Hostname: page-component-cd9895bd7-jkksz Total loading time: 0 Render date: 2024-12-22T17:16:11.308Z Has data issue: false hasContentIssue false
  • English
  • Français

Chronic stress in sheep: assessment tools and their use in different management conditions

Published online by Cambridge University Press:  11 January 2023

CM Dwyer  and
HLI Bornett
CM Dwyer*
Affiliation:
Animal Behaviour and Welfare, Sustainable Livestock Systems, Scottish Agricultural College, West Mains Road, Edinburgh EH9 3JG, UK
HLI Bornett
Affiliation:
Department of Animal Welfare and Veterinary Health, Moulton College, West Street, Moulton, Northampton NN3 7RR, UK
*
* Contact for correspondence and requests for reprints: [email protected]
Rights & Permissions [Opens in a new window]

Abstract

Core share and HTML view are not available for this content. However, as you have access to this content, a full PDF is available via the ‘Save PDF’ action button.

Chronic stress occurs when animals are unable to deal with a persistent stressor with species-typical responses, or when several stressors are present concurrently. Chronic stress is most frequently considered in intensive systems, but it may also be a welfare concern for extensively managed species, such as the sheep. Here we review behavioural and physiological responses of sheep to experimentally induced chronic stressors to determine relevant indicators of chronic stress. Neuroendocrine responses to chronic stress are difficult to interpret because initial responses are followed by an apparent normalisation. Thus, cortisol or catecholamines may be at or below pre-stress levels during chronic stress, but this varies with different stressors. Chronic stress can also affect reproductive function, impair body and wool growth and meat quality, reduce immune function, and is associated with greater parasite burdens in sheep. Chronic stress induces alterations in behaviour patterns, particularly activity and feeding, and circadian rhythms of behaviour. Stereotypic behaviours, however, are infrequent in sheep and may occur only in experimental conditions of social isolation. Behavioural and physiological data suggest that rough handling and sheepdogs may be sources of chronic stress for sheep. Social subordination and weaning also act as chronic stressors, leading to higher parasitism in these animals and a greater response to additional stressors. Lameness and parasitism are associated with physiological and behavioural responses indicating that these are severe forms of chronic stress in sheep. It is unclear whether environmental stressors, such as weather and food availability, induce chronic stress in sheep. Under-nutrition may, however, be a welfare concern through its impact on lamb survival. The existence of many sources of chronic stress in the management of sheep suggests that the welfare of this species requires more attention than it has currently received.

Keywords

animal welfarebehaviourchronic stresshandlinglamenessparasitismsheep
Type
Research Article
Information
Animal Welfare , Volume 13 , Issue 3 , August 2004 , pp. 293 - 304
Copyright
© 2004 Universities Federation for Animal Welfare

References

Abdel-Rahman, MA, Ahmed, MM and Sotohy, SA 2000 Acclimatization of sheep to housing stress of metabolism crates with reference to their behavioural and endocrinological aspects. Assiut Veterinary Medical Journal 43: 37–50Google Scholar
Adrichem, PWM and Vogt, JE 1993 The effect of isolation and separation on the metabolism of sheep. Livestock Production Science 33: 151–159CrossRefGoogle Scholar
Alexander, G, Lynch, JJ and Mottershead, BE 1979 Use of shelter and selection of lambing sites by shorn and unshorn ewes in paddocks with closely or widely spaced shelters. Applied Animal Ecology 5: 51–69CrossRefGoogle Scholar
Anderson, N, Barger, IA and Waller, PJ 1987 Impact of gastrointestinal parasitism on pasture utilization by grazing sheep. In: Wheeler, JL, Pearson, CJ and Robards, GE (eds) Temperate Pastures: Their Production, Use and Management pp 555–566. Commonwealth Scientific and Industrial Research Organisation (CSIRO): Victoria, AustraliaGoogle Scholar
Ansari-Renani, HR and Hynd, PI 2001 Cortisol-induced follicle shutdown is related to staple strength in Merino sheep. Livestock Production Science 69: 279–289CrossRefGoogle Scholar
Apple, JK, Dikeman, ME, Minton, JE, McMurphy, RM, Fedde, MR, Leith, DE and Unruh, JA 1995 Effects of restraint and isolation stress and epidural blockade on endocrine and blood metabolite status, muscle glycogen metabolism, and incidence of dark-cutting longissimus muscle of sheep. Journal of Animal Science 73: 2295–2307CrossRefGoogle ScholarPubMed
Apple, JK, Minton, JE, Parsons, KM and Unruh, JA 1993 Influence of repeated restraint and isolation stress and electrolyte administration on pituitary–adrenal secretions, electrolytes, and other blood constituents of sheep. Journal of Animal Science 71: 71–77CrossRefGoogle ScholarPubMed
Argo, CM and Smith, JS 1983 Relationship of energy requirements and seasonal cycles of food intake in Soay rams. Journal of Physiology 343: 23P–24PGoogle Scholar
Argo, CM, Smith, JS and Kay, RNB 1999 Seasonal changes of metabolism and appetite in Soay rams. Animal Science 69: 191–202CrossRefGoogle Scholar
Arnold, GW and Maller, RA 1974 Some aspects of competition between sheep for supplementary feed. Animal Production 19: 309–319Google Scholar
Arnold, GW and Pahl, PJ 1974 Some aspects of social behaviour in sheep. Animal Behaviour 22: 594–600CrossRefGoogle Scholar
Arnold, GW, Wallace, SR and Maller, RA 1979 Some factors involved in natural weaning processes in sheep. Applied Animal Ethology 5: 43–50CrossRefGoogle Scholar
Baldock, NM and Sibly, RM 1990 Effects of handling and transportation on the heart rate and behaviour of sheep. Applied Animal Behaviour Science 28: 15–39CrossRefGoogle Scholar
Bates, PG 1991a Recent advances in the biology and control of sheep scab. Proceedings of the Sheep Veterinary Society 15: 23–27Google Scholar
Bates, PG 1991b Ear mites in sheep. Veterinary Record 128: 555CrossRefGoogle ScholarPubMed
Bates, PG 1996 Epidemiology of sub-clinical ovine psoroptic otoacariasis in Great Britain. Veterinary Record 138: 388–393CrossRefGoogle Scholar
Bates, PG 1997 The pathogenesis and ageing of sheep scab lesions. Part 1. State Veterinary Journal 7: 11–15Google Scholar
Bates, P 2000 Sheep scab. In: Martin, WB and Aitkin, ID (eds) Diseases of Sheep pp 281–285. Blackwell Science: Oxford, UKGoogle Scholar
Berriatua, E, French, NP, Broster, CE, Morgan, KL and Wall, R 2001 Effect of infestation with Psoroptes ovis on the nocturnal rubbing and lying behaviour of housed sheep. Applied Animal Behaviour Science 71: 43–55CrossRefGoogle ScholarPubMed
Boivin, X, Nowak, R, Desprès, G, Tournadre, H and Le Neindre, P 1997 Discrimination between shepherds by lambs reared under artificial conditions. Journal of Animal Science 75: 2892–2898CrossRefGoogle ScholarPubMed
Bowers, CL, Friend, TH, Grissom, KK and Lay, DC 1993 Confinement of lambs (Ovis aries) in metabolism stalls increased adrenal function, thyroxine and motivation for movement. Applied Animal Behaviour Science 36: 149–158CrossRefGoogle Scholar
Bush, LF and Lind, RA 1973 Performance of ewes and lambs in confinement. Journal of Animal Science 36: 407–410CrossRefGoogle Scholar
Casamassima, D, Sevi, A, Palazzo, M, Ramacciato, R, Colella, GE and Bellitti, A 2001 Effects of two different housing systems on behaviour, physiology and milk yield of Comisana ewes. Small Ruminant Research 41: 151–161CrossRefGoogle ScholarPubMed
Cockram, MS, Imlah, P, Goddard, PJ, Harkiss, GD and Waran, NK 1993 The behavioural, endocrine and leucocyte response of ewes to repeated removal of lambs before the age of natural weaning. Applied Animal Behaviour Science 38: 127–142CrossRefGoogle Scholar
Cockram, MS, Kent, JE, Jackson, RE, Goddard, PJ, Doherty, OM, McGilp, IM, Fox, A, Studdert-Kennedy, TC, McConnell, TI and O'Riordan, T 1997 Effect of lairage during 24 h of transport on the behavioural and physiological responses of sheep. Animal Science 65: 391–402CrossRefGoogle Scholar
Cockram, MS and Lee, RA 1991 Some preslaughter factors affecting the occurrence of bruising in sheep. British Veterinary Journal 147: 120–125CrossRefGoogle ScholarPubMed
Cockram, MS, Ranson, M, Imlah, P, Goddard, PJ, Burrells, C and Harkiss, GD 1994 The behavioural, endocrine and immune responses of sheep to isolation. Animal Production 58: 389–400Google Scholar
Cook, CJ 1996 Basal and stress response cortisol levels and stress avoidance learning in sheep (Ovis aries). New Zealand Veterinary Journal 44: 162–163CrossRefGoogle Scholar
Coop, RL 1979 Production loss in sub-clinical helminth infections. Veterinary Record 105: 189Google Scholar
Coop, RL 1982 The impact of sub-clinical parasitism in ruminants. In: Mettrick, DF and Desser, SS (eds) Parasites — Their World and Ours. Proceedings of the 5th International Congress of Parasitology pp 439–450. Elsevier: Amsterdam, The NetherlandsGoogle Scholar
Coop, RL and Jackson, F 2000 Gastrointestinal helminthosis. In: Martin, WB and Aitkin, ID (eds) Diseases of Sheep pp 159–167. Blackwell Science: Oxford, UKGoogle Scholar
Cooper, JJ, Emmans, GE and Friggens, NC 1994 Effect of diet on behaviour of individually penned sheep. Animal Science 58: 44 (Abstract)Google Scholar
Cooper, J and Jackson, R 1996 A comparison of the feeding behaviour of sheep in straw yards and on slats. Applied Animal Behaviour Science 49: 99 (Abstract)CrossRefGoogle Scholar
Cooper, J, McCullam, J and Shanks, M 1995 The effect of fibrous diet on abnormal oral behaviour in stall housed lambs. In: Proceedings of the British Society for Animal Science Winter Meeting, March 1995, Scarborough, UK p 207 (Abstract). British Society for Animal Science: Penicuik, UKCrossRefGoogle Scholar
Coppinger, L and Coppinger, R 2000 Dogs for herding and guarding livestock. In: Grandin, T (ed) Livestock Handling and Transport pp 235–254. CAB International: Wallingford, UKCrossRefGoogle Scholar
Coppinger, TR, Minton, JE, Reddy, PG and Blecha, F 1991 Repeated restraint and isolation stress in lambs increases pituitary–adrenal secretion and reduces cell-mediated immunity. Journal of Animal Science 69: 2808–2814CrossRefGoogle ScholarPubMed
Corke, MJ and Broom, DM 1999 The behaviour of sheep with sheep scab, Psoroptes ovis, infestation. Veterinary Parasitology 83: 291–300CrossRefGoogle ScholarPubMed
Deag, JM 1996 Behavioural ecology and the welfare of extensively farmed animals. Applied Animal Behaviour Science 49: 9–22CrossRefGoogle Scholar
Degabriele, R and Fell, LR 2001 Changes in behaviour, cortisol and lymphocyte types during isolation and group confinement of sheep. Immunology and Cell Biology 79: 583–589CrossRefGoogle ScholarPubMed
Dobson, H and Smith, RF 1995 Stress and reproduction in farm animals. Journal of Reproduction and Fertility 49: 451–461 (Suppl)Google ScholarPubMed
Dobson, H and Smith, RF 2000 What is stress, and how does it affect reproduction? Animal Reproduction Science 60/61: 743-752CrossRefGoogle ScholarPubMed
Dobson, H, Tebble, JE, Phogat, JB and Smith, RF 1999 Effect of transport on pulsatile and surge secretion of LH in ewes in the breeding season. Journal of Reproduction and Fertility 116: 1–8CrossRefGoogle ScholarPubMed
Done-Currie, JR, Hecker, JF and Wodzicka-Tomaszewska, M 1984 Behaviour of sheep transferred from pasture to an animal house. Applied Animal Behaviour Science 12: 121–130CrossRefGoogle Scholar
Dwyer, CM and Lawrence, AB 2000 Effects of maternal genotype and behaviour on the behavioural development of their offspring in sheep. Behaviour 137: 1629–1654CrossRefGoogle Scholar
Dwyer, CM, Lawrence, AB and Bishop, SC 2001 Effects of selection for lean tissue content on maternal and neonatal lamb behaviours in Scottish Blackface sheep. Animal Science 72: 555–571CrossRefGoogle Scholar
Dwyer, CM, Lawrence, AB, Bishop, SC and Lewis, M 2003 Ewe–lamb bonding behaviours at birth are affected by maternal undernutrition in pregnancy. British Journal of Nutrition 89: 123–136CrossRefGoogle ScholarPubMed
Egerton, JR 2000 Foot-rot and other conditions. In: Martin, WB and Aitkin, ID (eds) Diseases of Sheep pp 243–249. Blackwell Science: Oxford, UKGoogle Scholar
Ehnert, K and Moberg, GP 1991 Disruption of estrous behavior in ewes by dexamethasone or management-related stress. Journal of Animal Science 69: 2988–2994CrossRefGoogle ScholarPubMed
Fell, LR, Lynch, JJ, Adams, DB, Hinch, GN, Munro, RK and Davies, HI 1991 Behavioural and physiological effects in sheep of a chronic stressor and a parasite challenge. Australian Journal of Agricultural Research 42: 1335–1346CrossRefGoogle Scholar
Fell, LR and Shutt, DA 1989 Behavioural and hormonal responses to acute surgical stress in sheep. Applied Animal Behaviour Science 22: 283–294CrossRefGoogle Scholar
Festa-Bianchet, M 1988 Nursing behaviour of bighorn sheep: correlates of ewe age, parasitism, lamb age, birthdate and sex. Animal Behaviour 36: 1445–1454CrossRefGoogle Scholar
Fleming, MW 1997 Cortisol as an indicator of severity of parasitic infections of Haemonchus contortus in lambs (Ovis aries). Comparative Biochemistry and Physiology 116B: 41-44CrossRefGoogle Scholar
Fordham, DP, Al-Gahtani, S, Durotoye, LA and Rodway, RG 1991 Changes in plasma cortisol and β-endorphin concentrations and behaviour in sheep subjected to a change in environment. Animal Production 52: 287–296Google Scholar
Fraser, AF and Broom, DM 1990 Farm Animal Behaviour and Welfare. Bailliere Tindall: London, UKGoogle Scholar
French, N, Wall, R, Cripps, PJ and Morgan, KL 1994 Blowfly strike in England and Wales: the relationship between prevalence and farm and management factors. Medical and Veterinary Entomology 8: 51–56CrossRefGoogle ScholarPubMed
Gonyou, HW 2000 Behavioural principles of animal handling. In: Grandin, T (ed) Livestock Handling and Transport pp 15–26. CAB International: Wallingford, UKCrossRefGoogle Scholar
Gonyou, HW, Stookey, JM and McNeal, LG 1985 Effects of double decking and space allowance on the performance and behavior of feeder lambs. Journal of Animal Science 60: 1110–1116CrossRefGoogle ScholarPubMed
Grandin, T 2000 Behavioural principles of handling cattle and other grazing animals under extensive conditions. In: Grandin, T (ed) Livestock Handling and Transport pp 63–86. CAB International: Wallingford, UKCrossRefGoogle Scholar
Hargreaves, AL and Hutson, GD 1990 Some effects of repeated handling on stress responses in sheep. Applied Animal Behaviour Science 26: 253–265CrossRefGoogle Scholar
Harlow, HJ, Thorne, ET, Williams, ES, Belden, EL and Gern, WA 1987 Adrenal responsiveness in domestic sheep (Ovis aries) to acute and chronic stressors as predicted by remote monitoring of cardiac frequency. Canadian Journal of Zoology 65: 2021–2027CrossRefGoogle Scholar
Hass, CC 1991 Social status in female bighorn sheep (Ovis canadensis): expression, development and reproductive correlates. Journal of Zoology 225: 509–523CrossRefGoogle Scholar
Heath, ACG, Bishop, DM and Tenquist, JD 1987 The effects of artificially induced fly-strike on food intake and liveweight gain in sheep. New Zealand Veterinary Journal 35: 50–52CrossRefGoogle ScholarPubMed
Hemsworth, PH and Coleman, GJ 1998 Human–Livestock Interactions: The Stockperson and the Productivity and Welfare of Intensively Farmed Animals. CAB International: Wallingford, UKGoogle Scholar
Hewson, R and Wilson, CJ 1979 Home range and movements of Scottish Blackface sheep in Lochaber, north-west Scotland. Journal of Applied Ecology 16: 743–751CrossRefGoogle Scholar
Hodge, RW, Watson, MJ, Butler, KL, Kelly, AJ, Beers, PJ, Bogdanovic, B and Winfield, CG 1991 Export of live sheep: nutritional studies and the failure to eat syndrome. In: Farrell, DJ (ed) Recent Advances in Animal Nutrition in Australia pp 113–120. University of New England: Armidale, AustraliaGoogle Scholar
Houpt, KA 1987 Abnormal behaviour. In: Price, EO (ed) Farm Animal Behaviour. The Veterinary Clinics of North America Volume 3(2) pp 357–367. WB Saunders: Philadelphia, USAGoogle Scholar
Hunter, RF and Milner, C 1963 The behaviour of individual, related and groups of South Country Cheviot hill sheep. Animal Behaviour 11: 507–513CrossRefGoogle Scholar
Hutchings, MR, Gordon, IJ, Robertson, E, Kyriazakis, I and Jackson, F 2000 Effects of parasitic status and level of feeding motivation on the diet selected by sheep grazing grass/clover swards. Journal of Agricultural Science 135: 65–75CrossRefGoogle Scholar
Hutchings, MR, Kyriazakis, I, Anderson, DH, Gordon, IJ and Coop, RL 1998 Behavioural strategies used by parasitized and non-parasitized sheep to avoid ingestion of gastro-intestinal nematodes associated with faeces. Animal Science 67: 97–106CrossRefGoogle Scholar
Hutchings, MR, Kyriazakis, I, Gordon, IJ and Jackson, F 1999 Trade-offs between nutrient intake and faecal avoidance in herbivore foraging decisions: the effect of animal parasitic status, level of feeding motivation and sward nitrogen content. Journal of Animal Ecology 68: 310–323CrossRefGoogle Scholar
Hutson, GD 1985 The influence of barley food rewards on sheep movement through a handling system. Applied Animal Behaviour Science 14: 263–273CrossRefGoogle Scholar
Hutson, GD 2000 Behavioural principles of sheep handling. In: Grandin, T (ed) Livestock Handling and Transport pp 175–200. CAB International: Wallingford, UKCrossRefGoogle Scholar
Jackson, RE, Cockram, MS, Goddard, PJ, Doherty, OM, McGilp, IM and Waran, NK 1999a The effects of 24 h water deprivation when associated with some aspects of transportation on the behaviour and blood chemistry of sheep. Animal Welfare 8: 229–241Google Scholar
Jackson, RE, Waran, NK and Cockram, MS 1999b Methods for measuring feeding motivation in sheep. Animal Welfare 8: 53–63Google Scholar
Jagusch, KT, Gumbrell, RC, Mobley, MC and Jay, NP 1977 Effect on growth of early weaning lambs onto pure stands of lucerne. New Zealand Journal of Experimental Agriculture 5: 15–18CrossRefGoogle Scholar
Janssens, CJJG, Helmond, FA and Wiegant, VM 1995 The effect of chronic stress on plasma cortisol concentrations in cyclic female pigs depends on the time of day. Domestic Animal Endocrinology 12: 167–177CrossRefGoogle ScholarPubMed
Jarvis, AM and Cockram, MS 1994 Effects of handling and transport on bruising of sheep sent directly from farms to slaughter. Veterinary Record 135: 523–527CrossRefGoogle ScholarPubMed
Jarvis, AM, Cockram, MS and McGilp, IM 1996 Bruising and biochemical measures of stress, dehydration and injury determined at slaughter in sheep transported from farms or markets. British Veterinary Journal 152: 719–722CrossRefGoogle ScholarPubMed
Jensen, KH, Hansen, SW and Pedersen, LJ 1996 The effect of long term stress on the hypothalamic–pituitary–adrenocortical axis and the role of the stressor. Acta Agriculturae Scandinavia, Section A, Animal Science 27: 40–45 (Suppl)Google Scholar
Johnson, EO, Kamilaris, TC, Chrousos, GP and Gold, PW 1992 Mechanisms of stress: a dynamic overview of hormonal and behavioural homeostasis. Neuroscience and Biobehavioral Reviews 16: 115–130CrossRefGoogle Scholar
Johnson, KG 1987 Shading behaviour of sheep: preliminary studies of its relation to thermoregulation, feed and water intakes, and metabolic rates. Australian Journal of Agricultural Research 38: 587–596CrossRefGoogle Scholar
Kiley-Worthington, M 1977 Behaviour Problems of Farm Animals. Oriel Press: Northumberland, UKGoogle Scholar
Kilgour, R and de Langen, H 1970 Stress in sheep resulting from management practices. Proceedings of the New Zealand Society of Animal Production 30: 65–76Google Scholar
Komesaroff, PA, Esler, M, Clarke, IJ, Fullerton, MJ and Funder, JW 1998 Effects of estrogen and estrous cycle on gluco-corticoid and catecholamine responses to stress in sheep. American Journal of Physiology 275: E671–E678Google Scholar
Knight, TW, Hall, DRH, Lynch, PR and Hockey, HUP 1988 Effects of pre-joining shearing, stress, pasture allowance, and haemoglobin type on reproductive performance of Romney and Marshall Romney ewes. New Zealand Journal of Agricultural Research 31: 249–258CrossRefGoogle Scholar
Lawrence, AB and Rushen, J 1993 Introduction. In: Lawrence, AB and Rushen, J (eds) Stereotypic Animal Behaviour pp 1–6. CAB International: Wallingford, UKGoogle Scholar
Lawrence, AB and Wood-Gush, DGM 1988 Home-range behaviour and social organization of Scottish Blackface sheep. Journal of Applied Ecology 25: 25–40CrossRefGoogle Scholar
Ley, SJ, Livingston, A and Waterhouse, AE 1989 The effect of chronic clinical pain on thermal and mechanical thresholds in sheep. Pain 39: 353–357CrossRefGoogle ScholarPubMed
Ley, SJ, Livingston, A and Waterhouse, AE 1991a Effects of chronic lameness on the concentrations of cortisol, prolactin and vasopressin in the plasma of sheep. Veterinary Record 129: 45–47CrossRefGoogle ScholarPubMed
Ley, SJ, Livingston, A and Waterhouse, AE 1992 Effects of clinically occurring chronic lameness in sheep on the concentrations of plasma noradrenaline and adrenaline. Research in Veterinary Science 53: 122–125CrossRefGoogle ScholarPubMed
Ley, S, Waterhouse, A and Livingston, A 1991b The influence of chronic pain on the analgesic effects of the alpha2-adrenoceptor agonists, xylazine, in sheep. Journal of Veterinary Pharmacology and Therapeutics 14: 141–144CrossRefGoogle Scholar
Ley, SJ, Waterhouse, AE and Livingston, A 1995 A field study of the effect of lameness on mechanical nociceptive thresholds in sheep. Veterinary Record 137: 85–87CrossRefGoogle ScholarPubMed
Ley, SJ, Waterhouse, AE, Livingston, A and Parkinson, TJ 1994 Effect of chronic pain associated with lameness on plasma cortisol concentrations in sheep: a field study. Research in Veterinary Science 57: 332–335CrossRefGoogle ScholarPubMed
Low, WA, Tweedie, RL, Edwards, CBH, Hodder, RM, Malapant, KWJ and Cunningham, RB 1981 The influence of environment on daily maintenance behaviour of free-ranging shorthorn cows in central Australia. Applied Animal Ethology 7: 11–56CrossRefGoogle Scholar
Lynch, JJ and Alexander, G 1973 Animal behaviour and the pastoral industries. In: Alexander, G and Williams, OB (eds) Pastoral Industries of Australia pp 371–400. Sydney University Press: Sydney, AustraliaGoogle Scholar
Lynch, JJ and Alexander, G 1976 The effect of gramineous windbreaks on behaviour and lamb mortality among shorn and unshorn Merino sheep during lambing. Applied Animal Ethology 2: 305–325CrossRefGoogle Scholar
Lynch, JJ, Hinch, GN and Adams, DB 1992 The Behaviour of Sheep: Biological Principles and Implications for Production. CAB International: Wallingford, UK, and the Commonwealth Scientific and Industrial Research Organisation (CSIRO): Victoria, AustraliaGoogle Scholar
Lynch, JJ, Hinch, GN, Bouissou, MF, Elwin, RL, Green, GC and Davies, HI 1989 Social organization in young Merino and Merino × Border Leicester ewes. Applied Animal Behaviour Science 22: 49–63CrossRefGoogle Scholar
Markowitz, TM, Dally, MR, Gursky, K and Price, EO 1998 Early handling increases lamb affinity for humans. Animal Behaviour 55: 573–587CrossRefGoogle ScholarPubMed
Marsden, D and Wood-Gush, DGM 1986a A note on the behaviour of individually penned sheep regarding their use for research purposes. Animal Production 42: 157–159Google Scholar
Marsden, D and Wood-Gush, DGM 1986b Abnormal behaviour versus production performance criteria in the assessment of welfare in housed lambs. Applied Animal Behaviour Science 15: 90–91CrossRefGoogle Scholar
Marshall, DJ, Walker, RI, Cullis, BR and Luff, MF 1991 The effect of footrot on body weight and wool growth of sheep. Australian Veterinary Journal 68: 45–49CrossRefGoogle ScholarPubMed
Mateo, JM, Estep, DQ and McCann, JS 1991 Effects of differential handling on the behaviour of domestic ewes (Ovis aries). Applied Animal Behaviour Science 32: 45–54CrossRefGoogle Scholar
Matteri, RL, Watson, JG and Moberg, GP 1984 Stress or acute adrenocorticotrophin treatment suppresses LHRH-induced LH release in the ram. Journal of Reproduction and Fertility 72: 385–393CrossRefGoogle ScholarPubMed
McBride, G, Arnold, GW, Alexander, G and Lynch, JJ 1967 Ecological aspects of the behaviour of domestic animals. Proceedings of the Ecological Society of Australia 2: 133–165Google Scholar
McNatty, KP and Young, A 1973 Diurnal changes of plasma cortisol levels in sheep adapting to a new environment. Journal of Endocrinology 56: 329–330CrossRefGoogle ScholarPubMed
Mears, GJ and Brown, FA 1997 Cortisol and β-endorphin responses to physical and psychological stressors in lambs. Canadian Journal of Animal Science 77: 689–694CrossRefGoogle Scholar
Mears, GJ, Brown, FA and Redmond, LR 1999 Effects of handling, shearing and previous exposure to shearing on cortisol and beta-endorphin responses in ewes. Canadian Journal of Animal Science 79: 35–38CrossRefGoogle Scholar
Minton, JE, Apple, JK, Parsons, KM and Blecha, F 1995 Stress-associated concentrations of plasma cortisol cannot account for reduced lymphocyte function and changes in serum enzymes in lambs exposed to restraint and isolation stress. Journal of Animal Science 73: 812–817CrossRefGoogle ScholarPubMed
Minton, JE and Blecha, F 1990 Effect of acute stressors on endocrinological and immunological functions in lambs. Journal of Animal Science 68: 3145–3151CrossRefGoogle ScholarPubMed
Minton, JE, Coppinger, TR, Reddy, PG, Davis, WC and Blecha, F 1992 Repeated restraint and isolation stress alters adrenal and lymphocyte functions and some leukocyte differentiation antigens in lambs. Journal of Animal Science 70: 1126–1132CrossRefGoogle ScholarPubMed
Moberg, GP and Wood, VA 1982 Effect of differential rearing on the behavioural and adrenocortical response of lambs to a novel environment. Applied Animal Ethology 8: 269–279CrossRefGoogle Scholar
Napolitano, F, Bragheir, A, Cifuni, GF, Pacelli, C and Girolami, A 2002 Behaviour and meat production of organically farmed unweaned lambs. Small Ruminant Research 43: 179–184CrossRefGoogle Scholar
Napolitano, F, Marino, V, de Rosa, G, Capparelli, R and Bordi, A 1995 Influence of artificial rearing on behavioural and immune response of lambs. Applied Animal Behaviour Science 45: 245–253CrossRefGoogle Scholar
Niezgoda, J, Wronska, D, Pierzchala, K, Bobek, S and Kahl, S 1987 Lack of adaptation to repeated emotional stress evoked by isolation of sheep from the flock. Zentralblatt Fur Veterinarmedizin Reihe A (Journal of Veterinary Medicine, Series A, Animal Physiology, Pathology and Clinical Veterinary Medicine) 34: 734–739Google ScholarPubMed
Orgeur, P, Bernard, S, Naciri, M, Nowak, R, Schaal, B and Lévy, F 1999 Psychobiological consequences of two different weaning methods in sheep. Reproduction, Nutrition, Development 39: 231–244CrossRefGoogle ScholarPubMed
Orgeur, P, Mavric, N, Yvore, P, Bernard, S, Nowak, R, Schaal, B and Levy, F 1998 Artificial weaning in sheep: consequences on behavioural, hormonal and immuno-pathological indicators of welfare. Applied Animal Behaviour Science 58: 87–103CrossRefGoogle Scholar
Parrott, RF, Thornton, SN, Forsling, ML and Delaney, CE 1987 Endocrine and behavioural factors affecting water balance in sheep subjected to isolation stress. Journal of Endocrinology 112: 305–310CrossRefGoogle ScholarPubMed
Pearson, RA and Mellor, DJ 1976 Some behavioural and physiological changes in pregnant goats and sheep during adaptation to laboratory conditions. Research in Veterinary Science 20: 215–217CrossRefGoogle ScholarPubMed
Pollard, JC, Littlejohn, RP and Suttie, JM 1992 Behaviour and weight change of red deer calves during different weaning procedures. Applied Animal Behaviour Science 35: 23–33CrossRefGoogle Scholar
Pollard, JC, Shaw, KJ and Littlejohn, RP 1999 A note on sheltering behaviour by ewes before and after lambing. Applied Animal Behaviour Science 61: 313–318CrossRefGoogle Scholar
Prichard, RK, Hennessy, DR and Griffiths, DA 1974 Endocrine responses of sheep to infection with Trichostrongylus colubriformis. Research in Veterinary Science 17: 182–187CrossRefGoogle ScholarPubMed
Przekop, F, Stupnicka, E, Wolriska-Witort, E, Mateusiak, K, Sadowski, B and Domariski, E 1985 Changes in circadian rhythm and suppression of the plasma cortisol level after prolonged stress in the sheep. Acta Endocrinologica 110: 540–545Google ScholarPubMed
Purchas, RW, Barton, RA and Kirton, AH 1980 Relationships of circulating cortisol levels with growth rate and meat tenderness of cattle and sheep. Australian Journal of Agricultural Research 31: 221–232CrossRefGoogle Scholar
Rasmussen, DD and Malven, PV 1983 Effects of confinement stress on episodic secretion of LH in ovariectomized sheep. Neuroendocrinology 36: 392–396CrossRefGoogle ScholarPubMed
Rhind, SM, Reid, HW, McMillen, SR and Palmarini, G 1998 The role of cortisol and β-endorphin in the response of the immune system to weaning in lambs. Animal Science 66: 397–402CrossRefGoogle Scholar
Rhodes, RC, Nippo, MM and Gross, WA 1994 Stress in lambs (Ovis aries) during a routine management procedure (tail docking): evaluation of acute and chronic responses. Comparative Biochemistry and Physiology, A — Comparative Physiology 107: 181–185CrossRefGoogle Scholar
Richards, RB, Norris, RT, Dunlop, RH and McQuade, NC 1989 Causes of death in sheep exported live by sea. Australian Veterinary Journal 66: 33–38CrossRefGoogle ScholarPubMed
Royal Veterinary College 1997 survey cited in: Department for Environment, Food and Rural Affairs (ed) Lameness in Sheep. DEFRA: London, UK. Available at: http://www.defra.gov.uk/animalh/welfare/pdf/sheeplameness.pdf.Google Scholar
Ruiz-de-la-Torre, JL, Velarde, A, Diestre, A, Gispert, M, Hall, SJG, Broom, DM and Manteca, X 2001 Effects of vehicle movements during transport on the stress responses and meat quality of sheep. Veterinary Record 148: 227–229CrossRefGoogle ScholarPubMed
Rushen, J 1990 Use of aversion learning techniques to measure distress in sheep. Applied Animal Behaviour Science 28: 3–14CrossRefGoogle Scholar
Rushen, J 1996 Using aversion learning techniques to assess the mental state, suffering and welfare of farm animals. Journal of Animal Science 74: 1990–1995CrossRefGoogle ScholarPubMed
Rushen, J and de Passillé, AMB 1992 The scientific assessment of the impact of housing on animal welfare: a critical review. Canadian Journal of Animal Science 72: 721–743CrossRefGoogle Scholar
Sabrh, HA, Shalash, MR, Ashoub, MR, Al-Wakeel, AM and Hassan, SG 1992 Effect of stocking density, age and body weight on some reproductive performance of Barki ewes. Egyptian Journal of Veterinary Science 29: 47–54Google Scholar
Sargison, ND 1995 Differential diagnosis and treatment of sheep scab. In Practice 17: 3–9CrossRefGoogle Scholar
Schlink, AC, Wynn, PC, Lea, JM, Briegel, JR and Adams, NR 2002 Effect of cortisol acetate on wool quality in sheep selected for divergent staple strength. Australian Journal of Agricultural Research 53: 183–189CrossRefGoogle Scholar
Schouten, WGP, Rushen, J and de Passillé, AMB 1991 Stereotypic behavior and heart rate in pigs. Physiology and Behavior 50: 617–624CrossRefGoogle ScholarPubMed
Schrader, L and Ladewig, J 1999 Temporal differences in the responses of the pituitary–adrenocortical axis, the sympathoad-renomedullary axis, heart rate, and behaviour to a daily repeated stressor in domestic pigs. Physiology and Behavior 66: 775–783CrossRefGoogle ScholarPubMed
Sevi, A, Massa, S, Annicchiarico, G, Dell'Aquila, S and Muscio, A 1999 Effect of stocking density on ewes' milk yield, udder health and microenvironment. Journal of Dairy Research 66: 489–499CrossRefGoogle ScholarPubMed
Sevi, A, Taibi, L, Albenzio, M, Muscio, A, Dell'Aquila, S and Napolitano, F 2001 Behavioural, adrenal, immune and productive responses of lactating ewes to regrouping and relocation. Journal of Animal Science 79: 1457–1465CrossRefGoogle ScholarPubMed
Sherwin, CM and Johnson, KG 1987 The influence of social factors on the use of shade by sheep. Applied Animal Behaviour Science 18: 143–155CrossRefGoogle Scholar
Shutt, DA, Smith, AI, Wallace, CA, Connell, R and Fell, LR 1988 Effect of myiasis and acute restraint stress on plasma levels of immunoreactive beta-endorphin, adrenocorticotrophin (ACTH) and cortisol in the sheep. Australian Journal of Biological Sciences 41: 297–301CrossRefGoogle ScholarPubMed
Silanikove, N 2000 Effects of heat stress on the welfare of extensively managed domestic ruminants. Livestock Production Science 67: 1–18CrossRefGoogle Scholar
Sotiraki, ST, Leontides, LS and Himonas, CA 1999 The effect of transportation and confinement stress on egg production by Dicrocoelium dendriticum in sheep. Journal of Helminthology 73: 337–339CrossRefGoogle ScholarPubMed
Stephens, DB and Baldwin, BA 1971 Observations of the behaviour of groups of artificially reared lambs. Research in Veterinary Science 12: 219–224CrossRefGoogle ScholarPubMed
Terlouw, EMC, Lawrence, AB and Illius, AW 1991 Influences of feeding level and physical restriction on the development of stereotypies in sows. Animal Behaviour 42: 981–992CrossRefGoogle Scholar
Terlouw, EMC, Schouten, WGP and Ladewig, J 1997 Physiology. In: Appleby, MC and Hughes, BO (eds) Animal Welfare pp 143–158. CAB International: Wallingford, UKGoogle Scholar
Thompson, K, Coleman, ES, Hudmon, A, Kemppainen, RJ, Soyoola, EO and Sartin, JL 1995 Effects of short-term cortisol infusion on growth hormone-releasing hormone stimulation of growth hormone release in sheep. American Journal of Veterinary Research 56: 1228–1231Google ScholarPubMed
Tilbrook, AJ, Cameron, AWN and Lindsay, DR 1987 The influence of ram mating preferences and social interaction between rams on the proportion of ewes mated at field joining. Applied Animal Behaviour Science 18: 173–184CrossRefGoogle Scholar
Tobler, I, Jaggi, K, Arendt, J and Ravault, JP 1991 Long-term 24-hour rest–activity pattern of sheep in stalls and in the field. Experientia 47: 744–749CrossRefGoogle ScholarPubMed
Uetake, K, Yamaguchi, S and Tanaka, T 2000 Psychological effects of early gentling on the subsequent ease of handling in lambs. Animal Science Journal 71: 515–519Google Scholar
Veissier, I, Boissy, A, Nowak, R, Orgeur, P and Poindron, P 1998 Ontogeny of social awareness in domestic herbivores. Applied Animal Behaviour Science 57: 233–245CrossRefGoogle Scholar
Veissier, I, le Neindre, P and Trillat, G 1989 The use of circadian behaviour to measure adaptation of calves to changes in their environment. Applied Animal Behaviour Science 22: 1–12CrossRefGoogle Scholar
Walkden-Brown, SW, Daly, BL, Crook, BJ and Colditz, IG 1999a Blowfly strike and wool growth. I. Role of anorexia in mediating effects on wool. Recent Advances in Animal Nutrition in Australia 12: 2A (Abstract)Google Scholar
Walkden-Brown, SW, Daly, B, Colditz, IG, McFarlan, JR and Kauter, KG 1999b Blowfly strike and wool growth. II. Potential mediators of effects, other than anorexia. Recent Advances in Animal Nutrition in Australia 12: 3A (Abstract)Google Scholar
Waller, PJ, Axelsen, A, Donald, AD, Morley, FHW, Dobson, RJ and Donnelly, JR 1987b Effects of helminth infection on the pre-weaning production of ewes and lambs: comparison between safe and contaminated pasture. Australian Veterinary Journal 64: 357–362CrossRefGoogle ScholarPubMed
Waller, PJ, Donnelly, JR, Dobson, RJ, Donald, AD, Axelsen, A and Morley, FHW 1987a Effects of helminth infection on the pre-weaning production of ewes and lambs: evaluation of pre- and post-lambing drenching and provision of safe lambing pasture. Australian Veterinary Journal 64: 339–343CrossRefGoogle ScholarPubMed
Waterhouse, A 1996 Animal welfare and sustainability of production under extensive conditions — A European perspective. Applied Animal Behaviour Science 49: 29–40CrossRefGoogle Scholar
Watson, DL 1991 Effect of weaning on antibody responses and nematode parasitism in Merino lambs. Research in Veterinary Science 51: 128–132CrossRefGoogle ScholarPubMed
Webster, AJF 1983 Environmental stress and the physiology, performance and health of ruminants. Journal of Animal Science 57: 1584–1593CrossRefGoogle ScholarPubMed
Wiepkema, PR and Koolhaas, JM 1993 Stress and animal welfare. Animal Welfare 2: 195–218Google Scholar
Yurtman, IY, Savas, T, Karaagac, F and Coskuntuna, L 2002 Effects of daily protein intake levels on the oral stereotypic behaviours in energy restricted lambs. Applied Animal Behaviour Science 77: 77–88CrossRefGoogle Scholar
Zenchak, JJ and Anderson, GC 1980 Sexual performance levels of rams (Ovis aries) as affected by social experiences during rearing. Journal of Animal Science 50: 167–174CrossRefGoogle Scholar