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Anim Cogn

DOI 10.1007/s10071-012-0538-2

ORIGINAL PAPER

Habituation and dishabituation during object play


in kennel-housed dogs
Anne J. Pullen • Ralph J. N. Merrill •

John W. S. Bradshaw

Received: 19 September 2011 / Revised: 18 June 2012 / Accepted: 10 July 2012


Ó Springer-Verlag 2012

Abstract Domestic dogs are reported to show intense Keywords Domestic dog  Habituation  Dishabituation 
but transient neophilia towards novel objects. Here, we Object play
examine habituation and dishabituation to manipulable
objects by kennel-housed dogs. Labrador retrievers (N =
16) were repeatedly presented with one toy for successive Introduction
30-s periods until interaction ceased. At this point (habitu-
ation), a different toy was presented that contrasted with the The domestic dog is renowned for routinely engaging in
first in both colour and odour (since the dog’s saliva would play even when adult, and object-orientated social play is
have accumulated on the first), colour alone, or odour alone. commonly used by owners to interact with dogs and enrich
No effect of the type of contrast was detected in the number their environment (Rooney et al. 2000). Provision of
of presentations to habituation, the difference in duration of manipulable objects can be beneficial for the welfare of
interaction between the first presentation of the first toy and individually housed dogs when preferred play objects
the presentation of the second toy (recovery), or the dura- (henceforth ‘toys’) are offered (Wells 2004b), but the
tion of interaction with the second toy (dishabituation). extent to which dogs interact with such toys appears to vary
Varying the time interval between successive presentations considerably. In laboratory kennels, beagles have been
of the first toy up to habituation between 10 s and 10 min observed to display more long-lasting interest in toys,
had no effect on the number of presentations to habituation, interacting with them for about a quarter of the observed
nor did it alter the extent of dishabituation. Varying the time (Hubrecht 1993), than did dogs in rehoming kennels,
delay from habituation to presentation of the second toy, which interacted for less than a tenth of the observed time
between 10 s and 15 min, affected neither the recovery nor (Wells 2004a). Dogs display powerful neophilia towards
the dishabituation. Overall, the study indicates that loss of novel toys (Kaulfuß and Mills 2008), and anecdotally, they
interest in the object during object-orientated play in this can also rapidly lose interest in particular toys. The
species is due to habituation to the overall stimulus prop- behavioural mechanisms that may lead to the latter, which
erties of the toy rather than to any single sensory modality are likely to include habituation, do not appear to have
and is also atypical in its insensitivity to the interval been investigated systematically.
between presentations. Habituation, ‘the response decrement as a result of
repeated stimulation’ (Harris 1943), is likely to impact on
the effectiveness of any inanimate enrichment (Tarou and
Bashaw 2007). For example, successive presentations of
A. J. Pullen  J. W. S. Bradshaw (&)
Anthrozoology Institute, School of Clinical Veterinary Sciences, enrichment objects to chimpanzees rapidly led to habitua-
University of Bristol, Langford BS40 5DU, UK tion towards those objects over a 3-day period (Celli et al.
e-mail: J.W.S.Bradshaw@bristol.ac.uk 2003). Habituation has even been observed when manipu-
lable and play-inducing objects were offered, affecting their
R. J. N. Merrill
WALTHAMÒ Centre for Pet Nutrition, Waltham-on-the-Wolds, usefulness as long-term enrichments (Line et al. 1991; Maki
Melton Mowbray, Leicestershire LE14 4RT, UK and Bloomsmith 1989 cited in Tarou and Bashaw 2007).

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Dishabituation is defined as any resumption of the behav- indoor pen (3.05 m2) with a raised, bed area (facing a
ioural response, which may be less than or equal to that seen central area and other dog pens) and an outdoor covered
prior to habituation (Holmes 1912 cited in Thompson area (4.15 m2); additional access to an outdoor paddock
2009), that occurs when the sensory characteristics of an was available during the day with access to large dog toys
object presented to the animal contrast sufficiently with in the paddocks such as Aussie hanging balls (Aussie dog
those of the object to which habituation has built up. Such products, AU), as well as platforms and staging to climb
dishabituation could potentially be used to restore interest in on. The indoor pen area could be sectioned off using a
object play and is therefore one of the key factors that could lockable dog flap. Overnight, pens were provided with
affect the value of enrichment using objects. fleece bedding and nylon chews, considered safe for
Object-orientated play in domestic cats is motivated as unsupervised interaction. In addition to exercise in pad-
if it were a type of predatory behaviour (Hall and Bradshaw docks with other dogs throughout the day, each dog
1998), with habituation rapidly inhibiting exploration of received at least one half hour of structured interaction with
the object unless its sensory characteristics change, as a person per day: this could consist of either a walk with off
would happen during successful predatory behaviour. In lead interaction in an enclosed field, interactive play in
this species, the dishabituation when the sensory charac- paddocks with kennel staff, or obedience training. The
teristics of the toy are changed can exceed that seen with kennel staff also spent time interacting individually with
the first presentation of the first toy (Hall et al. 2002), dogs in their pens or paddocks on an ad hoc basis.
termed post-inhibitory rebound (Kennedy 1985; Roper Destructible toys were only made available when the dog
1984). Since the domestic dog also evolved from a pred- was interacting with kennel staff due to the risk of
atory species (Coppinger and Schneider 1995), it is possi- destruction and ingestion during unsupervised interaction.
ble this species has a similar underlying motivation for The dogs could maintain visual contact with kennel staff
object-orientated play to that observed in cats (Hall et al. throughout the day. All the dogs were clicker trained for
2002). Social play in dogs is considered to be a means of basic obedience commands such as ‘sit’ and ‘wait’ on a
learning appropriate social interaction (Feddersen-Petersen daily basis. This continued throughout the study period.
2008). However, solitary play is less common, and
its construction and motivation are poorly documented Study area
(Feddersen-Petersen 2008; Horwitz et al. 2002).
From an animal welfare perspective, establishing the The experiments were undertaken in a room in one of the
motivation behind habituation to objects, and the nature of dog housing areas (approximately 4.2 m 9 4.0 m). The
any subsequent recovery in response that restores object- dogs were acclimatised to the room prior to the studies as
orientated play, should help to predict the effectiveness of part of their daily walks.
inanimate enrichments, the time interval needed between
successive presentations of any enrichment for effective or Study subjects
prolonged enrichment, and which cues lead to both habit-
uation and dishabituation. Sixteen adult (1–8 years) Labrador retrievers (LR) (9 neu-
The study described here aimed to determine the fol- tered males, 4 neutered females, 3 intact females) from 9
lowing: the time taken by dogs to habituate to a manipu- litters were randomly chosen from the dogs at the centre. The
lable, interactive, ‘play’ object over repeated presentations dogs were normally used for non-invasive feeding trials. All
(experiments 1 and 2); the extent of dishabituation when dogs were used for experiment 1. Seven of these dogs, all of
visual and/or olfactory characteristics of the play object which had interacted with and subsequently habituated to the
were changed (experiment 1); and the effect on dishabit- toy, were studied in experiment 2. Fifteen of the dogs were
uation of changing the interval between habituation and the used in experiment 3: one dog that had not shown any interest
presentation of the contrasting play object (experiment 3). in the toys in experiment 1 was not tested.

General methodology Experiment 1: The effect of visual and olfactory cues


on habituation
Husbandry
Procedure
The study was carried out at the WALTHAMÒ Centre for
Pet Nutrition, Leicestershire. The dogs were housed in Experiments were carried out on days 1, 3, and 5 of a five-
pairs, in pens arranged around an octagonal central court day period, with a distraction day on days 2 and 4 to reduce
(Loveridge 1998). The dogs had constant access to an the expectation of subsequent experimental days and to

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minimise any carryover of habituation due to previous between the last presentation of toy 1 (P1F) and the only
treatments. All dogs were presented with experiment 1a on presentation of toy 2 (P2), to the absence of the dogs’ own
day 1, half the dogs were given experiment 1b on day 3 and olfactory cues that were assumed to have accumulated on
then 1c on day 5, and the other half were given 1c on day 3 the toy through saliva, etc., during the repeated presenta-
and then 1b on day 5. tions of toy 1.

Experiment 1a: Visual and olfactory cues Experiment 1c: Visual cues

Two manipulable objects were used for the experiment on Experiment 1a was repeated, but the dogs were presented
day one: identical soft zoomorphic toys (20 cm 9 15 cm 9 with a different toy of the same colour every time toy 1 was
8 cm; Chubleez ‘Plush Squeaky Sniffer Dog Toy’, UK) presented, followed by a clean toy of the opposite colour
with an internal squeaker that sounded when mouthed by once habituation had been reached (P2).
the dog, varying only in colour: one was blue and one
brown/yellow. The dogs were taken to the test room indi- Distraction days The dog was taken to the test room and
vidually and given 2 min to settle. The experimenter, an given 2 min to settle. The experimenter entered the test room,
unfamiliar female, 26 years, then placed the first toy on the but no toy was given. After 30 s, the experimenter entered the
floor, 0.5 m from the door (Presentation 1.1, hereafter room in the same way as she had done when removing the toy
abbreviated P1.1) and left the room. After 30 s, she and then exited the room. This was repeated a further 10 times
removed the toy from the test room, ending P1.1. Ten to reduce the anticipation of a toy being presented every time
seconds later, the same toy was presented again in exactly the experimenter entered the room.
the same way as before (P1.2). P1.2 was then repeated until
the dog no longer interacted with the toy for more than the Data recording Interactions with the toys were recorded
first 10 s of the presentation: this was taken to indicate the using remote video cameras mounted above the test room.
point at which the dog had habituated to the first toy and An interaction was defined as anything other than sniffing
was designated the final presentation of toy 1, P1F. This or accidental contact, so included contact with the mouth or
toy was then removed; after a 10-s delay, the second toy paw, such as mouthing, chewing, and pawing at the toy.
was placed on the floor (P2). After 30 s, this toy was Intra-(AJP) and inter-observer reliability (AJP, JWSB) had
removed and the experiment ended. If any dog showed no been established for this measure in the previous studies
interest in toy 1 after the first three presentations, or the dog using dogs at the same establishment (Pullen 2011; Pullen
continued to interact with toy 1 after 10 presentations and et al. 2010, 2012).
therefore had failed to habituate, the experiment was ter- The video recordings were analysed using Observer 5.0
minated at that point. (Noldus, Wageningen) to determine total duration of
The experiment was performed on all 16 dogs on day 1, interaction with the toy during each presentation (seconds)
using fresh toys for each dog. Eight of the dogs were and the number of presentations to habituation.
presented with a blue toy first and eight dogs with a brown/
yellow toy to balance any effect of preference for toy Statistical analysis Data were analysed using SPSS 14.0
colour. The toys were washed prior to the start of the (SPSS Inc, Chicago). None of the data were normally
experiment (and between all subsequent test days) using distributed, so hypotheses were examined using nonpara-
the washing powder used to wash the dogs’ bedding. Any metric tests. The dog that failed to habituate was excluded
handling of the toys was carried out using rubber gloves to from further analysis.
minimise any preference due to deposited human scent. Proportions of dogs reaching criterion for habituation
The toys and test room were cleaned between every dog were compared between experiments by a Cochran Q-test.
studied to avoid any effects of scent from the preceding A Friedman test was applied to compare the effect of
dogs. treatments 1a, 1b, and 1c on the number of presentations of
The dogs were then randomly allocated to one of two toy 1 to habituation. Wilcoxon tests were then used to look
groups, so that half received experiment 1b on day 3 and at pairwise comparisons between the three treatments.
experiment 1c on day 5 while the other dogs received Recovery (defined as the difference in duration of inter-
experiment 1c on day 3 and experiment 1b on day 5. action between P1.1 and P2) and dishabituation (difference
in duration of interaction between P1F and P2) were
Experiment 1b: Olfactory cues compared between the three treatment groups (1a, 1b, and
1c) using Kruskal–Wallis tests. The magnitude of the
Experiment 1a was repeated, except that both toy 1 and toy recovery was tested against a null hypothesis of zero with
2 were the same colour. This restricted the contrast, Wilcoxon tests.

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Table 1 Habituation during the three trials in experiment 1 for all 16


dogs
Experiment
1a 1b 1c

Habituated 11 10 8
No habituation 4 5 7
No interest 1 1 1
‘Habituated’ dogs stopped interacting with the toy within the first 10 s
of a presentation between the second and the tenth presentation of toy
1. ‘No habituation’ dogs did not stop interacting within the first 10 s
of any of the 10 presentations. ‘No interest’ dogs did not interact with
the toy for more than 10 s during any of the first three presentations

Results

The number of dogs habituating within 10 presentations


was similar for all three treatments (69, 63, and 50 %:
Cochran Q = 0.5, P [ 0.50). One dog showed no interest
in the toys (Table 1).
For the dogs that habituated in one or more treatments
(N = 11), the median number of presentations to habitation
was 5.0 when colour and olfactory cues were altered together
(1a), and 4.0 when they were altered independently (1b and
1c). No overall difference was apparent between treatments
(Friedman chi-squared = 2.31, P \ 0.32). All pairs of
treatments were then compared to assess whether some were
more similar than others (noting that treatments 1a and 1b
were identical up to the point of habituation), but the distri-
bution of P values appeared to be random (experiment 1a vs.
1b, Wilcoxon T = 43, P \ 0.21; experiment 1a vs. 1c,
T = 36, P \ 0.42; experiment 1b vs. 1c, T = 48, P \ 0.10).
The degree of dishabituation (difference in duration of
interaction between P1F and P2) was similar across the
three treatment groups (Kruskal–Wallis = 0.81, P \ 0.67)
(Fig. 1). Recoveries, the differences between durations of
interaction during the first presentations of the two toys
(P1.1 and P2), were also similar for all three treatments
(Kruskal–Wallis = 0.67, P \ 0.72). Treatments could
therefore be combined by averaging within dogs. The
average dishabituation was highly significant (Wilcoxon
T = 65, P \ 0.001). The median of the average recovery
was 4.0 s, which is not significantly different from zero
(Wilcoxon T = 40, P \ 0.29), indicating that play returned
to approximately its original intensity after the toy was
changed.
Fig. 1 Boxplots of duration of interaction (s) with toys at presenta-
tion 1.1 (first presentation of toy 1), presentation 1F (final presen-
Discussion tation of toy 1 at the point of habituation), and presentation 2
(presentation of toy 2, following habituation to toy 1), by the dogs that
All three parts of this experiment confirm that dogs habituated within 10 presentations (N = 11, 10, and 8 for experiments
1a, 1b, and 1c, respectively) for visual and olfactory contrast between
habituate rapidly to manipulable objects and that changes
1F and 2 (a), visual contrast only (b), and olfactory contrast only (c).
in the sensory characteristics of the objects lead to imme- Heavy horizontal lines indicate medians, boxes indicate 25th and 75th
diate resumption of play. The extent of the dishabituation percentiles, and error bars indicate minima and maxima

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shows that the dogs’ motivation for object play had not Furthermore, it emphasises that consideration should be
diminished to any great extent over any of the treatments. given to interpretations based upon contrasts in odour in
Changes in colour and (presumed) odour cues appeared to experiments that involve objects that could have been
be equally effective in causing dishabituation; this suggests contaminated by odours that are detectable by dogs but not
that the alteration in any cue, whether visual or olfactory, by humans. Such changes in odour could result from
may be sufficient to induce dishabituation. Habituation, contact with either dogs or humans. For example, in the
therefore, appears to occur in parallel to each of the stim- movie clip presented by Kaminski et al. (2004), the subject
ulus properties of the toy, rather than any particular sensory dog repeatedly sniffs and mouths the novel object in the set
modality taking precedence. A remote possibility that has before being given the novel cue word to retrieve it. He
to be considered is that habituation was primarily driven by also sniffs the familiar objects in the set; presumably, these
cues other than colour or odour, although it is unclear what emit odours derived from himself and his owner. His
these could be, and this would not explain why dishabitu- behaviour suggests that the unfamiliar odour of the new
ation is almost complete when only colour and/or odour is object makes it particularly salient and therefore likely to
changed. be retrieved spontaneously, unless overridden by a com-
Thompson (2009) suggests that whenever regular pre- mand previously associated with a specific object.
sentation of a stimulus occurs, habituation will become
evident. All mammals able to exhibit habituation to a
stimulus appear capable of showing dishabituation (Harris Experiment 2: The effect of time interval
1943 cited Thompson 2009). By viewing dishabituation as between presentations on habituation
the ‘neutralisation’ of the habituation, a distinction can
then be made between that and ‘fatigue’ or loss of moti- Procedure
vation (Humphrey 1933 cited in Thompson 2009). In rats,
habituation was found to occur when presented with either The same two types of toy used in experiment 1 were used
auditory or tactile cues. However, generalised habituation for experiment 2. The method of presentation used in
was also apparent but not clearly distinguishable between experiment 1a (the same toy for Presentation 1 followed by
increased habituation and decreased sensitisation to the the other colour toy for P2) was used since it had caused
stimulus (Vogel and Wagner 2005). the greatest number of dogs to habituate in experiment 1
Over time, habituation can become more rapid through and was therefore considered the most effective combina-
training and recovery (Thompson 2009) or simply through tion to lead to habituation in future experiments.
repeated presentation of unchanging objects (Tarou and The dog was taken to the test room and given 2 min to
Bashaw 2007). This was not evident in the experiment settle. It was then presented with the first toy (P1.1). After
reported here, possibly because each dog was only sub- 30 s, the toy was removed from the room. After a fixed
jected to 3 experimental periods in experiment one, and a time interval that varied between treatments, the same toy
combination of distraction days, delays between experi- was presented again (P1.2). This continued until the dog no
ments (of a number of weeks), and randomisation of trial longer interacted with the toy for more than the first 10 s of
orders within experiments were all employed to negate the presentation (P1F) (up to 10 presentations). At this
these potential effects. point, the toy was removed. Following a 10-s interval, the
The almost complete dishabituation that occurred when second toy (P2) was presented. After 30 s, the toy was
the toy presented was visually identical to the toy to which removed, signalling the end of the experiment.
the dog had become habituated was unexpected. It is highly Each dog was studied on odd-numbered days over a
unlikely that any small differences in visual appearance 9-day period. The treatments consisted of five different
between the two toys provided the salient contrast, since delay intervals (either 10, 30 s, 1, 3, or 10 min) arranged in
short-range vision is poorer in dogs than it is in humans a balanced order for each dog using a Latin square design.
(Miller and Murphy 1995). While minor textural differ- On each day, the same delay was inserted between all
ences cannot be ruled out, the most likely explanation for presentations, until habituation occurred. Distraction, as for
the dishabituation is the contrast in odour that arose from a experiment 1, was performed on the days between each
clean toy being substituted for a toy that had been repeat- study day.
edly chewed by the dog and therefore would have smelled Comparisons between the number of presentations to
of its own saliva. This highlights both the dog’s highly habituation were made between the five treatments (time
sensitive olfaction (Walker et al. 2006) and the salience intervals) using a Friedman test. Recovery and dishabitu-
which dogs attach to what might seem to primates such as ation were compared between the five time intervals using
humans to be trivial differences in odour. Kruskal–Wallis tests.

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may apply to object play in another member of the Car-


nivora, the domestic cat Felis silvestris catus (Hall et al.
2002). Extending the time interval between presentations to
15 min or longer would presumably eventually increase the
number of exposures required to reach the habituation
criterion. However, if the time period between each pre-
sentation was very long, the dogs might not associate the
presentations together as belonging to the same play ses-
sion, therefore potentially introducing a different motiva-
tional basis for any change in response compared to the
shorter time periods. This might be possible to investigate
further using distractions (e.g. feeding or social play)
during the intervals between presentations of the toys.

Fig. 2 Boxplot of duration of interaction (s) with toys at presentation Experiment 3: The effect of delay on dishabituation
1.1 (first presentation of toy 1), presentation 1F (final presentation of
toy 1 at the point of habituation), and presentation 2 (presentation of
toy 2, following habituation to toy 1) (means for all 6 time intervals Procedure
combined) by the seven dogs that habituated within 10 presentations
in experiment 2 The procedure was identical to experiment 2, except that
the interval between each presentation of toy 1 (P1) was
Results fixed at 10 s, and the treatments consisted of varying
interval between the final presentation of toy 1 (P1F) and
The median number of presentations of toy 1 to habituation the presentation of toy 2 (P2) (10, 30 s, 1, 5, 10, or
was 4.0 for the 10, 30 s, 3, and 10 min between-presenta- 15 min).
tion intervals, and 3.0 for the 1-min interval: overall, no Each dog was studied on odd-numbered days over an
effect of interval was apparent (Friedman chi- 11-day period, with treatments arranged in a balanced order
squared = 2.48, P \ 0.65). The average durations of for each dog using a Latin square design. Each study day
interaction for habituation and dishabituation (Fig. 2) were was followed by a distraction day, as per experiment 1.
very similar to those in experiments 1a–c (Fig. 1). Recovery and dishabituation were also compared
between the six time intervals of the experiment 3 data
Discussion using Kruskal–Wallis tests.

Habituation over short time intervals appeared to be Results


insensitive to the time between presentations. From the
consistent median of four presentations to habituation for The median number of presentations of toy 1 to habituation
all time intervals of 10 min or less, dogs seem to have a was 4.5. Averaging across all intervals, the durations of
pattern of habituation that is insensitive to the temporary play for the first presentations of the two toys (P1.1 and P2)
absence of the stimuli that are becoming habituated, indi- were similar to each other and to those in experiment 1
cating that they remember those precise characteristics for (Fig. 3). Varying the time interval (10 s up to 15 min)
at least 10 min. Thompson (2009) suggests, from a review between the final presentation of toy 1, that is, once
of earlier studies by Harris (1943), Glanzer (1953), and habituation had been reached, and the presentation of P2
Welker (1961), that ‘the more rapid frequency of stimu- did not affect either the recovery (Kruskal–Wallis = 1.56,
lation, the more rapid and/or more pronounced the habit- P \ 0.91) or the dishabituation (Kruskal–Wallis = 6.05,
uation’. In this study, even when the inter-stimulus P \ 0.30).
intervals were brief, neither was found to be the case. This
lack of sensitivity to interruption is markedly different to Discussion
that observed in ‘classic’ habituation experiments. It is
unlikely that ‘ceiling’ effects played a part, since few dogs Dishabituation appeared to be insensitive to the duration of
played with the object for the full 30 s even on the first the delay since habituation. However, the relatively short
presentation. Thus, it is possible that object play behaviour maximum time interval used (15 min) may not have been
in dogs is terminated by mechanisms that are different sufficient to induce an observable effect. Dishabituation in
from, even if superficially similar to, habituation: the same cats did not diminish until the delay reached 25 min (Hall

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toys demonstrated by Kaulfuß and Mills (2008). However,


the unexpected lack of sensitivity to interruption of both
habituation and dishabituation suggests that more complex
inhibitory mechanisms may be involved in this species,
apparently reinforcing the effects of ‘classical’ habituation.
Since the results obtained here are broadly similar to
those found for domestic cats, it is clear that such habitu-
ation and dishabituation are common phenomena in the
object play of adult domesticated carnivores. Whether this
also applies to juveniles, which engage in more play than
adults do, remains to be tested.
More generally, our findings may be usefully applied to
environmental enrichment for kennelled dogs. It may be
possible to rekindle interest in objects provided to stimulate
play by altering any cue from the toy that is perceptible by
Fig. 3 Boxplot of duration of interaction (s) with toys at presentation the dog, so long as the toy remains a desired play object
1.1 (first presentation of toy 1), presentation 1F (final presentation of and motivation to play has not diminished (Celli et al.
toy 1 at the point of habituation), and presentation 2 (presentation of 2003; Tarou and Bashaw 2007). If the latter is the case,
toy 2, following habituation to toy 1) (means for all 5 time intervals
play is unlikely to restart, at least within 30 min or so,
combined) by the 12 dogs that habituated within 10 presentations in
experiment 3 regardless of the changes made.

et al. 2002), so the same response might occur in dogs if the Acknowledgments The authors would like to thank the BBSRC
and WALTHAMÒ Centre for Pet Nutrition for funding the project
time interval was extended beyond 15 min. The large post- (studentship to AJP). We are also grateful to WCPN for providing the
inhibitory rebound occurring in cats when the time interval facilities and dogs to enable the research to be undertaken.
was reduced to 5 min, such that play became more intense
than at the first presentation, was not observed in the dogs Conflict of interest Ralph Merrill is an employee of the WAL-
THAMÒ Centre for Pet Nutrition. Anne Pullen was in receipt of a
at this or any shorter time interval, suggesting that both studentship partially funded by the WALTHAMÒ Centre for Pet
habituation and dishabituation during object play may be Nutrition. John Bradshaw has no financial relationship with either of
qualitatively different in dogs as compared to cats (Hall the bodies that sponsored the research.
et al. 2002). However, several dogs played for almost the
whole of the 30 s available in P1.1 (see Fig. 3), and Ethical standards Approval was obtained from the University of
Bristol Ethics Committee for all the experiments described in this
therefore, any increase in play when P2 was presented paper. As part of this process, confirmation was obtained that none of
would have only been detectable for the dogs that did not the procedures used required a licence from the UK Home Office.
play throughout P1.1.
The motivation behind play is likely to be a contributing
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