Hostname: page-component-8448b6f56d-tj2md Total loading time: 0 Render date: 2024-04-16T13:37:38.374Z Has data issue: false hasContentIssue false

Minimising pain in farm animals: the 3S approach – ‘Suppress, Substitute, Soothe’

Published online by Cambridge University Press:  21 February 2012

R. Guatteo*
INRA, UMR 1300, Bio-Agression, Epidémiologie et Analyse de Risque, F-44307 Nantes, France Oniris, UMR 1300, Bio-Agression, Epidémiologie et Analyse de Risque, F-44307 Nantes, France Université Nantes Angers Le Mans, France
O. Levionnois
Anaesthesia Section, Department for Clinical Veterinary Medicine, Vetsuisse Faculty, University of Bern, Switzerland Allevia AG, The Bone CRO, Bern, Switzerland
D. Fournier
INRA, UAR378, Service Déconcentré d'Appui à la Recherche, Equipe Régionale d'Information Scientifique et Technique, F-34060 Montpellier, France INRA, Délégation et Partenariat aux Entreprises, F-75337 Paris Cedex 07, France
D. Guémené
INRA, UR83 Recherches Avicoles, F-37380 Nouzilly, France
K. Latouche
INRA, UR 1134 Laboratoire Etudes et Recherches Economiques, F-44300 Nantes, France
C. Leterrier
INRA, UMR 85 Physiologie de la Reproduction et des Comportements, F-37380 Nouzilly, France
P. Mormède
INRA, UMR 444 Génétique Cellulaire, F-31326 Castanet-Tolosan, France
A. Prunier
INRA, UMR 1079 Systèmes d'Elevage Nutrition Animale et Humaine, F-35590 Saint-Gilles, France
J. Servière
INRA, UMR 0791 Modélisation Systémique Appliquée aux Ruminants, F-75231 Paris, France
C. Terlouw
INRA, UR 1213, Herbivores, F-63122 St-Genès Champanelle, France
P. Le Neindre
INRA, CODIR, F-75338 Paris, France
Get access


Recently, the French National Institute for Agricultural Research appointed an expert committee to review the issue of pain in food-producing farm animals. To minimise pain, the authors developed a ‘3S’ approach accounting for ‘Suppress, Substitute and Soothe’ by analogy with the ‘3Rs’ approach of ‘Reduction, Refinement and Replacement’ applied in the context of animal experimentation. Thus, when addressing the matter of pain, the following steps and solutions could be assessed, in the light of their feasibility (technical constraints, logistics and regulations), acceptability (societal and financial aspects) and availability. The first solution is to suppress any source of pain that brings no obvious advantage to the animals or the producers, as well as sources of pain for which potential benefits are largely exceeded by the negative effects. For instance, tail docking of cattle has recently been eliminated. Genetic selection on the basis of resistance criteria (as e.g. for lameness in cattle and poultry) or reduction of undesirable traits (e.g. boar taint in pigs) may also reduce painful conditions or procedures. The second solution is to substitute a technique causing pain by another less-painful method. For example, if dehorning cattle is unavoidable, it is preferable to perform it at a very young age, cauterising the horn bud. Animal management and constraint systems should be designed to reduce the risk for injury and bruising. Lastly, in situations where pain is known to be present, because of animal management procedures such as dehorning or castration, or because of pathology, for example lameness, systemic or local pharmacological treatments should be used to soothe pain. These treatments should take into account the duration of pain, which, in the case of some management procedures or diseases, may persist for longer periods. The administration of pain medication may require the intervention of veterinarians, but exemptions exist where breeders are allowed to use local anaesthesia (e.g. castration and dehorning in Switzerland). Extension of such exemptions, national or European legislation on pain management, or the introduction of animal welfare codes by retailers into their meat products may help further developments. In addition, veterinarians and farmers should be given the necessary tools and information to take into account animal pain in their management decisions.

Full Paper
Copyright © The Animal Consortium 2012

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)


Allison, CP, Johnson, RC, Doumit, ME 2005. The effects of halothane sensitivity on carcass composition and meat quality in HAL-1843-normal pigs. Journal of Animal Science 83, 671678.CrossRefGoogle ScholarPubMed
Allison, CP, Marr, AL, Berry, NL, Anderson, DB, Ivers, DJ, Richardson, LF, Keffaber, K, Johnson, RC, Doumit, ME 2006. Effects of halothane sensitivity on mobility status and blood metabolites of HAL-1843-normal pigs after rigorous handling. Journal of Animal Science 84, 10151021.CrossRefGoogle ScholarPubMed
Anderson, DE, Muir, WW 2005. Pain management in cattle. Veterinary Clinics of North America – Food Animal Practice 21, 623635, v–vi.CrossRefGoogle ScholarPubMed
Anil, SS, Anil, L, Deen, J 2002. Challenges of pain assessment in domestic animals. Journal of the American Veterinary Medical Association 220, 313319.CrossRefGoogle ScholarPubMed
Backstrom, L, Kauffman, R 1995. The porcine stress syndrome – a review of genetics, environmental factors, and animal well-being implications. Agri-Practice 16, 2430.Google Scholar
Banner, M 1995. Report of the Committee to consider the ethical implications of the emerging technologies in the breeding of farm animals. ISBN 0 11 242965 3, 77pp.Google Scholar
Barillet, F, Rupp, R, Mignon-Grasteau, S, Astruc, J-M, Jacquin, M 2001. Genetic analysis for mastitis resistance and milk somatic cell score in French Lacaune dairy sheep. Genetics, Selection, Evolution 33, 397415.CrossRefGoogle ScholarPubMed
Bataille, G, Rugraff, Y, Chevillon, P, Meunier-Salaün, MC 2002. Caudectomie et section des dents chez le porcelet: conséquences comportementales, zootechniques et sanitaires. TechniPorc 25, 513.Google Scholar
Beilharz, RG 1998. Environmental limit to genetic change. An alternative theorem of natural selection. Journal of Animal Breeding and Genetics 115, 433437.Google Scholar
Bishop, SC, Morris, CA 2007. Genetics of disease resistance in sheep and goats. Small Ruminant Research 70, 4859.CrossRefGoogle Scholar
Blokhuis, HJ 1989. The effect of a sudden change in floor type on pecking behaviour in chicks. Applied Animal Behaviour Science 22, 6573.CrossRefGoogle Scholar
Bodin, L, Bolet, G, Garcia, M, Hgarreau, H, Larzul, C, David, I 2010. Robustesse et canalisation : vision de généticiens. INRA Productions Animales 23, 1122.CrossRefGoogle Scholar
Boesch, D, Steiner, A, Stauffacher, M 2006. Castration of calves: a survey among Swiss suckler beef farmers. Schweizer Archiv für Tierheilkunde 148, 231244.CrossRefGoogle Scholar
Brambell, FWR 1965. Report of the technical committee to enquire into the welfare of animals kept under intensive livestock husbandry systems. HMSO Cmmand 2836, London.Google Scholar
Breuer, K, Sutcliffe, MEM, Mercer, JT, Rance, KA, O'Connell, NE, Sneddon, IA, Edwards, SA 2005. Heritability of clinical tail-biting and its relation to performance traits. Livestock Production Science 93, 8794.CrossRefGoogle Scholar
Bretschneider, G 2005. Effects of age and method of castration on performance and stress response of beef male cattle: a review. Livestock Production Science 97, 89100.CrossRefGoogle Scholar
Breward, J, Gentle, MJ 1985. Neuroma formation and abnormal afferent nerve discharges after partial break amputation (beak trimming) in poultry. Experientia 41, 11321134.CrossRefGoogle Scholar
Buhk, HJ 1999. Rapport de la session Recherche. Conférences internationales du conseil de l'Europe sur les questions éthiques soulevées par l'application de la biotechnologie, Oviedo, pp. 45–51.Google Scholar
Buitenhuis, B, Hedegaard, J, Janss, L, Sorensen, P 2009. Differentially expressed genes for aggressive pecking behaviour in laying hens. BMC Genomics 10, 544.CrossRefGoogle ScholarPubMed
Chambers, C, Powell, L, Wilson, E, Green, LE 1995. A postal survey of tail biting in pigs in south-west England. Veterinary Record 136, 147148.CrossRefGoogle ScholarPubMed
Chapuis, H, Boulay, M, Retailleau, J-P, Arnould, C, Mignon-Grasteau, S, Berri, C, Besnard, J, Coudurier, B, Faure, J-M 2003. Sélection d'une souche de poulet label contre le picage: bilan après trois générations de sélection au picomètre. 5èmes Journées de la Recherche Avicole, 26–27 March 2003, Tours, France, pp. 363–366.Google Scholar
Cheng, H 2006. Morphopathological changes and pain in beak trimmed laying hens. World's Poultry Science Journal 62, 4152.CrossRefGoogle Scholar
Clapperton, M, Diack, AB, Matika, O, Glass, EJ, Gladney, CD, Mellencamp, MA, Hoste, A, Bishop, SC 2009. Traits associated with innate and adaptive immunity in pigs: heritability and associations with performance under different health status conditions. Genetics, Selection, Evolution 41, 5465.CrossRefGoogle ScholarPubMed
Colleau, JJ, Regaldo, D 2001. Définition de l'objectif de sélection dans les races bovines laitières. 8. Rencontres Recherches Ruminants, pp. 329–332.Google Scholar
Coopman, F, Gengler, N, Groen, AF, De Smet, S, Van Aalderen, WM 2004. Comparison of external morphological traits of newborns to inner morphological traits of the dam in the doublemuscled Belgian Blue beef breed. Journal of Animal Breeding and Genetics 121, 128134.CrossRefGoogle Scholar
Craig, JV, Muir, WM 1996. Group selection for adaptation to multiple-hen cages: beak-related mortality, feathering, and bodyweight responses. Poultry Science 75, 294302.CrossRefGoogle Scholar
D'Eath, RB, Turner, SP, Kurt, E, Evans, G, Tholking, L, Looft, H, Wimmers, K, Murani, E, Klont, R, Foury, A, Ison, SH, Lawrence, AB, Mormede, P 2010. Pigs’ aggressive temperament affects pre-slaughter mixing aggression, stress and meat quality. Animal 4, 604616.CrossRefGoogle ScholarPubMed
de Roest, K, Montanari, C, Fowler, T, Baltussen, W 2009. Resource efficiency and economic implications of alternatives to surgical castration without anaesthesia. Animal 3, 15221531.CrossRefGoogle ScholarPubMed
Desrochers, A 2004. Treatment of pathological diseased foot and digits – septic arthritis. In Farm animal surgery (ed. SL Fubini and NG Ducharme), pp. 330336. Saunders, St Louis, Missouri, USA.Google Scholar
Duncan, IJH, Slee, GS, Seawright, E, Breward, J 1989. Behavioural consequences of partial beak amputation (beak trimming) in poultry. British Poultry Science 30, 479488.CrossRefGoogle ScholarPubMed
EFSA 2004a. Opinion of the scientific panel on animal health and welfare on a request from the commission related to welfare aspects of the castration of piglets. The EFSA Journal 91, 18.CrossRefGoogle Scholar
EFSA 2004b. Scientific report of the scientific panel for animal health and welfare on a request from the commission related to welfare aspects of animal stunning and killing methods 91, 241.CrossRefGoogle Scholar
EFSA 2007a. Scientific report on the risks associated with tail biting in pigs and possible means to reduce the need for tail docking considering the different housing and husbandry systems 93, 96.Google Scholar
EFSA 2007b. The risks associated with tail biting in pigs and possible means to reduce the need for tail docking considering the different housing and husbandry systems. Scientific opinion of the Panel on Animal Health and Welfare. The EFSA Journal 93, 13.Google Scholar
Erskine, RJ, Wagner, S, De Graves, FJ 2003. Mastitis therapy and pharmacology. Veterinary Clinics of North America – Food Animal Practice 19, 109138.CrossRefGoogle ScholarPubMed
Farm Animal Welfare Council (FAWC) 2007. Opinion on beak trimming of laying hens, FAWC, London, UK, p. 12.Google Scholar
Fernandez, X, Neyraud, E, Astruc, T, Sante, V 2002. Effects of halothane genotype and pre-slaughter treatment on pig meat quality. Part 1. Post mortem metabolism, meat quality indicators and sensory traits of m. Longissimus lumborum . Meat Science 62, 429437.CrossRefGoogle ScholarPubMed
Fiks van Niekerk, T, de Jong, IC 2007. Mutilations in poultry European poultry production systems. Lohmann Information 42, 3546.Google Scholar
Flecknell, P 2002. Replacement, reduction and refinement. ALTEX 19, 7378.Google ScholarPubMed
Foury, A, Geverink, NA, Gil, M, Gispert, M, Hortos, M, Furnols, MFI, Carrion, D, Blott, SC, Plastow, GS, Mormede, R 2007. Stress neuroendocrine profiles in five pig breeding lines and the relationship with carcass composition. Animal 1, 973982.CrossRefGoogle ScholarPubMed
Fredriksen, B, Nafstad, O 2006. Surveyed attitudes, perceptions and practices in Norway regarding the use of local anaesthesia in piglet castration. Research in Veterinary Science 81, 293295.CrossRefGoogle ScholarPubMed
Fredriksen, B, Font i Furnols, M, Lundström, K, Prunier, A, Tuyttens, F, Migdal, W, Bonneau, M 2008. Practice on castration of piglets in Europe. Proceedings of 59th Annual Meeting of the European Association for Animal Production (EAAP), Vilnius, p. 93.Google Scholar
Fredriksen, B, Font i Furnols, M, Lundström, K, Migdal, W, Prunier, A, Tuyttens, FAM, Bonneau, M 2009. Practice on castration of piglets in Europe. Animal 3, 14801487.CrossRefGoogle ScholarPubMed
Fujii, J, Otsu, K, Zorzato, F, Deleon, S, Khanna, VK, Weiler, JE, Obrien, PJ, Maclennan, DH 1991. Identification of a mutation in porcine ryanodine receptor associated with malignant hyperthermia. Science 253, 448451.CrossRefGoogle ScholarPubMed
Fukawa, K, Kusuhara, S 2001. The genetic and non-genetic aspects of leg weakness and osteochondrosis in pigs – review. Asian-Australasian Journal of Animal Sciences 14, 4552.CrossRefGoogle Scholar
Fulponi, L 2006. Private voluntary standards in the food system: the perspective of major food retailers in OECD countries. Food Policy 31, 113.CrossRefGoogle Scholar
Gallois, M, Le Cozler, Y, Prunier, A 2005. Influence of tooth resection in piglets on welfare and performance. Preventive Veterinary Medicine 69, 1323.CrossRefGoogle ScholarPubMed
Garcia-Belenguer, S, Palacio, J, Gascon, M, Acena, C, Revilla, R, Mormede, P 1996. Differences in the biological stress responses of two cattle breeds to walking up to mountain pastures in the Pyrenees. Veterinary Research 27, 515526.Google ScholarPubMed
Gentle, MJ 1991. The acute effects of amputation on peripheral trigeminal afferents in Gallus gallus var domesticus . Pain 46, 97103.CrossRefGoogle ScholarPubMed
Gentle, MJ, Waddington, D, Hunter, LN, Jones, RB 1990. Behavioural evidence for persistent pain following partial beak amputation in chickens. Applied Animal Behaviour Science 27, 149157.CrossRefGoogle Scholar
Gentle, MJ, Hughes, BO, Fox, A, Waddington, D 1997. Behavioural and anatomical consequences of two beak trimming methods in 1- and 10-d-old domestic chicks. British Poultry Science 38, 453463.CrossRefGoogle ScholarPubMed
Gerritzen, M, Kluivers-Poodt, M, Reimert, H, Hindle, V, Lambooij, B 2009. Response to the letter to the editor on the surgical castration of piglets. Animal 3, 14761477.CrossRefGoogle Scholar
Glatz, PC, Lunam, CA 1994. Production and heart-rate responses of chickens beak-trimmed at hatch or at 10 or 42 days of age. Australian Journal of Experimental Agriculture 34, 443447.CrossRefGoogle Scholar
Glazier, DS 2008. Trade-offs. In Resource allocation theory applied to farm animal production (ed. WM Rauw), pp. 4460. CAB International, Cambridge.CrossRefGoogle Scholar
Goonewardene, LA, Price, MA, Liu, MF, Berg, RT, Erichsen, CM 1999. A study of growth and carcass traits in dehorned and polled composite bulls. Canadian Journal of Animal Science 79, 383385.CrossRefGoogle Scholar
Gross, WB 1976. Plasma steroid tendency, social environment and Eimeria necatrix infection. Poultry Science 55, 15081512.CrossRefGoogle ScholarPubMed
Guàrdia, MD, Estany, J, Balasch, S, Oliver, MA, Gispert, M, Diestre, A 2009. Risk assessment of skin damage due to pre-slaughter conditions and RYR1 gene in pigs. Meat Science 81, 745751.CrossRefGoogle ScholarPubMed
Guatteo, R, Holopherne, D, Whay, HR, Huxley, JN 2008. Attitudes et pratiques actuelles des vétérinaires praticiens dans la prise en charge de la douleur des bovins. Bulletin de la SNGTV 44, 6168.Google Scholar
Haga, HA, Ranheim, B 2005. Castration of piglets: the analgesic effects of intratesticular and intrafunicular lidocaine injection. Veterinary Anaesthesia and Analgesia 32, 19.CrossRefGoogle ScholarPubMed
Haga, HA, Lykkjen, S, Revold, T, Ranheim, B 2006. Effect of intratesticular injection of lidocaine on cardiovascular responses to castration in isoflurane-anesthetized stallions. American Journal of Veterinary Research 67, 403408.CrossRefGoogle ScholarPubMed
Hanzen, C, Laurent, Y, Ward, WR 1994. Comparison of reproductive performance in Belgian dairy and beef cattle. Theriogenology 41, 10991114.CrossRefGoogle ScholarPubMed
Hay, M, Rue, J, Sansac, C, Brunel, G, Prunier, A 2004. Long-term detrimental effects of tooth clipping or grinding in piglets: a histological approach. Animal Welfare 13, 2732.CrossRefGoogle Scholar
Heringstad, B, Klemetsdal, G, Ruane, J 2000. Selection for mastitis resistance in dairy cattle: a review with focus on the situation in the Nordic countries. Livestock Production Science 64, 95106.CrossRefGoogle Scholar
Herr, K, Titler, M, Fine, P, Sanders, S, Cavanaugh, J, Swegle, J, Forcucci, C, Tang, X 2010. Assessing and treating pain in hospices: current state of evidence-based practices. Journal of Pain and Symptom Management 39, 803819.CrossRefGoogle ScholarPubMed
Hickford, JGH, Zhou, H, Slow, S, Fang, Q 2004. Diversity of the ovine DQA2 gene. Journal of Animal Science 82, 15531563.CrossRefGoogle ScholarPubMed
Hoe, FGH, Ruegg, PL 2006. Opinions and practices of Wisconsin dairy producers about biosecurity and animal well-being. Journal of Dairy Science 89, 22972308.CrossRefGoogle ScholarPubMed
Huber-Eicher, B, Spring, P 2008. Attitudes of Swiss consumers towards meat from entire or immunocastrated boars: a representative survey. Research in Veterinary Science 85, 625627.CrossRefGoogle ScholarPubMed
Hughes, BO 1982. Feather pecking and cannibalism in domestic fowls. Disturbed behaviour of farm animals. Hohenheimer Arbeiten 121, 138146.Google Scholar
Hughes, BO 1985. Feather loss – How does it occur? Proceedings of the 2nd European Poultry Welfare Symposium, Edinburgh, pp. 178188.Google Scholar
Hughes, BO, Duncan, I 1972. The influence of strain and environmental factors upon feather pecking and cannibalism in fowls. British Poultry Science 13, 525547.CrossRefGoogle ScholarPubMed
Hughes, BO, Gentle, MJ 1995. Beak trimming of poultry: its implications for welfare. World's Poultry Science Journal 51, 5161.CrossRefGoogle Scholar
Hunter, EJ, Jones, TA, Guise, HJ, Penny, RHC, Hoste, S 1999. Tail biting in pigs 1: the prevalence at six UK abattoirs and the relationship of tail biting with docking, sex and other carcass damage. Pig Journal 43, 1832.Google Scholar
Hunter, EJ, Jones, TA, Guise, HJ, Penny, RHC, Hoste, S 2001. The relationship between tail biting in pigs, docking procedure and other management practices. Veterinary Journal 161, 7279.CrossRefGoogle ScholarPubMed
Julian, RJ 1998. Rapid growth problems: ascites and skeletal deformities in broilers. Poultry Science 77, 17731780.CrossRefGoogle ScholarPubMed
Kent, JE, Thrusfield, MV, Robertson, IS, Molony, V 1996. Castration of calves: a study of methods used by farmers in the United Kingdom. Veterinary Record 138, 384387.CrossRefGoogle Scholar
Kjaer, JB, Sorensen, P, Sub, G 2001. Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus). Applied Animal Behaviour Science 71, 229239.CrossRefGoogle ScholarPubMed
Knap, PW 2005. Breeding robust pigs. Proceedings of the 16th Association for the Advancement of Animal Breeding and Genetics (AAABG) Conference, 25–28 September 2005, Noosa, Queensland, Australia, pp. 763–773.Google Scholar
Knap, PW 2008. Robustness. In Resource allocation theory applied to farm animal production (ed. WM Rauw), pp. 288301. CAB International.CrossRefGoogle Scholar
Knap, PW, Rauw, WM 2008. Selection for high production in pigs. In Resource allocation theory applied to farm animal production (ed. WM Rauw), pp. 210229. CAB International.CrossRefGoogle Scholar
Krider, JL, Albright, JL, Plumlee, MP, Conrad, JH, Sinclair, CL, Underwood, L, Jones, RG, Harrington, RB 1975. Magnesium supplementation, space and docking effects on swine performance and behavior. Journal of Animal Science 40, 10271033.CrossRefGoogle Scholar
Lagerkvist, CJ, Carlsson, F, Viske, D 2006. Swedish consumer preferences for animal welfare and biotech: a choice experiment. AgBioForum 9, 5158.Google Scholar
Larrère, C, Larrère, R 2004. Actualité de l'animal-machine? Sens Public, Dossier: La représentation du vivant – du cerveau au comportement.Google Scholar
Larzul, C, le Roy, P, Gueblez, R, Talmant, A, Gogue, J, Sellier, P, Monin, G 1997. Effect of halothane genotype (NN, Nn, nn) on growth, carcass and meat quality traits of pigs slaughtered at 95 kg or 125 kg live weight. Journal of Animal Breeding and Genetics 114, 309320.CrossRefGoogle ScholarPubMed
Laven, RA, Lawrence, KE, Weston, JF, Dowson, KR, Stafford, KJ 2008. Assessment of the duration of the pain response associated with lameness in dairy cows, and the influence of treatment. New Zealand Veterinary Journal 56, 210217.CrossRefGoogle ScholarPubMed
Le Bihan-Duval, E, Beaumont, C, Colleau, JJ 1996. Genetic parameters of the twisted legs syndrome in broiler chickens. Genetics, Selection, Evolution 28, 177195.CrossRefGoogle Scholar
Le Neindre, P, Guatteo, R, Guémené, D, Guichet, JL, Latouche, K, Leterrier, C, Levionnois, O, Mormède, P, Prunier, A, Serrie, A, Servière, J 2009. Douleurs animales. Les identifier, les comprendre, les limiter chez les animaux d’élevage. Expertise scientifique collective, rapport d'expertise, p. 340. INRA, France.Google Scholar
Lefebvre, D, Lips, D, Ödberg, FO, Giffroy, JM 2007. Tail docking in horses: a review of the issues. Animal 1, 11671178.CrossRefGoogle ScholarPubMed
Lepkova, R, Sterc, J, Vecerek, V, Doubek, J, Kruzikova, K, Bedanova, I 2007. Stress responses in adult cattle due to surgical dehorning using three different types of anaesthesia. Berliner Und Munchener Tierarztliche Wochenschrift 120, 465469.Google ScholarPubMed
Levionnois, O, Guatteo, R 2008. Analgésie chez les bovins. Modalités de prise en charge de la douleur. Le point Vétérinaire Numéro Spécial.Google Scholar
Lunam, CA 2005. The anatomy and innervation of the chicken beak: effects of trimming and re-trimming. In Poultry welfare issues – beak trimming (ed. PC Glatz), pp. 5168. Nottingham University Press, UK.Google Scholar
Martrenchar, A, Guémené, D, Morisse, JP 2001. Evaluation de l'intérêt d'une anesthésie/analgésie locale ou générale lors du chaponnage. In 4th Journées de la recherche avicole, Nantes, pp. 119122.Google Scholar
Maxson, SC, Canastar, A 2007. Aggression: concepts and methods relevant to genetic analyses in mice and humans. In Neurobehavioral genetics: methods and applications (ed. BC Jones and P Mormède), pp. 281290. CRC, Boca Raton.Google Scholar
Mc Hugh, N, Evans, RD, Amer, PR, Fahey, AG, Berry, DP 2011. Genetic parameters for cattle price and body weight from routinely collected data at livestock auctions and commercial farms. Journal of Animal Science 89, 2939.CrossRefGoogle ScholarPubMed
McGlone, JJ, Sells, J, Hurst, RJ 1992. Cannibalism in growing pigs: effects of tail docking and housing system on behaviour, performance and immune function. Texas Tech University Health Science Technology.Google Scholar
McMeekan, C, Stafford, KJ, Mellor, DJ, Bruce, RA, Ward, RN, Gregory, NG 1999. Effects of a local anaesthetic and a non-steroidal anti-inflammatory analgesic on the behavioural responses of calves to dehorning. New Zealand Veterinary Journal 47, 9296.CrossRefGoogle Scholar
Megret, S, Rudeaux, F, Faure, J-M, Picard, M 1999. Rôles du bec chez les volailles. Conséquences du débecquage. INRA Productions Animales 9, 113119.CrossRefGoogle Scholar
Mellema, SC, Doherr, MG, Wechsler, B, Thuer, S, Steiner, A 2007. Influence of local anaesthesia on pain and distress induced by bloodless castration methods in young lambs. Schweizer Archiv für Tierheilkunde 149, 213225.CrossRefGoogle ScholarPubMed
Mellor, DJ, Thornber, AC, Bayvel, ACD, Kahn, S 2008. Scientific assessment and management of animal pain. OIE, Paris.Google Scholar
Mérour, I, Schwob, S, Hermesch, S, Larzul, C 2009. Effet du génotype halothane sur les performances de croissance, qualités de carcasse et de viande. TechniPorc 32, 913.Google Scholar
Minozzi, G, Guémené, D, Couty, M, Gourichon, D, Minvielle, F, Pinard-van der Laan, MH 2008. Circulating corticosterone reaction to restraint and adrenocorticotropin hormone administration in White Leghorns selected for immune response traits. Poultry Science 87, 22252230.CrossRefGoogle ScholarPubMed
Mirkena, T, Duguma, G, Haile, A, Tibbo, M, Okeyo, AM, Wurzinger, M, Sölkner, J 2010. Genetics of adaptation in domestic farm animals: a review. Livestock Science 132, 112.CrossRefGoogle Scholar
Misch, LJ, Duffield, TF, Millman, ST, Lissemore, KD 2007. An investigation into the practices of dairy producers and veterinarians in dehorning dairy calves in Ontario. Canadian Veterinary Journal 48, 12491254.Google ScholarPubMed
Moinard, C, Mendl, M, Nicol, CJ, Green, LE 2003. A case control study of on-farm risk factors for tail biting in pigs. Applied Animal Behaviour Science 81, 333355.CrossRefGoogle Scholar
Molony, V, Kent, JE 1997. Assessment of acute pain in farm animals using behavioral and physiological measurements. Journal of Animal Science 75, 266272.CrossRefGoogle ScholarPubMed
Molony, V, Kent, JE, Robertson, IS 1993. Behavioural responses of lambs of three ages in the first three hours after three methods of castration and tail docking. Research in Veterinary Science 55, 236245.CrossRefGoogle ScholarPubMed
Monin, G, Larzul, C, Roy, Pl, Culioli, J, Mourot, J, Rousset-Akrim, S, Talmant, A, Touraille, C, Sellier, P 1999. Effects of the halothane genotype and slaughter weight on texture of pork. Journal of Animal Science 77, 408415.CrossRefGoogle ScholarPubMed
Mormède, P, Foury, A, Terenina, E, Knap, PW 2011. Breeding for robustness: the role of cortisol. Animal 5, 651657.CrossRefGoogle ScholarPubMed
Morton, D 1992. Docking of dogs: practical and ethical aspects. The Veterinary Record 131, 301306.CrossRefGoogle ScholarPubMed
Mounier, L, Marie, M, Lensink, BJ 2007. Facteurs déterminants du bien-être des ruminants en élevage. INRA Productions Animales 20, 6572.CrossRefGoogle Scholar
Mrode, RA, Swanson, GJT, Winters, MS 1998. Genetic parameters and evaluations for somatic cell counts and its relationship with production and type traits in some dairy breeds in the United Kingdom. Animal Science 66, 569576.CrossRefGoogle Scholar
Muir, WM 1996. Group selection for adaptation to multiple-hen cages: selection program and direct responses. Poultry Science 75, 447458.CrossRefGoogle ScholarPubMed
Murray, AC, Johnson, CP 1998. Impact of the halothane gene on muscle quality and pre-slaughter deaths in Western Canadian pigs. Canadian Journal of Animal Science 78, 543548.CrossRefGoogle Scholar
Noonan, GJ, Rand, JS, Priest, J, Ainscow, J, Blackshaw, JK 1994. Behavioural observations of piglets undergoing tail docking, teeth clipping and ear notching. Applied Animal Behaviour Science 39, 203213.CrossRefGoogle Scholar
Penny, RHC, Hill, FWG 1974. Observations of some conditions in pigs at abattoir with particular reference to tail biting. Veterinary Record 94, 174180.CrossRefGoogle ScholarPubMed
Perez, MP, Palacio, J, Santolaria, MP, Acena, MC, Chacon, G, Gascon, M, Calvo, JH, Zaragoza, P, Beltran, JA, Garcia-Belenguer, S 2002. Effect of transport time on welfare and meat quality in pigs. Meat Science 61, 425433.CrossRefGoogle ScholarPubMed
PIGCAS 2008. Attitudes, practices and state of the art regarding piglet castration in Europe. Deliverable D4.1 Report on recommendations for research and policy support. 6. Framework Programme, p. 22.Google Scholar
Prayaga, KC 2007. Genetic options to replace dehorning in beef cattle: a review. Australian Journal of Agricultural Research 58, 18.CrossRefGoogle Scholar
Prunier, A, Mounier, AM, Hay, M 2005. Effects of castration, tooth resection, or tail docking on plasma metabolites and stress hormones in young pigs. Journal of Animal Science 83, 216222.CrossRefGoogle ScholarPubMed
Prunier, A, Gallois, M, Klouytten, A, Le Cozler, Y 2004. Effets de l’épointage des dents sur les performances, les lésions cutanées et le comportement des truies et des porcelets. Journées de la Recherche Porcine 36, 379388.Google Scholar
Prunier, A, Bonneau, M, VonBorell, EH, Cinotti, S, Gunn, M, Frediksen, B, Giersing, M, Morton, DB, Tuyttens, FAM, Velarde, A 2006. A review of the welfare consequences of surgical castration in piglets and the evaluation of non-surgical methods. Animal Welfare 15, 277289.CrossRefGoogle Scholar
Ranheim, B, Haga, HA, Ingebrigtsen, K 2005. Distribution of radioactive lidocaine injected into the testes in piglets. Journal of Veterinary Pharmacology and Therapeutics 28, 481483.CrossRefGoogle ScholarPubMed
Rauw, WM, Kanis, E, Noordhuizen-Stassen, EN, Grommers, FJ 1998. Undesirable side effects of selection for high production efficiency in farm animals: a review. Livestock Production Science 56, 1533.CrossRefGoogle Scholar
Ritter, MJ, Ellis, M, Bowman, R, Brinkmann, J, Curtis, SE, DeDecker, JM, Mendoza, O, Murphy, CM, Orellana, DG, Peterson, BA, Rojo, A, Schlipf, JM, Wolter, BF 2008. Effects of season and distance moved during loading on transport losses of market-weight pigs in two commercially available types of trailer. Journal of Animal Science 86, 31373145.CrossRefGoogle ScholarPubMed
Robertson, IS, Kent, JE, Molony, V 1994. Effect of different methods of castration on behaviour and plasma cortisol in calves of three ages. Research in Veterinary Science 56, 817.CrossRefGoogle ScholarPubMed
Rochard, O, Lubac, S, Aliner, A, Morin, H, Mirabito, L, Guéméné, D 2008. Traitement du bec par la technique infra rouge ou l’épointage aux ciseaux. Quelles conséquences ? 8. Journées de la Recherche sur les Palmipèdes à Foie Gras, 30–31 October 2008, Arcachon, pp. 165–169.Google Scholar
Rose, JD 2002. The neurobehavioral nature of fishes and the question of awareness and pain. Reviews in Fisheries Science 10, 138.CrossRefGoogle Scholar
Rose, JD 2007. Anthropomorphism and ‘mental welfare’ of fishes. Diseases of Aquatic Organisms 75, 139154.CrossRefGoogle ScholarPubMed
Rupp, R, Lagriffoul, G, Astruc, JM, Barillet, F 2003. Genetic parameters for milk somatic cell scores and relationships with production traits in French Lacaune dairy sheep. Journal of Dairy Science 86, 14761481.CrossRefGoogle ScholarPubMed
Russell, WMS, Burch, RL 1959. The principles of human experimental technique. Universities Federation for Animal Welfare (UFAW), Potters Bar, Herts, UK, 1992 (special edition).Google Scholar
Salak-Johnson, JL, McGlone, JJ 2007. Making sense of apparently conflicting data: stress and immunity in swine and cattle. Journal of Animal Science 85, E81E88.CrossRefGoogle ScholarPubMed
Schulz, C, Ritzmann, M, Palzer, A, Heinritzi, K, Zols, S 2007. Effect of isoflurane-anesthesia on postoperative pain due to castration of piglets. Berliner Und Munchener Tierarztliche Wochenschrift 120, 177182.Google ScholarPubMed
Sharma, V, Chand, D, Arneja, DV 1999. Feather pecking and cannibalism in poultry: causes and control measures. International Journal of Animal Science 14, 117121.Google Scholar
Sheridan, AK, Howlett, CR, Burton, RW 1978. The inheritance of tibial dyschondroplasia in broilers. British Poultry Science 19, 491499.CrossRefGoogle ScholarPubMed
Siegel, PB, Honaker, CF, Rauw, WM 2008. Selection for high production in poultry. In Resource allocation theory applied to farm animal production (ed. WM Rauw), pp. 230242. CAB International, Cambridge.CrossRefGoogle Scholar
Stafford, KJ, Mellor, DJ 2005. Dehorning and disbudding distress and its alleviation in calves. Veterinary Journal 169, 337349.CrossRefGoogle ScholarPubMed
Stafford, KJ, Mellor, DJ, Todd, SE, Bruce, RA, Ward, RN 2002. Effects of local anaesthesia or local anaesthesia plus a non-steroidal anti-inflammatory drug on the acute cortisol response of calves to five different methods of castration. Research in Veterinary Science 73, 6170.CrossRefGoogle ScholarPubMed
Stafford, KJ, Mellor, DJ, Todd, SE, Ward, RN, McMeekan, CM 2003. The effect of different combinations of lignocaine, ketoprofen, xylazine and tolazoline on the acute cortisol response to dehorning in calves. New Zealand Veterinary Journal 51, 219226.CrossRefGoogle ScholarPubMed
Star, L, Ellen, ED, Uitdehaag, K, Brom, FWA 2008. A plea to implement robustness into a breeding goal: poultry as an example. Journal of Agricultural & Environmental Ethics 21, 109125.CrossRefGoogle Scholar
Stewart, M, Stookey, JM, Stafford, KJ, Tucker, CB, Rogers, AR, Dowling, SK, Verkerk, GA, Schaefer, AL, Webster, JR 2009. Effects of local anesthetic and a nonsteroidal antiinflammatory drug on pain responses of dairy calves to hot-iron dehorning. Journal of Dairy Science 92, 15121519.CrossRefGoogle Scholar
Stilwell, G, Capitão, E, Nunes, T 2004. Effect of three different methods of dehorning on plasma cortisol levels and behaviour of calves. Proceedings of the 23rd World Buiatrics Congress, Quebec, 11–16 July 2004, Canada, p. 665.Google Scholar
Stilwell, G, Lima, MS, Broom, DM 2007. Comparing the effect of three different disbudding methods on behaviour and plasma cortisol of calves. Revista Portuguesa Ciencias Veteriniarias 102, 281288.Google Scholar
Stilwell, G, Lima, MS, Broom, DM 2008. Effects of nonsteroidal anti-inflammatory drugs on long-term pain in calves castrated by use of an external clamping technique following epidural anesthesia. American Journal of Veterinary Research 69, 744750.CrossRefGoogle ScholarPubMed
Stilwell, G, Campos de Carvalho, R, Caroline, N, Lima, MS, Broom, DM 2010. Effect of hot-iron disbudding on behaviour and plasma cortisol of calves sedated with xylazine. Research in Veterinary Science 88, 188193.CrossRefGoogle ScholarPubMed
Storskrubb, A, Sevón-Aimonen, M-L, Uimari, P 2010. Genetic parameters for bone strength, osteochondrosis and meat percentage in Finnish Landrace and Yorkshire pigs. Animal 4, 13191324.CrossRefGoogle ScholarPubMed
Stull, CL, Payne, MA, Berry, SL, Hullinger, PJ 2002. Evaluation of the scientific justification for tail docking in dairy cattle. Journal of the American Veterinary Medical Association 220, 12981303.CrossRefGoogle ScholarPubMed
Sylvester, SP, Mellor, DJ, Stafford, KJ, Bruce, RA, Ward, RN 1998. Acute cortisol responses of calves to scoop dehorning using local anaesthesia and/or cautery of the wound. Australian Veterinary Journal 76, 118122.CrossRefGoogle ScholarPubMed
Thuer, S, Doherr, MG, Wechsler, B, Mellema, SC, Nuss, K, Kirchhofer, M, Steiner, A 2007. Influence of local anaesthesia on short- and long-term pain induced in calves by three bloodless castration methods. Schweizer Archiv für Tierheilkunde 149, 201211.Google Scholar
Ting, AK, Earley, B, Veissier, I, Gupta, S, Crowe, MA 2005. Effects of age of Holstein–Friesian calves on plasma cortisol, acute-phase proteins, immunological function, scrotal measurements and growth in response to Burdizzo castration. Animal Science 80, 377386.CrossRefGoogle Scholar
Ting, ST, Earley, B, Crowe, MA 2003a. Effect of repeated ketoprofen administration during surgical castration of bulls on cortisol, immunological function, feed intake, growth, and behavior. Journal of Animal Science 81, 12531264.CrossRefGoogle ScholarPubMed
Ting, ST, Earley, B, Hughes, JM, Crowe, MA 2003b. Effect of ketoprofen, lidocaine local anesthesia, and combined xylazine and lidocaine caudal epidural anesthesia during castration of beef cattle on stress responses, immunity, growth, and behavior. Journal of Animal Science 81, 12811293.CrossRefGoogle ScholarPubMed
Ting, ST, Earley, B, Crowe, MA 2004. Effect of cortisol infusion patterns and castration on metabolic and immunological indices of stress response in cattle. Domestic Animal Endocrinology 26, 329349.CrossRefGoogle ScholarPubMed
Tucker, CB, Fraser, D, Weary, DM 2001. Tail docking dairy cattle: effects on cow cleanliness and udder health. Journal of Dairy Science 84, 8487.CrossRefGoogle ScholarPubMed
Turner, SP, D'Eath, RB, Roehe, R, Lawrence, AB 2010. Selection against aggressiveness in pigs at re-grouping: practical application and implications for long-term behavioural patterns. Animal Welfare 19, 123132.CrossRefGoogle Scholar
Turner, SP, Roehe, R, Mekkawy, W, Farnworth, MJ, Knap, PW, Lawrence, AB 2008. Bayesian analysis of genetic associations of skin lesions and behavioural traits to identify genetic components of individual aggressiveness in pigs. Behavior Genetics 38, 6775.CrossRefGoogle ScholarPubMed
Turner, SP, Farnworth, MJ, White, IMS, Brotherstone, S, Mendl, M, Knap, P, Penny, P, Lawrence, AB 2006a. The accumulation of skin lesions and their use as a predictor of individual aggressiveness in pigs. Applied Animal Behaviour Science 96, 245259.CrossRefGoogle Scholar
Turner, SP, White, IMS, Brotherstone, S, Farnworth, MJ, Knap, PW, Penny, P, Mendl, M, Lawrence, AB 2006b. Heritability of post-mixing aggressiveness in grower-stage pigs and its relationship with production traits. Animal Science 82, 615620.CrossRefGoogle Scholar
Turner, SP, Roehe, R, D'Eath, RB, Ison, SH, Farish, M, Jack, MC, Lundeheim, N, Rydhmer, L, Lawrence, AB 2009. Genetic validation of postmixing skin injuries in pigs as an indicator of aggressiveness and the relationship with injuries under more stable social conditions. Journal of Animal Science 87, 30763082.CrossRefGoogle ScholarPubMed
Vanhonacker, F, Verbeke, W, Tuyttens, FAM 2009. Belgian consumers’ attitude towards surgical castration and immunocastration of piglets. Animal Welfare 18, 371380.CrossRefGoogle Scholar
Veerkamp, RF, Windig, JJ, Calus, MPL, Ouweltjes, W, de Haas, Y, Beerda, B 2008. Selection for high production in dairy cattle. In Resource allocation theory applied to farm animal production (ed. WM Rauw), pp. 243260. CAB International.CrossRefGoogle Scholar
von Borell, E, Baumgartner, J, Giersing, M, Jäggin, N, Prunier, A, Tuyttens, FAM, Edwards, SA 2009. Animal welfare implications of surgical castration and its alternatives in pigs. Animal 3, 14881496.CrossRefGoogle ScholarPubMed
Webster, AJF 2002. Rendering unto Caesar: welfare problems in Belgian Blue cattle. Veterinary Journal 163, 228229.CrossRefGoogle ScholarPubMed
White, RG, DeShazer, JA, Tressler, CJ, Borcher, GM, Davey, S, Waninge, A, Parkhurst, AM, Milanuk, MJ, Clemens, ET 1995. Vocalization and physiological response of pigs during castration with or without a local anesthetic. Journal of Animal Science 73, 381386.CrossRefGoogle ScholarPubMed
Willam, A, Egger-Danner, C, Sölkner, J, Gierzinger, E 2002. Optimization of progeny testing schemes when functional traits play an important role in the total merit index. Livestock Production Science 77, 217225.CrossRefGoogle Scholar
Yazdi, MH, Rydhmer, L, Ringmar-Cederberg, E, Lundeheim, N, Johansson, K 2000. Genetic study of longevity in Swedish Landrace sows. Livestock Production Science 63, 255264.CrossRefGoogle Scholar
Zamaratskaia, G, Squires, EJ 2009. Biochemical, nutritional and genetic effects on boar taint in entire male pigs. Animal 3, 15081521.CrossRefGoogle ScholarPubMed
Zonderland, JJ, Wolthuis-Fillerup, M, Van Reenen, CG, Bracke, MBM, Kemp, B, Den Hartog, LA, Spoolder, HAM 2008. Prevention and treatment of tail biting in weaned piglets. Applied Animal Behaviour Science 110, 269281.CrossRefGoogle Scholar