paradigms,
is also disrupted by a change in the target stimulus at the
time of retrieval. In one study, Hayne, Boniface and Barr
(2000) found that 6-month-old infants who watched as
an experimenter performed a three-step sequence of
actions using a puppet failed to reproduce these actions
when they were tested 24 hours later if the color or the
form of the puppet was different from that encountered
during the original demonstration. Similarly, Herbert
and Hayne (2000) found that 18-month-old infants who
watched as an experimenter performed a three-step
sequence of actions to make a rattle or an animal failed
to reproduce those actions if the stimuli were changed
at the time of the test.
The highly specific nature of infant memory retrieval
also extends to cues associated with the environmental
context in which encoding originally occurred. In the
mobile conjugate reinforcement paradigm, for example,
changes to proximal contextual cues such as the color
and pattern of a crib liner that was present during
training (Borovsky & Rovee-Collier, 1990; Butler &
Rovee-Collier, 1989), or to more distal contextual cues
such as the room in which training occurs (Hayne,
Rovee-Collier & Borza, 1991) disrupts memory retrieval
by 3- to 6-month-olds. Similarly, in the deferred imitation
task, 6-month-old infants fail to retrieve a memory
acquired in one physical location (e.g. the laboratory)
when they are tested in a different location (e.g. their
own home) even when all other aspects of the target
objects remain the same (Hayne et al., 2000).
Over the course of human development, the high
degree of specificity that characterizes infant memory
retrieval gradually declines. In the deferred imitation
task, for example, 6-month-old infants fail to retrieve the
target memory when they are tested in a different con-
text, but 12-month-old infants exhibit the same level of
performance irrespective of the context at the time of the
test (Hayne et al., 2000). Furthermore, 12-month-olds
fail to retrieve the target memory if the characteristics
of the focal cue (e.g. the puppet) are changed, but when
faced with the same change in cue, 18-month-olds
exhibit excellent memory performance. A similar age-
related increase in the flexibility of memory retrieval is
observed in other infant memory tasks such as operant
conditioning paradigms (e.g. Hartshorn, Rovee-Collier,
Gerhardstein, Bhatt, Klein, Aaron, Wondolski & Wurtzel,
1998) and the visual paired comparison procedure (e.g.
© 2007 The Authors. Journal compilation © 2007 Blackwell Publishing Ltd.
Robinson & Pascalis, 2004), but the precise timetable for
change is unique to the task. Furthermore, within the
deferred imitation task in particular, making the memory
demands less difficult by allowing infants the opportu-
nity to practice the target actions prior to the test (Bauer
& Dow, 1994; Hayne, 2006; Hayne, Barr & Herbert, 2003)
or by increasing the similarity between the demonstration
and the test stimuli (Hayne, MacDonald & Barr, 1997)
contracts the timetable over which changes in flexibility
are typically observed (cf. Hayne et al., 2000; Herbert &
Hayne, 2000).
In addition to changes related to age, memory
retrieval is also facilitated by experiences provided by an
adult during the original learning episode. In Herbert
and Hayne (2000), for example, the experimenter pro-
vided a novel verbal label for the stimuli at the time of
the original demonstration. Infants were then re-presented
with the same verbal label when they were tested with
the novel stimulus. At 24 months of age, infants were
able to use this verbal information to facilitate memory
retrieval, but 18-month-olds were not. Subsequent
research has shown that although memory retrieval is
not facilitated by verbal information at 18 months of age
(Herbert & Hayne, 2000), it is facilitated by physical
experience with the stimuli during encoding (Hayne et al.,
2003). Similarly, flexible memory retrieval is enhanced in
3-month-olds by exposing them to different mobiles or
to different contexts during original encoding (Amabile
& Rovee-Collier, 1991; Fagen, Morrongiello, Rovee-
Collier & Gekoski, 1984; Greco, Hayne & Rovee-Collier,
1990; Rovee-Collier & DuFault, 1991).
Taken together, a number of studies have shown that
experiences provided by an adult during a learning
session can facilitate the flexibility of subsequent memory
retrieval by infant participants. In contrast, considerably
less is known about the effect of prior experience that
infants bring to the experimental setting. During the
course of their daily lives, infants undoubtedly accumul-
ate different experiences that might contribute to indi-
vidual differences in memory retrieval. One potential
source of individual differences in experience might be
the onset of independent locomotion. As infants begin
to crawl, their ability to explore the environment increases
dramatically. Many researchers have argued that this
increased opportunity for exploration may contribute to
advances in other areas of psychological development
(Adolph, 1997; Gibson, 1988). In fact, a growing body
of research has shown that infants’ social, emotional,
and cognitive skills do change once they learn to crawl
(for a review, see Campos, Anderson, Barbu-Roth,
Hubbard, Hertenstein & Witherington, 2000). With
respect to memory retrieval in particular, as infants
begin to crawl, they are faced with the new challenge of

Figure 1 The cow (left) and duck (right) stimuli that were used in the experiment.
transferring information acquired in one situation to
similar (but not identical) problems encountered in
another. In light of this new challenge, Rovee-Collier
(1996) has hypothesized that memory retrieval may
become more flexible as infants learn to crawl; in essence,
as infants begin to explore their environment on their
own, they rapidly learn a more general rule that some
changes are more (or less) important than others. The
present experiment was designed to test the hypothesis
that the onset of independent locomotion is associated
with more flexible memory retrieval. In order to provide
a particularly challenging test of the flexibility of memory
retrieval, we changed both the target stimulus and the
context at the time of the test.
Method
Participants
Ninety-six 9-month-old infants (48 male, 48 female)
who were born within 2 weeks of their due date were
recruited from public birth records and by word of
mouth. All infants were tested within 2 weeks of their
9-month birthdays. Infants were predominantly of Euro-
pean descent and came from a range of socioeconomic
backgrounds. Five additional infants were excluded
from the final sample due to scheduling difficulties,
maternal interference, failure to touch the experimental
stimulus, or crawling backwards as their primary means
of independent locomotion.
At the time of their participation, half of the infants
were crawling and half were not. The average age of the
infants in the crawling (M chronological age = 9.13
months, SD = .18; M gestational age = 78.99 weeks, SD
= 1.43) and noncrawling (M chronological age = 9.14,
SD = .13; M gestational age = 78.85 weeks, SD = 1.35)
© 2007 The Authors. Journal compilation © 2007 Blackwell Publishing Ltd.
Independent locomotion 185
groups did not differ. Infants’ crawling status was ini-
tially established by parental report and was confirmed
by experimenter observation at the time of the original
demonstration and again at the time of the test. Crawl-
ing was defined as the ability to continuously transverse
at least one meter using arms and/or knees. In no
instance was there a discrepancy between parents’
reports of their infant’s crawling status and what the
experimenter observed during the experimental session.
This high degree of reliability is consistent with recent
research by Bodnarchuk and Eaton (2004) showing that
parents provide reliable reports of their infants’ attain-
ment of motor milestones, including crawling.
Apparatus
The stimuli consisted of a black and white cow and an
orange duck that were constructed specifically for these
experiments and were not commercially available (see
Figure 1). Both stimuli were 60 cm in height and con-
sisted of a wooden head attached vertically to a wooden
base. A wooden button (6 cm in diameter) was recessed
into the center of the wooden base. The button was
either white or orange and matched the color of the cow
or duck respectively. The center of the button was
marked with a 1 cm in diameter circle made of black felt.
When the button was pressed, the cow stimulus ‘mooed’
and the duck stimulus ‘quacked’ for approximately 6 s.
A small LED in each eye also flashed for approximately
6 s. The stimuli (cow or duck) were counterbalanced
within and between groups.
Procedure
Infants were randomly assigned to one of two demon-
stration conditions (n = 64; 32 males, 32 females) or
to the non-demonstration control condition (n = 32;
186 Jane Herbert et al.
16 males, 16 females). All infants were tested at a time
of the day that the caregiver identified as an alert/play
period. During the initial visit, the purpose of the study
was explained to the caregiver, and informed consent
was obtained.
Demonstration condition
For half of the infants, the demonstration occurred in
the laboratory (n = 32; 16 males, 16 females) and for the
remaining half, the demonstration occurred in their own
home (n = 32; 16 males, 16 females). The female experi-
menter was positioned directly opposite the infant and
both were seated on the floor. At the beginning of the
demonstration and the test session, the experimenter
interacted with the infant for 5 minutes or until a smile
was elicited. If the infant failed to orient toward the
experimenter, the experimenter attracted his or her
attention by saying ‘look’ and using the infant’s name.
For infants in the demonstration condition, the experi-
menter demonstrated the novel action with one version
of the stimulus (cow or duck). The action was modeled
three times in succession out of reach of the infant. The
experimenter did not verbally describe the action or
label the stimulus. The entire demonstration session lasted
approximately 30 seconds.
Independent groups of crawling and non-crawling
infants were tested after a 24-hr delay (± 2 hours).
Within each crawling-status condition, infants were
randomly assigned to one of two test groups (8 males,
8 females per group). One group was tested with the
demonstration stimulus in the demonstration context
(No Change group). The other group was tested with a
different stimulus in a different context (Cue/Context
Change group). During the test, the female experimenter
was again positioned directly opposite the infant and
both were seated on the floor. The stimulus was placed
within the infant’s reach and his/her behavior was vide-
otaped for a 90-s period. The duration of the test period
was based on prior research on deferred imitation with
infants less than 1 year old (Barr, Dowden & Hayne,
1996; Hayne et al., 2000).
Non-demonstration control condition
To assess the spontaneous production of target actions
in the absence of adult demonstration, additional con-
trol groups of crawling and non-crawling infants were
exposed to the stimulus for the first time during the test.
For these 32 infants (16 males, 16 females), the test was
also scheduled within 2 weeks of their 9-month birthdays.
The test procedure was identical to that experienced by
infants in the demonstration condition. For half of the
© 2007 The Authors. Journal compilation © 2007 Blackwell Publishing Ltd.
infants, the test occurred in the laboratory ( n = 16;
8 males, 8 females) and for the remaining half, the test
occurred in their own home (n = 16; 8 males, 8 females).
Results
Each videotaped session was scored by two independent
observers, one of whom was unaware of the infant’s
experimental condition or crawling status. Each observer
scored the presence or absence of the target action
during a 90-s test period. A button press was coded as
present if it resulted in the stimulus producing an
audible sound. Infants were given credit only for the first
time that they produced the target action. Interobserver
reliability, calculated using percentage of agreement, was
100%.
Preliminary analyses indicated that there was no
difference in infants’ performance with the two sets of
stimuli; therefore data were averaged across both stimuli
for all subsequent analyses.
Figure 2 shows the number of infants who imitated
the target action as a function of crawling status and test
group (control, no change, cue/context change). Within
each crawling status, separate chi-square tests were con-
ducted across test group; significant group differences
were further analysed using Fisher’s Exact Test. Retention
was inferred when more infants in a given demonstration
condition performed the target action than did infants
in the non-demonstration control condition. As shown
Figure 2 The number of infants who performed the target
action as a function of crawling-status condition and
experimental group. An asterisk indicates that the performance
of a demonstration group is significantly above the performance
of the status-matched non-demonstration control group.

in Figure 2 (see asterisks), non-crawling infants only
exhibited retention when they were tested with the
same stimulus in the same context (X 2 [2] = 8.52, p < .05,
Fisher’s Exact Test, p < .05). Crawling infants, on the
other hand, exhibited retention when they were tested
with the same stimulus in the same context and when
they were tested with a different stimulus in a different
context (X 2 [2] = 14.40, p < .001, Fisher’s Exact Test,
p < .01).
Discussion
Taken together, these data provide the first empirical
support for Rovee-Collier’s (1996) hypothesis that
independent locomotion is associated with more flexible
memory retrieval. Although both crawling and non-
crawling infants exhibited excellent retention when tested
with the demonstration stimuli in the demonstration
context, only infants with crawling experience exhibited
retention when tested with a different stimulus in a
different context. The manipulation of both the cue and
the context at the time of the test provided a particularly
challenging memory retrieval problem, yet crawling
infants exhibited no difficulty at all. Whether crawling
infants would exhibit a similar retrieval advantage if
only the cue or the context (but not both) were changed
at the time of the test awaits further research. Similarly,
further research is also required to determine whether
crawling infants would exhibit a similar retrieval advantage
if the imitation task was even more difficult than the
one we used here (e.g. multiple target actions, a longer
delay).
Given that we have now shown that crawling is
associated with more flexible memory retrieval, the next
obvious question to answer is why? At this stage in our
research, we can rule out at least two simple explana-
tions for the relation between crawling and memory
retrieval. First, assuming that infants of the same chrono-
logical (and gestational) age are at the same level of
maturity, the differences observed in the present experi-
ment cannot be attributed to maturation alone because
the age of the infants was the same irrespective of their
crawling status. Second, although the beneficial effects
of active exploration within the context of an experiment
have been well established (Bensen & Uzgiris, 1985;
Feldman & Acredolo, 1979; for review, see Yan, Thomas
& Downing, 1998), this kind of proximal mechanism
cannot explain the effect of crawling reported here. In
the present experiment, and in others, infants were not
allowed to engage in exploratory behavior during the
task per se; rather, independent locomotion outside
the experimental context enhanced performance on the
© 2007 The Authors. Journal compilation © 2007 Blackwell Publishing Ltd.
Independent locomotion 187
target task. In the present experiment, for example,
infants were specifically prevented from crawling during
the demonstration and during the test.
The present data add to a growing body of research
showing that independent locomotion enhances at least
some aspects of cognitive development (Bertenthal, Campos
& Barrett, 1984; Clearfield, 2004; Gustafson, 1984; Ker-
moian & Campos, 1988; for a review, see Campos et al.,
2000). Across each of these cognitive domains, Campos
et al. (2000) have argued that crawling is neither a neces-
sary nor a sufficient condition to produce developmental
change; rather they propose that the onset of independent
locomotion yields a range of new experiences that help
push infants’ extant skills to a new level. Within the
domain of memory retrieval, independent locomotion
undoubtedly provides infants with new opportunities to
retrieve their memories in a range of different situations.
We hypothesize that these particular experiences facilitate
memory retrieval more generally. Partial support for
this hypothesis comes from experimental research in
which infants are given specific (albeit brief) experience
with novel test cues. Under these conditions, solving one
memory retrieval problem helps infants to solve addi-
tional, more difficult memory retrieval problems (Hayne,
2006). This research has shown that infants do not
require large amounts of additional experience to push
them to a new level of performance. As such, crawling
would provide more than enough additional experience
to push infants to a new level of memory flexibility.
In the final analysis, the answer to the question of
mechanism rests on the answers to at least two addi-
tional questions. First, how much crawling experience is
required for performance to improve on a particular
task? In the present experiment, as in others, crawling
was treated as a dichotomous variable – infants had
either achieved independent locomotion or they had
not. On the basis of maternal report, crawling experience
varied widely; some infants had been crawling for as
long as 4 months, others for only 1 week. We are currently
conducting a prospective study of the role of locomotor
experience in cognitive development in which caregivers
have kept detailed daily records of their infant’s locomotor
experience. This ongoing study will help us to answer
the question of how much crawling experience infants
require.
Finally, how long do these differences last? In other
words, at what point does the performance of the non-
crawling infants catch up with that of their crawling
counterparts? Furthermore, do the facilitative effects of
crawling extend to other forms of locomotor experience?
That is, do infants who walk early have a cognitive
advantage over infants who walk later when they are
tested with age-appropriate imitation tasks that are more
188 Jane Herbert et al.
difficult than those used here (cf. Herbert & Hayne,
2000)? If so, at what point do the late walkers catch up?
Again, our ongoing longitudinal research on the relation
between motor development and cognitive development
will help us to answer this question as well.
In conclusion, although our memories comprise a
record of the past, their most important function is to
guide behavior in the future. In order to achieve this
function, we must be able to retrieve our memories in
situations that are different from those in which they
were initially established. Young infants’ ability to
accomplish this task is extremely limited, but over the
course of development, older infants begin to exploit
retrieval cues that differ from those encountered at the
time of original encoding. Previous research has identi-
fied at least three variables that enhance the flexibility of
infant memory retrieval (Hayne et al., 2003; Herbert &
Hayne, 2000): the age of the infant, the complexity of
the task, and additional experience provided by an
adult. The results of the present experiment suggest that
crawling experience can now be added to this list.
Acknowledgements
This research was supported by a Marsden grant from
the Royal Society of New Zealand to Harlene Hayne.
The authors would like to thank Jackie Clearwater and
Bridget McDonald for help with data collection, Brendon
Sturgeon for constructing the stimuli, and the infants and
parents who participated in this project.
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Received: 4 July 2005
Accepted: 17 March 2006