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REVIEW

Advances in genetic selection and breeder practice improve commercial layer hen welfare

Greg Underwood https://orcid.org/0000-0002-5391-9884 A B D , Daniel Andrews B and Tin Phung C
+ Author Affiliations
- Author Affiliations

A Aves Consulting, PO Box 837, Gisborne, Vic. 3437, Australia.

B Bioproperties Pty Ltd, 36 Charter Street, Ringwood, Vic. 3134, Australia.

C Keans Free Range Eggs, Midland Hwy, Huntly, Vic. 3551, Australia.

D Corresponding author. Email: greg.underwood@bigpond.com

Animal Production Science 61(10) 856-866 https://doi.org/10.1071/AN20383
Submitted: 26 July 2021  Accepted: 23 September 2020   Published: 26 April 2021

Abstract

Modern commercial layer breeds represent the culmination of ~7000 years of natural genetic selection. This selection was driven in former times by a combination of genetic-shift and -drift events, that led to chickens being favoured as domesticated species for meat and egg production. More recently, in the early 20th century, the concept of hybrid vigour was discovered and accelerated the natural breeding progress that delivered new genetic lines and more favourable production traits. In the mid-20th century, the broiler-type and egg layer-type lines diverged and, in the 21st century, genetic analysis has further accelerated the progress made towards extended primary breeding characteristics such as egg quality, production and feed-intake traits, together with secondary breeding characteristics such as behavioural traits that have improved robustness in different housing systems, climates and feed types, which together have significantly improved welfare traits. Most recently, there has been the adoption of higher-powered computational analytics together with quantitative trait loci and single-nucleotide polymorphism assessment, which have further improved the uniformity of production traits within breeds. Most importantly, this has provided the primary breeding companies with improved and broader basis of selection of the modern commercial layer breeds, which also improved the alignment of layer strains with market requirements, and diverse variations in housing, nutritional and environmental conditions. This is also testament to the speed with which the commercial layer geneticists can respond to changing welfare policy on factors such as stocking density and beak treatment. The present paper reviews the modern approaches to genetic selection, including considerations of and benefits to the welfare state of commercial layers.

Keywords: animal breeding, egg production, genetics, poultry, welfare.


References

Appleby M, Walker AW, Nichol CJ, Lindberg AC, Friere R, Hughes BO, Elson HA (2002) Development of furnished cages for laying hens. British Poultry Science 43, 489–500.
Development of furnished cages for laying hens.Crossref | GoogleScholarGoogle Scholar | 12365505PubMed |

Basheer A, Haley CS, Law A, Windsor D, Morrice D, Talbot R, Wilson PW, Sharp PJ, Dunn IC (2015) Genetic loci inherited from hens lacking maternal behaviour both inhibit an paradoxically promote this behaviour. Genetics, Selection, Evolution 47, 100
Genetic loci inherited from hens lacking maternal behaviour both inhibit an paradoxically promote this behaviour.Crossref | GoogleScholarGoogle Scholar | 26718134PubMed |

Bedford A, Gong J (2018) Impications of butyrate and its derivatives for gut health and animal production. Animal Nutrition 4, 151–159.
Impications of butyrate and its derivatives for gut health and animal production.Crossref | GoogleScholarGoogle Scholar | 30140754PubMed |

Begka C, Marsland BJ (2018) Make (no) bones about butyrate. Immunity 49, 994–996.
Make (no) bones about butyrate.Crossref | GoogleScholarGoogle Scholar | 30566888PubMed |

Bell DJ, Siller W (1962) Cage layer fatigue in Brown Leghorns. Research in Veterinary Science 3, 219–230.
Cage layer fatigue in Brown Leghorns.Crossref | GoogleScholarGoogle Scholar |

Bennewitz J, Bogelein S, Stratz P, Rodehutscord M, Piepho HP, Kjaer JB, Bessei W (2014) Genetic parameters for feather pecking and aggressive behavior in a large F2-cross of laying hens using generalized linear mixed models. Poultry Science 93, 810–817.
Genetic parameters for feather pecking and aggressive behavior in a large F2-cross of laying hens using generalized linear mixed models.Crossref | GoogleScholarGoogle Scholar | 24706957PubMed |

Bessei W, Bauhaus H, Bogelein S (2013a) The effect of selection for high and low feather pecking on aggression-related behaviors of laying hens. Archiv für Geflügelkunde 77, 10–14.

Bessei W, Bogelein S, Bennewitz J, Kjaer JB (2013b) The relationships between social behavior and feather pecking in laying hens. In ‘Proceedings of the XIth European symposium’, Uppsala, Sweden. (Swedish Poultry Welfare)

Bestman MWP, Wagenaar JP (2003) Farm level factors associated with feather pecking in organic laying hens. Livestock Production Science 80, 133–140.
Farm level factors associated with feather pecking in organic laying hens.Crossref | GoogleScholarGoogle Scholar |

Bishop SC, Fleming RH, McCormack HA, Flock DK, Whitehead CC (2000) Inheritance of bone characteristics affecting osteoporosis in laying hens. British Poultry Science 41, 33–40.
Inheritance of bone characteristics affecting osteoporosis in laying hens.Crossref | GoogleScholarGoogle Scholar | 10821520PubMed |

Buitenhuis AJ, Rodenburg TB, van Hierden YM, Siwek M, Cornelissen SJ, Nieuwland MG, Crooijmans RP, Groenen MA, Koene P, Korte SM, Bovenhuis H, van der Poel JJ (2003) Mapping quantitative trait loci affecting feather pecking behavior and stress response in laying hens. Poultry Science 82, 1215–1222.
Mapping quantitative trait loci affecting feather pecking behavior and stress response in laying hens.Crossref | GoogleScholarGoogle Scholar | 12943291PubMed |

Campbell DLM (2020) Skeletal health of layers across all housing systems and future research directions for Australia. Animal Production Science.
Skeletal health of layers across all housing systems and future research directions for Australia.Crossref | GoogleScholarGoogle Scholar |

Campbell DLM, Bari MS, Rault J-L (2020) Free-range egg production: its implications for animal welfare. Animal Production Science.
Free-range egg production: its implications for animal welfare.Crossref | GoogleScholarGoogle Scholar |

Carter CT (1964) Modern trends in animal health and husbandry: poultry breeding. The British Veterinary Journal 120, 506–517.
Modern trends in animal health and husbandry: poultry breeding.Crossref | GoogleScholarGoogle Scholar |

Cavero D, Schmutz M, Philipp HC, Preisinger R (2009) Breeding to reduce susceptibility to Escherichia coli in layers. Poultry Science 88, 2063–2068.
Breeding to reduce susceptibility to Escherichia coli in layers.Crossref | GoogleScholarGoogle Scholar | 19762857PubMed |

Channing CE, Hughes BO, Walker AW (2001) Spatial distribution and behaviour of laying hens housed in an alternative system. Applied Animal Behaviour Science 72, 335–345.
Spatial distribution and behaviour of laying hens housed in an alternative system.Crossref | GoogleScholarGoogle Scholar | 11348682PubMed |

Cheng HH, Perumbakkam S, Pyrkosz AB, Dunn JR, Legarra A, Muir WM (2015) Fine mapping of QTL and genomic prediction using allele-specific expression SNPs demonstrates that the complex trait of genetic resistance to Marek’s disease is predominantly determined by transcriptional regulation. BMC Genomics 16, 816
Fine mapping of QTL and genomic prediction using allele-specific expression SNPs demonstrates that the complex trait of genetic resistance to Marek’s disease is predominantly determined by transcriptional regulation.Crossref | GoogleScholarGoogle Scholar | 26481588PubMed |

Christensen JP, Dietz HH, Bisgaard M (1998) Phenotypic and genotypic characters of isolates of Pasturella multocida obtained from back-yard poultry and from two outbreaks of avian cholera in avifauna in Denmark. Avian Pathology 27, 373–381.
Phenotypic and genotypic characters of isolates of Pasturella multocida obtained from back-yard poultry and from two outbreaks of avian cholera in avifauna in Denmark.Crossref | GoogleScholarGoogle Scholar | 18484015PubMed |

Couch JR (1955) Cage layer fatigue. Feed Age 5, 55–57.

Crawford RD (1990) Origin and history of poultry species. In ‘Poultry breeding and genetics’. (Ed. RD Crawford) pp. 1–42. (Elsevier)

Cronin GM, Glatz PC (2020) Causes of feather pecking and subsequent welfare issues for the laying hen: a review. Animal Production Science.
Causes of feather pecking and subsequent welfare issues for the laying hen: a review.Crossref | GoogleScholarGoogle Scholar |

Dawkins MS, Hardi S (1989) Space needs of laying hens. British Poultry Science 30, 413–416.
Space needs of laying hens.Crossref | GoogleScholarGoogle Scholar |

Edgar J, Held S, Jones C, Troisi C (2016) Influences of maternal care on chicken welfare. Animals (Basel) 6, 2
Influences of maternal care on chicken welfare.Crossref | GoogleScholarGoogle Scholar |

5m Editor (2011) ‘Celebrating 75 Years! The early years of Hy-line.’ Available at https://thepoultrysite.com/articles/celebrating-75-years-the-early-years-of-hyline [Verified January 2021]

Estevez I, Newberry RC, Keeling LJ (2002) Dynamics of aggression in the domestic fowl. Applied Animal Behaviour Science 76, 307–325.
Dynamics of aggression in the domestic fowl.Crossref | GoogleScholarGoogle Scholar |

Fathi MM, Ebeid TA, Al-Homidan I, Soliman NK, Abou-Emera OK (2017) Influence of probiotic supplementation on immune response in broilers raised under hot climate. British Poultry Science 58, 512–516.
Influence of probiotic supplementation on immune response in broilers raised under hot climate.Crossref | GoogleScholarGoogle Scholar | 28521530PubMed |

Freire R, Wilkins LJ, Short F, Nichol CJ (2003) Behaviour and welfare in of individual laying hens in a non-cage system. British Poultry Science 44, 22–29.
Behaviour and welfare in of individual laying hens in a non-cage system.Crossref | GoogleScholarGoogle Scholar | 12737221PubMed |

Fumihito A, Miyake T, Sumi S, Takada M, Ohno S, Kondo N (1994) One subspecies of the red junglefowl (Gallus gallus gallus) suffices as the matriarchic ancestor of all domestic breeds. Proceedings of the National Academy of Sciences of the United States of America 91, 12505–12509.
One subspecies of the red junglefowl (Gallus gallus gallus) suffices as the matriarchic ancestor of all domestic breeds.Crossref | GoogleScholarGoogle Scholar | 7809067PubMed |

Glatz PC, Underwood G (2020) Current methods and techniques of beak trimming layer hens, welfare issues and alternative approaches. Animal Production Science.
Current methods and techniques of beak trimming layer hens, welfare issues and alternative approaches.Crossref | GoogleScholarGoogle Scholar |

Godwin RM, Morgan JA (2015) A molecular survey of Eimeria in chickens across Australia. Veterinary Parasitology 214, 16–21.
A molecular survey of Eimeria in chickens across Australia.Crossref | GoogleScholarGoogle Scholar | 26467277PubMed |

Gregory NG, Wilkins LJ (1989) Broken bones in domestic fowl: handling and processing damage in end-of-lay battery hens. British Poultry Science 30, 555–562.
Broken bones in domestic fowl: handling and processing damage in end-of-lay battery hens.Crossref | GoogleScholarGoogle Scholar | 2819499PubMed |

Gross WB, Siegel PB (1985) Selective breeding of chickens for corticosterone response to social stress. Poultry Science 64, 2230–2233.
Selective breeding of chickens for corticosterone response to social stress.Crossref | GoogleScholarGoogle Scholar | 4095060PubMed |

Groves P (2021) Impact of parasites on Australian laying hen welfare. Animal Production Science
Impact of parasites on Australian laying hen welfare.Crossref | GoogleScholarGoogle Scholar | (In press).

Guo J, Sun C, Qu L, Shen M, Dou T, Ma M, Wang K, Yang N (2017) Genetic architecture of bone quality variation in layer chickens revealed by a genome-wide association study. Scientific Reports 7, 45317
Genetic architecture of bone quality variation in layer chickens revealed by a genome-wide association study.Crossref | GoogleScholarGoogle Scholar | 28383518PubMed |

Harlander-Matauschek A, Bessei W (2005) Feather eating and crop filling in laying hens. Archiv für Geflügelkunde 69, 241–244.

Hemsworth PH (2021) Cage production and laying hen welfare Animal Production Science.
Cage production and laying hen welfareCrossref | GoogleScholarGoogle Scholar |

Horsted K, Hermansen JE (2007) Whole wheat versus mixed layer diet as supplementary feed to layers foraging a sequence of different crops. Animal 1, 575–585.
Whole wheat versus mixed layer diet as supplementary feed to layers foraging a sequence of different crops.Crossref | GoogleScholarGoogle Scholar | 22444414PubMed |

Hughes BO (1983) Conventional and shallow cages: a summary of research from welfare and production aspects. World’s Poultry Science Journal 39, 218–228.
Conventional and shallow cages: a summary of research from welfare and production aspects.Crossref | GoogleScholarGoogle Scholar |

Hughes BO, Black AJ (1976) Battery cage shape: ITs effect on diurnal feeding pattern, egg-shell cracking and feather pecking. British Poultry Science 17, 327–336.
Battery cage shape: ITs effect on diurnal feeding pattern, egg-shell cracking and feather pecking.Crossref | GoogleScholarGoogle Scholar |

Hughes AL, Buitenhuis AJ (2010) Reduced variance of gene expression at numerous loci in a population of chickens selected for high feather pecking. Poultry Science 89, 1858–1869.

Icken W, Cavero D, Schmutz M (2017) Selection on beak shape to reduce feather pecking in laying hens. Lohmann Information 51, 22–27.

International Chicken Genome Sequencing Consortium (2004) Sequence and comparative analysis of the chicken genome provide unique perspectives on vertebrate evolution. Nature 432, 695–716.
Sequence and comparative analysis of the chicken genome provide unique perspectives on vertebrate evolution.Crossref | GoogleScholarGoogle Scholar | 15592404PubMed |

Jensen P (2014) Behaviour genetics and the domestication of animals. Annual Review of Animal Biosciences 2, 85–104.
Behaviour genetics and the domestication of animals.Crossref | GoogleScholarGoogle Scholar | 25384136PubMed |

Jensen P, Andersson L (2005) Genomics meets ethology: a new route to understanding domestication, behaviour, and sustainability in animal breeding. Ambio 34, 320–324.
Genomics meets ethology: a new route to understanding domestication, behaviour, and sustainability in animal breeding.Crossref | GoogleScholarGoogle Scholar | 16092263PubMed |

Johnsson M, Jonsson KB, Andersson L, Jensen P, Wright D (2015) Genetic regulation of bone metabolism in the chicken: similarities and differences to mammalian systems. PLOS Genetics 11, e1005250
Genetic regulation of bone metabolism in the chicken: similarities and differences to mammalian systems.Crossref | GoogleScholarGoogle Scholar | 26023928PubMed |

Kaiser P, Underwood G, Davison F (2003) Differential cytokine responses following Marek’s disease virus infection of chickens differing in resistance to Marek’s disease. Journal of Virology 77, 762–768.
Differential cytokine responses following Marek’s disease virus infection of chickens differing in resistance to Marek’s disease.Crossref | GoogleScholarGoogle Scholar | 12477883PubMed |

Kaiser P, Howell MM, Fife M, Sadeyen JR, Salmon N, Rothwell L, Young J, Poh TY, Stevens M, Smith J, Burt D, Swaggerty C, Kogut M (2009) Towards the selection of chickens resistant to Salmonella and Campylobacter infections. Bulletin et Memoires de l’Academie Royale de Medecine de Belgique 164, 17–25, discussion 25–26.

Karlsson AC, Svemer F, Eriksson J, Darras VM, Andersson L, Jensen P (2015) The effect of a mutation in the thyroid stimulating hormone receptor (TSHR) on development, behaviour and TH levels in domesticated chickens. PLoS One 10, e0129040
The effect of a mutation in the thyroid stimulating hormone receptor (TSHR) on development, behaviour and TH levels in domesticated chickens.Crossref | GoogleScholarGoogle Scholar | 26402620PubMed |

Ketelaars, EH (1992) ‘Historia van de Nederlandse pluimveehouderji; van kippenboer tot specialist.’

Kim WK, Ford BC, Mitchell AD, Elkin RG, Leach RM (2004) Comparative assessment of bone among wild-type, restricted ovulator and out-of-production hens. British Poultry Science 45, 463–470.
Comparative assessment of bone among wild-type, restricted ovulator and out-of-production hens.Crossref | GoogleScholarGoogle Scholar | 15484719PubMed |

Kjaer JB, Bessei W (2013) The interrelationship of nutrition and feather pecking in the domestic fowl: a review. Archiv für Geflügelkunde 77, 1–9.

Kjaer JB, Sorensen P, Su G (2001) Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus). Applied Animal Behaviour Science 71, 229–239.
Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus).Crossref | GoogleScholarGoogle Scholar | 11230903PubMed |

Lagerström MC, Hellstrom AR, Gloriam DE, Larsson TP, Schioth HB, Fredriksson R (2006) The G protein-coupled receptor subset of the chicken genome. PLoS Computational Biology 2, e54
The G protein-coupled receptor subset of the chicken genome.Crossref | GoogleScholarGoogle Scholar | 16741557PubMed |

Leenstra F, Ten Napel J, Van Sambeek F (2016) Layer breeding programmes in changing production environments: a historic perspective. World’s Poultry Science Journal 72, 21–36.
Layer breeding programmes in changing production environments: a historic perspective.Crossref | GoogleScholarGoogle Scholar |

Liu L, Lin L, Zheng L, Tang H, Fan X, Xue N, Li M, Liu M, Li X (2018) Cecal microbiome profile altered by Salmonella enterica, serovar Enteritidis inoculation in chicken. Gut Pathogens 10, 34
Cecal microbiome profile altered by Salmonella enterica, serovar Enteritidis inoculation in chicken.Crossref | GoogleScholarGoogle Scholar | 30087697PubMed |

Loudovaris T, Yoo BH, Fahey KJ (1991) Genetic resistance to infectious laryngotracheitis in inbred lines of White Leghorn chickens. Avian Pathology 20, 357–361.
Genetic resistance to infectious laryngotracheitis in inbred lines of White Leghorn chickens.Crossref | GoogleScholarGoogle Scholar | 18680029PubMed |

Macia L, Tan J, Vieira AT, Leach K, Stanley D, Luong S, Maruya M, Ian McKenzie C, Hijikata A, Wong C, Binge L, Thorburn AN, Chevalier N, Ang C, Marino E, Robert R, Offermanns S, Teixeira MM, Moore RJ, Flavell RA, Fagarasan S, Mackay CR (2015) Metabolite-sensing receptors GPR43 and GPR109A facilitate dietary fibre-induced gut homeostasis through regulation of the inflammasome. Nature Communications 6, 6734
Metabolite-sensing receptors GPR43 and GPR109A facilitate dietary fibre-induced gut homeostasis through regulation of the inflammasome.Crossref | GoogleScholarGoogle Scholar | 25828455PubMed |

McKeegan DEF, Savory CJ (2001) Feather eating in individually caged hens which differ in their propensity to feather peck. Applied Animal Behaviour Science 73, 131–140.
Feather eating in individually caged hens which differ in their propensity to feather peck.Crossref | GoogleScholarGoogle Scholar |

Meyer B, Zentek J, Harlander-Matauschek A (2013) Differences in intestinal microbial metabolites in laying hens with high and low levels of repetitive feather-pecking behavior. Physiology & Behavior 110–111, 96–101.
Differences in intestinal microbial metabolites in laying hens with high and low levels of repetitive feather-pecking behavior.Crossref | GoogleScholarGoogle Scholar |

Mika M, Nita I, Morf L, Qi W, Beyeler S, Bernasconi E, Marsland BJ, Ott SR, von Garnier C, Hilty M (2018) Microbial and host immune factors as drivers of COPD. ERJ Open Research 4, 00015-2018
Microbial and host immune factors as drivers of COPD.Crossref | GoogleScholarGoogle Scholar | 30519565PubMed |

Miyazaki T, Tokimura F, Tanaka S (2014) A review of denosumab for the treatment of osteoporosis. Patient Preference and Adherence 8, 463–471.
A review of denosumab for the treatment of osteoporosis.Crossref | GoogleScholarGoogle Scholar | 24748775PubMed |

Moinard C, Morisse JP, Faure JM (1998) Effect of cage area, cage height and perches on feather condition, bone breakage and mortality of laying hens. British Poultry Science 39, 198–202.
Effect of cage area, cage height and perches on feather condition, bone breakage and mortality of laying hens.Crossref | GoogleScholarGoogle Scholar | 9649871PubMed |

Muir WM (1996) Group selection for adaptation to multiple hen cages: selection program and direct responses. Poultry Science 75, 447–458.
Group selection for adaptation to multiple hen cages: selection program and direct responses.Crossref | GoogleScholarGoogle Scholar | 8786932PubMed |

Newberry RC (2004) Cannibalism. In ‘Welfare of the laying hen’. (Ed. GC Perry) pp. 239–258. (CABI Publishing: Wallingford, UK)

Newberry RC, Estevez I, Keeling LJ (2001) Group size and perching behaviour in young domestic fowl. Applied Animal Behaviour Science 73, 117–129.
Group size and perching behaviour in young domestic fowl.Crossref | GoogleScholarGoogle Scholar | 11358609PubMed |

Nicol CJ (1987) Effect of cage height and area on the behaviour of hens housed in battery cages. British Poultry Science 28, 327–335.
Effect of cage height and area on the behaviour of hens housed in battery cages.Crossref | GoogleScholarGoogle Scholar | 3607556PubMed |

Nicol CJ, Gregory NJ, Knowles T, Parkman I, Wilkins LJ (1999) Differential effects of increased stocking density, mediated by increased flock size, on feather pecking and aggression in laying hens. Applied Animal Behaviour Science 65, 137–152.
Differential effects of increased stocking density, mediated by increased flock size, on feather pecking and aggression in laying hens.Crossref | GoogleScholarGoogle Scholar |

Noormohammadi AH (2020) Welfare implications of bacterial and viral infectious diseases for laying hens. Animal Production Science.
Welfare implications of bacterial and viral infectious diseases for laying hens.Crossref | GoogleScholarGoogle Scholar |

Preisinger R (2018) Innovative layer genetics to handle global challenges in egg production. British Poultry Science 59, 1–6.
Innovative layer genetics to handle global challenges in egg production.Crossref | GoogleScholarGoogle Scholar | 29129115PubMed |

Psifidi A, Russell KM, Matika O, Sanchez-Molano E, Wigley P, Fulton JE, Stevens MP, Fife MS (2018) The genomic architecture of fowl typhoid resistance in commercial layers. Frontiers in Genetics 9, 519
The genomic architecture of fowl typhoid resistance in commercial layers.Crossref | GoogleScholarGoogle Scholar | 30510562PubMed |

Riedstra B, Groothuis TGG (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.

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. Poultry Science 82, 861–867.
Heritability of feather pecking and open-field response of laying hens at two different ages.Crossref | GoogleScholarGoogle Scholar | 12817438PubMed |

Rubin CJ, Zody MC, Eriksson J, Meadows JR, Sherwood E, Webster MT, Jiang L, Ingman M, Sharpe T, Ka S, Hallbook F, Besnier F, Carlborg O, Bed’hom B, Tixier-Boichard M, Jensen P, Siegel P, Lindblad-Toh K, Andersson L (2010) Whole-genome resequencing reveals loci under selection during chicken domestication. Nature 464, 587–591.
Whole-genome resequencing reveals loci under selection during chicken domestication.Crossref | GoogleScholarGoogle Scholar | 20220755PubMed |

Schmutz J, Grimwood J (2004) Genomes: fowl sequence. Nature 432, 679–680.
Genomes: fowl sequence.Crossref | GoogleScholarGoogle Scholar | 15592391PubMed |

Singh M, Groves PJ (2020) Welfare implications for barn (and aviary) egg production systems. Animal Production Science.
Welfare implications for barn (and aviary) egg production systems.Crossref | GoogleScholarGoogle Scholar |

Su G, Kjaer JB, Sorensen P (2005) Variance components and selection response for feather-pecking behavior in laying hens. Poultry Science 84, 14–21.
Variance components and selection response for feather-pecking behavior in laying hens.Crossref | GoogleScholarGoogle Scholar | 15685937PubMed |

Swaggerty CL, Pevzner IY, He H, Genovese KJ, Nisbet DJ, Kaiser P, Kogut MH (2009) Selection of broilers with improved innate immune responsiveness to reduce on-farm infection by foodborne pathogens. Foodborne Pathogens and Disease 6, 777–783.
Selection of broilers with improved innate immune responsiveness to reduce on-farm infection by foodborne pathogens.Crossref | GoogleScholarGoogle Scholar | 19737057PubMed |

Swaggerty CL, McReynolds JL, Byrd JA, Pevzner IY, Duke SE, Genovese KJ, He H, Kogut MH (2016) Selection for pro-inflammatory mediators produces chickens more resistant to Clostridium perfringens-induced necrotic enteritis. Poultry Science 95, 370–374.
Selection for pro-inflammatory mediators produces chickens more resistant to Clostridium perfringens-induced necrotic enteritis.Crossref | GoogleScholarGoogle Scholar | 26706357PubMed |

Swaggerty CL, Pevzner IY, He H, Genovese KJ, Kogut MH (2017) Selection for pro-inflammatory mediators produces chickens more resistant to Campylobacter jejuni. Poultry Science 96, 1623–1627.
Selection for pro-inflammatory mediators produces chickens more resistant to Campylobacter jejuni.Crossref | GoogleScholarGoogle Scholar | 28339707PubMed |

Tauson R (1998) Health and production in improved cage designs. Poultry Science 77, 1820–1827.
Health and production in improved cage designs.Crossref | GoogleScholarGoogle Scholar | 9872585PubMed |

Ugalde XA (2019) Path to the 100 week age layer hen in cage free systems. In ‘30th Annual Australian poultry science symposium’, Sydney, New South Wales, Australia. (Ed. J Roberts) (The University of Sydney: Sydney, NSW, Australia)

Underwood GJ, Andrews D, Phung T, Edwards L (2021) Incubation, hatchery practice and the welfare of layer hens. Animal Production Science
Incubation, hatchery practice and the welfare of layer hens.Crossref | GoogleScholarGoogle Scholar | (In press).

Uni Z, Yadgary L, Yair R (2012) Nutritional limitations during poultry embryonic development. Journal of Applied Poultry Research 21, 175–184.
Nutritional limitations during poultry embryonic development.Crossref | GoogleScholarGoogle Scholar |

West B, Zhou B-X (1988) Did chickens go north? New evidence for domestication. Journal of Archaeological Science 15, 515–533.
Did chickens go north? New evidence for domestication.Crossref | GoogleScholarGoogle Scholar |

Whitehead CC, Wilson S (1992) Characterisitcs of osteopenia in hens. In ‘Bone biology and skeletal disorders in poultry’. (Ed. CC Whitehead) pp. 265–280. (Carfax Publishers: Abdingdon, UK)

Widowski T, Duncan IJH (2000) Working form a dustbath: are hens increasing pleasure rather than reducing suffering? Applied Animal Behaviour Science 68, 39–53.
Working form a dustbath: are hens increasing pleasure rather than reducing suffering?Crossref | GoogleScholarGoogle Scholar | 10771314PubMed |

Wilkins LJ, Brown SN, Zimmerman PH, Leeb C, Nicol CJ (2004) Investigation of palpation as a method for determining the prevelance of keel and furculum damage in laying hens. The Veterinary Record 155, 547–549.
Investigation of palpation as a method for determining the prevelance of keel and furculum damage in laying hens.Crossref | GoogleScholarGoogle Scholar | 15559420PubMed |

Wysocki M, Bessei W, Kjaer JB, Bennewitz J (2010) Genetic and physiological factors influencing feather pecking in chickens. World’s Poultry Science Journal 66, 659–671.
Genetic and physiological factors influencing feather pecking in chickens.Crossref | GoogleScholarGoogle Scholar |

Xu L, He Y, Ding Y, Liu GE, Zhang H, Cheng HH, Taylor RL, Song J (2018) Genetic assessment of inbred chicken lines indicates genomic signatures of resistance to Marek’s disease. Journal of Animal Science and Biotechnology 9, 65
Genetic assessment of inbred chicken lines indicates genomic signatures of resistance to Marek’s disease.Crossref | GoogleScholarGoogle Scholar | 30221000PubMed |

Zeuner FE (1963) ‘A history of domesticated animals.’ (Hutchinson and Co.: London, UK)