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Reproduction in Domestic Animals

Reproduction in Domestic Animals

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12 D Blache, GB Mart<strong>in</strong> and SK MaloneyVoluntary feed<strong>in</strong>takeMetabolic fuelEnergy reserveEmotional reactivityHPA axisMetabolicHormones:lept<strong>in</strong>, orex<strong>in</strong>s<strong>in</strong>sul<strong>in</strong>,IGF-1, GH,T3, T4ThermogenesisImmune functionCellular ma<strong>in</strong>tenanceGrowth, locomotion<strong>Reproduction</strong>Fig. 3. Central role of metabolic hormones <strong>in</strong> the <strong>in</strong>teractions amongemotion, reproductive function, immune function, metabolic status,thermogenesis and other bodily functions. The sympathetic pathwayshave been omitted for clarityand limit the risk of conflict<strong>in</strong>g results s<strong>in</strong>ce, bydef<strong>in</strong>ition, a group of animals show<strong>in</strong>g a homogenousresponse to any stimulus will have less variability <strong>in</strong> anyparameter associated with that response.As described previously, the amount of availablemetabolic fuel needs to be ‘assessed’ by a control systemand then expenditure ‘directed’ towards reproductivefunction if and when required. For all systems, theassessment of energy levels and their partition<strong>in</strong>g isunder the control of metabolic hormones (Fig. 3).Thyroid concentrations were higher <strong>in</strong> a sub-populationof ‘calm’ than <strong>in</strong> a sub-population of ‘nervous’ animals<strong>in</strong> our selection l<strong>in</strong>es (Blache et al. 2002). We have alsoshown that the cortisol response to a novel object ishigher <strong>in</strong> nervous ewes than <strong>in</strong> calm ewes (Bickell et al.2008). These considerations apply equally to stressassociated with experimental protocols (Mart<strong>in</strong> et al.1981; Adams et al. 1993). The effects of stress andcortisol on the reproductive axis are numerous anddiverse – <strong>in</strong> the female, they can either <strong>in</strong>hibit orstimulate ovulation (Dobson and Smith 2000). Theeffect of temperament genetics on reproductive processeslikely <strong>in</strong>volves hormonal pathways. These l<strong>in</strong>ksare still to be demonstrated experimentally, but temperamentaffects the hormonal systems <strong>in</strong>volved <strong>in</strong> thecontrol of energy partition<strong>in</strong>g and this may be how it<strong>in</strong>fluences the outcomes of experimentation target<strong>in</strong>g therelationship between nutrition and reproduction.Aside from improv<strong>in</strong>g the experimental design andoutcomes, the welfare of experimental animals could beimproved by selection for temperament. Less reactiveanimals will have an improved freedom from fear anddistress s<strong>in</strong>ce animals that are less reactive to novelty willpresent reduced physiological and behavioural reactionsto novel experiences (Korte 2001), <strong>in</strong>clud<strong>in</strong>g exposure toa research environment.ConclusionIn <strong>in</strong>ference science, hypothesis test<strong>in</strong>g <strong>in</strong>volves manipulat<strong>in</strong>gone or a few parameters, while other factors thatcould <strong>in</strong>fluence the dependent parameters are controlled.In this paper, we have shown that multiplefactors that affect and control reproduction could affectthe outcomes of animal experimentation because theyaffect reproductive physiology, over both short and longtime scales, and even across generations. These factorsneed to be controlled. In addition, emotional reactivityhas the potential to alter experimental outcomes, just asgenotype can affect the outcomes of experimentation <strong>in</strong>which photoperiod or nutrition is manipulated. Byconsider<strong>in</strong>g the nutritional history and emotional characteristicsof the <strong>in</strong>dividual subjects, the variabilitywith<strong>in</strong> experimental groups can be reduced and two ofthe 3Rs (reduction, ref<strong>in</strong>ement) would be applied,lead<strong>in</strong>g to more ethical experimentation. This creates aw<strong>in</strong>–w<strong>in</strong> situation for researchers and experimentalanimals because more ethical science results <strong>in</strong> betterscience.AcknowledgementsWe would like to thank the generous assistance of the students andstaff of Animal Science (University of WA). Fund<strong>in</strong>g was supplied bythe National Health and Medical Research Council, the AustralianResearch Council, the Ada Bartholomew Medical Research Trust, andthe University of Western Australia.ReferencesAdams NR, Atk<strong>in</strong>son S, Mart<strong>in</strong> GB, Briegel JR, Boukhliq R,Sanders MR, 1993: Frequent blood sampl<strong>in</strong>g changes theplasma concentration of LH and FSH and the ovulationrate <strong>in</strong> Mer<strong>in</strong>o ewes. J Reprod Fertil 99, 689–694.Archer GS, D<strong>in</strong>dot S, Friend TH, Walker S, Zaunbrecher G,Lawhorn B, Piedrahita J, 2003a: Hierarchical phenotypicand epigenetic variation <strong>in</strong> cloned sw<strong>in</strong>e. Biol Reprod 69,430–436.Archer GS, Friend TH, Piedrahita J, Nevill CH, Walker S,2003b: Behavioural variation among cloned pigs. ApplAnimal Behav Sci 82, 151–161.Bacci ML, 2007: A brief overview of transgenic farm animals.Veter<strong>in</strong> Res Commun 31(Suppl. 1), 9–14.Bickell SL, Ferguson DM, Boissy A, Blache D, 2008: Sheepselected on their behavioural response to a social and humanchallenge differ also <strong>in</strong> their response to novelty. PhysiolBehav (<strong>in</strong> press).Bielli A, Pérez R, Pedrana G, Milton JTB, Lopez A,Blackberry M, Duncombe G, Rodriguez-Mart<strong>in</strong>ez H, Mart<strong>in</strong>GB, 2002: Low maternal nutrition dur<strong>in</strong>g pregnancyreduces the numbers of Sertoli cells <strong>in</strong> the newborn lamb.Reprod Fertil Dev 14, 333–337.Biggers JD, 1986: The potential use of artificially producedmonozygotic tw<strong>in</strong>s for comparative experiments. Theriogenology26, 1–25.Blache D, Tjondronegoro S, Blackberry MA, Anderson ST,Curlewis JD, Mart<strong>in</strong> GB, 1997: Gonadotroph<strong>in</strong> and prolact<strong>in</strong>secretion <strong>in</strong> castrated male sheep follow<strong>in</strong>g subcutaneousor <strong>in</strong>tracranial treatment with testicular hormones.Endocr 7, 235–243.Blache D, Blackberry MA, Van Cleeff JK, 2002: Does thethyroid axis play a role <strong>in</strong> domestication? 56th AnnualMeet<strong>in</strong>g of the American Institute of Biological Sciences,Wash<strong>in</strong>gton, DC, 22–24 March 2002, Abstract 27.Blache D, Zhang S, Mart<strong>in</strong> GB, 2006: Dynamic and <strong>in</strong>tegrativeaspects of the regulation of reproduction by metabolicstatus <strong>in</strong> male sheep. Reprod Nutr Dev 46, 379–390.Blache D, Chagas LM, Mart<strong>in</strong> GB, 2007: Nutritional <strong>in</strong>puts<strong>in</strong>to the reproductive neuroendocr<strong>in</strong>e control system – amultidimensional perspective. <strong>in</strong>: Juengel JI, Murray JF,Smith MF (eds), <strong>Reproduction</strong> <strong>in</strong> <strong>Domestic</strong> Rum<strong>in</strong>ants VI,Vol. Suppl 64. Nott<strong>in</strong>gham University Press, Nott<strong>in</strong>gham,UK, pp. 123–139.Ó 2008 The Authors. Journal compilation Ó 2008 Blackwell Verlag

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