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http://dx.doi.org/10.5536/KJPS.2015.42.2.169

A Review of Laying Hens' Behavior  

Cheon, Si-Nae (National Institute of Animal Science, R. D. A.)
Lee, Jun-Yeob (National Institute of Animal Science, R. D. A.)
Choi, Hee-Chul (National Institute of Animal Science, R. D. A.)
Jeon, Jung-Hwan (National Institute of Animal Science, R. D. A.)
Publication Information
Korean Journal of Poultry Science / v.42, no.2, 2015 , pp. 169-180 More about this Journal
Abstract
Domestic fowl still tend to exhibit very similar behavior such as perching, nesting and dust-bathing to their ancestor, Red Jungle Fowl (Gallus gallus), even though they have been domesticated for 8,000 years. If given the opportunity, they choose high place for staying at night, a dark place for laying eggs and do dust bath in litter area. Therefore, these behaviors are considered for very important role in improving the welfare of laying hens. In EU and some countries, farmers must provide perches, nests and litter for the housing of laying hens. An understanding of behavior can lead to enhance physical health, emotional condition and welfare of laying hens. Unfortunately, there is little information on behavior of laying hens in Korea. Therefore, we researched the literature on certain behavior of laying hens.
Keywords
behavior; laying hens; animal welfare;
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  • Reference
1 Gentle MJ, Hunter LN 1990 Physiological and behavioural responses associated with feather removal in Gallus gallus var domesticus. Research in veterinary science 50(1):95-101.
2 Gentle MJ, Hunter LN, Waddington D 1991 The onset of pain related behaviours following partial beak amputation in the chicken. Neuroscience letters 128(1):113-116.   DOI
3 Gentle MJ, Waddington D, Hunter LN, Jones RB 1990 Behavioural evidence for persistent pain following partial beak amputation in chickens. Appl Anim Behav Sci 27(1):49-157.
4 Glatz PC, Murphy LB, Reston A 1992 Analgesic therapy of beak trimmed chickens. Australian veterinary journal 69(1):18.   DOI
5 Green LE, Lewis K, Kimpton A, Nicol CJ 2000 Cross-sectional study of the prevalence of feather pecking in laying hens in alternative systems and its associations with management and disease. The Veterinary Record 147(9):233-238.   DOI
6 Guesdon V, Ahmed AMH, Mallet S, Faure JM, Nys Y 2006 Effects of beak trimming and cage design on laying hen performance and egg quality. Br Poult Sci 47(1):1-12.   DOI
7 Gunnarsson S 1999 Effect 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(1):12-18.   DOI
8 Hafez ESE 1962 The behaviour of domestic animals. Williams and Wilkins, Baltimore, USA.
9 Hansen I, Braastad BO, Storbraten J, Tofastrud M 1993 Differences in fearfulness indicated by tonic immobility between laying hens in aviaries and in cages. Animal Welfare 2(2):105-112.
10 Harrison R 1964 Animal machines: The new factory farming industry. Vincent Stuart, London, UK.
11 Hogan JA, Van Boxel F 1993 Causal factors controlling dustbathing in Burmese red jungle fowl: Some results and a model. Anim Behav 46(4):627-635.   DOI
12 Huber-Eicher B 2004 The effect of early colour preference and of a colour exposing procedure on the choice of nest colours in laying hens. Appl Anim Behav Sci 86(1):63-76.   DOI
13 Huber-Eicher B, Audige L 1999 Analysis of risk factors for the occurrence of feather pecking in laying hen growers. Br Poult Sci 40(5):599-604.   DOI
14 Huber-Eicher B, Sebo F 2001a Reducing feather pecking when raising laying hen chicks in aviary systems. Appl Anim Behav Sci 73(1):59-68.   DOI
15 Huber-Eicher B, Sebo F 2001b The prevalence of feather pecking and development in commercial flocks of laying hens. Appl Anim Behav Sci 74(3):223-231.   DOI
16 Huber-Eicher B, Wechsler B 1997 Feather pecking in domestic chicks: Its relation to dustbathing and foraging. Anim Behav 54(4):757-768.   DOI
17 Huber-Eicher BEAT, Wechsler B 1998 The effect of quality and availability of foraging materials on feather pecking in laying hen chicks. Anim Behav 55(4):861-873.   DOI
18 Huber HU, Folsch DW, Stahli U 1985 Influence of various nesting materials on nest site selection of the domestic hen. Br Poult Sci 26(3):367-373.   DOI
19 Hughes BO 1973 The effect of implanted gonadal hormones on feather pecking and cannibalism in pullets. Br Poult Sci 14(4):341-348.   DOI
20 Hughes BO 1993 Choice between artificial turf and wire floor as nest sites in individually caged laying hens. Appl Anim Behav Sci 36(4):327-335.   DOI
21 Hughes BO, Appleby MC 1989 Increase in bone strength of spent laying hens housed in modified cages with perches. The Veterinary Record 124(18):483-484.   DOI
22 Hughes BO, Duncan IJH, Brown MF 1989 The performance of nest building by domestic hens: Is it more important than the construction of a nest? Anim Behav 37:210-214.   DOI
23 Jendral MJ, Korver DR, Church JS, Feddes JJR 2008 Bone mineral density and breaking strength of white leghorns housed in conventional, modified, and commercially available colony battery cages. Poult Sci 87(5):828-837.   DOI
24 Jiang S, Hester PY, Hu JY, Yan FF, Dennis RL, Cheng HW 2014 Effect of perches on liver health of hens. Poult Sci 93(7):1618-1622.   DOI
25 Johnsen PF, Vestergaard KS, Norgaard-Nielsen G 1998 Influence of early rearing conditions on the development of feather pecking and cannibalism in domestic fowl. Appl Anim Behav Sci 60(1):25-41.   DOI
26 Kjaer JB, Sorensen P 1997 Feather pecking behaviour in White Leghorns, a genetic study. Br Poult Sci 38(4):333-341.   DOI
27 Kjaer JB, Vestergaard KS 1999 Development of feather pecking in relation to light intensity. Appl Anim Behav Sci 62(2):243-254.   DOI
28 Klein T, Zeltner E, Huber-Eicher B 2000 Are genetic differences in foraging behaviour of laying hen chicks paralleled by hybrid-specific differences in feather pecking? Appl Anim Behav Sci 70(2):43-155.
29 Knowles TG, Broom DM 1990 Limb bone strength and movement in laying hens from different housing systems. The Veterinary Record 126(15):354-356.   DOI
30 Klinger G 1982 Das Verhalten von Junghennen unter dem Einfluss verschiedener Zuechtungs-, Haltungs-und Testbedingungen. Deutsche Gefluegelwirtschaft und Schweineproduktion.
31 Koene P 2001 Animal welfare and genetics in organic farming of layers: The example of cannibalism. In Proc. of the 4th NAHWOA Workshop, Wageningen, pp. 62-85.
32 Koene P, Wiepkema PR 1991 Pre-dustbath vocalizations as an indicator of need in domestic hens. In: Bohncke E, Molkenthin V Eds., Alternatives in Animal Husbandry. Agrarkultur Verlag, Witzenhausen, pp. 95-103.
33 Lambton SL, Knowles TG, Yorke C, Nicol CJ 2010 The risk factors affecting the development of gentle and severe feather pecking in loose housed laying hens. Appl Anim Behav Sci 123(1):32-42.   DOI
34 Lay DC, Fulton RM, Hester PY, Karcher DM, Kjaer JB, Mench JA, Porter RE 2011 Hen welfare in different housing systems. Poult Sci 90(1):278-294.   DOI
35 Lentfer TL, Gebhardt-Henrich SG, Frohlich EK, von Borell E 2011 Influence of nest site on the behaviour of laying hens. Appl Anim Behav Sci 135(1):70-77.   DOI
36 Leyendecker M, Hamann H, Hartung J, Kamphues J, Neumann U, Surie C, Distl O 2005 Keeping laying hens in furnished cages and an aviary housing system enhances their bone stability. Br Poult Sci 46(5):536-544.   DOI
37 Lindberg AC 1999 Effects of vacuum and real dustbathing bouts on dustbathing motivation in domestic hens. In 33rd International Congress of the International Society for Animal Ethology (Boe KE, M Bakken, BBO Bakken, eds). As, Norway: Agricultural University of Norway (Vol. 91).
38 McAdie TM, Keeling LJ 2002 The social transmission of feather pecking in laying hens: Effects of environment and age. Appl Anim Behav Sci 75(2):147-159.   DOI
39 Lindberg AC, Nicol CJ 1997 Dustbathing in modified battery cages: Is sham dustbathing an adequate substitute? Appl Anim Behav Sci 55(1):113-128.   DOI
40 Lundberg A, Keeling LJ 1999 The impact of social factors on nesting in laying hens (Gallus gallus domesticus). Appl Anim Behav Sci 64(1):57-69.   DOI
41 McCoy MA, Reilly GAC, Kilpatrick DJ 1996 Density and breaking strength of bones of mortalities among caged layers. Research in veterinary science 60(2):185-186.   DOI
42 Mills AD, Wood Gush DGM 1985 Pre laying behaviour in battery cages. Br Poult Sci 26(2):247-352.   DOI
43 Muiruri HK, Harrison PC, Gonyou HW 1990 Preferences of hens for shape and size of roosts. Appl Anim Behav Sci 27(1):141-147.   DOI
44 Newberry RC, Estevez I, Keeling LJ 2001 Group size and perching behaviour in young domestic fowl. Appl Anim Behav Sci 73(2):117-129   DOI
45 Nicol CJ, Bestman M, Gilani AM, De Haas EN, De Jong IC, Lambton S, Rodenburg TB 2013 The prevention and control of feather pecking: Application to commercial systems. World's Poult Sci J 69(04):775-788.   DOI
46 Nicol CJ, Gregory NG, Knowles TG, Parkman ID, Wilkins LJ 1999 Differential effects of increased stocking density, mediated by increased flock size, on feather pecking and agg ression in laying hens. Appl Anim Behav Sci 65(2):137-152.   DOI
47 Olsson IAS, Keeling LJ 2000 Night-time roosting in laying hens and the effect of thwarting access to perches. Appl Anim Behav Sci 68(3):243-256.   DOI
48 Nicol CJ, Lindberg AC, Phillips AJ, Pope SJ, Wilkins LJ, Green LE 2001 Influence of prior exposure to wood shavings on feather pecking, dustbathing and foraging in adult laying hens. Appl Anim Behav Sci 73(2):141-155.   DOI
49 Norgaard-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(1):81-89.   DOI
50 Oden K, Keeling LJ, Algers B 2002 Behaviour of laying hens in two types of aviary systems on 25 commercial farms in Sweden. Br Poult Sci 43(2):169-181.   DOI
51 Olsson IAS, Keeling LJ 2005 Why in earth? Dustbathing behaviour in jungle and domestic fowl reviewed from a Tinbergian and animal welfare perspective. Appl Anim Behav Sci 93(3):259-282.   DOI
52 Olsson IAS, Keeling LJ, Duncan IJ 2002 Why do hens sham dustbathe when they have litter? Appl Anim Behav Sci 76(1):53-64.   DOI
53 Pagel M, Dawkins MS 1997 Peck orders and group size in laying hens: Futures contracts' for non-aggression. Behav Proc 40(1):13-25.   DOI
54 Petherick JC, Duncan IJH 1989 Behaviour of young domestic fowl directed towards different substrates. Br Poult Sci 30(2):229-238.   DOI
55 Petherick JC, Seawright E, Waddington D 1993 Influence of quantity of utter on nest box selection and nesting behaviour of domestic hens. Br Poult Sci 34(5):857-872.   DOI
56 Preston AP 1987 Restricted feeding time and the behaviour of caged laying hens. Br Poult Sci 28(3):387-396.   DOI
57 Petherick JC, Seawright E, Waddington D, Duncan IJ, Murphy LB 1995 The role of perception in the causation of dustbathing behaviour in domestic fowl. Anim Behav 49(6):1521-1530.   DOI
58 Pickel T, Scholz B, Schrader L 2010 Perch material and diameter affects particular perching behaviours in laying hens. Appl Anim Behav Sci 127(1):37-42.   DOI
59 Potzsch CJ, Lewis K, Nicol CJ, Green LE 2001 A cross-sectional study of the prevalence of vent pecking in laying hens in alternative systems and its associations with fea ther pecking, management and disease. Appl Anim Behav Sci 74(4):259-272.   DOI
60 Riber AB 2010 Development with age of nest box use and gregarious nesting in laying hens. Appl Anim Behav Sci 123(1):24-31.   DOI
61 Riber AB 2012 Gregarious nesting-An anti-predator response in laying hens. Appl Anim Behav Sci 138(1):70-78.   DOI
62 Riedstra B, Groothuis TG 2002 Early feather pecking as a form of social exploration: The effect of group stability on feather pecking and tonic immobility in domestic chicks. Appl Anim Behav Sci 77(2):127-138.   DOI
63 Rietveld-Piepers B, Blokhuis HJ, Wiepkema PR 1985 Egglaying behaviour and nest-site selection of domestic hens kept in small floor-pens. Appl Anim Behav Sci 14(1):75-88.   DOI
64 Rodenburg TB, Buitenhuis AJ, Ask B, Uitdehaag KA, Koene P, Van der Poel JJ, Bovenhuis H 2003 Heritability of feather pecking and open-field response of laying hens at two different ages. Poult Sci 82(6):861-867.   DOI
65 Rodenburg TB, Van Hierden YM, Buitenhuis AJ, Riedstra B, Koene P, Korte SM, Blokhuis HJ 2004 Feather pecking in laying hens: New insights and directions for research? Appl Anim Behav Sci 86(3):291-298.   DOI
66 Rodenburg TB, Koene P 2004 Feather pecking and feather Ioss. Welfare of the Laying Hen 27:227.
67 Rodenburg TB, Komen H, Ellen ED, Uitdehaag KA, van Arendonk JA 2008 Selection method and early-life history affect behavioural development, feather pecking and cannibalism in laying hens: A review. Appl Anim Behav Sci 110(3):217-228.   DOI
68 Rodenburg TB, Reu KD, Tuyttens FAM, Sandilands V, Hocking P 2012 Performance, welfare, health and hygiene of laying hens in non-cage systems in comparison with cage systems. In Proceedings of the 30th Poultry Science Symposium, University of Strathclyde, Glasgow, Scotland, 2011. (pp. 210-224). CABI.
69 Rushen J 1982 The peck orders of chickens: How do they develop and why are they linear? Anim Behav 30(4):1129-1137.   DOI
70 Sanotra GS, Vestergaard KS, Agger JF, Lawson LG 1995 The relative preferences for feathers, straw, wood-shavings and sand for dustbathing, pecking and scratching in domestic chicks. Appl Anim Behav Sci 43(4):263-277.   DOI
71 Savory CJ 1995 Feather pecking and cannibalism. World's Poult Sci J 51(02):215-219.   DOI
72 Schenk PM, Meysser FM, Limpens HJG 1983 Gakeln als indikator für frustration bei Legehennen. K.T.B.L Schrift 299:65-81.
73 Shields SJ, Garner JP, Mench JA 2004 Dustbathing by broiler chickens: A comparison of preference for four different substrates. Appl Anim Behav Sci 87(1):69-82.   DOI
74 Scholz B, Ronchen S, Hamann H, Surie C, Neumann U, Kamphues JOSEF, Distl OTTMAR 2008 Evaluation of bone strength, keel bone deformity and egg quality of laying hens housed in small group housing systems and furnished cages in comparison to an aviary housing system. Archiv fur Tierzucht 51(2):179.
75 Scott GB, MacAngus G 2004 The ability of laying hens to negotiate perches of different materials with clean or dirty surfaces. Animal Welfare, 13(3):361-365.
76 Sherwin CM, Nicol CJ 1993 A descriptive account of the prelaying behaviour of hens housed individually in modified cages with nests. Appl Anim Behav Sci 38(1):49-60.   DOI
77 Shimmura T, Eguchi Y, Uetake K, Tanaka T 2007 Differences of behavior, use of resources and physical conditions between dominant and subordinate hens in furnished cages. Ani Sci J 78(3):307-313.   DOI
78 Simonsen HB, Vestergaard K, Willeberg P 1980 Effect of floor type and density on the integument of egg layers. Poult Sci 59:2002-2006.   DOI
79 Singh R, Cheng KM, Silversides FG 2009 Production performance and egg quality of four strains of laying hens kept in conventional cages and floor pens. Poult Sci 88(2):256-264.   DOI
80 Stampfli K, Buchwalder T, Frohlich EKF, Roth BA 2012 Influence of front curtain design on nest choice by laying hens. Br Poult Sci 53(5):553-560.   DOI
81 Struelens E, Tuyttens FAM, Ampe B, Odberg F, Sonck B, Duchateau L 2009 Perch width preferences of laying hens. Br Poult Sci 50(4):418-423.   DOI
82 Tauson R 1984 Effects of a perch in conventional cages for laying hens. Acta Agric Scand Sect A Anim Sci 34(2):193-209.   DOI
83 Struelens E, Van Nuffel A, Tuyttens FA, Audoorn L, Vranken E, Zoons J, Sonck B 2008 Influence of nest seclusion and nesting material on pre-laying behaviour of laying hens. Appl Anim Behav Sci 112(1):106-119.   DOI
84 Struelens E, Tuyttens FAM, Janssen A, Leroy T, Audoorn L, Vranken E, Sonck B 2005 Design of laying nests in furnished cages: Influence of nesting material, nest box position and seclusion. Br Poult Sci 46(1):9-15.   DOI
85 Tactacan GB, Guenter W, Lewis NJ, Rodriguez-Lecompte JC, House JD 2009 Performance and welfare of laying hens in conventional and enriched cages. Poult Sci 88(4):698-707.   DOI
86 Tauson R, Abrahamsson P 1996 Foot and keel bone disorders in laying hens: Effects of artificial perch material and hybrid. Acta Agric Scand Sect A Anim Sci 46(4):239-246.   DOI
87 Taylor AA, Hurnik JF 1996 The long-term productivity of hens housed in battery cages and an aviary. Poult Sci 75(1):47-51.   DOI
88 Van Hierden YM, Korte SM, Ruesink EW, van Reenen CG, Engel B, Koolhaas JM, Blokhuis HJ 2002 The development of feather pecking behaviour and targeting of pecking in chicks from a high and low feather pecking line of laying hens. Appl Anim Behav Sci 77(3):183-196.   DOI
89 Van Horne PLM 1996 Production and economic results of commercial flocks with white layers in aviary systems and battery cages. Br Poult Sci 37(2):255-261.   DOI
90 Van Krimpen MM, Kwakkel RP, Reuvekamp BFJ, Van Der Peet-Schwering CMC, Den Hartog LA, Verstegen MWA 2005 Impact of feeding management on feather pecking in laying hens. World's Poult Sci J 61(04):663-686.   DOI
91 Van Liere DW 1991 Function and organization of dustbathing in laying hens (Doctoral dissertation, Landbouwuniversiteit te Wageningen.).
92 Van Liere DW 1992 Dustbathing as related to proximal and distal feather lipids in laying hens. Behav Pro 26(2):177-188.   DOI
93 Van Liere DW, Bokma S 1987 Short-term feather maintenance as a function of dust-bathing in laying hens. Appl Anim Behav Sci 18(2):197-204.   DOI
94 Van Liere DW, Kooijman J, Wiepkema PR 1990 Dustbathing behaviour of laying hens as related to quality of dustbathing material. Appl Anim Behav Sci 26(1):127-141.   DOI
95 Van Liere DW, Wiepkema PR 1992 Effects of long-term deprivation of sand on dustbathing behaviour in laying hens. Anim Behav 43(4):549-558.   DOI
96 Van Rooijen J 2005 Dust bathing and other comfort behaviours of domestic hens. Welfare of laying hens in Europereports, analyses and conclusions (eds. G. Martin, HH Sambraus & A. Steiger), 110-123.
97 Vestergaard K 1982 Dust-bathing in the domestic fowl-diurnal rhythm and dust deprivation. Appl Anim Ethol 8(5):487-495.   DOI
98 Vestergaard KS, Damm BI, Abbott UK, Bildso EM 1999 Regulation of dustbathing in feathered and featherless domestic chicks: The Lorenzian model revisited. Anim Behav 58(5):1017-1025.   DOI
99 Appleby MC 1991 Do Hens Suffer in Battery Cages? A Review of the Scientific Evidence. The Athene Trust, Petersfield, UK.
100 Abrahamsson P, Tauson R 1997 Effects of group size on performance, health and birds' use of facilities in furnished cages for laying hens. Acta Agric Scand Sect A Anim Sci 47(4):254-260.   DOI
101 Appleby MC 1995 Perch length in cages for medium hybrid laying hens. Br Poult Sci 36(1):23-31.   DOI
102 Appleby MC, Duncan IJH 1989 Development of perching in hens. Biology of behaviour 14(2):157-168.
103 Appleby MC, Hughes BO, Cdonald MM, Cordiner LS 1998 Factors affecting the use of perches in cages by laying hens. Br Poult Sci 39(2):186-190.   DOI
104 Weeks CA, Nicol CJ 2006 Behavioural needs, priorities and preferences of laying hens. World's Poult Sci J 62(02):296-307.   DOI
105 Vestergaard KS, Kruijt JP, Hogan JA 1993 Feather pecking and chronic fear in groups of red junglefowl: Their relations to dustbathing, rearing environment and social status. Anim behav 45(6):1127-1140.   DOI
106 Wall H, Tauson R, Elwinger K 2002 Effect of nest design, passages, and hybrid on use of nest and production performance of layers in furnished cages. Poult Sci 81(3):333-339.   DOI
107 Wechsler B, Huber-Eicher B 1998 The effect of foraging material and perch height on feather pecking and feather damage in laying hens. Appl Anim Behav Sci 58(1):131-141.   DOI
108 Whitehead CC 2004 Overview of bone biology in the egglaying hen. Poult Sci 83(2):193-199.   DOI
109 Whitehead CC, Fleming RH 2000 Osteoporosis in cage layers. Poult Sci 79(7):1033-1041.   DOI
110 Wichman A, Heikkila M, Valros A, Forkman B, Keeling LJ 2007 Perching behaviour in chickens and its relation to spatial ability. Appl Anim Behav Sci 105(1):165-179.   DOI
111 Wilson S, Hughes BO, Appleby MC, Smith SF 1993 Effects of perches on trabecular bone volume in laying hens. Research in Veterinary Science 54(2):207-211.   DOI
112 Wood-Gush DGM 1971 The behaviour of the domestic fowl (No. 7). Heinemann Educational Books Ltd.
113 Wood-Gush DGM, Duncan IJH 1976 Some behavioural observations on domestic fowl in the wild. Appl Anim Ethol 2(3):255-260.   DOI
114 Appleby MC, Smith SF, Hughes BO 1992 Individual perching behaviour of laying hens and its effects in cages. Br Poult Sci 33(2):227-238.   DOI
115 Appleby MC, McRae HE 1986 The individual nest box as a super-stimulus for domestic hens. Appl Anim Behav Sci 15(2):169-176.   DOI
116 Appleby MC, McRae HE, Duncan IJH, Bisazza A 1984a Choice of social conditions by laying hens. Br Poult Sci 25(1):111-117   DOI
117 Appleby MC, McRae HE, Peitz BE 1984b The effect of light on the choice of nests by domestic hens. Appl Anim Ethol 11(3):249-254.   DOI
118 Appleby MC, Smith SF, Hughes BO 1993 Nesting, dust bathing and perching by laying hens in cages: Effects of design on behaviour and welfare. Br Poult Sci 34(5):835-847.   DOI
119 Barnett JL, Glatz PC, Newman EA, Cronin GM 1997 Effects of modifying layer cages with perches on stress physiology, plumage, pecking and bone strength of hens. Anim Pro Sci 37(5):523-529.   DOI
120 Bilcik B, Keeling LJ 1999 Changes in feather condition in relation to feather pecking and aggressive behaviour in laying hens. Br Poult Sci 40(4):444-451.   DOI
121 Bilcik B, Keeling LJ 2000 Relationship between feather pecking and ground pecking in laying hens and the effect of group size. Appl Anim Behav Sci 68(1):55-66.   DOI
122 Blokhuis HJ 1984 Rest in poultry. Appl Anim Behav Sci 12(3):289-303.   DOI
123 Blokhuis HJ 1986 Feather-pecking in poultry: Its relation with ground-pecking. Appl Anim Behav Sci 16(1):63-67.   DOI
124 Zimmerman PH, Koene P 1998 The effect of frustrative nonreward on vocalisations and behaviour in the laying hen, Gallus gallus domesticus. Behav Proc 44(1):73-79.   DOI
125 Wood-Gush DGM, Gilbert AB 1973 Some hormones involved in the nesting behaviour of hens. Anim Behav 21(1):98-103.   DOI
126 Yue S, Duncan IJH 2003 Frustrated nesting behaviour: Relation to extra-cuticular shell calcium and bone strength in White Leghorn hens. Br Poult Sci 44(2):175-181.   DOI
127 Zimmerman PH, Lindberg AC, Pope SJ, Glen E, Bolhuis JE, Nicol CJ 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(1):111-124.   DOI
128 Zimmerman PH, Koene P, van Hooff JA 2000 The vocal expression of feeding motivation and frustration in the domestic laying hen, Gallus gallus domesticus. Appl Anim Behav Sci 69(4):265-273.   DOI
129 Zupan M, Kruschwitz A, Buchwalder T, Huber-Eicher B, Stuhec I 2008 Comparison of the prelaying behavior of nest layers and litter layers. Poult Sci 87(3):399-404.   DOI
130 Zupan M, Kruschwitz A, Huber-Eicher B 2007 The influence of light intensity during early exposure to colours on the choice of nest colours by laying hens. Appl Anim Behav Sci 105(1):154-164.   DOI
131 Zupan M, Kruschwitz A, Huber-Eicher B, Witkowski A 2005 The influence of exposure to light and colour on the choice of nest colours in laying hens. In: Animal Science Papers and Reports. Polish Academy of Sciences, Institute of Genetics and Animal Breeding. 129-133.
132 Blokhuis HJ, Fiks van Niekerk T, Bessei W, Elson A, Guemene D, Kjaer JB, Van De Weerd HA 2007 The LayWel project: welfare implications of changes in production systems for laying hens. World's Poult Sci J 63(01):101-114.   DOI
133 Blokhuis HJ, Arkes JG 1984 Some observations on the development of feather-pecking in poultry. Appl Anim Behav Sci 12(1):145-157.   DOI
134 Blokhuis HJ, Beutler A 1992 Feather pecking damage and tonic immobility response in two lines of White Leghorn hens. J Anim Sci 70(Suppl. 1):170.
135 Blokhuis HJ, Beuving G 1993 Feather pecking and other characteristics in two lines of laying hens. In: Proceedings of the Fourth European Symposium on Poultry Welfare, Edinburgh, UK, pp. 266 - 267.
136 Brake J, Keeley TP, Jones RB 1994 Effect of age and presence of perches during rearing on tonic immobility fear reactions of broiler breeder pullets. Poult Sci 73(9):1470-1474.   DOI
137 Brendler C, Kipper S, Schrader L 2014 Vigilance and roosting behaviour of laying hens on different perch heights. Appl Anim Behav Sci 157:93-99.   DOI
138 Brunberg E, Jensen P, Isaksson A, Keeling L 2011 Feather pecking behavior in laying hens: Hypothalamic gene expression in birds performing and receiving pecks. Poult Sci 90(6):1145-1152.   DOI
139 Channing CE, Hughes BO, Walker AW 2001 Spatial distribution and behaviour of laying hens housed in an alternative system. Appl Anim Behav Sci 72(4):335-345.   DOI
140 Chen DH, Bao J, Meng FY, Wei CB 2014 Choice of perch characteristics by laying hens in cages with different group size and perching behaviours. Appl Anim Behav Sci 150:37-43.   DOI
141 Cooper JJ, Appleby MC 1995 Nesting behaviour of hens: Effects of experience on motivation. Appl Anim Behav Sci 42(4):283-295.   DOI
142 Chow A, Hogan JA 2005 The development of feather pecking in Burmese red junglefowl: The influence of early experience with exploratory-rich environments. Appl Anim Behav Sci 93(3):283-294.   DOI
143 Clausen T, Riber AB 2012 Effect of heterogeneity of nest boxes on occurrence of gregarious nesting in laying hens. Appl Anim Behav Sci 142(3):168-175.   DOI
144 Collias NE, Collias EC 1967 A field study of the red jungle fowl in north-central India. Condor. 360-386.
145 Cooper JJ, Appleby MC 1996 Demand for nest boxes in laying hens. Behav Proc 36(2):171-182.   DOI
146 Colson S, Arnould C, Michel V 2008 Influence of rearing conditions of pullets on space use and performance of hens placed in aviaries at the beginning of the laying period. Appl Anim Behav Sci 111(3):286-300.   DOI
147 Council directive 99/74/EC. laying down minimum standards for the protection of laying hens. Official Journal of the European Communities (L203/53), 19 July.
148 Dawkins MS 1983 Cage size and flooring preferences in litter reared and cage reared hens. Br Poult Sci 24(2):177-182.   DOI
149 Dawkins MS 1999 The role of behaviour in the assessment of poultry welfare. World's Poult Sci J 55(03):295-303.   DOI
150 De Jong IC, Wolthuis-Fillerup M, van Reenen CG 2007 Strength of preference for dustbathing and foraging substrates in laying hens. Appl Anim Behav Sci 104(1):24-36.   DOI
151 Duncan IJH, Kite VG 1989 Nest site selection and nest-building behaviour in domestic fowl. Anim Behav 37:215-231.   DOI
152 Dixon LM, Duncan IJH, Mason G 2008 What's in a peck? Using fixed action pattern morphology to identify the motivational basis of abnormal feather-pecking behaviour. Anim Behav 76(3):1035-1042.   DOI
153 Drake KA, Donnelly CA, Dawkins MS 2010 Influence of rearing and lay risk factors on propensity for feather damage in laying hens. Br Poult Sci 51(6):725-733.   DOI
154 Duncan ET, Appleby MC, Hughes BO 1992 Effect of perches in laying cages on welfare and production of hens. Br Poult Sci 33(1):25-35.   DOI
155 El-Lethey H, Aerni V, Jungi TW, Wechsler B 2000 Stress and feather pecking in laying hens in relation to housing conditions. Br Poult Sci 41(1):22-28.   DOI
156 Faure JM, Jones RB 1982 Effects of sex, strain and type of perch on perching behaviour in the domestic fowl. Appl Anim Behav Sci 8(3):281-293.
157 Folsch DW, Muller A, Dolf C 1986 Die bedeutung der einstreu fur huhner in den funktionsbereichen der nahrungssuche und korperpflege. KTBL-Schrift 311:168-176.
158 Freire R, Appleby MC, Hughes BO 1996 Effects of nest quality and other cues for exploration on pre-laying behaviour. Appl Anim Behav Sci 48(1):37-46.   DOI
159 Freire R, Appleby MC, Hughes BO 1998 Effects of social interactions on pre-laying behaviour in hens. Appl Anim Behav Sci 56(1):47-57.   DOI
160 Freire R, Walker A, Nicol CJ 1999 The relationship between trough height, feather cover and behaviour of laying hens in modified cages. Appl Anim Behav Sci 63(1):55-64.   DOI