Hen welfare in different housing systems

Poultry Science

Volumn 90, Issue 1, 2011, Pages 278-294

  Lay, D C ^a   Fulton, R M ^b   Hester, P Y ^c   Karcher, D M ^b   Kjaer, J B ^d   Mench, J A ^e   Mullens, B A ^f   Newberry, R C ^g   Nicol, C J ^h   O'Sullivan, N P ⁱ   Porter, R E ^j  

^a AGRICULTURAL RESEARCH SERVICE   (United States)

^b MICHIGAN STATE UNIVERSITY   (United States)

^c PURDUE UNIVERSITY   (United States)

^d FEDERAL RESEARCH INSTITUTE FOR ANIMAL HEALTH   (Germany)

^e UNIVERSITY OF CALIFORNIA   (United States)

^f UNIVERSITY OF CALIFORNIA   (United States)

^g WASHINGTON STATE UNIVERSITY   (United States)

^h UNIVERSITY OF BRISTOL   (United Kingdom)

ⁱ Hy Line International   (United States)

^j UNIVERSITY OF MINNESOTA   (United States)

more..

Author keywords

Alternative; Health; Housing; Poultry; Welfare

Indexed keywords

EID: 78650599953     PISSN: 00325791     EISSN: 15253171     Source Type: Journal    
DOI: 10.3382/ps.2010-00962     Document Type: Article

References (203)

1. Abasht, B., E. Beach, J. Arango, P. Settar, J. E. Fulton, N. P. O'Sullivan, A. Hassen, D. Habier, R. L. Fernando, J. C. M. Dekkers, and S. J. Lamont. 2009. Extent and consistency of linkage disequilibrium and identification of DNA markers for production and egg quality traits in commercial layer chicken populations. Proc. PAG XVII Conference, San Diego, CA. Scherago International, Jersey City, NJ.
2. Abrahamsson, P., and R. Tauson. 1995. Aviary systems and conventional cages for laying hens: Effects on production, egg quality, health and bird location in three hybrids. Acta Agric. Scand. A 45:191-203.
3. Abrahamsson, P., and R. Tauson. 1997. Effects of group size on performance, health and birds' use of facilities in furnished cages for laying hens. Acta Agric. Scand. A 45:191-203.
4. Abrahamsson, P., R. Tauson, and M. C. Appleby. 1996. Behaviour, health and integument of four hybrids of laying hens in modified and conventional cages. Br. Poult.Sci. 37:521-540.
5. Aerni, V., H. El-Lethey, and B. Wechsler. 2000. Effect of foraging material and food form on feather pecking in laying hens. Br. Poult.Sci. 41:16-21.
6. Ahmad, A., T. G. Nagaraja, and L. Zurek. 2007. Transmission of Escherichia coli O157-H7 to cattle by house flies. Prev. Vet. Med. 80:74-81.
7. American Veterinary Medical Association. 2009. A comparison of conventional cage, furnished cage, and non-cage (barn and outdoor/free range) systems for housing laying hens. Accessed June 2010. http://www.avma.org/issues/animal_welfare/cage_non-cage_systems.asp.
8. Anderson, K. E. 2002. Final report of the thirty fourth North Carolina layer performance and management test. Cooperative Extension Service, North Carolina State University, Raleigh.
9. Appleby, M. C., I. J. H. Duncan, and H. E. McCrae. 1988. Perching and floor laying by domestic hens: Experimental results and their commercial application. Br. Poult.Sci. 29:351-357.
10. Appleby, M. C., H. E. McRae, and B. E. Peitz. 1984. The effect of light on the choice of nests by domestic hens. Appl. Anim. Ethol. 11:249-254.
11. Appleby, M. C., J. A. Mench, and B. O. Hughes. 2004. Poultry Behaviour and Welfare. CABI Publishing, Wallingford, UK.
12. Appleby, M. C., A. W. Walker, C. J. Nichol, A. C. Lindberg, R. Freire, B. O. Hughes, and H. A. Elson. 2002. Development of furnished cages for laying hens. Br. Poult.Sci. 43:489-500.
13. Axtell, R. C., and J. J. Arends. 1990. Ecology and management of arthropod pests of poultry. Annu. Rev. Entomol. 35:101-126.
14. Barnett, J. L., P. C. Glatz, E. A. Newman, and G. M. Cronin. 1997. Effects of modifying layer cages with perches on stress physiology, plumage, pecking and bone strength of hens. Aust. J. Exp. Agric. 37:523-529.
15. Bestman, M. W. P., and J. P. Wagenaar. 2003. Farm level factors associated with feather pecking in organic laying hens. Livest. Prod. Sci. 80:133-140.
16. Bishop, S. C., R. H. Fleming, H. A. McCormack, D. K. Flock, and C. C. Whitehead. 2000. Inheritance of bone characteristics affect ing osteoporosis in laying hens. Br. Poult.Sci. 41:33-40.
17. Black, H., and N. Christensen. 2009. CO206/2006 Comparative assessment of laying hen welfare in New Zealand-Final Report. AsureQuality Independent Quality Assurance. Avivet, Palmer-ston North, New Zealand.
18. Bubier, N. E. 1996. The behavioural priorities of laying hens: The effect of cost/no cost multi-choice tests on time budgets. Behav. Processes 37:225-238.
19. Buil, T., G. Chacon, G. Maria, and R. Cepero. 2006. Manuscript 4. The effects of cage model, genotype and rearing treatment upon faecal corticosterone. Pages 21-22 in LayWel Deliverable 5.4. Welfare Implications of Changes in Production Systems for Laying Hens. Final Report for WP4, Physiology and Stress Indicators, Annexes. Accessed Jul. 2010. http://www.laywel.eu/web/pdf/deliverable%2054%20annexes-2.pdf.
20. Buitenhuis, A. J., T. B. Rodenburg, M. Siwek, S. J. B. Cornelissen, M. G. B. Nieuwland, R. P. M. A. Crooijmans, M. A. M. Groenen, P. Koene, H. Bovenhuis, and J. J. van der Poel. 2003. Identification of quantitative trait loci for receiving pecks in young and adult laying hens. Poult.Sci. 82:1661-1667.
21. Campo, J. L., M. T. Prieto, and S. G. Dávila. 2008. Effects of housing system and cold stress on heterophil-to-lymphocyte ratio, fluctuating asymmetry, and tonic immobility duration of chickens. Poult.Sci. 87:621-626.
22. Chakrabarti, S., D. J. King, C. Afonso, D. Swayne, C. J. Cardona, D. R. Kuney, and A. C. Gerry. 2007. Detection and isolation of exotic Newcastle disease virus from field-collected flies. J. Med. Entomol. 44:840-844.
23. Channing, C. E., B. O. Hughes, and A. W. Walker. 2001. Spatial distribution and behaviour of laying hens housed in an alternative system. Appl. Anim. Behav. Sci. 72:335-345.
24. Chirico, J., H. Eriksson, O. Fossum, and D. Jansson. 2003. The poultry red mite, Dermanyssus gallinae, a potential vector of Eryspelothrix rhusiopathiae causing erysipelas in hens. Med. Vet. Entomol. 17:232-234.
25. Christensen, J. P., H. H. Dietz, and M. Bisgaard. 1998. Phenotypic and genotypic characters of isolates of Pasteurella multocida obtained from back-yard poultry and from two outbreaks of avian cholera in avifauna in Denmark. Avian Pathol. 27:373-381.
26. Cloutier, S., R. C. Newberry, K. Honda, and J. R. Alldredge. 2002. Cannibalistic behaviour spread by social learning. Anim. Behav. 63:1153-1162.
27. Colson, S., C. Arnould, D. Guémené, and V. Michel. 2005. Bien-être de poules pondeuses logées en cage ou en volière: Paramètres physiologiques et comportementaux. 6èmes Journées de la Recherche Avicole, St Malo, France. ITAVI, Paris, France.
28. Cooper, J. J., and M. J. Albentosa. 2003. Behavioural priorities of laying hens. Avian Poult. Biol. Rev. 14:127-149.
29. Cooper, J. J., and M. C. Appleby. 2003. The value of environmental resources to domestic hens: A comparison of the work-rate for food and for nests as a function of time. Anim. Welf. 12:39-52.
30. Cordiner, L. S., and C. J. Savory. 2001. Use of perches and nestboxes by laying hens in relation to social status, based on examination of consistency of ranking orders and frequency of interaction. Appl. Anim. Behav. Sci. 71:305-317.
31. Couch, J. R. 1955. Cage layer fatigue. Feed Age 5:55-57, 88.
32. Craig, J. V. 1982. Behavioral and genetic adaptation of laying hens to high density environments. Bioscience 32:33-37.
33. Craig, J. V., and A. W. Adams. 1984. Behavior and well-being of hens (Gallus domesticus) in alternative housing environments. World's Poult.Sci. J. 40:221-240.
34. Craig, J. V., J. A. Craig, and J. Vargas Vargas. 1986. Corticoster-oids and other indicators of hens' well-being in four laying-house environments. Poult.Sci. 65:856-863.
35. Craig, J. V., and W. M. Muir. 1993. Selection for reduction of beak-inflicted injuries among caged hens. Poult.Sci. 72:411-420.
36. Craig, J. V., and W. M. Muir. 1996. Group selection for adaptation to multiple-hen cages: Beak-related mortality, feathering, and body weight responses. Poult.Sci. 75:294-302.
37. Dawkins, M. S., and S. Hardie. 1989. Space needs of laying hens. Br. Poult.Sci. 30:413-416.
38. DEFRA. 2006. The welfare effects of different methods of depopulation on laying hens. Accessed Dec. 2008. http://randd.defra.gov.uk/Default.aspx?Menu=Menu&Module=More&Location=None&ProjectID=11925&FromSearch=Y&Publisher=1&SearchText=aw0231&SortString=ProjectCode&SortOrder=Asc&Pagin g=10#Description.
39. DEFRA. 2007. The effects of stocking rate on the welfare of laying hens in non-cage systems. Accessed Jul. 2010. http://randd.de-fra.gov.uk/Document.aspx?Document=AW0223_2420_FRP.doc.
40. De Reu, K., K. Grijspeerdt, M. Heyndrickx, J. Zoons, K. De Baere, M. Uyttendaele, J. Debevere, and L. Herman. 2005. Bacterial eggshell contamination in conventional cages, furnished cages, and aviary housing systems for laying hens. Br. Poult.Sci. 46:149-155.
41. Drakley, C., and A. Walker. 2002. Monitoring of red mite habitat preference and distribution in a barn egg production system. Br. Poult.Sci. 43:553-554.
42. Duncan, E. T., M. C. Appleby, and B. O. Hughes. 1992. Effect of perches in laying cages on welfare and production of hens. Br. Poult.Sci. 33:25-35.
43. Esquenet, C., P. De Herdt, H. De Bosschere, S. Ronsmans, R. Duca-telle, and J. Van Erum. 2003. An outbreak of histomoniasis in free-range layer hens. Avian Pathol. 32:305-308.
44. Estevez, I., R. C. Newberry, and L. J. Keeling. 2002. Dynamics of aggression in the domestic fowl. Appl. Anim. Behav. Sci. 76:307-325.
45. Faure, J. M., and R. B. Jones. 1982. Effects of age, access and time of day on perching behavior in the domestic fowl. Appl. Anim. Ethol. 8:357-364.
46. Fickenwirth, A., D. W. Fölsch, and C. Dolf. 1985. Sand shortens the claws and beak of hens-Prevents injuries. Pages 288-290 in Second European Symposium on Poultry Welfare, Celle, Germany.
47. World's Poultry Science Association, printed by Federal Agricultural Research Centre, Braunschweig-Volkenrode, Germany.
48. Fleming, R. H., D. Korver, H. A. McCormack, and C. C. Whitehead. 2004. Assessing bone mineral density in vivo: Digitized fluoros-copy and ultrasound. Poult.Sci. 83:207-214.
49. Fleming, R. H., H. A. McCormack, L. McTeir, and C. C. Whitehead. 2006. Relationships between genetic, environmental and nutritional factors influencing osteoporosis in laying hens. Br. Poult.Sci. 47:742-755.
50. Fleming, R. H., C. C. Whitehead, D. Alvey, N. G. Gregory, and L. J. Wilkins. 1994. Bone structure and breaking strength in laying hens housed in different husbandry systems. Br. Poult.Sci. 35:651-662.
51. Flisikowski, K., H. Schwarzenbacher, M. Wysocki, S. Weigend, R. Preisinger, J. B. Kjaer, and R. Fries. 2009. Variation in neighbouring genes of the dopaminergic and serotonergic systems affects feather pecking behaviour of laying hens. Anim. Genet. 40:192-199.
52. Flock, D. K., and L. Norman. 2008. Criteria for the competitiveness of different management systems and strain differences in the adaptability of layer hens to non-cage environments. Proc. 16th Baltic and Finnish Poultry Conference, Vantaa, Finland. World's Poultry Science Association, Finland Branch, Jokioinen, Finland.
53. Fossum, O., D. S. Jansson, P. E. Etterline, and I. Vagsholm. 2009. Cause of mortality in laying hens in different systems in 2001-2004. Acta Vet. Scand. 51:3.
54. Freire, R., L. J. Wilkins, F. Short, and C. J. Nicol. 2003. Behaviour and welfare of individual laying hens in a non-cage system. Br. Poult.Sci. 44:22-29.
55. Gentle, M., and S. Wilson. 2004. Pain and the laying hen. Pages 165-175 in Welfare of the Laying Hen. G. C. Perry, ed. CABI Publishing, Wallingford, UK.
56. Glatz, P. C. 2002. Claw abrasives in layer cages-A review. Int. J. Poult.Sci. 1:1-5.
57. Glofcheskie, B. D., and G. A. Surgeoner. 1993. Efficacy of muscovy ducks as an adjunct for house-fly (Diptera: Muscidae) control in swine and dairy operations. J. Econ. Entomol. 86:1686-1692.
58. Graml, C., K. Niebuhr, and S. Waiblinger. 2008. Reaction of laying hens to humans in the home or a novel environment. Appl. Anim. Behav. Sci. 113:98-109.
59. Gregory, N. G., and L. J. Wilkins. 1989. Broken bones in domestic fowl: Handling and processing damage in end-of-lay battery hens. Br. Poult.Sci. 30:555-562.
60. Gregory, N. G., and L. J. Wilkins. 1992. Skeletal damage and bone defects during catching and processing. Pages 313-328 in Bone Biology and Skeletal Disorders in Poultry. C. C. Whitehead, ed. Carfax Publishing Co., Abington, Oxfordshire, UK.
61. Gregory, N. G., L. J. Wilkins, S. D. Austin, C. G. Belyavin, D. M. Alvey, and S. A. Tucker. 1992. Effect of catching method on the prevalence of broken bones in end of lay hens. Avian Pathol. 21:717-722.
62. Gregory, N. G., L. J. Wilkins, S. D. Eleperuma, A. J. Ballantyne, and N. D. Overfield. 1990. Broken bones in domestic fowls: Effects of husbandry system and stunning method in end-of- lay hens. Br. Poult.Sci. 31:59-69.
63. Gregory, N. G., L. J. Wilkins, S. C. Kestin, C. G. Belyavin, and D. M. Alvey. 1991. Effect of husbandry system on broken bones and bone strength in hens. Vet. Rec. 128:397-399.
64. Gross, W. B. 1984. Staphylococcus. Pages 263-266 in Diseases of Poultry. 8th ed. M. S. Hofstad, H. J. Barnes, B. W. Calnek, W. M. Reid, and H. W. Yoder Jr., ed. Iowa State University Press, Ames.
65. Guarino, M., A. Caroli, and P. Navarotto. 1999. Dust concentration and mortality distribution in an enclosed laying house. Trans. ASAE 42:1127-1133.
66. Guémené, D., V. Guesdon, R. O. Moe, V. Michel, and J. M. Faure. 2004. Production and stress parameters in laying hens beak-trimmed or not, housed in standard or furnished cages. Page 321 in Proc. 22nd World's Poultry Congress, Istanbul, Turkey. World's Poultry Science Association, Turkish Branch, Istanbul, Turkey. (Abstr.)
67. Guesdon, V., C. Leterrier, P. Constantin, D. Guémené, M. Couty, and J. M. Faure. 2004. Humeral quality and adrenal respon-siveness in laying hens reared in standard and furnished cages. Anim. Res. 53:235-243.
68. Gunnarsson, S., L. J. Keeling, and J. Svedberg. 1999. Effects of rearing factors on the prevalence of floor eggs, cloacal cannibalism and feather pecking in commercial flocks of loose housed laying hens. Br. Poult.Sci. 40:12-18.
69. Hane, M. 1999. Legehennenhaltung in der Scheiz. Schlussbericht BVET Forschungsprojekt. 2.29.1. 1-164.
70. Hansen, I., B. O. Braastad, J. Storbråten, and M. Tofastrud. 1993. Differences in fearfulness indicated by tonic immobility between laying hens in aviaries and in cages. Anim. Welf. 2:105-112.
71. Hansen, R. S. 1976. Nervousness and hysteria of mature female chickens. Poult.Sci. 55:531-543.
72. Harner, J. P., III, and J. H. Wilson. 1985. Effect of body size and cage profile on the shear strength of bones of caged layers. Br. Poult.Sci. 26:543-548.
73. Hegelund, L., J. T. Sorensen, J. B. Kjaer, and I. S. Kristensen. 2005. Use of the range area in organic egg production systems: Effect of climatic factors, flock size, age and artificial cover. Br. Poult.Sci. 46:1-8.
74. Hester, P. Y., and M. Shea-Moore. 2003. Beak trimming egg-laying strains of chickens. World's Poult.Sci. J. 59:458-474.
75. Hetland, H., and B. Svihus. 2007. Inclusion of dust bathing materials affects nutrient digestion and gut physiology of layers. J. Appl. Poult. Res. 16:22-26.
76. Hocking, P. M., C. E. Channing, G. W. Robertson, A. Edmond, and R. B. Jones. 2004. Between breed genetic variation for welfare-related behavioural traits in domestic fowl. Appl. Anim. Behav. Sci. 89:85-105.
77. Höglund, J., H. Nordenfors, and A. Uggla. 1995. Prevalence of the poultry red mite, Dermanyssus gallinae, in different types of production systems in Sweden. Poult.Sci. 74:1793-1798.
78. Horsted, K., and J. E. Hermansen. 2007. Whole wheat versus mixed layer diet as supplementary feed to layers foraging a sequence of different forage crops. Animal 1:575-585.
79. Huber-eicher, B., and B. Weschler. 1997. Feather pecking in domestic chicks: Its relation to dustbathing and foraging. Anim. Behav. 54:757-768.
80. Hudson, H. A., W. M. Britton, G. N. Rowland, and R. J. Buhr. 1993. Histomorphometric bone properties of sexually immature and mature White Leghorn hens with evaluation of fluorochrome injection on egg production traits. Poult.Sci. 72:1537-1547.
81. Hughes, B. O. 1983. Conventional and shallow cages: A summary of research from welfare and production aspects. World's Poult.Sci. J. 39:218-228.
82. Hughes, B. O., and M. C. Appleby. 1989. Increase in bone strength of spent laying hens housed in modified cages with perches. Vet. Rec. 124:483-484.
83. Hughes, B. O., and A. J. Black. 1976. Battery cage shape-Its effect on diurnal feeding pattern, egg-shell cracking and feather pecking. Br. Poult.Sci. 17:327-336.
84. Hughes, B. O., I. J. H. Duncan, and M. F. Brown. 1989. The performance of nest building by domestic hens-Is it more important than the construction of a nest? Anim. Behav. 37:210-214.
85. Hughes, B. O., S. Wilson, M. C. Appleby, and S. F. Smith. 1993. Comparison of bone volume and strength as measures of skeletal integrity in caged laying hens with access to perches. Res. Vet. Sci. 54:202-206.
86. Icken, W., S. Thurner, D. Cavero, M. Schmutz, G. Wendl, and R. Preisinger. 2008. Analysis of the free range behaviour of laying hens and the genetic and phenotypic relationships with laying performance. Br. Poult.Sci. 49:533-541.
87. Icken, W., S. Thurner, D. Cavero, M. Schmutz, G. Wendl, and R. Preisinger. 2009. Analysis of the nesting behaviour from laying hens in a floor system. Arch. Geflügelkd. 73:253-257.
88. Janczak, A. M., P. Torjesen, R. Palme, and M. Bakken. 2007. Effects of stress in hens on the behaviour of their offspring. Appl. Anim. Behav. Sci. 107:66-77.
89. Jendral, M. J., D. R. Korver, J. S. Church, and J. J. R. Feddes. 2008. Bone mineral density and breaking strength of White Leghorns housed in conventional, modified, and commercially available colony battery cages. Poult.Sci. 87:828-837.
90. Jensen, P., L. Keeling, K. Schütz, L. Andersson, P. Mormède, H. Brändström, B. Forkman, S. Kerje, R. Fredriksson, C. Ohlsson, S. Larsson, H. Mallmin, and A. Kindmark. 2005. Feather pecking in chickens is genetically related to behavioural and developmental traits. Physiol. Behav. 86:52-60.
91. Jones, R. B. 1996. Fear and adapatability in poultry: Insights, implications, and imperatives. World's Poult.Sci. J. 52:131-174.
92. Jones, R. B., H. J. Blokhuis, and G. Beuving. 1995. Open-field and tonic immobility responses in domestic chicks of two genetic lines differing in their propensity to feather peck. Br. Poult.Sci. 36:525-530.
93. Kaufmann-Bart, M., and R. K. Hoop. 2009. Diseases in chicks and laying hens during the first 12 years after battery cages were banned. Vet. Rec. 164:203-207.
94. Keeling, L., L. Andersson, K. E. Schütz, S. Kerje, R. Fredriksson, O. Carlborg, C. K. Cornwallis, T. Pizzari, and P. Jensen. 2004. Chicken genomics: Feather-pecking and victim pigmentation. Nature 431:645-646.
95. Kilpinen, O., A. Roepstorff, A. Permin, G. Norgaard-Nielsen, L. G. Lawson, and H. B. Simonsen. 2005. Influence of Dermanyssus gallinae and Ascaridia galli infections on behaviour and health of laying hens (Gallus gallus domesticus). Br. Poult.Sci. 46:26-34.
96. Kinde, H., D. M. Castellan, D. Kerr, J. Campbell, R. Breitmeyer, and A. Ardans. 2005. Longitudinal monitoring of two commercial layer flocks and their environments for Salmonella enterica serovar enteritidis and other salmonellae. Avian Dis. 49:189-194.
97. Kirkwood, A. C. 1967. Anemia in poultry infested with the red mite Dermanyssus gallinae. Vet. Rec. 80:514-516.
98. Kjaer, J. B., B. M. Hjarvard, K. H. Jensen, J. Hansen-Møller, and O. N. Larsen. 2004. Effects of haloperidol, a dopamine D2 receptor antagonist, on feather pecking behaviour in laying hens. Appl. Anim. Behav. Sci. 86:77-91.
99. Kjaer, J. B., and P. M. Hocking. 2004. The genetics of feather pecking and cannibalism. Pages 102-121 in Welfare of the Laying Hen. G. C. Perry, ed. CABI, Abingdon, Oxfordshire, UK.
100. Kjaer, J. B., and P. K. Isaksen. 1998. Individual use of the free-range area by laying hens and effect of genetic strain. Page 88 in Proc. 32nd Congress of the International Society for Applied Ethology, Clermont-Ferrand, France. I. Veissier and A. Boissy, ed. INRA, Paris France.
101. Kjaer, J. B., and P. Sørensen. 1997. Feather pecking behaviour in White Leghorns-A genetic study. Br. Poult.Sci. 38:333-341.
102. Kjaer, J. B., P. Sørensen, and G. Su. 2001. Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesti-cus). Appl. Anim. Behav. Sci. 71:229-239.
103. Knowles, T. G., and D. M. Broom. 1990. Limb bone strength and movement in laying hens from different housing systems. Vet. Rec. 126:354-356.
104. Koelkebeck, K. W., M. S. Amoss Jr., and J. R. Cain. 1987. Production, physiological, and behavioral responses of laying hens in different management environments. Poult.Sci. 66:397-407.
105. Koelkebeck, K. W., and J. R. Cain. 1984. Performance, behavior, plasma corticosterone, and economic returns of laying hens in several management alternatives. Poult.Sci. 63:2123-2131.
106. Koelkebeck, K. W., J. R. Cain, and M. S. Amoss. 1986. Corticos-terone sampling of laying hens in different management systems. Poult.Sci. 65:183-185.
107. Kreienbrock, L., B. Schneider, J. Schal, and S. Glaser. 2003. Epileg- Orienterende epidemiologische Untersuchung zum Leistungsniv-eau und Gesundheitsstatus in Legehennenhaltungen verscheiden-er Haltungssysteme. Zwischenbericht: Deskriptive Auswertung. 1. Institue fur Biometrie, Epidemiologie und Informationsverarbei-tung (IBEI), Hannover, Germany.
108. Laboriau, R., J. B. Kjaer, G. C. G. Abreu, J. Hedegaard, and A. J. Buitenhuis. 2009. Analysis of severe feather pecking behavior in a high feather pecking selection line. Poult.Sci. 88:2052-2062.
109. Lagadic, H., and J.-M. Faure. 1987. Preferences of domestic hens for cage size and floor types as measured by operant conditioning. Appl. Anim. Behav. Sci. 19:147-155.
110. Lay Wel. 2006a. Welfare implications of changes in production systems for laying hens: Deliverable 7.1.Overall strengths and weaknesses of each defined housing system for laying hens, and detail-ing the overall welfare impact of each housing system http:// www.laywel.eu/web/pdf/deliverable%2071%20welfare%20assess-ment.pdf Accessed Mar. 2010.
111. Lay Wel. 2006b. Welfare implications of changes in production systems for laying hens: Deliverable 5.4. Physiology and Stress Indicators. Accessed Mar. 2010. http://www.laywel.eu/web/pdf/ deliverable%2054%20physiology.pdf.
112. Leone, E. H., and I. Estevez. 2008. Use of space in the domestic fowl: Separating the effects of enclosure size, group size and density. Anim. Behav. 76:1673-1682.
113. Lervik, S., R. O. Moe, C. M. Mejdell, and M. Bakken. 2007. Challenges in different housing systems for laying hens. Nor. Veteri-naertidsskr. 119:5-14.
114. Leyendecker, M. 2003. Einfluss verschiedener Legehennenhaltungs-systeme (konventionelle Käfige, ausgestaltete Käfige, intensive Auslauf- und Volierenhaltung) auf die Legeleistung, Eiqualität und Knochenfestigkeit von Legehennen. PhD. Diss. Univ. Osn-abrück, Fachbereich Biologie, Osnabrück, Germany.
115. Lindberg, A. C., and C. J. Nicol. 1997. Dustbathing in modified battery cages: Is sham dustbathing an adequate substitute? Appl. Anim. Behav. Sci. 55:113-128.
116. Lindqvist, C., A. M. Janczak, D. Nätt, I. Baranowska, N. Lindqvist, A. Wichman, J. Lundeberg, J. Lindberg, P. A. Torjesen, and P. Jensen. 2007. Transmission of stress-induced learning impairment and associated brain gene expression from parents to offspring in chickens. PLoS ONE 2:e364.
117. Lindqvist, C. E. S., K. E. Schutz, and P. Jensen. 2002. Red jungle-fowl have more contrafreeloading than White Leghorn layers: Effects of food deprivation and consequences for information gain. Behaviour 139:1195-1209.
118. Maurer, V., J. Baumgartner, M. Bieri, and D. W. Folsch. 1993. The occurrence of the chicken mite Dermanyssus gallinae (Acari: Der-manyssidae) in Swiss poultry houses. Mitt. Schweiz. Entomol. Ges. 66:87-97.
119. Maxwell, M. H. 1993. Avian blood leucocyte responses to stress. World's Poult.Sci. J. 54:155-178.
120. Mazaheri, A., M. Lierz, and H. M. Hafez. 2005. Investigation on the pathogenicity of Erysipelothrix rhusiopathiae in laying hens. Avian Dis. 49:574-576.
121. McAllister, J. C., C. D. Steelman, and J. K. Skeeles. 1994. Reservoir competence of the lesser mealworm (Coleoptera: Tenebrionidae) for Salmonella typhimurium (Eubacteriales: Enterobacteriaceae). J. Med. Entomol. 31:369-372.
122. McCoy, M. A., G. A. C. Reilly, and D. J. Kilpatrick. 1996. Density and breaking strength of bones of mortalities among caged layers. Res. Vet. Sci. 60:185-186.
123. McGibbon, W. H. 1976. Floor laying-A heritable and environmentally influenced trait of the domestic fowl. Poult.Sci. 55:765-771.
124. McLean, K. A., M. R. Baxter, and W. Michie. 1986. A comparison of the welfare of laying hens in battery cages and in a perchery. Res. Dev. Agric. 3:93-98.
125. Mench, J. A., A. van Tienhoven, J. A. Marsh, C. C. McCormick, D. L. Cunningham, and R. C. Baker. 1986. Effects of cage and floor pen management on behavior, production, and physiological stress responses of laying hens. Poult.Sci. 65:1058-1069.
126. Meyer, W. A., and M. L. Sunde. 1974. Bone breakage as affected by type housing or an exercise machine for layers. Poult.Sci. 53:878-885.
127. Meyer-Kuhling, B., J. Heine, J. Muller-Lindloff, and K. Pfister. 2007. Epidemiology of Dermanyssus gallinae and acaricidal efficacy of phoxim 50% in alternative housing systems during the laying periods of hens. Parasitol. Res. 101:S1-S12.
128. Michel, V., and D. Huonnic. 2003. A comparison of welfare, health and production performance of laying hens reared in cages or in aviaries. Br. Poult.Sci. 44:775-776.
129. Moe, R. O., D. Guémené, H. J. S. Larsen, M. Bakken, S. Lervick, H. Hetland, and R. Tauson. 2004. Effect of pre-laying rearing conditions in laying hens housed in standard or furnished cages on various indicators of animal welfare. Page 4 in 22nd World's Poultry Congress, Istanbul, Turkey. World's Poultry Science As-socication, Turkish Branch, Izmir, Turkey. (Abstr.)
130. Moinard, C., J. P. Morisse, and J. M. Faure. 1998. Effect of cage area, cage height and perches on feather condition, bone breakage and mortality of laying hens. Br. Poult.Sci. 39:198-202.
131. Muir, W. M. 1996. Group selection for adaptation to multiple-hen cages: Selection program and direct responses. Poult.Sci. 75:447-458.
132. Mullens, B. A., D. R. Kuney, N. C. Hinkle, and C. E. Szijj. 2004. Producer attitudes and control practices for northern fowl mites in southern California. J. Appl. Poult. Res. 13:488-492.
133. Mullens, B. A., J. P. Owen, D. R. Kuney, C. E. Szijj, and K. A. Klingler. 2009. Temporal changes in distribution, prevalence and intensity of northern fowl mite (Ornithonyssus sylviarum) parasitism in commercial caged laying hens, with a comprehensive economic analysis of parasite impact. Vet. Parasitol. 160:116-133.
134. Newberry, R. C. 2004. Cannibalism. Pages 239-258 in Welfare of the Laying Hen. G. C. Perry, ed., CABI Publishing, Wallingford, UK.
135. Newberry, R. C., I. Estevez, and L. J. Keeling. 2001. Group size and perching behaviour in young domestic fowl. Appl. Anim. Behav. Sci. 73:117-129.
136. Newberry, R. C., A. B. Webster, N. J. Lewis, and C. Van Arnam. 1999. Management of spent hens. J. Appl. Anim. Welf. Sci. 2:13-29.
137. Newman, S., and S. Leeson. 1997. Skeletal integrity in layers at completion of egg production. World's Poult.Sci. J. 53:265-286.
138. Newman, S., and S. Leeson. 1998. Effect of housing birds in cages or an aviary system on bone characteristics. Poult.Sci. 77:1492-1496.
139. Nicol, C. J. 1987a. Effect of cage height and area on the behaviour of hens housed in battery cages. Br. Poult.Sci. 28:327-335.
140. Nicol, C. J. 1987b. Behavioural responses of laying hens following a period of spatial restriction. Anim. Behav. 35:1709-1719.
141. Nicol, C. J., S. N. Brown, E. Glen, S. J. Pope, F. J. Short, P. D. Warriss, P. H. Zimmerman, and L. J. Wilkins. 2006. Effects of stocking density, flock size and management on the welfare of laying hens in single-tier aviaries. Br. Poult.Sci. 47:135-146.
142. Nicol, C. J., G. Caplen, J. Edgar, and W. J. Browne. 2009. Associations between welfare indicators and environmental choice in laying hens. Anim. Behav. 78:413-424.
143. Nicol, C. J., N. G. Gregory, T. Knowles, I. Parkman, and L. Wilkins. 1999. Differential effects of increased stocking density, mediated by increased flock size, on feather pecking and aggression in laying hens. Appl. Anim. Behav. Sci. 65:137-152.
144. Nicol, C. J., A. C. Lindberg, A. J. Phillips, S. J. Pope, L. J. Wilkins, and L. E. Green. 2001. Influences of prior exposure to wood shavings on feather pecking, dustbathing and foraging in adult laying hens. Appl. Anim. Behav. Sci. 73:141-155.
145. Nicol, C. J., C. Potzsch, K. Lewis, and L. E. Green. 2003. Matched concurrent case-control study of risk factors for feather pecking in hens on free-range commercial farms in the UK. Br. Poult.Sci. 44:515-523.
146. Nørgaard-Nielsen, G. 1990. Bone strength of laying hens kept in an alternative system, compared with hens in cages and on deep-litter. Br. Poult.Sci. 31:81-89.
147. Odén, K., S. Gunnarsson, C. Berg, and B. Algers. 2005. Effects of sex composition on fear measured as tonic immobility and vigilance behaviour in large flocks of laying hens. Appl. Anim. Behav. Sci. 95:89-102.
148. Olsson, I. A. S., and L. J. Keeling. 2002. The push-door for measuring motivation in hens: Laying hens are motivated to perch at night. Anim. Welf. 11:11-19.
149. Olsson, I. A. S., and L. J. Keeling. 2005. Why in earth? Dustbathing behaviour in jungle and domestic fowl reviewed from a Tinber-gian and animal welfare perspective. Appl. Anim. Behav. Sci. 93:259-282.
150. Owen, J. P., M. E. Delany, and B. A. Mullens. 2008. MHC haplotype involvement in avian resistance to an ectoparasite. Immunoge-netics 60:621-631.
151. Pedersen, S., M. Nonnenmann, R. Rautiainen, T. G. Demmers, T. Banhazi, and M. Lyngbye. 2000. Dust in pig buildings. J. Agric. Saf. Health 6:261-274.
152. Permin, A., M. Bisgaard, F. Frandsen, M. Pearman, J. Kold, and P. Nansen. 1999. Prevalence of gastrointestinal helminthes in different poultry production systems. Br. Poult.Sci. 40:439-443.
153. Pohle, K., and H.-W. Cheng. 2009. Comparative effects of furnished and battery cages on egg production and physiological parameters in White Leghorn hens. Poult.Sci. 88:2042-2051.
154. Rodenburg, T. B., F. A. M. Tuyttens, K. De Reu, L. Herman, J. Zoons, and B. Sonck. 2008. Welfare assessment of laying hens in furnished cages and non-cage systems: An on-farm comparison. Anim. Welf. 17:363-373.
155. Rodenburg, T. B., F. A. M. Tuyttens, B. Sonck, K. De Reu, L. Herman, and J. Zoons. 2005. Welfare, health and hygiene of laying hens housed in furnished cages and in alternative housing systems. J. Appl. Anim. Welf. Sci. 8:211-226.
156. Rowland, L. O., Jr., and R. H. Harms. 1970. The effect of wire pens, floor pens and cages on bone characteristics of laying hens. Poult.Sci. 49:1223-1225.
157. Rowland, L. O., Jr., and R. H. Harms. 1972. Time required to develop bone fragility in laying hens. Poult.Sci. 51:1339-1341.
158. Savory, C. J. 1995. Feather pecking and cannibalism. World's Poult.Sci. J. 51:215-219.
159. Savory, C. J., M. C. Jack, and V. Sandilands. 2006. Behavioural responses to different floor space allowances in small groups of laying hens. Br. Poult.Sci. 47:120-124.
160. Schreiweis, M. A., P. Y. Hester, and D. E. Moody. 2005. Identification of quantitative trait loci associated with bone traits and BW in an F2 resource population of chickens. Genet. Sel. Evol. 37:677-698.
161. Scott, G. B., N. R. Lambe, and D. Hitchcock. 1997. Ability of laying hens to negotiate horizontal perches at different heights, separated by different angles. Br. Poult.Sci. 38:48-54.
162. Settar, P., J. Arango, N. P. O'Sullivan, and J. A. Arthur. 2006. Evidence of genetic variability for floor and nest egg laying behavior in floor pens. Proc. XII European Poultry Conference, Verona, Italy. World's Poultry Science Association, Beekbergen, the Netherlands.
163. Sherwin, C. M., G. J. Richards, and C. J. Nicol. 2010. A comparison of the welfare or layer hens in four housing systems in the UK. Br. Poult.Sci. In press.
164. Shimmura, T., Y. Eguchi, K. Uetake, and T. Tanaka. 2008a. Effects of separation of resources on behaviour of high-, medium- and low-ranked hens in furnished cages. Appl. Anim. Behav. Sci. 113:74-86.
165. Shimmura, T., T. Suzuki, S. Hirahara, Y. Eguchi, K. Uetake, and T. Tanaka. 2008b. Pecking behavior of laying hens in single-tiered aviaries with and without outdoor area. Br. Poult.Sci. 49:396-401.
166. Shini, S. 2003. Physiologic responses of laying hens to the alternative housing systems. Int. J. Poult.Sci. 2:357-360.
167. Sørensen, P. 1992. Selection, environments of layers and response to nesting behaviour. Proc. XIX Worlds's Poult. Congr. 2:409-412.
168. Struelens, E., F. A. M. Tuyttens, L. Duchateau, T. Leroy, M. Cox, E. Vranken, J. Buyse, J. Zoons, D. Berckmans, F. Odberg, and B. Sonck. 2008a. Perching behaviour and perch height preference of laying hens in furnished cages varying in height. Br. Poult.Sci. 49:381-389.
169. Struelens, E., A. Van Nuffel, F. A. M. Tuyttens, L. Audoorn, E. Vranken, J. Zoons, D. Berckmans, F. Odberg, S. Van Dongen, and B. Sonck. 2008b. Influence of nest seclusion and nesting material on pre-laying behaviour of laying hens. Appl. Anim. Be-hav. Sci. 112:106-119.
170. Swarbrick, O. 1986. Clinical problems in 'free-range' layers. Vet. Rec. 118:363.
171. Tactacan, G. B., W. Guenter, N. J. Lewis, J. C. Rodriguez-Lecompte, and J. D. House. 2009. Performance and welfare of laying hens in conventional and enriched cages. Poult.Sci. 88:698-707.
172. Tauson, R. 1998. Health and production in improved cage designs. Poult.Sci. 77:1820-1827.
173. Tauson, R. 2002. Furnished cages and aviaries: Production and health. World's Poult.Sci. J. 58:49-63.
174. Tauson, R., and P. Abrahamsson. 1994a. Effects on production, health and behaviour in three SCWL strains in an EMC model in comparison with other modified and conventional cages. Pages 41-53 in Modified Cages for Laying Hens. C. M. Sherwin, ed. Universities Federation for Animal Welfare, Potters Bar, UK.
175. Tauson, R., and P. Abrahamsson. 1994b. Foot and skeletal disorders in laying hens: Effects of perch design, hybrid, housing system and stocking density. Acta Agric. Scand. A 44:110-119.
176. Tauson, R., K. Elwinger, K.-E. Holm, and H. Wall. 2006. Analyses of a data base for health parameters in different housing systems. Deliverables D.3.2-D.3.3. Accessed Nov. 2007. http://www.lay-wel.eu/web/pdf/deliverables%2031-33%20health-2.pdf.
177. Tauson, R., A. Wahlstrom, and P. Abrahamsson. 1999. Effect of two floor housing systems and cages on health, production, and fear response in layers. J. Appl. Poult. Res. 8:152-159.
178. Taylor, A. A., and J. F. Hurnik. 1996. The long-term productivity of hens housed in battery cages and an aviary. Poult.Sci. 75:47-51.
179. Templeton, J. M., A. J. DeJong, P. J. Blackall, and J. K. Miflin. 2006. Survival of Campylobacter spp. in darkling beetles (Alphi-tobius diaperinus) and their larvae in Australia. Appl. Environ. Microbiol. 72:7909-7911.
180. Thogerson, C. M., P. Y. Hester, J. A. Mench, R. C. Newberry, C. M. Okura, E. A. Pajor, P. N. Talaty, and J. P. Garner. 2009a. The effect of feeder space allocation on productivity and physiology of Hy-Line W-36 hens housed in conventional cages. Poult.Sci. 88:1793-1799.
181. Thogerson, C. M., P. Y. Hester, J. A. Mench, R. C. Newberry, E. A. Pajor, and J. P. Garner. 2009b. The effect of feeder space allocation on behavior of Hy-Line W-36 hens housed in conventional cages. Poult.Sci. 88:1544-1552.
182. United States Animal Health Association. 2007. Report of the Committee on Transmissible Diseases of Poultry and Other Avian Species. Accessed Jul. 2010. http://www.usaha.org/committees/ reports/2007/report-pad-2007.pdf.
183. Valiente Moro, C. V., C. Chave, and L. Zenner. 2007. Experimental infection of Salmonella enteritidis by the poultry red mite, Der-manyssus gallinae. Vet. Parasitol. 146:329-336.
184. Van Liere, D. W. 1991. Function and organisation of dustbathing in laying hens. PhD Thesis. Agricultural University, Wageningen, the Netherlands.
185. Van Niekerk, T. G. C. M., and B. F. J. Reuvekamp. 1994. Husbandry factors and bone strength in laying hens. Pages 133-136 in Proc. 9th European Poultry Conference, Glasgow, UK. World's Poultry Association, United Kingdom Branch, Norfolk, UK.
186. Vestergaard, K. S., J. P. Kruijt, and J. A. Hogan. 1993. Feather pecking and chronic fear in groups of red junglefowl: Their relationships to dustbathing, rearing environment and social status. Anim. Behav. 45:1127-1140.
187. Vits, A., D. Weitzenburger, H. Hamann, and O. Distl. 2005. Production, egg quality, bone strength, claw length, and keel bone deformities of laying hens housed in furnished cages with different group sizes. Poult.Sci. 84:1511-1519.
188. Walker, A. W., D. M. Alvey, and S. A. Tucker. 1997. Effect of an elevated food trough on bone strength and ease of catching laying hens. Br. Poult.Sci. 38:S14-S15.
189. Wang, G., C. Ekstrand, and J. Svedberg. 1998. Wet litter and perches as risk factors for the development of foot-pad dermatitis in floor housed hens. Br. Poult.Sci. 39:191-197.
190. Webster, A. B. 2004. Welfare implications of avian osteoporosis. Poult.Sci. 83:184-192.
191. Weeks, C. A., and C. J. Nicol. 2006. Behavioural needs, preferences and priorities of laying hens. World's Poult.Sci. J. 62:297-308.
192. Weitzenbürger, D., A. Vits, H. Hamman, and O. Distl. 2005. Effect of furnished small group housing systems and furnished cages on mortality and causes of death in two layer strains. Br. Poult.Sci. 46:553-559.
193. Weitzenbürger, D., A. Vits, H. Hamann, M. Hewicker-Trautwein, and O. Distl. 2006. Macroscopic and histopathological alterations of foot pads of laying hens kept in small group housing systems and furnished cages. Br. Poult.Sci. 47:533-543.
194. Whitehead, C. C. 2004. Overview of bone biology in the egg-laying hen. Poult.Sci. 83:193-199.
195. Whitehead, C. C., and R. H. Fleming. 2000. Osteoporosis in cage layers. Poult.Sci. 79:1033-1041.
196. Whitehead, C. C., and S. Wilson. 1992. Characteristics of osteopenia in hens. Pages 265-280 in Poultry Science Symposium 23: Bone Biology and Skeletal Disorders in Poultry. C. C. Whitehead, ed. Carfax Publishing Co., Abington, Oxfordshire, UK.
197. Widowski, T., and I. J. H. Duncan. 2000. Working for a dustbath: Are hens increasing pleasure rather than reducing suffering? Appl. Anim. Behav. Sci. 68:39-53.
198. Wilkins, L. J., S. N. Brown, P. H. Zimmerman, C. Leeb, and C. J. Nicol. 2004. Investigation of palpation as a method for determining the prevalence of keel and furculum damage in laying hens. Vet. Rec. 155:547-549.
199. Wilson, S., S. R. I. Duff, and C. C. Whitehead. 1992. Effects of age, sex, and housing on the trabecular bone of laying strain domestic fowl. Res. Vet. Sci. 53:52-58.
200. Yue, S., and I. J. H. Duncan. 2003. Frustrated nesting behaviour: Relation to extra-cuticular shell calcium and bone strength in White Leghorn hens. Br. Poult.Sci. 44:175-181.
201. Zeltner, E., and H. Hirt. 2008. Factors involved in the improvement of the use of hen runs. Appl. Anim. Behav. Sci. 114:395-408.
202. Zimmerman, P. H., A. C. Lindberg, S. J. Pope, E. Glen, J. E. Bolhu-is, and C. J. Nicol. 2006. The effect of stocking density, flock size and modified management on laying hen behaviour and welfare in a non-cage system. Appl. Anim. Behav. Sci. 101:111-124.
203. Zupan, M., A. Kruschwitz, T. Buchwalder, B. Huber-eicher, and I. Stuhec. 2008. Comparison of the prelaying behavior of nest layers and litter layers. Poult.Sci. 87:399-404.