exercise, manipulative activities, and cognitive challenges according to species-specific characteristics (NRC 1998a; Young 2003). Examples of enrichment include structural additions such as perches and visual barriers for nonhuman primates (Novak et al. 2007); elevated shelves for cats (Overall and Dyer 2005; van den Bos and de Cock Buning 1994) and rabbits (Stauffacher 1992); and shelters for guinea pigs (Baumans 2005), as well as manipulable resources such as novel objects and foraging devices for nonhuman primates; manipulable toys for nonhuman primates, dogs, cats, and swine; wooden chew sticks for some rodent species; and nesting material for mice (Gaskill et al. 2009; Hess et al. 2008; Hubrecht 1993; Lutz and Novak 2005; Olsson and Dahlborn 2002). Novelty of enrichment through rotation or replacement of items should be a consideration; however, changing animals’ environment too frequently may be stressful.

Well-conceived enrichment provides animals with choices and a degree of control over their environment, which allows them to better cope with environmental stressors (Newberry 1995). For example, visual barriers allow nonhuman primates to avoid social conflict; elevated shelves for rabbits and shelters for rodents allow them to retreat in case of disturbances (Baumans 1997; Chmiel and Noonan 1996; Stauffacher 1992); and nesting material and deep bedding allow mice to control their temperature and avoid cold stress during resting and sleeping (Gaskill et al. 2009; Gordon 1993, 2004).

Not every item added to the animals’ environment benefits their wellbeing. For example, marbles are used as a stressor in mouse anxiety studies (De Boer and Koolhaas 2003), indicating that some items may be detrimental to well-being. For nonhuman primates, novel objects can increase the risk of disease transmission (Bayne et al. 1993); foraging devices can lead to increased body weight (Brent 1995); shavings can lead to allergies and skin rashes in some individuals; and some objects can result in injury from foreign material in the intestine (Hahn et al. 2000). In some strains of mice, cage dividers and shelters have induced overt aggression in groups of males, resulting in social stress and injury (e.g., Bergmann et al. 1994; Haemisch et al. 1994). Social stress was most likely to occur when resources were monopolized by dominant animals (Bergmann et al. 1994).

Enrichment programs should be reviewed by the IACUC, researchers, and veterinarian on a regular basis to ensure that they are beneficial to animal well-being and consistent with the goals of animal use. They should be updated as needed to ensure that they reflect current knowledge. Personnel responsible for animal care and husbandry should receive training in the behavioral biology of the species they work with to appropriately monitor the effects of enrichment as well as identify the development of adverse or abnormal behaviors.

Like other environmental factors (such as space, light, noise, temperature, and animal care procedures), enrichment affects animal phenotype



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