Accepted Article

Accepted Article *Author for correspondence([email protected]) *Author for Kingdom United 8QQ, G12 Glasgow of University Sciences, Life & Veterinary Medical, of College Building, Kerr Graham Medicine, Comparative & Health Animal ofBiodiversity, Institute Killen* S. Shaun Growth trajectory influences temperature preference Section : Physiological Ecology Editor : Murray Humphries Article type Paper : Standard 15-Apr-2014 : Date Accepted : 24-Jan-2014 Date Revised Received: Date 27-Nov-2013 This article is protected by copyright. All rights reserved. reserved. rights All copyright. by protected is article This 10.1111/1365-2656.12244 doi: as article this cite Please Record. of Version the and this version between differences to lead may which process, proofreading and pagination typesetting, copyediting, the through been not has but review peer full undergone and forpublication beenaccepted has article This Summary compensatory growth ecology, physiological nutrition, variation, intraspecific choice, habitat scope, aerobic keywords: fish in rate metabolic and selection Temperature headline: Running in fish through an effect on metabolic rate effect an through infish Accepted Article This article This protectedis by Allcopyright. rights reserved. 5) 4) 3) 2) 1)

are both plastic in response to feeding histor tofeeding response in bothplastic are thatvary temperatures prefer fish Individual wascorrelatedtemperature. positively withpreferred probably allow individuals to reduce mainten reduce to individuals allow probably reduction in activity. In growth-compensated fish and controls, activity measured at 10 at measured activity controls, and fish growth-compensated In activity. in reduction alarge displayed also growth compensatory experiencing Fish study. the throughout ration preferred temperatures that were on average 2.85 average on were that temperatures preferred fish growth-compensated these so and growth, compensatory of effects tothe due probably controls, to TheSMRofwaselevated the compared temperature. food-deprived the fish SM fish, control and food-deprived both In measured. was also regime temperature 10 shuttl a using temperatures preferred select days ; or 2) fed fed then days, 21 for food-deprived 1) either: were Individuals preference. temperature affect to interact standar history, feeding which to extent the This study examined i these growth history feeding trajectory. and linked to thermal preference andwhether such relationships are affected by factors such are traits physiological how on information is lacking Especially of individuals. temperature well-studied the at species level, lessis While choice. habitat on influence a strong has temperature particular, in ectotherms For variatio Most animals temperature experience o C, Acti oxygen estimated of uptake. by rates ad libitum ad throughout the entire period. All animals were then allowed to to allowed then were animals All period. entire the throughout ssues in the common minnow common minnow ssues inthe known about factors affecting the preferred preferred the affecting factors about known R was negatively correlated with preferred preferred with correlated negatively was R d metabolic rate (SMR) and aerobic scope (AS), (AS), scope aerobic and (SMR) rate metabolic d e-box, and then measured for SMR and AS at and AS SMR for thenmeasured and e-box, y growth trajectories.y and Cooler temperatures predictably with SMR and activity level, which which level, activity and SMR with predictably ance costs and divert more energy towards towards energy more divert and costs ance ns as they movethro they as ns vity within the shuttle-box under a constant a constant under theshuttle-box within vity o C cooler than controls fed a maintenance maintenance a fed controls than cooler C Phoxinus phoxinus Phoxinus ad libitum ad ugh the environment. theenvironment. ugh fornext the 74 , to determine , todetermine o C Accepted Article standard metabolic rate (SMR) in ectotherms) shows a large degree of variability among individuals individuals among variability of degree a large shows ectotherms) in rate(SMR) metabolic standard physiological traits (Angilletta, Niewiarowski Navas 2002). Baseline & metabolic demand (i.e. va species of thesame individuals which to extent (Angil level species atthe well-studied been have ectotherms of preferences thermal and thermoregulation behavioural While 2009). Porter & Shine microhabitatskeep to body temperature within relatively a narrow range (Reynolds 1978; Kearney, specific select or behaviours alter often ectotherms demands, physiological or fluctuations & Kingsolver 1989; Angilletta, Niewiarowski &Navas 2002). In response toexternal temperature lo including performance of measures various on For enviro ectotherms temperature. particular, in INTRODUCTION This article This protectedis by Allcopyright. rights reserved. changing environmental conditions (Killen conditions environmental changing determinants important are traits physiological and in behavioural differences individual selection, natural for requirement basic a being to addition al. et 2003;Stapley 2006; Clusella Trullas Nearly all animals are exposed to spatial and temporal variation in environmental habitats. cooler preferred to access limited having simultaneously while growth, compensatory to due experience frequent fluctuations in food supp clim warming In stressors. environmental with coping for scope aerobic surplus toincrease individuals allow also would activity, temperat cooler at SMR in reduction A growth. It is possible that animals may show differences in thermal preference in relation to to relation in preference thermal in differences mayshow animals that is possible It etal. et al. et 2007; Artacho, Jouanneau & Galliard 2013). InGalliard 2013). & Artacho, Jouanneau 2007; 2013; Humphries 2013; &McCann 2014). of how individuals and populations respond to to respond populations and individuals how of comotion, growth, and repr and growth, comotion, letta 2009), the examined fewstudieshave only letta a ates, however, aquatic ectotherms could could ectotherms aquatic however, ates, nmental temperature can have a profound impact impact aprofound can have temperature nmental ry with respect to temperature preference (Pulgar preference respect totemperature with ry ly with long-lasting effects on metabolic rate metabolic on effects long-lasting with ly ures, coupled with a decrease in spontaneous spontaneous in decrease a with coupled ures, oductive output (Huey output(Huey oductive

Accepted Article among individuals with regard to temperature preference. Examples of such plasticity have been been have plasticity of such Examples preference. temperature to regard with individuals among potential. this realise fully can throughput of energy preferen temperature and AS, SMR, among correlations This Glazier 2010). & Atkinson (Killen, rest at mitochondrial density) torequired support an lifestyle willactive maintained be need also to while mass, muscle increased (e.g. machinery” “metabolic the because andAS, SMR between correlation (Careau model theproduction andAS, SMR between correlation negative a predict would which model, allocation the to contrast In 2013). Clark & Malte Norin, therefore influence individual temperature preference and habitat selection (Khan &Herbert 2012; could &Malte2011), and timeframes (Norin medium-term short- and over least at be repeatable compensation model, CareauGarland and 2012), which states that basal metabolic demand variationwithin species (Norin &Malte Killen 2011; ectothermic species (Portner &Knust Pörtner2007; 2008). &Farrell Aerobic scope also shows of distribution thegeographic on influence alarge has AS that beensuggested recently has it 1993), Taper & Marquet (Brown, fitness increased with associated is AS increased that assumption requirements (e.g. 2008). activity,growth, Farrell & digestion; Pörtner Combined with the maintenance above ananimal within tasks physiological oxygen-consuming simultaneous for capacity the and sets (MMR) rate metabolic maximal its and SMR individual’s an between difference physiological determining variable in thermal preference absolute is aerobic scope (AS), which the is competesother with physiological processes fo This scenario is predicted by the allocation model of energy budgeting (Careau energy budgeting of model theallocation by predicted is scenario This cooler temperatures, particularlywhen food availability lowis (van Dijk, Staaks & Hardewig 2002). energetic their support to requirements foraging elev relatively a with individuals that speculated of thesame species (Burton This article This protectedis by Allcopyright. rights reserved. In addition to genetic sources of variation, environmental factors may generate variability variability generate may factors environmental variation, of sources genetic to addition In etal. 2011) and varies with body temperature. It has previously been previously has It temperature. body with and varies 2011) form of energy budgeting could generate positive energycould generate budgeting of form r acquired energy. Another potentially important ated SMR at a given temperature will have increased increased have will temperature given a at SMR ated demand, and may therefore preferentially select preferentially therefore may and demand, etal. ce, so that individuals capable of a high rates rates high of a capable individuals that so ce, 2012;Marras et al. et 2008) predicts positive a et al. et 2013), that appears to to that appears 2013), et al. et 2008; or Accepted Article Although it feeds on a range of prey items (e.g. algae, detritus, macrobenthos) this species species this macrobenthos) detritus, algae, (e.g. items prey of range a on feeds it Although lake environments tospatia they areexposed where ahabi is It Asia. in Europe in zones temperate across bygr affected be can of individuals temperature thepreferred whether 2) and animals; in individual AS) and SMR (i.e. traits physiological activity. as such processes, demanding energetically decrease in available AS if there is noconcomitant increase in MMR or offsettingchanges otherin a be could growth compensatory dueto SMR elevated an of effect adverse Anadditional 1990). Neill requirements and increase the surplus energy av maintenance to decrease attempt individuals as temperature preferred in decrease a cause poor diets can decrease the preferred temperatur thepreferred decrease can diets poor longer2006). Over the term,th meal (Wallman &Bennett temperatur select individuals as thermoregulation mostlyhave overlooked. been feed Recent common minnow the in questions these studied I traits. physiological on effects via or independently either growth, (Criscuolo concluded has growth rapid of period the evenafter persists that rate metabolic basal increased and 2006), Metcalfe & Lindstrom (Royle, exercise during endurance 2007), Metcalfe & (Álvarez ability locomotor including traits numerous to alterations long-lasting a cause to beenobserved –has growth reduced of period a following after extension, SMR and available AS. For example,comp by and, trajectories growth byinfluencing preference temperature alter could history Dietary 2002; Pulgar Fangue (Blouin-Demers level population the at observed This article This protectedis by Allcopyright. rights reserved. etal. The aims of this study were to determine: 1) todetermine: were 1) thisstudy of aims The et al. et 2009), but 2009), environmental effects on temper 2003) but the mechanisms that could underlie such a shift remain unknown. unknown. remain shift a such underlie could that mechanisms but the 2003) Phoxinus phoxinus Phoxinus . This is a freshwater teleost fish that has a wide distribution distribution wide a has fish that teleost afreshwater is This . ing, for example, may affect behavioural behavioural affect may example, ing, for etal. owth trajectories, sp and owth et al. ailable for growth (Priede 1985; Bryan, Kelsch & e Diof ectothermse (van es which digesttheir them toeffectively allow tat generalist, living in both flowing streams and and streams flowing both in living generalist, tat l and temporal fluctuations in thermal regimes. regimes. thermal in fluctuations temporal and l ensatory growth – the rapid growth that occurs occurs that growth rapid the – growth ensatory if individual temperature preference is preference to linked individual temperature if 2008). Increased maintenance costs could 2000; Mortensen,& Lund Ugedal 2007; ature preference at the individual level level individual atthe preference ature ere is evidence that fa ere isevidence that ecifically compensatory compensatory ecifically jk, Staaks Hardewig & sting or energeticallysting or Accepted Article feed for the first 21 daysof the study; or (2) a food-deprived (FD) group, in which fishwere = 13fish each control in treatment): (1)a group, which fedwere in fish two assigned of intoone treatments (n were changed week.waterrandomly Tanks each of being the 30% with daily thetanks from siphoned was Waste water. tap dechlorinated with filled were Tanks Wootton 1992), each containing aplastic plant and smal placed were fish recovery After individual 2012). leng of standard measurement later for photographed Diet Treatments throughout the experiment. a12h on were commercial troutFish using feed. fed were fish time this During experiments. beginning before cm) 30 x cm x40 cm (100 nets and minnow traps. Fish were held in the laboratory at 10 at thelaboratory in held were Fish traps. minnow and nets Animals Experimental METHODS AND MATERIALS how individualswill respond aspectsto of environmental change. better to us help will ectotherms in preference Increased intrinsic1992). influencetemperature knowledge ofextrinsic that factors the and Wootton & (Russell food-deprivation of period a following growth compensatory exhibit to observed previously been has and items, prey of abundance and quality the in variability seasonal experiences This article This protectedis by Allcopyright. rights reserved. Fish were netted from stock tanks then light Wild minnows were caught in December 2011 from the River Endrick in Scotland using dip- using Scotland in Endrick fromtheRiver 2011 December in caught were minnows Wild understand intraspecific variation in habitat use and and use habitat in variation intraspecific understand into 10-L tanks (one fish per tank; Russell & & Russell pertank; fish tanks (one 10-L into light: 12 h dark photoperiod during holding and and holding during photoperiod light:h 12 dark ly anaesthetised, weighed (±0.001 g), and and g), (±0.001 weighed anaesthetised, ly th (ImageJ, Rasband, Schneider, Eliceiri & l filter that also bubbled air into the water. o C for three weeks in two stock tanks ad libitum with commercial commercial with ad libitum

Accepted Article degree days in the current study compared to 16 days x 15 x days 16 to compared study current the in days degree buffer tank as required. required. as tank buffer within each temperature alterthe eitherreservoir to coils steel in through and tanks buffer the whileheater chilling the cooled was with reservoir rese heating The reservoir. cooling anda reservoir heating a were tanks buffer the and chambers choice the both to External one. cold the as other chambers choice the of One tank. buffer external atta was cm and 7 of depth a to water contained chamber choice Each shape. “dumbbell” a form cm) to (10 way a passage by edge one of centre the at joined diameter) cm (40 chambers choice plastic circular opaque two of consisted shuttle-box The temperature experienced by animal the controlled is by behaviour its & Steffensen (Petersen 2003). environmental the which in Denmark) Tjele, Systems, (Loligo apparatus ashuttlebox using fish Preference Temperature thestudy. of 95 and 42, phase), of thefasting end (atthe 21 days on photographed and measured anaesthetised, again were Fish phase”. “growth the as to referred 2007), Metcalfe Wootton this 1992). After point both fedtreatments were the effects of fasting on subsequent growth trajectories incommon minnows (21days x10 current in the used fasting of period the days), in x time temperature (water degreedays and in low ambient temperatures, whenperiods at fasting of models, figureof 1) > 0.05; p treatment, effect linear (general significant not was difference this treatments, the to assigned randomly were fish the when groups the between and mass length in difference initial small a was there Although phase”. “fasting the as to referred study, of the days 21 thefirst for food of deprived completely This article This protectedis by Allcopyright. rights reserved. At the end of the 95-day diet treatments, temp treatments, 95-day the diet of At theend . Temperate fish can readily withstand prolonged prolonged withstand can readily fish Temperate . ched by silicone tubing to its own dedicated dedicated own its to tubing bysilicone ched was designated as the warm chamber and the and chamber warm the as designated was study was close to that previously used to study tostudy used previously tothat close was study rvoir was supplied with an electric aquarium aquarium electric an with supplied was rvoir an external chilling unit. Water was pumped from from pumped was Water unit. chilling external an erature preference was tested in individual individual in tested was preference erature ad libitum o C = 240 degree days in Russell and for the next 74 days (Álvarez & & (Álvarez days 74 next the for o C = 210 C =210 Accepted Article maximum of24 temperature would increase in both side inboth wouldtemperature increase the chamber, warm the into moved fish a if chambers. Thus cold and thewarm between differential 9 box that had been set to the static mode, with the warm chamber at 11 at chamber warm with the mode, static the to beenset thathad box between trials, 0.0005). p =0.89, = r correlation Pearson individuals, (n=10 temperature acclimation same the at period week one temperatur preferred final for repeatability a high that temperature could regulate theenvironmental in theshuttle-box of chambers cold and warm the by2 decrease to begin would box chambe cold the to moved fish when a Conversely, with the mounted camera and associated software and camera mounted with the persecond) (once thefish of coordinates x-y the tracking by calculated was cm) in moved distance chamber but a 2 buta chamber “static”, When mode. or dynamic static a either Specifically,accordingly. temper the chambers thechoice in temperature the change to tanks reservoir the from flow adjusted and thefish of position the detected tank, the above mounted Germany) Obersulm, GmbH, Systems Development Imaging (uEye, camera a from display video via a which, Denmark) Tjele, Systems, controlledLoligo (Shuttlesoft, was software choice within by chamber eachultimately temperature controller temperature system Loligo (DAQ-M, acquisition data Denmark). The and Systems, Tjele, prob temperature probes. The line temperature would change within the shuttle-box depending on on depending the shuttle-box within change would This article This protectedis by Allcopyright. rights reserved. o C. The fish was then left overnight. The next morning, activity within the shuttle-box (total (total the shuttle-box within activity morning, next The overnight. left then was Thefish C. During each testing run, a single fish that had been fasted for 24 h was placed in the shuttle- the in placed h was for 24 been fasted had that fish a single run, testing each During The temperature within each of the choice chambers was continually monitored using in- using monitored continually was chambers choice the of each within temperature The o o C) but the warm chamber would always remain 2 remain always would chamber warm the but C) C differential between them. When the system was “dynamic” the temperature temperature was the system “dynamic” C differential When them. between the o C h ature within either choice chamber could be set to be either in in set tochambereither be could within be eitherchoice ature -1 (to a pre-set minimum of 3 minimum of pre-set a (to s of the shuttle-box at a rate of 2 of rate a at theshuttle-box of s es were connected to a computer-driven computer-driven a to connected were es there was a constant temperature in each choice ineach constant temperature there was a es es in common minnows using this systemover a response to changing temperatures, an individual individual an temperatures, changing to response r, the temperature in both sides of sides inboth theshuttle- temperature the r, between the hours of 08:00-10:00. At this point point Atthis 08:00-10:00. of hours the between the location of the fish, while maintaining a 2 a maintaining while fish, of the the location it experienced. Pilot experiments demonstrated demonstrated experiments Pilot it experienced. o o C warmer than the cold chamber. chamber. cold the than warmer C C). Thus, by alternating between between alternating by Thus, C). o C and the cold chamber at chamber at cold the C and o C h -1 (to a pre-set pre-set (toa o C Accepted Article k sucker white approximated using the relationship between between relationship the using approximated placed into individual cylindrical 110 ml PCV ml PCV 110 cylindrical individual into placed respirometer back controlrespirometer for was to empty left varies with body size. Aspecificvalue for and (min), readings temperature between time T following equation: second using the once was every estimated h.body temperature Core 8 thenext for temperature preferred its select to left was fish the and mode, dynamic the to switched was system the This article This protectedis by Allcopyright. rights reserved. Where T Where Killen each fish asoxygen rates of uptake using interm Scope Aerobic and Rate Metabolic of Measurement provided was food No day. thefollowing tested be to individual, another with replaced and tank individual to its returned then was fish the protocol fortemperatur chambers alternate into of fish is temperature preferred thefinal temperature, water the taking than simply rather body temperature, core estimating By individual. of each temperature preferred final the as taken was h 2 thefinal during experienced temperature core body mean The =3.69 * b = T etal. a +(T Following choicewere metabolic for estimated after), trials(4-7 temperature days rates b is core body temperature, T temperature, is body core m i –T 2012). Each day at approximately 10:00 h, three fish that had been fasted for 24 h were were h for24 fasted been had fish that three h, 10:00 atapproximately day Each 2012). -0.574 Catostomus commersoni Catostomus a )*e (eq. 2) - kt (eq. 1) a (Stevens &Fry1974): (Stevens is ambient water temperature, T temperature, water isambient k is not available for thecommon minnow, and so respirometers and left undisturbed overnight. One One overnight. undisturbed andleft respirometers k e “sampling”. Following the temperature selection temperature Following the “sampling”. e and mass ( during the period spent in shuttle-box. spent in the period the during ground bacterial oxygen consumption. For all all For consumption. oxygen bacterial ground k ittent stopped-flow respirometry (Steffensen 1989; 1989; (Steffensen respirometry stopped-flow ittent is the rate of change of core temperature, which which core temperature, of change of rate the is less likely influenced tobe bybrief excursions m ) for another freshwater teleost, the freshwater another ) for i is initial temperature, temperature, isinitial k was was t is is Accepted Article h 1 respirometers for approximately 22 h in total. Whole-animal standard metabolic rate (SMR; O mg in (SMR; rate metabolic standard Whole-animal total. hin 22 approximately for respirometers the in remained having day, following hthe 08:00 around at respirometer the removed from were h. Fish 19:00 at off hand 07:00 at on turn to set was Lighting water. aerated with be flushed of rate to indicate be analysed could content oxygen in decrease andthe sealed were respirometers the off, When off. or on switch would pump flush an automated 15-min Every tubing. gas-impermeable of circuit external an round and chamber respirometer a withinwith was each achieved mixing Water blind. plastic opaque an by lighting direct and disturbance external from shielded were and the maximal rate of oxygen uptake measured during this time was taken as MMR (in mg (in O MMR as taken was time this during measured uptake oxygen rateof maximal and the durin intervals 3-min in measured then were uptake cessation of swimming and closureof the respirometer was always less than Rates s. of oxygen15 immediatel were fish chasing, of min 3-5 within (Killen circular a in fish the chasing manually by exercise the bout of exhaustive anaerobic exercise (e.g. as as (e.g. exercise anaerobic of exhaustive bout the in several fish species, this method has been demonstrated to elicit rates of oxygen uptake equal to to equal uptake oxygen of rates toelicit demonstrated been method has this species, fish several in were located within an aerated, rectangul aerated, an within located were (PyroScien sensors associated and meter oxygen quantifie was content oxygen water measurements This article This protectedis by Allcopyright. rights reserved. metabolic rate(RMR; mgO dur chambers inthe confinement (Dupont-Prinet period measurement ).This method assumes that maximal rates of oxygen of maximal method).This rates assumesthat -1 ) was estimated as the lowest 10th 10th percenti asthelowest ) was estimated etal. At this were fromstage fish the removed 2012; Clark, Sandblom & Jutfelt 2013). After complete &Jutfelt 2013). Clark, Sandblom occurred exhaustion, which 2012; 2 h -1 ) was measured as the mean level of oxygen uptake during this time. time. this during uptake oxygen of mean level the as measured was ) ing which oxygen consumption was often elevated. Routineconsumption elevated. often was oxygen which ing etal. ar, temperature regulated water bath(10 water regulated temperature ar, 2010; Killen 2010; oxygen uptake. When open uptake. oxygen When le of measurements taken throughout takenthroughout the measurements ofle peristaltic pump that moved water through the y returned to the respir to the y returned ce GmbH,Germany). respirometersce The Aachen, tank (50 cm diameter) with a water depth of 10 cm cm 10 of depth water a with diameter) cm (50 tank exhaustive to subjected and respirometers stores of glycogen and ATP are replenished), and and replenished), ATPare and ofglycogen stores d once every 5 s using a Firesting 4-channel 4-channel Firesting a using s every 5 once d g a 15-min closed phase in the respirometers, closed in the a 15-min g phase uptake are achieved during the recovery from from recovery the during areachieved uptake etal. 2012), excluding thefirst 5hof ometers. Time between the ometers. the between Time , the respirometers would, therespirometers o C; 11.5 L) and 2 h - 2

Accepted Article the dependent variable, treatment as a categorical variable, and either mass (for (for mass either and variable, categorical a as treatment variable, dependent the length ( length ( and and G included as a covariate to correct for the effects theeffects for correct to covariate a as included treatments using modelslinear general with trea factor. arepeated as period measurement and factor random line a using accommodated were measures repeated were log-transformed.were (Cleasby &Nakagawa 2011). use For in models, measures of SMR, RMR, MMR, mass,AS, and activity treatments diet between variances of homoscedasticity to paid attention particular with plots, of residua and homoscedasticity linearity, normality, (Systat Software Inc.). of The level significance for all tests was Analyses Statistical and Data 2013). Jutfelt & Sandblom (Reidy swimming sustained during observed that than or greater This article This protectedis by Allcopyright. rights reserved. Where s Where compared during the starvation phase using general linear models with growth (either (either growth with models linear general using phase starvation the during compared =100* (ln( =100* d is the time elapsed in days (Hopkins 1992). Growth time elapsed 1992). days(Hopkins were in control ratesof treatments is the and FD G G Measures of SMR, MMR, AS, FAS, and activity level were compared between control and FD FD and control between compared were level activity and FAS, AS, MMR, ofSMR, Measures Rates Rates of mass loss or growth were estimatedof in terms body mass ( Statistics were performedwith SPSSstatistics v20.0 Inc. (SPSS and IBM)and SigmaPlot v11.0 t is the body mass or standard length at time at length or standard mass body the is L L ) at the beginning of each measurement period as a covariate. During the growth phase, growth phase, the During covariate. a as period measurement each of beginning the ) at equation: the to according periods measurement between calculated were and ), s t ) – ln( s i ))/ d (eq. 4) of body size. Changes in body temperature during during temperature body in Changes size. body of tment as acategorical variable.Mass was also ls were verified by inspection of residual-fit residual-fit of inspection verifiedby were ls ar mixed model with fish identity added as a a as added identity fish with model ar mixed t , s i is the initial body mass or standard length length or standard mass body theinitial is α = 0.05. When required the =0.05.Whenrequired etal. 1995; Killen 1995; G M ) and standard standard and ) G etal. M ) or standard or standard ) G 2007;Clark, M or G L ) as Accepted Article treatment effect on effect treatment of beginning the at size body in differences for accounting after controls than rapidly more length and mass gained FD fish phase growth the Effects of Food Deprivation on Growth Rate Growth on Deprivation ofFood Effects RESULTS linear model, effect of treatment on treatment of linear model, effect significant interactions were sequent were interactions significant each between interactions as well as variables,

categorical variable, and SMR, AS, AS, SMR, and variable, categorical a as treatment with temperature, preferred thefinal on variables of these effects combined the examine to used were correlations. with models Pearson linear General explored initiallywere and Rohlf 2003). The inclusion of random slopes accounted for any individual differences in changes changes in differences individual for any slopes accounted random of inclusion The 2003). and Rohlf t body mean in change the of nature curvilinear time, and diet and time and diet time, and with time. temperature in preferred a repeated factor. A factor. a repeated time as and factor random a as identity fish covariate, a mass as variable, acategorical as treatment diet with slopes and intercepts random with model mixed alinear using treatments between in min 30 each at taken (means trials shuttle-box This article This protectedis by Allcopyright. rights reserved. During the food-deprivation phase thefood-deprivation During Effects of SMR, AS, AS, ofSMR, Effects G quadratic term(time quadratic L : F 2 . 1,25 1,25 G = 4.322, p = 0.044; 4.322, = = p L during the growth phase, and activity on final preferred temperature temperature preferred final on activity and phase, thegrowth during G L Initial models also included the interactions between diet and and diet between interactions the included also models Initial G during the growth phase, activity, and mass as continuous continuous as mass and activity, phase, growth the during ially dropped and the models re-run. re-run. models the and dropped ially L : F 1,25 2 ) was also added to the model to account for the for account to model the to added also was ) treatments differed in both both in differed treatments each measurement period (linear mixed models,mixed period (linear each measurement = 18.638,p <0.001; factor and diet treatment. In all models, non- models, all In treatment. and diet factor G emperature with time during each interval (Sokal interval (Sokal each during time with emperature M tervals throughout the h 8 trial) were compared : F 1,25 =11.199, p<0.002). G M : F 1,25 1,25 G L = 85.549, 0.001). During p < and G M (figure 1; general (figure Accepted Article = 0.005). While values showed for RMR an opposite trend, with RMRbeing 19.1% highercontrol in linear model, linear model, F mass (general for correcting after significant not was difference this fish, FD to compared as fish degrees lower than that the of control group (16.79±1.16 model, effect of treatment: F ofmodel, treatment: effect movingtemperatures, on 59.97% average thetotal of static two the at held when apparatus shuttle-box the in fish control than active less much were fish linear model, linear model, F (general fish FD for than higher 19.8% average on were that forAS values had fish control However, with the mean final preferred core body temperature of the FD group (13.94±1.27 group theFD of temperature body core preferred mean final the with of FD fish, but this difference was also not significant (general was F linear model, also significant not this (general difference of but fish, FD effect ofeffect time: F (5.25±0.64) as compared controls to (8.08±0.71; general linear model, F toward temperatures warmer than the10 warmer than toward temperatures Preference Temperature p=0.0004). -0.646, = r correlation, Pearson s2; (figure individuals among andAS SMR either SMR ormodels, AS(general linear > 0.05) p temperatures than controls (linear mixed model, effect of model, treatment: F controls effect mixed than (linear temperatures individual variation in body mass (figure 2; general linear model, effect of treatment: F treatment: of effect model, linear general 2; (figure mass body in variation individual and group control in the fish of thanthat higher Phenotype Metabolic on Diet of Effects This article This protectedis by Allcopyright. rights reserved. During the 8 h dynamic temperature selection peri selection temperature dynamic h the8 During At the end of the 95-day study the whole-animal SMR of FD fish was on average 51.9% average on was FDfish 95-day of thewhole-animal SMR the of study At theend 1,25 1,25 1,25 = = 0.778,p =0.384). Whole-animal of MMR controlwas fish higher 8.08% than that = 5.416 p =0.029). Factorial scope was also significantly lower FDfishin = 11.184,= However, p=0.003). throughout this time FD fish preferred colder 1,25 = 14.753, p=0.001). Among individuals, activitywas not related to o C acclimation temperature (figure 3; linear mixed model, model, mixed linear 3; (figure temperature C acclimation remained significantly elevated after accounting for for accounting after elevated significantly remained . Overall, there was a .correlation therewas Overall, negative between a distance moved by control fish (general linear linear (general fish control by moved distance o C). Interactions C). diettreatment between od fish in both treatments tended to shift treatments in both fish od 1,345.93 1,25 = 4.923, p=0.037).FD = 11.306,= 0.001), p = 1,25 =3.021,p 0.096). = o C) being two over 1,25 1,25 = 9.697, p Accepted Article activity: F prefer to tended active more were that Individuals cooler temperatures (figure 4; table 1; (figure4; cooler temperatures table preferring SMR ahigher had that individuals with SMR, by influenced was preference temperature treatment x treatment time DISCUSSION time time and of and both This article This protectedis by Allcopyright. rights reserved. to simply massrestoring lost (Nicieza 2009) &Álvarez experiencing were they indicating mass, as well as growth. compensatory as such by factors affected being plastic, indi of preferences thermal the demand influences preference. thermal influencing for be important to revealed were level andactivity SMR only model, same the in included were variables explanatory cooler a relatively temper SMRpreferred high feedin normal of the resumption several weeks after compensatory Thuswhile growth compensa on SMR. of effect the is this link of cause underlying the but temperatures, cooler toprefer tended significant effect on temperature preference (table 1; figure 4). 1;figure 4). (table preference temperature on significant effect into weretaken andactivity onceSMR temperature did it 3), (figures independently examined when When all potential explanatory variables were combined into a single model, final final model, single combinedvariables intoa potentialexplanatory allWhen were During the growth phase FD individuals displayed increased growth rate in terms of length length of terms ratein growth increased displayed individuals FD phase thegrowth During growth compensatory subsequent and deprivation food earlier experienced that Individuals 1,25 =4.469, 0.049). p = Although diet treat 2 interactions, p> 0.05) and were not included in the final model. 2 were not significant (linear mixed model, treatment x time time and x treatment model, mixed (linear significant not were general linear model, effect of SMR: F model,of SMR: effectgenerallinear atures. As a consequence, when all potential all potential when As atures. a consequence, not have a significant effect on final preferred preferred on final effect asignificant have not Overall these results demonstrate that metabolic metabolic that demonstrate results these Overall increased rates of structural growth as opposed opposed as growth ofstructural rates increased vidual fish, and that preferred temperature is temperature preferred that and fish, vidual warmer temperatures (figure 4; table 1; effect of of 4;table1; effect (figure temperatures warmer ment was linked to temperature preference preference to temperature was linked ment account (table 1). Aerobic scope did not have a have not did scope Aerobic 1). (table account g, even individuals in the control treatment withcontrol treatment the in evenindividuals g, tory growth resulted in an elevation of SMR elevation SMR of growth an tory resultedin . It is likely that fasted fish were undergoing were . fasted fish likely Itis that 1,25 = 9.076, p=0.007). Accepted Article they were observed to have an increased SMR as compared to the control individuals. This finding is is finding This individuals. control the to compared as SMR increased an have to observed were they For FDtreatment,effect ofthe this SMRon th growth efficiency decreasing by baseline maintenance costs (Ware 1982; Mehner &Wieser1994). 1992). Itazawa & Takemori (Oikawa, individuals compensated relativethe metabolically sizesof org active change and organ sizes of theallometry affect can maintenance metabolism (Houlihan 1991). Itmay also be possible that altered growth trajectories lead to increased rates of protein turnover or quality tissue Decreased 2008). etal. (Criscuolo compensatory of earlierperiod an metabolic rateof freel became food after manyweeks of SMR levels elevated relatively had growth compensatory experienced had that fish study, current Inthe 1992). Wootton & (Russell deprivation food of period prolonged a after minnows common in to occur of phase thegrowth during growth compensatory This article This protectedis by Allcopyright. rights reserved. 10 at that, observation the is hypothesis this for support Added growth. including functions, physiological additional for available the energy from detract necessarily costs maintenance increased which in (2008), etal. Careau by proposed budgeting of energy model theallocation with compatible routinely operate near MMR experience an incr an experience MMR near operate routinely (masking and limiting factors 1971;in e.g. Fry disease, environmental stressors).Animals that incr which arise may that conditions with coping SMR. lower a with those to compared tasks physiological oxygen-demanding among trade-offs corre compensate with a do not relativelySMR high o C, there was a negative correlation between SMR and AS. This indicates that individuals with a a with individuals that indicates AS. This and SMR between correlation negative a was C, there By selecting a cooler temperature, individuals with a relatively high SMR could increase could increase SMR high relatively a with individuals temperature, cooler a By selecting A major additional benefit of a reduced SMR, however, is an increase in surplus AS for AS surplus in increase is an however, SMR, reduced of a benefit additional A major ans and tissues (e.g. liver, muscle) in growthin muscle) liver, (e.g. tissues and ans repair (Johnston 1999), so elevating of rates ermal preference may have been exaggerated since since exaggerated been have may ermal preference eased likelihood of mortality (Wood, Turner & & Turner (Wood, mortality of likelihood eased stemming from rapid growth could subsequently subsequently could growth rapid from stemming ease baselineease metabolic costs that or limit MMR s in body composition that cause differences in in differences cause that composition body in s the study, as has been previously demonstrated demonstrated been previously has as study, the y available; asimilar long-lasting effect on spondingly high MMR, and thus will face greater greater face will thus and MMR, high spondingly growth has been previously noted in other taxa taxa other in noted previously been has growth Accepted Article chose to occupy a wide range of temperatures (~6-23 temperatures of wide range occupy a chose to increased statistical powerwould reveal such an effect, but even in this case, the effect size of SMR study detected no significant effect main of AS may be optimisationthe 1971;Pörtner (Fry 2008;of AS Farrell & Jørgensen stressors. environmental with deal to capacity the increasing the and growth accommodating better temperatures, warmer ata available that activity as such costs, energetic other by modulating relative thermal sensitivities ofand SMR MMRover temperature the range in question). However, should therefore remain unchanged or more likely AS being total equal, 2008). All factors Pörtner & Farrell other 2007; Lefrancois (Claireaux & and SMR below which that optimises further AS, in a decrease temperature will cause drop in MMR both a maintaining surplus of availablea AS.Importantly,maximum a of surplus while growth in engage to animal an allow could – activity to due that as –such costs energetic Bryan, Kelsch &Neill 1990). Decreasing (by selectingcoolerSMR waters) and also reducing other 1985; Priede 1982; (Ware digestion and activity of costs the for accounting after AS surplus increase Graham 1983; Priede 1985), and there should be st This article This protectedis by Allcopyright. rights reserved. attained below the theoreti below the attained studywere measured formetabolic traits at current the in fish that note to is important It activity. decreasing by costs maintenance increased activity) equal was twobetween treatments.the Th spontaneous of costs the includes (which RMR residual fish, control and FD between different was SMR though even Indeed, available. freely became food after weeks several fish FD among observed accommodating limiting and masking factors (Fry 1971). This likely explains reduced the activity thus activity, and digestion of costs associated the and foraging increased necessitates temperatures athigher SMR elevated is because This It has been hypothesised that a primary driver for habitat selection in aquatic ectotherms ectotherms aquatic in selection habitat for driver aprimary that hypothesised been has It cal optimum temperature for optimum for cal temperature a constant intermediate temperature(10 intermediate constant a on temperature preference. It is possible that possible that It preference. is temperature on diminishing the surplus AS actually available for for available actually AS thesurplus diminishing decrease as temperatures cool (depending on the on cool astemperatures (depending decrease rong selection toprefer environments which is suggests that FD fish effectively offset offset suggests fisheffectively thatFD is , individuals could increase surplus AS relative to to relative AS surplus increase could individuals , o C). Assuming the original temperature is at or ator is temperature theoriginal Assuming C). total AS ( sensu Portner and Portner and 2008).Farrell available et al. 2012). current The AS could be could AS o C), while they they while C), Accepted Article optimum (Clark, Sandblom & Jutfelt 2013). Alternatively, some individuals may simply minimise SMR SMR minimise simply may individuals some Alternatively, 2013). Jutfelt & Sandblom (Clark, optimum thermal own its having function with each function), immune performance, swim (e.g. functions different prioritise individuals different that is possibility exclusive non-mutually a costs, energetic 2009; Schram and within species do not always coincide with optima the specificfor physiological traits (Angilletta activity and digestion accounted are will for temperatures).(which vary across that which maximises surplus ASactually available after coststhe of required processes such as with overlaps preference temperature individual which to extent the be determining will research future for area Akey populations. within AS and SMR as such traits for repeatability of stability thethermal examine andto etal. 2013), (Norin regimes thermal in variation and temporal spatial to exposed are and environment the through move individuals as shift dynamically AS and SMR how measuring of tasks challenging the undertake relative ranking of metabolic traits when exposed to the altering thus sensitivity, thermal in vary may individuals that possible Itis temperature. single cu the in that fish is consideration important an Again, traits. specific of optimisation tothe opposed as thermoregulation, behavioural during on thermal preference exceeds that of that AS(table on preference exceeds thermal This article This protectedis by Allcopyright. rights reserved. of size, G and 1990), and so cooler temperatures may still be preferred when possible. Indeed, Van Djik et al. etal. Djik Van Indeed, possible. when bepreferred may still cooler temperatures andso 1990), couldconstrain thecope ability with environmental to stressors (Priede 1985; Bryan, Kelsch Neill & activity) reducing by (e.g. scope aerobic surplus the balance to mechanisms countering without maximum growth rateis positivelyoften correla is growthin fishes rate correlati apositive be theremay circumstances, in ex engaging of actively preference individuals At the time of measurement for SMR and AS, FD fi and FD AS, SMR for measurement At thetimeof L et al. was not elevated at time.this at not Additional elevated was 2013). In addition to increasing surplus AS by minimising SMR and other SMRand ASbyminimising increasing to surplus In addition 2013). achieved; Jobling 1995). Even inthis scenario,however,maximising growth rrent study were measured for SMR and MMR at a at MMR and SMR for were measured rrent study ted with temperature (until an optimum at which optimum at an (until temperature with ted on between SMR and preferred temperature, as as temperature, andpreferred SMR between on Under these compensatoryor growth. rapid tremely 1).Interestingly, pref the temperature thatopti temperature the different temperatures. More More to is work temperatures. needed different research is needed to research needed is sh had “caught up” to control fish in terms terms fish in control to up” had “caught sh erred temperatures among among erred temperatures mises total AS, versus versus AS, total mises examine thermal the Accepted Article environment, then at least a portion of this activitymay have also been due to exploration, which is (Careau species same the of individuals prev has Activity preference. temperature influence also tendencies behavioural in variation traits, physiological in variation with conjunction in that, that these personality traits can be plastic in response to factors such as growth trajectory and and trajectory growth such as factors to response in plastic be can traits personality these that (Dingemanse personality animal of dimension a considered also while same atenvironmental timeany the reducewarming themay availability of preferred comp of occurrence the increase could phenomenon fluctuations in food availability freshwater temperate in habitats (Winder &Schindler This 2004). microclimates will lessbe suitable. Future chan perturbations earlier However, if of foraging. rates and activity increased facilitate these since AS), or high SMR low a with those (i.e. them occupy to afford can that individuals ectothermic for beneficial be can microclimates warm Relatively st are links these SMR, in variation by underpinned functions. oxygen-consuming additional for AS of available asurplus and efficiency growth increasing for strategy a is likely even fish, individual in metabolism maintenance co studypresent the of results The evenwhiledisplayi cooler temperatures preferred roach juvenile starvation, of period a following immediately that observed (2002) This article This protectedis by Allcopyright. rights reserved. predator avoidance (Álvarez &Metcalfe 2007). ecology behavioural the for implications profound microhabitats. Further, effects on activity leve Fish that were more activeat 10 are preference temperature and AS, history, dietary between relationships Although nfirm that this preference for cooler temperatures is related to is related cooler temperatures for this nfirm preference that et al. et o C tended to prefer warmer temperatures, and it is possible possible is it and temperatures, warmer toprefer C tended 2008), and if shuttle-boxthe treated is as novela to growth cause an increase in SMR then these to growth thenthese incauseincrease an SMR ls stemming from compensatory growth could have could growth compensatory from stemming ls ges in climate are predicted to cause increasedcause predictedclimate to are ges in in those not undergoing compensatory growth, and and growth, compensatory undergoing not those in ng evidence of a compen evidence a of ng of fish, possibly affecting foraging abilityor ill likely to be of key ecological importance. importance. key ecological be of to likely ill iously been noted as an intrinsic trait within within trait intrinsic an notedas been iously ensatory growth among freshwater ectotherms ensatory among growth freshwater etal. 2002). The results here show satory growth response. Rutilus rutilus

Accepted Article http://doi.org/10.5061/dryad.5h2sd Repository: Digital Dryad the from available Data ACCESSIBILITY DATA availability. food and risk predator as such wild, the in variables environmental for future research will be to examine how such sh avenue additional An metabolism. aerobic for scope the reduces which SMR, of elevation an through mediated are effects cooler thatthese toprefer and temperatures, compensatorytend growth the allocation modelof energy budgeting (Careau for support additional provides instead study current the in activity and SMR between association etal. Careau 2008; of increased models intake musclemass, mitochondrial density) will need tobe maintained productioneven rest(the at or to needed morphology the because rates, metabolic resting increased possess mayalso individuals active more that been hypothesised previously has it Interestingly, environments. in these SMR ahigher support to berequired would that foraging may be willingoccupymore to warmer areas accept theand risks associated with increased of rates positively correlated among individuals (Réale individuals among correlated positively be may also boldness and Activity warmertemperatures. preferred lizards males aggressive active more are that (those scope maximise aerobic or lifestyle active amore facilitate that habitats warmer relatively or individuals) active (less expenditure energy minimise that habitats cooler occupy either to opting selection, to temperature comes it when strategies differing have to individuals cause may types personality differing traits, metabolic to addition in However, 1993). Wieser & (Méndez history dietary This article This protectedis by Allcopyright. rights reserved. In summary, the results of this study suggest that fish that have gone through a period of of period a through gone have that fish that suggest study this of results the summary, In ). In support of this view, Stapley (2006) observed that more more that observed (2006) Stapley view, this of support In ). et al. et Careau and Garland 2012). However, the lack of an However,of 2012). Garland thelack and Careau et al. et ifts in temperature preference interact with other other interact with preference temperature ifts in 2010), and so bolder and more activeindividuals support high levels of activity (e.g. increased increased (e.g. activity of levels high support 2008). 2008). Accepted Article Claireaux,G. &Lefrancois, C. (2007) Linking environmental variability and fish performance: Careau, V.,Thomas, Humphries,D., M.M. & Réale, D.(2008) Energy metabolism and animal Careau,Garland, V.& (2012)Performance, T., Jr. personality, and energetics: correlation,causation, Burton, T., Killen, Armstrong,S.S., J.D. Metcalfe, & What(2011) N.B. causes intraspecific variation in Bryan, J.D., Kelsch,Neill, & The S.W. W.H.Maximum (1990) Power Principle Behavioral in Brown,J.H., Marquet,& Taper,P.A. Evolution M.L. (1993) of Body Size:Consequences of an Blouin-Demers,P.J.G., & Kissner, K.J., Weatherhead, Sensitivity of Thermal in Interindividual Variation (2013) J.-F.L. & Galliard, Jouanneau, I. P., Artacho, Angilletta, M.J., Niewiarowski, P.H. &Navas, evolution The C.A.(2002) of thermal physiology in Angilletta, Thermal M.J. Adaptation: (2009) trade The (2007) N.B. Metcalfe, & D. Álvarez, LITERATURE CITED guidelines approvedwithin the UK. care animal with comply paper this in described procedures The maintenance. fish with assistance for Ryan andMary Kirk, Alistair Law, Graham to also Thanks manuscript. this of versions on earlier feedback constructive for reviewers anonymous two thank also I study. this of results the regarding Guy Claireaux and Holt, Rebecca Jorgensen, Christian with conversations theinformative appreciate greatly also I collection. fish with assisted also Metcalfe Prof. appreciated. much very were Marras Stefano and Dr. Metcalfe Neil Prof. by manuscript this of version earlier an on Comments research fellowship. (NERC) Council Research Environmental Natural UK a by funded was study This ACKNOWLEDGEMENTS This article This protectedis by Allcopyright. rights reserved. personality. personality. mechanism. and Sciences, Society B: Biological consequences? ecological its are what and rate metabolic resting Fishes. Thermoregulation by Energetic Definition Fitness. of 2000, Development. during Temperature to Response in Snakes Young of Temperature Society B: Sciences, Society B: Biological activity. for concept scope of integration the through Physiological and Biochemical Zoology, Biochemical and Physiological Maximal Sprint Speed, Thermal Behavior, RestingMetabolicand Lizard. in Rate a ectotherms. Oxford. Press, variation between habitats in the cost of compensation. compensation. of cost the in habitats between variation 841-845. Oikos, Journal of Thermal Biology, Physiological and Biochemical Zoology, Biochemical and Physiological

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Accepted Article Killen, S.S., Atkinson, D. & Glazier, D.S. (2010) The intraspecific scaling of metabolic rate with body body with rate metabolic of scaling Theintraspecific (2010) D.S. Glazier, & D. Atkinson, S.S., Killen, Jobling, M.(1995) Humphries, M.M. & McCann, Metabolic (2014) K.S. ecology. Huey, R.B.&Kingsolver, Evolution J.G. (1989) of Killen, S.S. (2014) Growth trajectory influences temperature preference in fish through an effect on on aneffect through fish in preference temperature influences trajectory Growth (2014) S.S. Killen, Killen, S.S., Marras, Steffensen, S., McKenzie, J.F. & D.J. Aerobic(2012) capacity influences the spatial Khan, J.R. & Herbert, N.A. (2012) The behaviour The (2012) N.A. Herbert, & J.R. Khan, The (2009) W.P. Porter, & R. Shine, M., Kearney, Antognarelli, Azzurr F., M.A., Peck, Jørgensen, C., Fry, F.E.J. EffectThe (1971) of Environmental Factors on thePhysiology of Fish. Killen, S.S., Marras, S., Metcalfe, N.B., McKenzie, D.J. & Domenici, P. (2013) Environmental stressors stressors Environmental (2013) P. Domenici, & D.J. McKenzie, N.B., Metcalfe, S., Marras, S.S., Killen, Criscuolo, F., Monaghan, P., Nasir, L. & Metcalfe, N.B. (2008) Early nutrition and phenotypic phenotypic and nutrition Early (2008) N.B. Metcalfe, & L. Nasir, P., Monaghan, F., Criscuolo, Fangue, N.A., Podrabsky, J.E., Crawshaw, L.I. & Grima, Vande B., L., A., Chatain, Dupont-Prinet, and Repeatability (2002) A.J. Noordwijk, van & K. Oers, van P.J., Drent, C., Both, N.J., Dingemanse, Killen, I.,Gamperl, Costa, S.S., A.K.metabolic J.A. thetank: in & Brown, Little leftin scaling (2007) Clusella Trullas, S., Terblanche, &Spotila,J. J., (2007) Wyk, J.Low repeatability preferred of body theresiduals. in patterns biological Neglected (2011) S. Nakagawa, & I. Cleasby, of an era in fishes of measurements scope Aerobic (2013) F. Jutfelt, E. & Sandblom, T.D., Clark, This article This protectedis by Allcopyright. rights reserved. distribution. distribution. its of limit thermal the upper optima at scope aerobic tracks lapillum) (Forsterygion Evolution, & Ecology in Trends metabolic rate. Dryad Digital Repository doi:10.5061/dryad.5h2sd doi:10.5061/dryad.5h2sd Repository Digital Dryad rate. metabolic Sciences, schools. fish within of individuals position 651-658. Sciences, warming. climate against animals “cold-blooded” predictivefishes: models. capacity of the advancing Domenici, & J., L.R. M., Teal, Steffensen, Fleng Holt, R.E., Huebert, K.B.,Marras, S., McKenzie, D., Metcalfe, J., Perez-Ruzafa, A.,Sinerchia, 1-98. Zoology, Biochemical Populations in Preference Temperature alter relationships between physiologybehaviour. and deprivation in the European sea bass ( bass sea theEuropean in deprivation atrade-off underlying mechanisms Physiological 938. the wild. from tits great in behaviour exploratory of heritability Sciences, Biological B: Society Royal of the adulthood. in rate metabolic elevated to leads growth ‘catch-up’ development: Biological Sciences, Biological scope. aerobic for implications and its teleosts marine andtemperature. on lifestyle depends fishes in mass adaptive significance. significance. adaptive temperature in four species of Cordylid lizards: Temporal variation and implications for and Sociobiology, Biology, recommendations. and relevance respirometry, change: climate

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of Killifish Fundulus heteroclitus. heteroclitus. Fundulus Killifish of 37, potential for behavioral thermoregulation to buffer buffer to thermoregulation behavioral for potential o, E., Burrows, M.T., Cheung, W.W.L., Cucco, A.,

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The Journal Journal Accepted Article acclimated to 10 to acclimated Table 1. of food deprivation in the food-deprived treatment. duration the represent bars shaded Horizontal point. data previous the since plotted, are they which on data the until up rates growth represent points data D, Band panels In circles). treatment; dark deprived for 21 days before being fed either fed Wood, C.D., Turner, J.D. & Graham, M.S. (1983) Why do fish die after severe exercise? exercise? severe after die Why do fish (1983) M.S. Graham, & J.D. Turner, C.D., Wood, Ware, D.M.Power (1982) and Evolutionary Fitness ofTeleosts. scope; G activity AS 1 1 66.956 22.417 SMR 4.469 1 1.632 diet 124.698 9.076 1 0.049 0.218 16.449 Term 0.007 3.771 -0.015 df -21.785 1.197 mean a treatment diet between two-way interactions 2.621 8.781 0.288 7.487 square -2.522 F p parameter 2.192 s.e.m growth rate growth( rate FIGURE 1. error 18 mass 11.291 1 21.645 1.575 0.225 5.933 4.845 Wallman, H. & Bennett, W. (2006) Effects of Parturition and Feeding on Thermal Preference of Preference Thermal on Feeding and Parturition of Effects (2006) W. Bennett, H. & Wallman, This article This protectedis by Allcopyright. rights reserved. L 1 2.632 0.192 0.667 -11.858 27.092 G General linear results model for effects on Biology, Aquatic Sciences, Aquatic Atlantic Stingray, Dasyatis sabina (Lesueur). (Lesueur). sabina Dasyatis Stingray, Atlantic L = growth rate during the growth phase (days 21-95 of the current study). Non-significant Non-significant study). current ofthe 21-95 (days phase thegrowth during rate growth = commonGrowth trajectories in minnows. Chan ad libitum G M ); C) standard length; and D) length-specific growth rate ( rate growth length-specific D) and length; C)standard );

22, o C, n = 13 per diet treatment. SMR = standard metabolic rate; AS = absolute aerobic aerobic absolute = AS rate; metabolic standard = SMR treatment. diet per 13 = n C, 189-201. throughout the entire 95 day study study entire95day (contr throughout the

39, 3-13. 3-13. ad libitum ad nd other factors weredr factors nd other for the remainder of the study (food-deprived (food-deprived study the of remainder the for preferred temperature in common minnows common in minnows temperature preferred Environmental Biology Fishes, of Biology Environmental ges total in: wet A) mass-specificmass; B) ol treatment; open circles), or food Canadian Journal of Fisheries and opped from the model. model. the from opped G L ) in common minnows ) in

75, Journal of Fish Fish of Journal 259-267. Accepted Article Results for details). SMR = standard metabolic rate; RMR = routine metabolic rate; MMR = maximal =maximal rate; MMR metabolic =routine RMR rate; metabolic standard = SMR fordetails). Results food-deprived control and betweenthe difference asignificant *indicates boxes). dark treatment; (food-deprived study the of remainder the for treatment; fi grey boxes), food deprived while for raw per animal data (mg O (mg data animal per raw for s1 figure see size; body for correction after residuals are Values scope. =aerobic rate;AS metabolic before being fed days 21 for deprived orfood bars), open treatment; (control study day 95 entire the throughout interv min 30 a given during mean temperature temperature. preferred their select could individuals FIGURE 3. deprivation. Contro deprivation. FIGURE 2. This article This protectedis by Allcopyright. rights reserved. common minnows fed either common either minnows fed in C)activity, and (AS); scope B)aerobic (SMR); rate metabolic A)standard traits. and behavioural FIGURE 4. s.e.m. Error = bars temperature. temperature. no was There 1. Table in described temperature preferred final on C) (panel andactivity A) (panel SMR of effects main thesignificant represent point circles). dark (food-deprivedEach treatment; deprived or food circles), Mean body temperatures for common minnows held in a shuttle-box apparatus in in apparatus shuttle-box a in held minnows forcommon temperatures body Mean Metabolic traits for common minnows a cont fed forcommon minnows traits Metabolic Relationships between preferred body temp body preferred between Relationships ad libitum ad l minnows were fed were minnows l for days21 before being fed for the remainder of the study (food-deprived treatment; dark bars). dark bars). treatment; (food-deprived study of the remainder the for ad libitum ad 2 h -1 ) throughout the entire 95 day study (control treatment; open treatment; (control open day entire95 study the throughout ad libitum sh were fasted for 21 days before being fed being before days for 21 fasted were sh al. Treatments were either fed either either eitherfed were al. Treatments effect of aerobic scope (panel B) on preferred on preferred B) (panel scope aerobic of effect throughout the entire 95 day study (control (control study day 95 theentire throughout treatments (general linea treatments represents one individual. Solid regression lines lines regression Solid individual. one represents Values at each point were calculated as the the as were calculated point each at Values ad libitum ad erature in individual minnows and metabolic metabolic and minnows individual in erature rol diet or that experienced earlier food- earlier thatexperienced or diet rol for the remainder of the study of remainder the for the r models, p < 0.05, see 0.05, < models,r p ad libitum ad libitum

Accepted Article This article This protectedis by Allcopyright. rights reserved.