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The prevention and control of feather pecking in laying hens: identifying the underlying principles

Published online by Cambridge University Press:  28 June 2013

T.B. RODENBURG ,
M.M. VAN KRIMPEN ,
I.C. DE JONG ,
E.N. DE HAAS ,
M.S. KOPS ,
B.J. RIEDSTRA ,
R.E. NORDQUIST ,
J.P. WAGENAAR ,
M. BESTMAN  and
C.J. NICOL
T.B. RODENBURG*
Affiliation:
Animal Breeding and Genomics Centre, Wageningen University, PO Box 338, 6700 AH Wageningen, The Netherlands Behavioural Ecology Group, Wageningen University, PO Box 338, 6700 AH Wageningen, The Netherlands
M.M. VAN KRIMPEN
Affiliation:
Livestock Research, Wageningen UR, PO Box 65, 8200 AB, Lelystad, The Netherlands
I.C. DE JONG
Affiliation:
Livestock Research, Wageningen UR, PO Box 65, 8200 AB, Lelystad, The Netherlands
E.N. DE HAAS
Affiliation:
Adaptation Physiology Group, Wageningen University, PO Box 338, 6700 AH Wageningen, The Netherlands
M.S. KOPS
Affiliation:
Department of Psychopharmacology, Utrecht Institute for Pharmaceutical Sciences (UIPS) and Rudolf Magnus Institute of Neuroscience, Utrecht University, PO Box 80.082, 3508 TB Utrecht, The Netherlands
B.J. RIEDSTRA
Affiliation:
Behavioural Biology, University of Groningen, Nijenborgh 7, 9747 AG Groningen, The Netherlands
R.E. NORDQUIST
Affiliation:
Emotion & Cognition Group, Department of Farm Animal Health, Faculty of Veterinary Medicine and Rudolf Magnus Institute of Neuroscience, Utrecht University, Yalelaan 7, 3584 CL Utrecht, The Netherlands
J.P. WAGENAAR
Affiliation:
Louis Bolk Institute, Hoofdstraat 24, 3972 LA Driebergen, The Netherlands
M. BESTMAN
Affiliation:
Louis Bolk Institute, Hoofdstraat 24, 3972 LA Driebergen, The Netherlands
C.J. NICOL
Affiliation:
Animal Welfare & Behaviour, School of Veterinary Sciences, Bristol University, Langford House, Langford, Bristol, BS40 5DU, United Kingdom
*
Corresponding author: bas.rodenburg@wur.nl
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Abstract

Feather pecking (FP) in laying hens remains an important economic and welfare issue. This paper reviews the literature on causes of FP in laying hens. With the ban on conventional cages in the EU from 2012 and the expected future ban on beak trimming in many European countries, addressing this welfare issue has become more pressing than ever. The aim of this review paper is to provide a detailed overview of underlying principles of FP. FP is affected by many different factors and any approach to prevent or reduce FP in commercial flocks should acknowledge that fact and use a multifactorial approach to address this issue. Two forms of FP can be distinguished: gentle FP and severe FP. Severe FP causes the most welfare issues in commercial flocks. Severe FP is clearly related to feeding and foraging behaviour and its development seems to be enhanced in conditions where birds have difficulty in coping with environmental stressors. Stimulating feeding and foraging behaviour by providing high-fibre diets and suitable litter from an early age onwards, and controlling fear and stress levels through genetic selection, reducing maternal stress and improving the stockmanship skills of the farmer, together offer the best prospect for preventing or controlling FP.

Keywords

laying henfeather peckingontogenyfearfulnessenvironmental enrichmentforaging
Type
Review Article
Information
World's Poultry Science Journal , Volume 69 , Issue 2 , June 2013 , pp. 361 - 374
Copyright
Copyright © World's Poultry Science Association 2013

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References

AL BUSTANY, Z. and ELWINGER, K. (1987a) Comparison between barley/fish meal- and maize/soybean meal-based diets with various lysine and protein levels fed to different strains of laying hens. Acta Agriculturae Scandinavica 37: 41-49.CrossRefGoogle Scholar
AL BUSTANY, Z. and ELWINGER, K. (1987b) Response of laying hens to different dietary lysine intakes. A comparison of some commercial hybrids with strains selected on a low protein diet. Acta Agriculturae Scandinavica 37: 27-40.CrossRefGoogle Scholar
ALBENTOSA, M.J., KJAER, J.B. and NICOL, C.J. (2003) Strain and age differences in behaviour, fear response and pecking tendency in laying hens. British Poultry Science 44: 333-344.CrossRefGoogle ScholarPubMed
AMBROSEN, T. and PETERSEN, V.E. (1997) The influence of protein level in the diet on cannibalism and quality of plumage of layers. Poultry Science 76: 559-563.CrossRefGoogle ScholarPubMed
BLOKHUIS, H.J. (1989) The development and causation of feather pecking in the domestic fowl. - Vakgroep Ethologie, Thesis Landbouw Universiteit Wageningen, Wageningen.Google Scholar
BLOKHUIS, H.J. and BEUTLER, A. (1992) Feather pecking damage and tonic immobility response in two lines of white leghorn hens. Journal of Animal Science 70: 170.Google Scholar
BLOKHUIS, H.J. and VAN DER HAAR, J.W. (1992) Effects of pecking incentives during rearing on feather pecking of laying hens. British Poultry Science 33: 17-24.CrossRefGoogle ScholarPubMed
BOLHUIS, J.E., ELLEN, E.D., VAN REENEN, C.G., DE GROOT, J., TEN NAPEL, J., KOOPMANSCHAP, R., DE VRIES REILINGH, G., UITDEHAAG, K.A., KEMP, B. and RODENBURG, T.B. (2009) Effects of genetic group selection against mortality on behaviour and peripheral serotonin in domestic laying hens with trimmed and intact beaks. Physiology & Behavior 97: 470-475.CrossRefGoogle ScholarPubMed
CAMPO, J.L., PRIETO, M.T. and DÁVILA, S.G. (2008) Association between vent pecking and fluctuating asymmetry, heterophil to lymphocyte ratio, and tonic immobility duration in chickens. Applied Animal Behaviour Science 113: 87-97.CrossRefGoogle Scholar
CHOW, A. and HOGAN, J.A. (2005) The development of feather pecking in Burmese red junglefowl: the influence of early experience with exploratory-rich environments. Applied Animal Behaviour Science 93: 283-294.CrossRefGoogle Scholar
DE HAAS, E.N., NIELSEN, B.L., BUITENHUIS, A.J. and RODENBURG, T.B. (2010) Selection on feather pecking affects response to novelty and foraging behaviour in laying hens. Applied Animal Behaviour Science 124: 90-96.CrossRefGoogle Scholar
DE JONG, I.C., GUNNINK, H., ROMMERS, J.M. and BRACKE, M.B.M. (2013) Effect of substrate during early rearing on floor- and feather pecking behaviour in young and adult laying hens. Archiv fur Geflugelkunde 77: 15-22.Google Scholar
DE JONG, I.C., REUVEKAMP, B.M., GUNNINK, H. (submitted) Exposure to substrate in early rearing and the development of (feather) pecking behaviour in laying hens.Google Scholar
DIXON, L.M. and DUNCAN, I.J.H. (2010) Changes in substrate access did not affect early feather-pecking behavior in two strains of laying hen chicks. Journal of Applied Animal Welfare Science 13: 1-14.CrossRefGoogle Scholar
EL-LETHEY, H., JUNGI, T.W. and HUBER-EICHER, B. (2001) Effects of feeding corticosterone and housing conditions on feather pecking in laying hens (Gallus gallus domesticus). Physiology & Behavior 73: 243-251.CrossRefGoogle ScholarPubMed
ELLEN, E.D., MUIR, W.M. and BIJMA, P. (2007) Genetic improvement of traits affected by interactions among individuals: sib selection schemes. Genetics 176: 489-499.CrossRefGoogle ScholarPubMed
ELWINGER, K., TAUSON, R., TUFVESSON, M. and HARTMANN, C. (2002) Feeding of layers kept in an organic feed environment. 11th. European Poultry Conference, Bremen.Google Scholar
ESMAIL, S.H.M. (1997) Fibre nutrition. Poultry International 36: 31-34.Google Scholar
FREIRE, R., VAN DORT, S. and ROGERS, L.J. (2006) Pre- and post-hatching effects of corticosterone treatment on behavior of the domestic chick. Hormones and Behavior 49: 157-165.CrossRefGoogle ScholarPubMed
GLATZ, P.C. and BOURKE, M. (2006) Chapter 1. Pecking problems, in: Beak trimming handbook for egg producers: best practices for minimising cannibalism in poultry, pp. 1-5. (Collingwood Victoria, Australia, CSIRO Publishing).Google Scholar
GROOTHUIS, T.G.G., MULLER, W., VON ENGELHARDT, N., CARERE, C. and EISING, C. (2005) Maternal hormones as a tool to adjust offspring phenotype in avian species. Neuroscience and Biobehavioral Reviews 29: 329-352.CrossRefGoogle ScholarPubMed
GRUSS, M. and BRAUN, K. (1997) Distinct activation of monoaminergic pathways in chick brain in relation to auditory imprinting and stressful situations: A microdialysis study. Neuroscience 76: 891-899.CrossRefGoogle ScholarPubMed
HARLANDER-MATAUSCHEK, A., BENDA, I., LAVETTI, C., DJUKIC, M. and BESSEI, W. (2007) The relative preferences for wood shavings or feathers in high and low feather pecking birds. Applied Animal Behaviour Science 107: 78-87.CrossRefGoogle Scholar
HARLANDER-MATAUSCHEK, A. and HÄUSLER, K. (2009) Understanding feather eating behaviour in laying hens. Applied Animal Behaviour Science 117: 35-41.CrossRefGoogle Scholar
HARLANDER-MATAUSCHEK, A., PIEPHO, H.P. and BESSEI, W. (2006) The effect of feather eating on feed passage in laying hens. Poultry Science 85: 21-25.CrossRefGoogle ScholarPubMed
HEMSWORTH, P.H. and BARNETT, J.L. (1989) Relationships between fear of humans, productivity and cage position of laying hens. British Poultry Science 30: 505-518.CrossRefGoogle ScholarPubMed
HETLAND, H., CHOCT, M. and SVIHUS, B. (2004) Role of insoluble non-starch polysaccharides in poultry nutrition. World's Poultry Science Journal 60: 415-422.CrossRefGoogle Scholar
HOCKING, P.M., CHANNING, C.E., WADDINGTON, D. and JONES, R.B. (2001) Age-related changes in fear, sociality and pecking behaviours in two strains of laying hen. British Poultry Science 42: 414-423.CrossRefGoogle ScholarPubMed
HUBER-EICHER, B. and WECHSLER, B. (1997) Feather pecking in domestic chicks: its relation to dustbathing and foraging. Animal Behaviour 54: 757-768.CrossRefGoogle ScholarPubMed
HUGHES, A.L. and BUITENHUIS, A.J. (2010) Reduced variance of gene expression at numerous loci in a population of chickens selected for high feather pecking. Poultry Science 89: 1858-1869.CrossRefGoogle Scholar
HUGHES, B.O. (1973) The effect of implanted gonadal hormones on feather pecking and cannibalism in pullets. British Poultry Science 14: 341-348.CrossRefGoogle ScholarPubMed
HUGHES, B.O. and DUNCAN, I.J.H. (1972) The influence of strain and environmental factors upon feather pecking and cannibalism in fowls. British Poultry Science 13: 525-547.CrossRefGoogle ScholarPubMed
HUGHES, B.O. and WHITEHEAD, C.C. (1974) Sodium deprivation, feather pecking and activity in laying hens. British Poultry Science 15: 435-439.CrossRefGoogle ScholarPubMed
JANCZAK, A.M., TORJESEN, P., PALME, R. and BAKKEN, M. (2007) Effects of stress in hens on the behaviour of their offspring. Applied Animal Behaviour Science 107: 66-77.CrossRefGoogle Scholar
JENSEN, P., KEELING, L., SCHUTZ, K., ANDERSSON, L., MORMEDE, P., BRANDSTROM, H., FORKMAN, B., KERJE, S., FREDRIKSSON, R. and OHLSSON, C. (2005) Feather pecking in chickens is genetically related to behavioural and developmental traits. Physiology & Behavior 86: 52-60.CrossRefGoogle ScholarPubMed
JOHNSEN, P.F., VESTERGAARD, K.S. and NORGAARD NIELSEN, G. (1998) Influence of early rearing conditions on the development of feather pecking and cannibalism in domestic fowl. Applied Animal Behaviour Science 60: 25-41.CrossRefGoogle Scholar
JONES, R.B. (1996) Fear and adaptability in poultry: insights, implications and imperatives. World's Poultry Science Journal 52: 131-174.CrossRefGoogle Scholar
JONES, R.B., BLOKHUIS, H.J. and BEUVING, G. (1995) Open-field and tonic immobility responses in domestic chicks of two genetic lines differing in their propensity to feather peck. British Poultry Science 36: 525-530.CrossRefGoogle ScholarPubMed
KJAER, J.B. and GUÉMENÉ, D. (2009) Adrenal reactivity in lines of domestic fowl selected on feather pecking behavior. Physiology & Behavior 96: 370-373.CrossRefGoogle ScholarPubMed
KJAER, J.B., HJARVARD, B.M., JENSEN, K.H., HANSEN-MOLLER, J. and NAESBYE LARSEN, O. (2004) Effects of haloperidol, a dopamine D2 receptor antagonist, on feather pecking behaviour in laying hens. Applied Animal Behaviour Science 86: 77-91.CrossRefGoogle Scholar
KJAER, J.B. and JORGENSEN, H. (2011) Heart rate variability in domestic chicken lines genetically selected on feather pecking behavior. Genes Brain and Behavior 10: 747-755.CrossRefGoogle ScholarPubMed
KJAER, J.B., SØRENSEN, P. and SU, G. (2001) Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus). Applied Animal Behaviour Science 71: 229-239.CrossRefGoogle ScholarPubMed
KORTE, S.M. (2001) Corticosteroids in relation to fear, anxiety and psychopathology. Neuroscience & Biobehavioral Reviews 25: 117-142.CrossRefGoogle ScholarPubMed
KORTE, S.M., BEUVING, G., RUESINK, W. and BLOKHUIS, H.J. (1997) Plasma catecholamine and corticosterone levels during manual restraint in chicks from a high and low feather pecking line of laying hens. Physiology & Behavior 62: 437-441.CrossRefGoogle ScholarPubMed
KORTE, S.M., RUESINK, W. and BLOKHUIS, H.J. (1999) Heart rate variability during manual restraint in chicks from high- and low-feather pecking lines of laying hens. Physiology & Behavior 65: 649-652.CrossRefGoogle ScholarPubMed
LAMBTON, S.L., KNOWLES, T.G., YORKE, C. and NICOL, C.J. (2007) Gentle and severe feather pecking: one problem or two? Proceedings of the 41st International Congress of the ISAE, Merida. Mexico, pp. 48Google Scholar
LEONARD, M.L., HORN, A.G. and FAIRFULL, R.W. (1995) Correlates and consequences of allopecking in White Leghorn chickens. Applied Animal Behaviour Science 43: 17-26.CrossRefGoogle Scholar
MAHBOUB, H.D.H., MÜLLER, J. and VON BORELL, E. (2004) Outdoor use, tonic immobility, heterophil/lymphocyte ration and feather condition in free-range laying hens of different genotype. British Poultry Science 45: 738-744.CrossRefGoogle Scholar
MCADIE, T.M. and KEELING, L.J. (2002) The social transmission of feather pecking in laying hens: effects of environment and age. Applied Animal Behaviour Science 75: 147-159.CrossRefGoogle Scholar
MCADIE, T.M., KEELING, L.J., BLOKHUIS, H.J. and JONES, R.B. (2005) Reduction in feather pecking and improvement of feather condition with the presentation of a string device to chickens. Applied Animal Behaviour Science 93: 67-80.CrossRefGoogle Scholar
MCKEEGAN, D.E.F. and SAVORY, C.J. (1999) Feather eating in layer pullets and its possible role in the aetiology of feather pecking damage. Applied Animal Behaviour Science 65: 73-85.CrossRefGoogle Scholar
METZGER, M., TOLEDO, C. and BRAUN, K. (2002) Serotonergic innervation of the telencephalon in the domestic chick. Brain Research Bulletin 57: 547-551.CrossRefGoogle ScholarPubMed
NEWBERRY, R.C. (2004) Cannibalism, in: PERRY, G.C. (Ed.) Welfare of the laying hen, pp. 239-258 (Wallingford, United Kingdom, CABI Publishing).Google Scholar
NEWBERRY, R.C., KEELING, L.J., ESTEVEZ, I. and BILCIK, B. (2007) Behaviour when young as a predictor of severe feather pecking in adult laying hens: The redirected foraging hypothesis revisited. Applied Animal Behaviour Science 107: 262-274.CrossRefGoogle Scholar
NICOL, C.J., LINDBERG, A.C., PHILLIPS, A.J., POPE, S.J., WILKINS, L.J. and GREEN, L.E. (2001) Influence of prior exposure to wood shavings on feather pecking, dustbathing and foraging in adult laying hens. Applied Animal Behaviour Science 73: 141-155.CrossRefGoogle ScholarPubMed
NRC, (1994) Nutrient Requirements of Poultry. 9th rev. ed. Natl. Acad. Press, Washington, DC.Google Scholar
RAMADAN, S.G.A. and VON BORELL, E. (2008) Role of loose feathers on the development of feather pecking in laying hens. British Poultry Science 49: 250-256.CrossRefGoogle ScholarPubMed
RIEDSTRA, B. and GROOTHUIS, T.G.G. (2002) Early feather pecking as a form of social exploration: the effect of group stability on feather pecking and tonic immobility in domestic chicks. Applied Animal Behaviour Science 77: 127-138.CrossRefGoogle Scholar
RODENBURG, T.B., BOLHUIS, J.E., KOOPMANSCHAP, R.E., ELLEN, E.D. and DECUYPERE, E. (2009a) Maternal care and selection for low mortality affect post-stress corticosterone and peripheral serotonin in laying hens. Physiology & Behavior 98: 519-523.CrossRefGoogle ScholarPubMed
RODENBURG, T.B., BUITENHUIS, A.J., ASK, B., UITDEHAAG, K.A., KOENE, P., VAN DER POEL, J.J., VAN ARENDONK, J.A.M. and BOVENHUIS, H. (2004a) Genetic and phenotypic correlations between feather pecking and open-field response in laying hens at two different ages. Behavior Genetics 34: 407-415.CrossRefGoogle ScholarPubMed
RODENBURG, T.B., DE HAAS, E.N., NIELSEN, B.L. and BUITENHUIS, A.J. (2010) Fearfulness and feather damage in laying hens divergently selected for high and low feather pecking. Applied Animal Behaviour Science 128: 91-96.CrossRefGoogle Scholar
RODENBURG, T.B., DE REU, K. and TUYTTENS, F.A.M. (2012) Performance, welfare, health and hygiene of laying hens in non-cage systems in comparison with cage systems, in: SANDILANDS, V. & HOCKING, P. (Eds) Alternative Systems for Poultry – Health, Welfare and Productivity, pp. 210-224 (Glasgow, United Kingdom).Google Scholar
RODENBURG, T.B. and KOENE, P. (2003) Comparison of individual and social feather pecking tests in two lines of laying hens at ten different ages. Applied Animal Behaviour Science 81: 133-148.CrossRefGoogle Scholar
RODENBURG, T.B., KOMEN, H., ELLEN, E.D., UITDEHAAG, K.A. and VAN ARENDONK, J.A.M. (2008) Selection method and early-life history affect behavioural development, feather pecking and cannibalism in laying hens: A review. Applied Animal Behaviour Science 110: 217-228.CrossRefGoogle Scholar
RODENBURG, T.B., UITDEHAAG, K.A., ELLEN, E.D. and KOMEN, J. (2009b) The effects of selection on low mortality and brooding by a mother hen on open-field response, feather pecking and cannibalism in laying hens. Animal Welfare 18: 427-432.CrossRefGoogle Scholar
RODENBURG, T.B., VAN HIERDEN, Y.M., BUITENHUIS, A.J., RIEDSTRA, B., KOENE, P., KORTE, S.M., VAN DER POEL, J.J., GROOTHUIS, T.G.G. and BLOKHUIS, H.J. (2004b) Feather pecking in laying hens: new insights and directions for research? Applied Animal Behaviour Science 86: 291-298.CrossRefGoogle Scholar
SAVORY, C.J. (1980) Meal occurrence in Japanese quail in relation to particle size and nutrient density. Animal Behaviour 28: 160-171.CrossRefGoogle Scholar
SAVORY, C.J. (1995) Feather pecking and cannibalism. World's Poultry Science Journal 51: 215-219.CrossRefGoogle Scholar
SCHAIBLE, P.J., DAVIDSON, J.A. and BANDEMER, S.L. (1947) Cannibalism and feather pecking in chicks as influenced by certain changes in a specific ration. Poultry Science 26: 651-656.CrossRefGoogle Scholar
SCHWABL, H. (1993) Yolk is a source of maternal testosterone for developing birds. Proceedings Of The National Academy Of Sciences Of The United States Of America 90: 11446-11450.CrossRefGoogle Scholar
SUNDE, M.L. (1972) Zinc requirement for normal feathering of commercial Leghorn-type pullets. Poultry Science 51: 1316-1322.CrossRefGoogle ScholarPubMed
UITDEHAAG, K., KOMEN, H., RODENBURG, T.B., KEMP, B. and VAN ARENDONK, J. (2008a) The novel object test as predictor of feather damage in cage-housed Rhode Island Red and White Leghorn laying hens. Applied Animal Behaviour Science 109: 292-305.CrossRefGoogle Scholar
UITDEHAAG, K.A., RODENBURG, T.B., BOLHUIS, J.E., DECUYPERE, E. and KOMEN, H. (2009) Mixed housing of different genetic lines of laying hens negatively affects feather pecking and fear related behaviour. Applied Animal Behaviour Science 116: 58-66.CrossRefGoogle Scholar
UITDEHAAG, K.A., RODENBURG, T.B., KOMEN, H., KEMP, B. and VAN ARENDONK, J.A.M. (2008b) The association of response to a novel object with subsequent performance and feather damage in adult, cage-housed, pure-bred Rhode Island Red laying hens. Poultry Science 87: 2486-2492.CrossRefGoogle ScholarPubMed
VAN DER LEE, A.G., HEMKE, G. and KWAKKEL, R.P. (2001) Low density diets improve plumage condition in non-debeaked layers. Proceedings of the 13th European Symposium on Poultry Nutrition, Blankenbergen, Belgium, pp. 244-245Google Scholar
VAN HIERDEN, Y.M., DE BOER, S.F., KOOLHAAS, J.M. and KORTE, S.M. (2004a) The control of feather pecking by serotonin. Behavioral Neuroscience 118: 575-583.CrossRefGoogle ScholarPubMed
VAN HIERDEN, Y.M., KOOLHAAS, J.M. and KORTE, S.M. (2004b) Chronic increase of dietary -tryptophan decreases gentle feather pecking behaviour. Applied Animal Behaviour Science 89: 71-84.CrossRefGoogle Scholar
VAN HIERDEN, Y.M., KOOLHAAS, J.M., KOST'AL, L.U., VYBOH, P., SEDLACKOVA, M., RAJMAN, M., JURANI, M. and MECHIEL KORTE, S. (2005) Chicks from a high and low feather pecking line of laying hens differ in apomorphine sensitivity. Physiology & Behavior 84: 471-477.CrossRefGoogle ScholarPubMed
VAN HIERDEN, Y.M., KORTE, S.M., RUESINK, E.W., VAN REENEN, C.G., ENGEL, B., KOOLHAAS, J.M. and BLOKHUIS, H.J. (2002a) The development of feather pecking behaviour and targeting of pecking in chicks from a high and low feather pecking line of laying hens. Applied Animal Behaviour Science 77: 183-196.CrossRefGoogle Scholar
VAN HIERDEN, Y.M., KORTE, S.M., RUESINK, E.W., VAN REENEN, C.G., ENGEL, B., KORTE-BOUWS, G.A.H., KOOLHAAS, J.M. and BLOKHUIS, H.J. (2002b) Adrenocortical reactivity and central serotonin and dopamine turnover in young chicks from a high and low feather-pecking line of laying hens. Physiology & Behavior 75: 653-659.CrossRefGoogle ScholarPubMed
VAN HORNE, P.L.M. and ACHTERBOSCH, T.J. (2008) Animal welfare in poultry production systems: impact of EU standards on world trade. World's Poultry Science Journal 64: 40-52.CrossRefGoogle Scholar
VAN KRIMPEN, M.M., BINNENDIJK, G.P. and VAN DIEPEN, J.T.M. (2012) Effect of diluted practical diets on plumage condition, behaviour and performance of rearing and laying hens. Report 534 Wageningen UR Livestock Research Lelystad The Netherlands. pp. 1-47.Google Scholar
VAN KRIMPEN, M.M., DE BRUIN, W.W., DE VEER, R. and BINNENDIJK, G.P. (2010) Effect of diluted NSP-high diets under feather pecking prone conditions on the development of feather damage and performance of rearing and laying hens. Report 420. Wageningen UR Livestock Research, Lelystad, The Netherlands. pp. 1-46.Google Scholar
VAN KRIMPEN, M.M., KWAKKEL, R.P., ANDRÉ, G., VAN DER PEET-SCHWERING, C.M.C., DEN HARTOG, L.A. and VERSTEGEN, M.W.A. (2007) Effect of nutrient dilution on feed intake, eating time and performance of hens in early lay. British Poultry Science 48: 389-398.CrossRefGoogle ScholarPubMed
VAN KRIMPEN, M.M., KWAKKEL, R.P., REUVEKAMP, B.F.J., VAN DER PEET-SCHWERING, C.M.C., DEN HARTOG, L.A. and VERSTEGEN, M.W.A. (2005) Impact of feeding management on feather pecking in laying hens. World's Poultry Science Journal 61: 663-685.CrossRefGoogle Scholar
VAN KRIMPEN, M.M., KWAKKEL, R.P., VAN DER PEET-SCHWERING, C.M.C., DEN HARTOG, L.A. and VERSTEGEN, M.W.A. (2009) Effects of nutrient dilution and nonstarch polysaccharide concentration in rearing and laying diets on eating behavior and feather damage of rearing and laying hens. Poultry Science 88: 759-773.CrossRefGoogle ScholarPubMed
VAN KRIMPEN, M.M., KWAKKEL, R.P., VAN DER PEET-SCHWERING, C.M.C., DEN HARTOG, L.A. and VERSTEGEN, M.W.A. (2011) Effects of dietary energy concentration, nonstarch polysaccharide concentration, and particle sizes of nonstarch polysaccharides on digesta mean retention time and gut development in laying hens. British Poultry Science 52: 730-741.CrossRefGoogle ScholarPubMed
VESTERGAARD, K.S., KRUIJT, J.P. and HOGAN, J.A. (1993) Feather pecking and chronic fear in groups of red junglefowl: their relations to dustbathing, rearing environment and social status. Animal Behaviour 45: 1127-1140.CrossRefGoogle Scholar
VESTERGAARD, K.S. and LISBORG, L. (1993) A model of feather pecking development which relates to dustbathing in the fowl. Behaviour 126: 291-308.CrossRefGoogle Scholar
VON ENGELHARDT, N. and GROOTHUIS, T.G.G. (2011) Maternal hormones in avian eggs, in: NORRIS, D.O. & LOPEZ, K.H. (Eds) Hormones and Reproduction of Vertebrates, Volume 4: Birds, pp. 91-127 (Academic Press).Google Scholar
WEBSTER, A.B. (1995) Immediate and subsequent effects of a short fast on the behavior of laying hens. Applied Animal Behaviour Science 45: 255-266.CrossRefGoogle Scholar