Relación Cortisol Personalidad Canina

Applied Animal Behaviour Science 146 (2013) 96 106

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Applied Animal Behaviour Science

journa l h omepa g e: www.elsev ier .com/ locate /applan im

Explori weebehavi mpa

Lydia Ott rsonDepartment of Psychology, Memorial University of Newfoundland, St. Johns, NL, A1B 3X9, Canada

a r t i c l e i n f o

Article history:Accepted 2 ApAvailable onlin

Keywords:Domestic dog CortisolPersonalitySocial behavioDog park

a b s t r a c t

1. Introdu

The relacortisol, sochuman ani

CorresponE-mail add

0168-1591/$ http://dx.doi.oril 2013e 28 April 2013

(Canis familiaris)

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The relationships between behaviour, owner-rated personality, and cortisol were exam-ined in companion dogs that visited a local off-leash dog park. In Study 1, salivary cortisolincreased signicantly from baseline levels following 20 min in the dog park (P = 0.013),but not in the same dogs following a 20 min on-leash walk. In Study 2, cortisol was cor-related with dog park visit frequency, such that dogs which visited the park least oftenhad higher cortisol levels (r = 0.34, P = 0.013). Hunched posture in dogs was associatedwith higher cortisol, even after the effect of park visit frequency was removed. Cortisolappeared to be independent of all other measured behaviours and signals indicative of play,stress, agonism, and mounting, as well as dog time budgets. Nor was cortisol related to dogpersonality scores as measured by the Monash Canine Personality Questionnaire-Revised(MCPQ-R). Scores on the Extraversion, Amicability, and Neuroticism scales predicted someobservations in the park: more extraverted dogs showed higher activity (measured astime budget state changes; R2 = 0.21, P < 0.001) and spent more time in conspecic dyads(R2 = 0.083, P = 0.033), more amicable dogs showed more behaviours indicative of play(R2 = 0.10, P = 0.014), and more neurotic dogs showed higher frequencies of hunched pos-ture (R2 = 0.10, P = 0.008). Time budget states correlated with specic behaviours, e.g., focaldogs time spent in dyads correlated highly with total play signals/behaviours in the session(r = 0.69, P < 0.001). Thus, in a social context such as an off-leash dog park, changes in corti-sol may be largely independent of social behaviour/signalling (with the possible exceptionof postural changes), and personality scores may predict some social behaviours, but notnecessarily changes in cortisol. Given that this dog park sample contains dogs which appearto score higher than average for Extraversion, additional relationships between personal-ity, behaviour and cortisol may be detected in broader dog populations and/or other socialcontexts. As the popularity of off-leash dog parks is increasing in North America, under-standing factors related to individual dogs experiences in such parks may be important forwelfare reasons.

2013 Elsevier B.V. All rights reserved.

ction

tionships between the stress-related hormoneial behaviours, and personality traits in non-mals have rarely been considered together.

ding author. Tel.: +1 709 864 4738; fax: +1 709 864 2430.ress: [email protected] (C.J. Walsh).

Studies in various species have examined two-way com-ponents of these relationships: i.e., behaviour and cortisol(e.g., birds, Garamszegi et al., 2012; baboons, Sapolsky andRay, 1989; dogs, Hiby et al., 2006 and Blackwell et al.,2010), behaviour and personality (e.g., horses, Lloyd et al.,2007; langur monkeys, Konecn et al., 2008), and person-ality and cortisol (e.g., rhesus macaques, Capitanio et al.,2004). However, these three factors likely inuence oneanother in specic contexts, particularly in highly arousing

see front matter 2013 Elsevier B.V. All rights reserved.rg/10.1016/j.applanim.2013.04.002ng the dog park: Relationships betours, personality and cortisol in co

enheimer Carrier, Amanda Cyr, Rita E. Anden socialnion dogs

, Carolyn J. Walsh

L. Ottenheimer Carrier et al. / Applied Animal Behaviour Science 146 (2013) 96 106 97

social situations with conspecics, such as is found amongdogs at off-leash dog parks. With the increasing popularityof such parks in North America (cited in Lee et al., 2009),greater numbers of companion dogs are being exposed todog parks. Kand social bcations for tindividual such parks.factors thatignored by

Cortisol the hypothato externalmeasured n2005, 2009cortisol andin kennel seand on wormay be stre2011). In tlack of relateven when Beerda et aet al., 2001;

In non-hpersonalitying styles, consistent stressors asles (Koolhcharacterizing style shposture, sna cortisol (Horvth etsimilarly toviduals thattime and acoping stylobservationoped for npersonalityfearfulnessrated for spedgeable wpersonalityBorchelt, 1Svartberg asensus on t(reviewed analysis of moderatelysonality scopersonalitybehaviouraa few attemto which probserved bfrom those2003; Svart

The Monash Canine Personality Questionnaire-Revised(MCPQ-R) is an owner-based survey that uses a ratingscale for 26 traits that cluster into ve broad personal-ity dimensions: Extraversion, Motivation, Training Focus,

ability dimenversioivationgs (Le

both ib) and, 2009ruct vother

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exhibnder utionadiscipl10/11nowing how dog personality, stress responses,ehaviour interact in this setting may have impli-he welfare of pet dogs which visit dog parks, asdogs may have highly variable experiences in

To date, dog behaviour at dog parks, and the may inuence this behaviour, has been largelyresearchers (cf. Shyan et al., 2003).is a glucocorticoid hormone that is released bylamic-pituitary-adrenal (HPA) axis in response

and internal stimuli (Nelson, 1995). It can beon-invasively via saliva (Dreschel and Granger,). In dogs, research on the relationship between

behaviour has primarily focused on individualsttings (Coppola et al., 2006; Rooney et al., 2007)king or assistance dogs in work situations thatssful (e.g., Haverbeke et al., 2008; King et al.,

hese contexts, there appears to be an overallionship between the hormone and behaviours,a change in cortisol has been demonstrated (e.g.,l., 1998; Dreschel and Granger, 2005; Hennessy

Rooney et al., 2007).umans, the relationships between cortisol and

have emerged mainly in the context of cop-dened as individual differences in temporallyresponses to physiological and behaviouralsociated with particular neuroendocrine pro-aas et al., 1999). For example, police dogs

ed behaviourally as having an ambivalent cop-owed more signs of acute stress (e.g., low bodyout-licking, and paw-lifting) and demonstratedsurge in response to a threatening stimulus

al., 2007). In broad terms, personality is dened coping style, e.g., the characteristics of indi-

describe consistent patterns of behaviour overcross situations (Gosling, 2008). In contrast toes that are derived from direct behaviourals and analyses, many personality scales devel-on-humans are based on cluster analyses of

traits or adjectives (e.g., exploration, boldness,, aggression; Gosling and Harley, 2009) that areecic animals by experts or individuals knowl-ith the animals behaviour. In dogs, several

scales have been developed (e.g., Goodloe and998; Hsu and Serpell, 2003; Ley et al., 2008;nd Forkman, 2002), but there is not yet con-he dimensions that comprise dog personalityin Jones and Gosling, 2005). A recent meta-data from 31 studies determined that there is

high temporal consistency (R = 0.43) in dog per-res, with no differences in consistency between

scores based on behavioural ratings versusl coding (Fratkin et al., 2013). However, onlypts have been made to evaluate the degree

oposed canine personality dimensions predictehaviour of individuals in contexts different

in which they were developed (Gosling et al.,berg and Forkman, 2002; Svartberg et al., 2005).

AmictheseExtra(Mottic dohave2009et al.constover

InpersoindicdogsfreelySociastatemay examconspwe ausedfocalpostuMariand mounin doerancworkbehatisol aroussonalevelcismand hdogsimpaon th

2. M

Twber osampologyafterowneexamlevelsionsout uInstitInter(# 20, and Neuroticism (Ley et al., 2008). Some ofsions appear to be shared by many species (e.g.,n, Amicability, and Neuroticism), while others

and Training Focus) may be unique to domes-y et al., 2008). The MCPQ-R has been shown tonter-rater and testretest reliability (Ley et al.,

validity across purebred dog breed groups (Leya). Given that the MCPQ-R has been tested foralidity and is easy to administer, it was preferredcanine personality scales for this study.rrent study, the relationships between cortisol,, and specic behaviours and postural changesf play, agonism, and stress were explored in

as companion animals, while they interacted conspecic social setting, i.e., the dog park.ls reect the internal motivational or emotionale individual producing the signals, and, in turn,ce conspecics (Kelley, 1981). Play signals, fore used to initiate and continue interactions withs (Bekoff, 1995; Horowitz, 2009). In this paper,oth play signals and attention-getting signals

context of play (Horowitz, 2009) generated byn the dog park. We also assess behaviours andanges that indicate stress (Beerda et al., 1999a,b;., 2012) and those that indicate agonism (Scott

1965). Finally, we examine the incidence ofehaviour, as it is a behaviour often observeds for which owners appear to have low tol-lsh et al., 2011). Despite the lack of empirical

relationships between cortisol, personality and, we expected to detect: (1) an increase in cor-in dogs at the dog park, given the potentiallycial setting, and (2) relationships between per-d behaviours/signals that are related to cortisole dog park, e.g., dogs scoring high on Neuroti-

be expected to show higher cortisol levels frequencies of stress-related behaviours thanower scores. We also investigated the potentialother demographic and environmental factorsbles measured.

ls and methods

dies were performed. In the rst, a small num- was examined to determine whether saliva

dogs in a dog park setting was a viable method-whether dogs showed a difference in cortisolpark visit compared to after a walk with theirhe second study, a larger sample of dogs wasto investigate relationships between cortisoll behaviours or signals, and personality dimen-ited in the dog park. Both studies were carriedthe ethical approval of Memorial Universitysl Animal Care Committee (#11-01-CW) and theinary Committee on Ethics in Human Research-045-SC; #2010/11-151-SC).

98 L. Ottenheimer Carrier et al. / Applied Animal Behaviour Science 146 (2013) 96 106

2.1. Study 1

2.1.1. SubjectsEleven owners and dogs (ve female) were recruited

from the coyears (4.49 spayed or nited the dogthe remainpark.

2.1.2. ProceOwners

were askeddogs: one 2bourhood aa local off-ledog park wto see activseveral bengated. The dogs and twnated. The pgrass.

Owners take four saengaging in2030 min the dog parin the dog pshown to in1998), we aing to earlypark visit aday. In all bpark visit, awalk saliva

Writtenin the kit,on-line vidple (http:/were takenmer spongLLC., Pennsdescribed bkeep the swit immediaage Tubes, Sthe samplewere brougsaliva sampsamples we(A.C.) met tin taking th

Dog parnition usinElectric Co. were savedrial Universto code spevideos wer

measures extracted from the videos in Study 2; thus,behavioural data for Study 1 are not reported.

2.1.3. Cortisol analysesliva sses weresched usiwith asylvanhen a

verage

tudy 2

Subjexty comnthss (34 Dog Pags (67wners, and 8wners

timesthat wxcept

commcal dor (10%)

Matee dog17:0

have on by o enteable (uogs tact andded thagreed

basicex, br

housog parher thwhich

consests of 2lly doe(Ley etg the s) for 2vely saping ssertin.1.2) fe to eby the

the swmmunity. Dog ages ranged from 8 months to 11 3.83 years; mean SD); most dogs (73%) wereeutered. Three owners reported that they vis-

park more than three times per month, whileing owners reported less frequent visits to the

durewere provided with a saliva sampling kit and

to perform two separate activities with their030 min on-leash walk in their usual neigh-nd one 20-min visit to the Quidi Vidi Dog Park,ash enclosed park measuring 45 m 65 m. Thisas enclosed by wire fencing, enabling dogsity occurring outside the park, and containedches at which dog owners invariably congre-park also contained a water fountain for theo re hydrants on which dogs frequently uri-ark substrate was composed of ne gravel and

were informed that they would be required toliva samples from their dog: (1) 30 min prior to

a walk (pre-walk), (2) immediately following awalk (post-walk), (3) 30 min prior to leaving fork session (pre-park), and (4) 20 min after arrivalark (post-park). While time of day has not beenuence canine cortisol levels (e.g., Beerda et al.,sked owners do these activities from late morn-

evening, and to perform the walk and the dogt approximately the same time on a differentut one case, the walk event occurred before thes it was convenient for owners to deliver their

samples to the researcher at the dog park. instructions for saliva sampling were included

and owners were asked to view a shorteo demonstrating how to take a saliva sam-/youtu.be/9QdJLm9Udro). All saliva samples

with Salimetrics Childrens Swabs inert poly-es, measuring 8 mm 125 mm (Salimetricsylvania, USA) and subsequently analysed, aselow. For each sample, owners were asked toab in the dogs mouth for 4560 s, then to placetely in the polypropylene cryovial (Swab Stor-alimetrics LLC., Pennsylvania, USA) and freeze

immediately. Pre-walk and post-walk samplesht to scheduled dog park visits. At the dog park,les were immediately placed on ice, and allre frozen at 20 C within 2 h. The researcherhe owner and dog at the dog park and assistede post-park sample only if the owner requested.k sessions were video recorded in high de-g a Sanyo VPC-HD1010 digital camera (SanyoLtd., Osaka, Japan) at 60 frames per s, and videos

in .mp4 format. Logger.app ( A. Earle, Memo-ity) was used as the behavioural event recordercic behaviours from these video les. Thesee used as pilot data to rene the behavioural

Saanalysee Dassaytisol Penncate wthe a

2.2. S

2.2.1.Si

6 mobreedVidi of doing oyearsdog othreedogs one emostthe foBoxe

2.2.2.Th

12:00may chosedog tavailous dprojeproviwho videdage, sin thethe dwhetwith Afterconsi1 (readog) (usin2.1.2relatithe tby intion 2choicseen Onceamples were sent to Salimetrics LLC, wherere carried out (for detailed procedural details,

el and Granger, 2009). In brief, samples wereng a standard enzyme immunoassay kit for cor-

sensitivity of


stored in a portable cooler containing ice packs, and frozenat 20 C, with no more than 3 h between sample acqui-sition and freezer storage. A maximum of three focal dogswere videotaped during each park visit.

2.2.3. CortiCortisol

1.

2.2.4. TimeThe gen

in the dog Time budgefrequency atime alone,group (withand time intime budgedistance behumans. Spin a dyad, cwhen it waslength for a1.01.5 m) tance indicin time. Thethe dog was1.5 m fromting with thchanges wadog switchmeasure separk.

The freqindicative othe operatiosignals werfocal dog, wof any type

2.2.5. MCPQPersona

were calcuafter videobehaviouradog. As perwithin eachand dividedscale. The rcreating a psions for ev

2.2.6. StatisAll statis

Statistics 1and not n2005), log-for all statirithmic tra(percent) sand focal

square-root transformed. If data transformations did notnormalize the distribution for a given variable, conrmedby one-sample Kolmogorov-Smirnov tests (as was thecase for mounting and agonistic behaviour), nonpara-

ic testucted reseneporteo allow

1, a le acol. In ses wo-tail

correcishedariabl

madeonferris stud

Inter assecategoed obsted vidlations inteon-amting,

s for thvioursg almoever, fo

lower2. Becd wel

re-codvioursles forreed usess ts dec

viour d behnsus p

sults

Study 1

x of thsis (twles). Inof thertisol og pas for sol chmeansol analysesanalyses were carried out as described for Study

budget and behavioural codingeral activity and behaviours of the focal dogspark were coded from the video recordings.ts for each focal dog were quantied via thend duration of ve mutually exclusive states:

time in dyad (with one conspecic), time in dog two or more conspecics), time with humans,

mixed groups (with dogs and humans). Thet state of the focal dog was determined by thetween it and any surrounding conspecics orecically, the focal dog was viewed as being

onspecic group, mixed group, or with humans judged to be within one to one and a half of then average Labrador Retriever (approximatelyof the appropriate social stimulus. Greater dis-ated that the focal dog was alone at that point

only exception to this distance rule was when engaged in a game of fetch and moved beyond

the person, but was obviously still interac-e person. Total frequency of behavioural states quantied by adding the number of times aed from one time budget state to another. Thisrved as a proxy index of overall activity in the

uency of mounting, and behaviours or signalsf play, agonism and stress were quantied usingnal denitions in Table 1. These behaviours ande assessed for the entire 20 min session for eachithout restricting the behaviour coding to bouts

of activity (e.g., play or aggression).

-R scoringlity scores on the ve MCPQ-R dimensionslated either by an independent researcher ortape coding was completed to ensure thatl coders were blind to the MCPQ-R scores of each

Ley et al. (2009a), raw scores for each adjective personality dimension subscale were summed

by the maximum score possible for the sub-esult was converted to a percentage, therebyercent score for each of ve personality dimen-ery focal dog.

tical analysestics were conducted using the program PASW8. Since cortisol levels are positively skewedormally distributed (Dreschel and Granger,transformed values of cortisol data were usedstical analyses involving the hormone. Loga-nsformations were also used for personalitycores. Behavioural events (frequency data)dog time budgets (proportional data) were

metrcondare pare rate, tStudysampcortisanalyas twroni we wthe visonsthe Bfor th

2.2.7.To

four trainseleccorreassesthe nmounraterbehacatinHowtiallyof 0.4agreeraterbehathe disagto assensubeharelateconse

3. Re

3.1.

Sianalysampeach

Cothe dmentCortipark equivalents (e.g., Spearmans rho, rs) wereon non-transformed data. Transformed datated in gures, and back-transformed meansd in the text and in tables, where appropri-

for biologically-meaningful interpretation. Inrepeated measures ANOVA (pre/post-activitytivity type) was used to examine changes inStudy 2, multiple regression and correlationalere conducted. All probabilities are reporteded, and alpha level was set at 0.05. Bonfer-tions to probability values were not applied, as

to explore the potential relationships amonges of interest. Given the number of compar-

(e.g., time budget states social behaviours),oni correction was considered too conservativey (Bender and Lange, 2001).

-rater reliabilityss the inter-rater reliability of each of theries coded and listed in Table 1, a seconderver assessed 12 (20% of sample) randomly-eos. As the data were continuous, intra-class

s (ICCs) were calculated for each category tor-rater reliability (Rousson et al., 2002). Duebiguity and rarity of agonistic behaviours andthere was absolute agreement between theese categories (ICC = 1). For frequency of play

/signals, intraclass correlation was 0.99, indi-st perfect agreement among the two raters.r stress-related behaviours, there was substan-

agreement, with an initial intraclass correlationause of this poor agreement among raters whol in other behavioural categories, the seconded the remaining 48 videos for stress-related

only. Upon completion, the rst rater examined instances in which the two raters agreed andpon behaviours. The videos were re-watchedhose instances of disagreement, and a con-ision was made as to whether the particularwas or was not present. Thus, for all stress-aviours, the frequencies reported reect thisrocess.

e 44 samples had insufcient saliva for hormoneo post-walk, one pre-park, and three post-park

total, six of 11 dogs had cortisol measures at four saliva sampling times.levels increased signicantly after 20 min inrk, relative to the pre-park cortisol measure-the same dogs (F1,5 = 14.34, P < 0.013; Fig. 1).anged an average of 0.06 g/dL (from pre-

= 0.14 g/dL to post-park mean = 0.20 g/dL;


Table 1Categories of behavioural activities and denitions for specic behaviours and signals. Play signal denitions are modied from Horowitz (2009); ps = playsignal, att = attention-getting signal.

Behaviour Description

Play/attentionExaggerated approach (ps) Slow, running approach in sightline of partner; loose, rolling nExaggerated retreat (att) Backwards leap; head up towards partner.Play bow (ps) Forelimbs down; hind legs raised; tail erect or wagging.Chase-me (ps) Withdraw with looks backward; at a reduced pace or with lop

for 5 s.Open-mouth (ps) Frontal display with teeth and lips showing; no biting.Bow head (ps) Nod head below shoulder level; maintain or nod up.Play slap (ps) Usually simultaneous slap of ground with two forelimbs, occuLeap-on (ps) On hind legs, with front paws around partners head; tail up.Bump (att) Use part of body to knock into partner.Nose (att) Put nose and closed mouth to other; non-investigatory.Bite (att) Make rm mouth contact (of scruff, rump, face, or body); forcBite-at (att) Can have no clear object; biting at air in the direction of, but nPaw (att) Paw at others face or body.

AgonismGrowl A low frequency but audible rumbling produced in the throatBare teeth Lips curled upward, possible exposure of teeth.Snap Sudden biting motion in direction of a conspecic.Bite Firm mouth contact where mouth and teeth have rm grip onLunge Sudden angular leap towards conspecic.Chase Driving away conspecic.

StressTucked tail hen taiHunched po nd; newPaw lift t.Snout lick nose.Run away cation w

ack), buYawn Pull away ical inte

Mounting

back-transfnot signic

3.2. Study 2

3.2.1. TimeIn ve in

park or only

Fig. 1. Salivar20 min walk asignicant incments (transfoTail positioned between the back legs; new instance wsture Back curved upward, body and head lower to the grou

One front paw is lifted off the ground and slightly benTongue runs over top of snout, usually going over theRemoving or attempting to remove oneself from alterchance to interact with other dog (stopping, looking bMouth open wide with large intake of breath.Removing or attempting to remove oneself from physAttempting to clasp or successfully clasping front legs around

ormed means). Post-walk cortisol levels wereantly different from pre-walk levels.

budget and behaviourstances, focal dogs were either alone in the dog

one other dog was present for more than half

y cortisol measures for dogs (N = 6) taken prior to and after and prior to and after a 20 min dog park visit. Dogs showed arease in cortisol between pre-park and post-park measure-rmed mean sem).

of the sessian opportutaneously. Dall analysesber of dogs with approaverage, dotheir time with other bined). Perhmore than 4vicinity of hhuman-dog

Frequenseveral timchange freqtion of timegroups (r = time with hysis showea larger prfemales of a

As showplay behaviP = 0.003). Dtended to eP = 0.052), aature to run.

ing stride; new instance of behaviour when it persisted

rs in play bow position.

e is tempered.ot touching, partner; can be partial or repeated.

.

conspecic.

l remained tucked for 5 s. instance when back stayed curved for 5 s.

ith conspecic; new instance counted when dog hadt removed itself.

raction with human. conspecics body and performing pelvic thrusts.

on and, hence, these focal dogs did not havenity to interact with two or more dogs simul-ata from these ve dogs were removed from

of time budgets (remaining N = 55). The num-in the park for each session was relatively low,ximately 7 (3; mean SD) dogs present. Ongs were alone for approximately one-third ofin the dog park, and spent 23% of their timedogs exclusively (i.e., in dyads and groups com-aps due to the physical parameters of the park,0% of focal dogs time budget was spent in theumans, either solely with humans, or in mixed

groups (Fig. 2).cy of activity state changes was related toe budget states. As shown in Table 2, stateuency was positively correlated with propor-

in dyads (r = 0.43, P < 0.001) and in conspecic0.58, P = 0.001), and negatively correlated withumans (r = 0.36, P = 0.007). Regression anal-

d an age sex interaction; young males spentoportion of their time budget in dyads thanll ages (F2,53 = 3.50, R2 = 0.12, P = 0.038).n in Table 2, the frequencies of mounting andour/signals were positively correlated (rs = 0.38,ogs that displayed more agonistic behaviours

xhibit more stress-related behaviours (rs = 0.26,nd total number of stress-related behaviours


Table 2Correlations between (a) focal dogs time budget states, frequency of specic social behaviours/signals, (b) Monash Canine Personality Questionnaire-Revised (MCPQ-R) personality dimension scores, and (c) activity level (*P < 0.05, **P < 0.01, ***P < 0.001).

Time budget Behaviours

Alone Dyad Group Human Mixed Total PlaySignals

Total Stress-related

HunchedPosture

TotalAgonism

Mounting

(a) BehaviourTotal Play 0.23 0.69*** 0.43** 0.40** 0.01 0.05 0.05 0.01 0.38**Total stress-related 0.13 0.09 0.07 0.10 0.24 0.62*** 0.26 0.06Hunched posture 0.11 0.06 0.03 0.03 0.16 0.13 0.02Total agonism 0.03 0.02 0.38** 0.07 0.11 0.02Mounting 0.24 0.41** 0.34* 0.29* 0.16 (b) MCPQ-R DimensionsExtraversion 0.23 0.29* 0.19 0.22 0.34* 0.25 0.13 0.05 0.01 0.03Motivation 0.11 0.05 0.11 0.17 0.01 0.04 0.01 0.21 0.16 0.07Training focus 0.18 0.14 0.08 0.09 0.16 0.12 0.002 0.09 0.20 0.08Amicability 0.16 0.24 0.03 0.03 0.06 0.32* 0.16 0.09 0.09 0.06Neuroticism 0.06 0.13 0.04 0.03 0.14 0.12 0.09 0.32** 0.09 0.04(c) Frequency of

state changes0.05 0.43** 0.58*** 0.36** 0.01 0.29* 0.03 0.06 0.17 0.10

shown washunched postress-relatture correlaand run awtail and ruP < 0.001). behaviours(data not shatively comcorrelated o

RelationbehavioursTotal play portion of tdog groupswith time frequency atime budgebehaviour fin dog groucorrelated (r = 0.41, P =

Fig. 2. Proporbudget states

egatively correlated with time spent with humans0.29, P = 0.033).unger dogs displayed play signals more frequently0.36, P = 0.006). Older dogs were found to be gener-

ess active (age and state change frequency: r = 0.37,006). The only sex difference in the coded behavioursounting; only males mounted and of the seven malesid so, three were sexually intact.

verall, 98% of dogs displayed at least one stress-relatedviour, 83% displayed play behaviours or signals,

3tage of focal dogs (N = 55) exhibiting total and specicttention-getting signals and stress-related behaviours, andgonistic behaviours and mounting. The back-transformed meanncy of fohe 20 m

l play p

bow geratede-me gerated-at

correlated signicantly with the frequency ofsture (r = 0.62, P < 0.001). Within the category ofed behaviours, the frequency of hunched pos-ted highly with tucked tail (r = 0.78, P < 0.001),ay behaviours (r = 0.75, P < 0.001), while tuckedn away also correlated signicantly (r = 0.54,Not surprisingly, most of the coded play/signals correlated signicantly with each otherown). A notable exception to this was the rel-mon behaviour exaggerated approach, whichnly with play bow (r = 0.38, P < 0.004).ships were also found between/signals and time budget, as shown in Table 2.frequency positively correlated with the pro-ime dogs spent in dyads (r = 0.69, P < 0.001) and

(r = 0.43, P = 0.001), and negatively correlatedspent with humans (r = 0.40, P = 0.003). Playlso correlated positively with the frequency oft state changes (r = 0.29, P = 0.037). Agonisticrequency positively correlated with time spentps (r = 0.38, P = 0.005). Mounting was positivelywith the amount of time dogs spent in dyads

0.002) and in dog groups (r = 0.34, P = 0.012)

and n(r =

Yo(rs = ally lP = 0.was mthat d

Obeha

Table Percenplay/atotal afrequeover t

TotaBumPlayExagChasExagBitePawtion of time dogs (N = 55) were engaged in various timein the dog park (mean sd).

Leap-on Open mouthNose Bite Play slap Bow headTotal stressSnout lick Paw lift Hunched poRun away Tucked tail Yawn Pull away Total agoniTotal moun

The bolded vathe non-boldethe totals. Thecal dogs specic play signals and stress-related behavioursin session is reported.

Percent Focal dogs Mean Frequency

83% 14.9154% 1.9551% 1.28

approach 49% 0.5849% 0.63

retreat 40% 0.7133% 0.5032% 0.41

28% 0.51

26% 0.2621% 0.0919% 0.087% 0.010%

98% 13.1283% 5.3476% 2.09

sture 36% 0.4531% 0.2527% 0.3717% 0.053% 0.01

sm 22% 0.33ting 12% 0.18

lues are for the total behaviours in each category, whereasd numbers indicate the specic behaviours that comprise

bolding was used as a way of differentiating the totals.


Table 4Descriptive statistics for the ve Monash Canine Personality Questionnaire-Revised (MCPQ-R) personality dimensions measured by owner ratings for focaldogs in the dog park (N = 59).

MCPQ-R Personality Dimensions

aining Focus Amicability Neuroticism

Mean (%) 6.89 80.90 46.97S.D. 2.75 11.44 14.90Minimum (% 3.30 53.30 14.70Maximum (% 0.00 100.00 83.30

22% showemounted awhich showtive of playaverage frein Table 3.

3.2.2. PersoThe mea

ve personobtained bwhich was Ley study (T

Multiplecability anand behavicability sigplay behavP = 0.014). Eof time dogP = 0.033). time in mixits positivethat scoredber of stateR2 = 0.21, Pspecically(r = 0.32; F1quency of t

The dimcorrelate wstates.

3.2.3. CortiTwo of t

testing. Thethe values edata from tyses, as it iscontaminatet al., 2007was 0.22 gvalue obtai

Cortisol ber of time(r = 0.34, Pof park visithan four vpattern, as tfrequency b

dogs were removed from the sample. Cortisol andequency of hunched posture correlated signicantly.31, P = 0.023), even after the effect of dog park visitency was removed via partial correlation (r = 0.29,04).e number of days since a dog last visited the park

cantly correlated with the frequency of total stress-d behaviours (r = 0.30, P = 0.024), but was unrelatedrtisol levels. Specically, focal dogs that had last vis-he park within 7 days (N = 30) had signicantly fewers-relatark 7

groulevels.

Otherher ponality

variabiarity h of timived inr foca

signi unfamat the

who w to doed no equens.Extraversion Motivation Tr

76.11 66.38 716.51 14.62 1

) 25.00 33.33 4) 100.00 90.00 10

d one or more agonistic behaviours, and 12%t least once (Table 3). The proportion of dogsed the specic behaviours and signals indica-

and stress, as well as the back-transformedquencies of these behaviours/signals, is shown

nality dimensionsn (non-transformed) percentages across theality dimensions were comparable to thosey Ley et al. (2009a) except for Extraversion,13.1% higher than the scores reported in theable 4).

regression analyses showed that both Ami-d Extraversion correlated with time budgetoural frequencies (shown in Table 2). Ami-nicantly predicted the frequency of totaliours/signals (r = 0.32; F1,58 = 6.49; R2 = 0.10,xtraversion signicantly predicted the amounts spent in dyads (r = 0.29; F1,58 = 4.81; R2 = 0.083,Extraversion was negatively correlated withed groups (r = 0.34, P = 0.012), likely due to

correlation with time spent in dyads. Dogs high in Extraversion also had a higher num-

changes (activity level; r = 0.46; F1,58 = 15.25; < 0.001). Interestingly, Neuroticism scores

predicted the frequency of hunched posture,58 = 7.54; R2 = 0.10, P = 0.008), but not the fre-otal stress-related behaviours.ensions Training Focus and Motivation did notith any behavioural measures or time budget

solhe 60 saliva samples had insufcient saliva for

cortisol levels for two dogs were outliers, asxceeded the mean by two standard deviations;hese dogs were removed from all cortisol anal-

possible that these high values resulted fromion by blood in the dogs mouths (e.g., Granger). The average back-transformed cortisol level/dL (N = 56), slightly higher than the post-park

ned in Study 1 (0.20 g/dL).

ve the fr(r = 0frequP = 0.

Thsignirelateto coited tstresthe pThesetisol

3.2.4.Ot

persosuredfamillengtdog lage, oto bewerepark Dogsparedshowthe frscorelevels were negatively correlated with the num-s a dog visited the park in the past 365 days

= 0.013; Fig. 3). Five dogs in the 10th percentilets (i.e., the least frequent visitors with fewerisits in the last year) were responsible for thishe relationship between cortisol and park visitecome statistically non-signicant when these

Fig. 3. Salivarfrequency in ted behaviours than those dogs that had been inor more days ago (N = 23; t51 = 2.15, P = 0.04).ps did not show a signicant difference in cor-

variablesssible relationships between each of cortisol,

, behaviour, and time budgets, and other mea-les, specically the number of dogs in the park,

of conspecics, focal dog reproductive status,e focal dog lived with owner, whether the focal

a single dog or multi-dog household, focal dogl dog sex, were examined and none were foundcant. For example, 57% of the dogs in the sampleiliar with any of the other dogs present in thetime of their visit, as reported by their owners.ere familiar with other dogs in the park com-

gs who were unfamiliar with their conspecicssignicant differences in cortisol, time budgets,cy of any behaviours measured, or personalityy cortisol levels negatively correlated with dog park visithe past year (N = 56).


4. Discussion

In Study 1, cortisol increased signicantly in dogs after20 min in the dog park, but not following an on-leash walkwith ownerthe dog parand cortisomeasured inhunched ortiple democortisol levspecics in was only nvisited the due to the fdog park assince whenwere remocortisol andble results hdogs expossubstantialstress (e.g., number of unrelated toof total strepark withinfewer behavisited mor

The expbehavioursported, in cortisol lev(Beerda et aand van dera social signstatus (e.g.,estingly, poto inuenceet al., 2010power posecortisol relapowered poincreased cthat posturstate of mabe reectedand cortisopersonalitymore frequbetween Nthe frequen

Taken tocortisol seeprocesses: iologically familiar andronment, reare not freqaroused, orting, thereb

that visit rarely), or to higher stress-related behaviour fre-quencies (in dogs which are not rare visitors, but which hadnot visited the park within the past week). An additionalreason for increased cortisol in some dogs in the dog park

be relalness-R Nef hunmay bdogs irtisol tsm anrguablcy of hency o

the bee failu

behabehavehaviot comes exabehavited an

and ol ha; Henna et and tected serve havio

(2009 a newure asver, shsed frolus, darly, indepenlated rns di

vs. wan et

viourscortisoduresol in gs (Stol has

capuchay, 19ated t, 2010)viour acollaree sole,is likeviour r whicviourss. In Study 2, dogs cortisol levels after 20 min ink were similar to the post-park levels in Study 1,l was independent of all behavioural activities

that time frame, with the singular exception of lowered posture. Despite also examining mul-graphic and other factors that might inuenceels in the dogs (e.g., age, sex, density of con-the park, presence of familiar dogs, etc.), cortisolegatively related to the number of times dogspark in the last year. This relationship may beact that infrequent park visitors experience the

a physically and socially novel environment, dogs that were the most infrequent visitorsved from the data, the relationship between

frequency of park visits disappeared. Compara-ave been found in previous research involving

ed to novelty, in which cortisol levels becamely lower between early and later exposures toHennessy et al., 2001; Rooney et al., 2007). Thedays since focal dogs last visited the park was

cortisol levels, but did inuence the frequencyss-related behaviours; dogs that had visited the

1 week of the test session showed signicantlyvioural indicators of stress than those which hade than a week prior to the session.ectation that cortisol would be related to

shown by dogs in the park was partially sup-that low or hunched posture correlated withels. Low posture in dogs is indicative of stressl., 1998, 1999b; Haverbeke et al., 2008; Schilder

Borg, 2004; van den Berg et al., 2003) and can beal of deference to conspecics of higher social

Bonanni et al., 2010; Handleman, 2008). Inter-stural changes in humans have been shown

both testosterone and cortisol levels (Carney). Participants instructed to maintain a high showed increased testosterone and decreasedtive to baseline levels, while maintaining a low-sition resulted in decreased testosterone andortisol levels (Carney et al., 2010). It is likelyal changes are reliable signals of the internalny species; in the dog park context, this may

by a relationship between lowered posturel. Additionally, dogs that scored higher in the

dimension Neuroticism showed signicantlyent hunched posture, despite no relationshipeuroticism scores and either cortisol levels orcy of total stress-related behaviours.gether, these data suggest that the increase inn in dogs in the dog park is caused by at least tworst, most dogs are likely emotionally and phys-

aroused by the presence of conspecics, both unfamiliar, and by the dog parks physical envi-sulting in increased cortisol; second, dogs whouent or recent visitors are likely additionally

stressed, by the novelty of the dog park set-y contributing to higher cortisol levels (in dogs

may fearfuMCPQcies oThis rotic in coroticiare aquenfrequwith

Thotherplay tic bis nostudicic repormonecortis2005Beerdtisol (subjto obcal beet al.tle inmeashoweassesstimuSimiltime-correpattedogsHekmbehavary procecortisof docortis(e.g.,and Ris relet al.beha(e.g. gales

It behaundebehated to their underlying predisposition towards. In this study, dogs which scored high in theuroticism dimension showed higher frequen-ched posture, but not higher levels of cortisol.e due to the fact that less fearful/low neu-n this setting show arousal-induced increaseshat may mask any relationship between neu-d cortisol. Those dogs showing hunched posturey the most stressed dogs in the park, as the fre-unched posture also correlated highly with thef total stress behaviours, and more specically,haviours of tucked tail and run away.re to nd relationships between cortisol and

viours measured (e.g., frequency of mounting,iours, stress-related behaviours, and agonis-urs), or the dogs time budgets in the park,pletely surprising. In fact, results from othermining cortisol and its relationship to spe-ours have been equivocal. Many studies have

overall lack of relationship between the hor-behaviours in dogs, even when a change ins been demonstrated (Dreschel and Granger,essy et al., 2001; Rooney et al., 2007). Whileal. (1999a) found correlations between cor-hree behaviours in chronically stressed dogsto social and spatial restriction), they faileda correlation between cortisol and the typi-ural expressions of chronic housing stress. Batt) found that young guide dogs ability to set-

kennel environment was the only behaviouralsociated with cortisol. Horvth et al. (2007),owed that the coping styles of police dogs, asm their behavioural responses to a threateningid signicantly correlate with cortisol changes.

dogs that recently entered a re-homing shelter,dent changes in urinary cortisol/creatine ratioswith dog activity/locomotion, but relationshipffered in direction when examined betweenithin the same individuals (Hiby et al., 2006).al. (2012) investigated the relationship of three- head resting, panting, and lip licking- to sali-l in dogs hospitalized for elective veterinary

. These behaviours signicantly correlated withtheir Study 1 dogs, but not in a second groupudy 2). In primates and various bird species,

been shown to predict some social behavioursins, Byrne and Suomi, 2002; baboons, Sapolsky89; grey-lag geese, Kralj-Fiser et al., 2010) ando coping styles in birds (e.g., great tits, Stwe. However, a lack of clear relationships betweennd cortisol has been reported in other animalsd ycatchers, Garamszegi et al., 2012; Sene-

Silva et al., 2010).ly that the relationships between cortisol andin any species are mediated by the contexth testing is carried out and by the particular

measured. It could be that certain behaviours


are reliable measures of arousal or stress in specic con-texts only (e.g., settling in a new kennel situation, Battet al., 2009). Alternatively, the relationships between cor-tisol and behaviours may be only reliably discerned atthe level obehaviourschanges in tet al., 1999)tially fear- olikely impacto the stimstrated thaincrease in tied as feaas fearless. of oors didin cortisol, stressor didogs. We dcerning themay have hcortisol-behbe because lated to corthis dog padog park sebehavioursarousal or environmenspecics. Inafter dogs hanticipation

Dog timeage. Youngedogs showalone to dyastates may social activcies were asignals, a gand conspespent near older dogs behavioursdog activitymost specieByers, 1998as domestic2007; Cordomay be thephysical ab

Positive and in consplay behaviing; these context. Dohigh levels omounting inphysical anpark settinging is an ele

(Leca et al., 2012). Play is composed of many behaviours,some of which are agonistic in other contexts, such asbiting, barking, chasing and growling (Bekoff, 1995). Asthere were no sexually receptive females (the appropriate

lus) inviour wl cont

the -R, Ex

d somark. Din Exty greatnalityversioy leveGoslinness a

act thaively tcores g behafound r theireys, Kan, 2

sed th may exclusversiole, likeor theose orir samighly ers as h

and rrs of sash pa

with o, dogsvertedo be p

on th and bighly ag playis dims the

and een d

et al., 2viours

this rendepealiditygs giveency oark. T

this di (Ley ef coping styles, in which clusters of specic that dene a coping style are associated withhe hormone (e.g., Horvth et al., 2007; Koolhaas. In addition, dogs past experiences with poten-r stress-inducing stimuli or environments willt their physiological and behavioural responsesuli. Hydbring-Sandberg et al. (2004) demon-t exposure to gunshot noise caused a drasticplasma cortisol in collies that had been iden-rful of gunshots, but not in collies describedIn that same study, collies which were afraid

not respond to the stressor with an increasealthough other physiological responses to theffered between oor-fearful and non-fearfulid not systematically collect information con-

types of past experiences that the focal dogsad in dog parks. Of course, the lack of strongaviour relationships in the present study may

the particular behaviours examined were unre-tisol (e.g., Garamszegi et al., 2012), at least inrk context. As well, an increase in cortisol in atting may not be related to most stress-related

in many dogs, because the increase may reectexcitement induced by the conspecic socialt, or anticipation of social interaction with con-

sled dogs, cortisol levels increased signicantlyad been harnessed for pulling, presumably in

of the activity (Angle et al., 2009). budgets in the park were inuenced by sex andr males spent the most time in dyads and older

ed less frequent changes in states (e.g., fromd, dyad to group, etc.). Changes in time budgetbe a useful proxy measure of overall dog-dogity in the park, as higher state change frequen-ssociated with higher frequency of total playreater proportion of time spent in both dyadscic groups, and a lower proportion of timehumans. Age was related to some behaviours;played less and showed fewer stress-related. Mirk et al. (2012) similarly reported that

level was negatively related to dog age. Ins, play is associated with juveniles (Bekoff and); however, adult wolves (Canis lupus) as well

dogs are known to play (e.g., Bauer and Smuts,ni, 2009). Decreased play activity in older dogs

result of decreased interest, motivation, orility.correlations between the time spent in dyadspecic groups with both the frequency of totalours/signals and of mounting are not surpris-behaviours must occur in a conspecic socialgs that showed mounting behaviour also hadf play behaviour. The co-occurrence of play and

these dogs may be a product of the heightenedd emotional arousal, which is elicited by the dog

or by play itself. It is also possible that mount-ment of play, as it is in juvenile male macaques

stimubehasexua

OfMCPQdictedog phigh cantlpersoExtraenergand ProneThe fpositthat sof doalso undemonkForkm

Bastudypled Extrasampings fto thin oui.e., hownelivelyowneoff-lethemtivelyextraties tstudyterns

Hcatinin thpassedogs,has b(Ley behaplay,and ithe vratinfrequdog psent timid the dog park during the times that mountingas recorded, mounting occurred outside of a

ext.ve personality dimensions assessed by thetraversion, Amicability, and Neuroticism pre-e behaviours and time budget states in theogs which were assessed by their owners asraversion were more active and spent signi-er proportions of time in dyads. In other canine

scales, dimensions similar to the MCPQ-Rsn are also highly associated with activity andl (Energy, Gosling et al., 2003; Activity, Jonesg, 2005; Activity, Mirk et al., 2012; Chase-nd Playfulness, Svartberg and Forkman, 2002).t owner-reported ratings for Extraversion relateo observed activity level in the park suggestson this dimension accurately describe this facetviour. Personality models in other species havetraits related to activity and sociability to t

respective Extraversion dimensions (languronecn et al., 2008; humans, Svartberg and002).e MCPQ-R results, the local dog park in thisattract a distinct group of dogs. Dogs sam-ively from the dog park scored 13.1% higher inn than the Ley et al. (2009a) community-basedly a more heterogeneous population. Our rat-

four other personality dimensions were similarginally reported. This may imply that the dogsple represent a subset of the dog population,xtraverted dogs, which are described by theirighly active, energetic, excitable, hyperactive,

estless (Ley et al., 2009a). It is possible that theuch dogs are more likely to bring their dogs to anrk than are owners of less active dogs to providepportunities to socialize and exercise. Alterna-

that attend the dog park may become more through that process, as they have opportuni-hysically active and to socialize. A prospectivee factors that relate to dog park attendance pat-ehaviour outcomes is warranted.micable dogs showed more behaviours indi-

compared to dogs which were rated lowerension. The dimension of Amicability encom-overall level of friendliness and sociability inparallels a social interaction component thatescribed in previous dog personality studies008; Svartberg, 2002). As the play signals and

measured in this study encompassed sociallationship between MCPQ-R Amicability ratingsndently scored behaviours and signals supports

of this dimension. The MCPQ-R Neuroticismn to dogs by their owners correlated with thef lowered or hunched posture exhibited in thehe adjectives used in the MCPQ-R to repre-mension are fearful, nervous, submissive, andt al., 2009a). Again, owner perception of their


dogs personality and independent observation of the dogsbehaviour correlated. It is possible that owner perception ofthe dogs level of extraversion, amicability, or neuroticismwas inuenced by their observations of their dogs in thedog park; hwhile owneof the sessiunderlying accurately dog park re

In a pracpark is a hialmost all dthe park (e.age also diswere fairly rappeared mbehaviourathe dog pawas associarelated behneurotic shThus, whiledog park, pimpaired wshowing lowto reconsidfare reasonowners rathave overalbenet fromthat such a

5. Conclus

Relationsignals, andpark settinare inuencparticular dand the beple, individsocial behaand hormodogs inuesol responssuch as howinteractionbe further ewere recruireect pattDifferent resocial behaparks, and/

Acknowled

This stuto CJW. ThCroucher, C

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Exploring the dog park: Relationships between social behaviours, personality and cortisol in companion dogs1 Introduction2 Materials and methods2.1 Study 12.1.1 Subjects2.1.2 Procedure2.1.3 Cortisol analyses

2.2 Study 22.2.1 Subjects2.2.2 Materials and procedure2.2.3 Cortisol analyses2.2.4 Time budget and behavioural coding2.2.5 MCPQ-R scoring2.2.6 Statistical analyses2.2.7 Inter-rater reliability

3 Results3.1 Study 13.2 Study 23.2.1 Time budget and behaviour3.2.2 Personality dimensions3.2.3 Cortisol3.2.4 Other variables

4 Discussion5 ConclusionAcknowledgmentsReferences

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Relación Cortisol Personalidad Canina