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

Despite several decades of research, the epigenesis of behavioural and brain lateralization is still elusive, although its knowledge is important in understanding developmental plasticity, function and evolution of lateralization, and its relationship with developmental disorders. Over the last decades, it has become clear that behavioural lateralization is not restricted to humans, but a fundamental principle in the organization of behaviour in vertebrates. This has opened the possibility of extending descriptive studies on human lateralization with descriptive and experimental studies on other vertebrate species. In this review, we therefore explore the evidence for the role of genes and environment on behavioural lateralization in humans and other animals. First, we discuss the predominant genetic models for human handedness, and conclude that their explanatory power alone is not sufficient, leaving, together with ambiguous results from adoption studies and selection experiments in animals, ample opportunity for a role of environmental factors. Next, we discuss the potential influence of such factors, including perinatal asymmetrical perception induced by asymmetrical head position or parental care, and social modulation, both in humans and other vertebrates, presenting some evidence from our own work on the domestic chick. We conclude that both perinatal asymmetrical perception and later social modulation are likely candidates in influencing the degree or strength of lateralization in both humans and other vertebrates. However, in most cases unequivocal evidence for this is lacking and we will point out further avenues for research.