Hamster wheel

Like other rodents, hamsters are highly motivated to run in wheels.

Hamster wheels or running wheel are exercise devices used primarily by hamsters and other rodents, but also by other cursorial animals when given the opportunity. Most of these devices consist of a runged or ridged wheel held on a stand by a single or pair of stub axles. Hamster wheels allow rodents to run even when their space is confined. The earliest dated use of the term "hamster wheel", located by the Oxford English Dictionary, is in a 1949 newspaper advertisement.[1]

Preferences

Most wheels are constructed of steel or plastic, both of which have advantages and disadvantages. Solid plastic wheels are safer for some types of animals, such as hamsters and hedgehogs, because the animal's feet or legs cannot get trapped and injured between rungs. However, some rodents (e.g. gerbils) will quickly chew and destroy plastic wheels but not steel versions.

Choice tests with Syrian hamsters (Mesocricetus auratus) have shown that they prefer larger wheels; the animals chose a wheel diameter of 35 cm (14 in) over 23 cm (9 in),[2] which itself was preferred over 17.5 cm (7 in).[3]

Hamsters showed no preference between a relatively uniform running surface made of plastic mesh and a surface made of rungs spaced 9 mm apart, although they did prefer the mesh compared to rungs spaced 12 mm apart, most likely because the wider space between the rungs allowed their legs to slip through.[3] The hamsters neither preferred nor avoided wheels that had small "speed bumps" installed along the running surface to provide environmental enrichment.[2]

Choice tests with mice have also shown a preference for larger wheels (17.5 cm over 13 cm in diameter) and a preference for plastic mesh over rungs and over solid plastic as a running surface.[4] More acrobatic species, such as the canyon mouse (Peromyscus crinitus) and the deer mouse (Peromyscus maniculatus) can develop preferences for wheels that force the animals to jump, such as square wheels or wheels with hurdles along the running surface.[5][6][7]

Use by animals

Like other rodents, hamsters are highly motivated to run in wheels; it is not uncommon to record distances of 9 km (5.6 mi) being run in one night. Other 24-h records include 43 km (27 mi) for rats, 31 km (19 mi) for wild mice, 19 km (12 mi) for lemmings, 16 km (9.9 mi) for laboratory mice, and 8 km (5.0 mi) for gerbils.[8] Hypotheses to explain such high levels of running in wheels include a need for activity, substitute for exploration, and stereotypic behaviour. However, free wild mice will run on wheels installed in the field, which speaks against the notion of stereotypic behaviour induced by captivity conditions.[9] Alternatively, various experimental results strongly indicate that wheel-running, like play or the endorphin or endocannabinoid[10] release associated with the 'runner's high', is self-rewarding.[8][11][12] Wheel use is highly valued by several species as shown in consumer demand studies which require an animal to work for a resource, i.e. bar-press or lift weighted doors.[8][13] This makes running wheels a popular type of enrichment to the captivity conditions of rodents.

Captive animals continue to use wheels even when provided with other types of enrichment. In one experiment, Syrian hamsters that could use tunnels to access five different cages each containing a toy, showed no more than a 25% reduction in running-wheel use compared to hamsters housed in a single cage without toys (except for the running wheel).[14] In another study, female Syrian hamsters housed with a nestbox, bedding, hay, paper towels, cardboard tubes, and branches used a wheel regularly, and benefitted from it as indicated by showing less stereotypic bar-gnawing and producing larger litters of young compared to females kept under the same conditions but without a wheel.[15] Laboratory mice were prepared to perform more switch presses to enter a cage containing a running wheel compared to several metres of Habitrail tubing or a torus of Habitrail tubing.[16]

Running in wheels can be so intense in hamsters that it may result in foot lesions, which appear as small cuts on the paw pads or toes.[17][18] Such paw wounds rapidly scab over and do not prevent hamsters from continuing to run in their wheel.

Use in science

Voluntary wheel running is one of the most widely used indicators of activity and wake-time in research on circadian rhythms and other aspects of chronobiology.[19] Miniature running wheels have even been used to measure the circadian locomotor activity of cockroaches.[20][21] For rodents, running wheels are easier to set up and automate than other techniques of activity recording such as bar-gnawing[22] and spring-suspended or knife-edge balanced cages.[23]

In rodents, voluntary exercise is almost always measured by use of wheels. This makes running wheels the tool of choice in research on the effects of exercise and voluntary activity on metabolism, obesity, and pain.[24][25][26][27][28]

The neurotransmitter systems involved in wheel-running behavior have received considerable study.[29] Recent evidence suggests that changes in both dopaminergic and serotonergic tone alter running-wheel activity. For example, one study in mice has shown that several antidepressant medications (all of which directly or indirectly enhance serotonergic tone) suppress running-wheel activity without suppressing general locomotion.[30] The endocannabinoid system also contributes to wheel running in a sex-specific manner in rodents.[31] Mice from lines that have been selectively bred for high levels of voluntary wheel running have altered responsiveness to drugs that alter dopamine and endocannabinoid signalling, and enlarged midbrains.[32]

Hamster ball

A mouse in green hamster ball

A related exercise device, the hamster ball, is a hollow plastic ball into which a pet can be temporarily placed. The ball allows the pet to freely roll around on the floor to explore and exercise, while preventing escape.

Running disc

A related exercise device is the running disc. This is a rotatable shallow bowl, or slightly concave disc, which is set an angle to the horizontal. Some commercial refuges for caged rodents have a disc mounted on the roof at a slight angle. The rodents run on the rim of the disc in a similar way to running in hamster wheels.

See also

References

  1. ↑ Los Angeles Times, 9 Oct. 1949, p. B20/4 (advt.), referenced in Oxford English Dictionary, Draft Additions, Sep. 2007, under entry “Hamster.”
  2. 1 2 Reebs, S.G. and St-Onge, P., 2005, Running wheel choice by Syrian hamsters, Laboratory Animals, 39: 442–451
  3. 1 2 Mrosovsky, N., Salmon, P.A. and Vrang, N., 1998, Revolutionary science: an improved running wheel for hamsters, Chronobiology International, 15: 147–158.
  4. ↑ Banjanin, S., and Mrosovsky, N., 2000, Preferences of mice, Mus musculus, for different types of running wheel, Laboratory Animals, 34: 313–318.
  5. ↑ Kavanau, J.L., and Brant, D.H., 1965, Wheel-running preferences of Peromyscus, Nature, 208: 597–98
  6. ↑ Kavanau, J.L., 1966, Wheel-running preferences of mice, Zeitschrift für Tierpsychologie 23: 858–66
  7. ↑ Kavanau, J.L., 1967, Behavior of captive white-footed mice, Science, 155: 1623–39.
  8. 1 2 3 Sherwin, C.M., 1998, Voluntary wheel running: A review and novel interpretation, Animal Behaviour, 56: 11–27
  9. ↑ Meijer, Johanna H.; Robbers, Yuri, "Wheel running in the wild", Proceedings of the Royal Society B, 281 (1786), doi:10.1098/rspb.2014.0210
  10. ↑ Raichlen, D. A., A. D. Foster, G. L. Gerdeman, A. Seillier, and A. Giuffrida. 2012. Wired to run: exercise-induced endocannabinoid signaling in humans and cursorial mammals with implications for the 'runner's high'. Journal of Experimental Biology 215:1331-1336.
  11. ↑ Novak, C.M., Burghardt, P.R. and Levine, J.A., 2012, The use of a running wheel to measure activity in rodents: Relationship to energy balance, general activity, and reward, Neuroscience and Biobehavioral Reviews, 36: 1001–1014
  12. ↑ Belke, T.W., and Wagner, J.P. 2005. The reinforcing property and the rewarding aftereffect of wheel running in rats: a combination of two paradigms. Behavioural Processes 68: 165-172.
  13. ↑ Belke, T.W. and Garland, T., Jr., 2007, A brief opportunity to run does not function as a reinforcer for mice selected for high daily wheel-running rates, Journal of the Experimental Analysis of Behavior, 88: 199-213
  14. ↑ Reebs, S.G., and Maillet, D., 2003, Effect of cage enrichment on the daily use of running wheels by Syrian hamsters, Chronobiology International, 20: 9-20.
  15. ↑ Gebhardt-Henrich, S.G., Vonlanthen, E.M., and Steiger, A., 2005, How does the running wheel affect the behaviour and reproduction of golden hamsters kept as pets, Applied Animal Behaviour Science, 95: 199-203.
  16. ↑ Sherwin, C.M., 1998, The use and perceived importance of three resources which provide caged laboratory mice the opportunity of extended locomotion, Applied Animal Behaviour Science, 55: 353-367.
  17. ↑ Beaulieu, A., and Reebs, S.G., 2009, Effects of bedding material and running wheel surface on paw wounds in male and female Syrian hamsters, Laboratory Animals, 43: 85-90.
  18. ↑ Veillette, M., Guitard, J., and Reebs, S.G., 2010, Cause and possible treatments of foot lesions in captive Syrian hamsters (Mesocricetus auratus), Veterinary Medicine International, article ID 951708, doi:10.4061/2010/951708 .
  19. ↑ Dunlap, J.C., Loros, J.J., and DeCoursey, P.J. 2004. Chronobiology: Biological Timekeeping. Sinauer Associates, Sunderland MA.
  20. ↑ Binkley, S. 1990. The clockwork sparrow: time, clocks, and calendars in biological organisms. Prentice-Hall, Englewood Cliffs NJ.
  21. ↑ Roberts. S. 1965. Photoreception and entrainment of cockroach activity rhythms. Science 148: 958-959.
  22. ↑ Morin, L. 1978. Rhythmicity of hamster gnawing: Ease of measurement and similarity to running activity. Physiology and Behavior 21: 317-320
  23. ↑ Aschoff, J., Figala, J., and Poppel, E. 1973. Circadian rhythms of locomotor activity in the golden hamster (Mesocricetus auratus) measured with two different techniques. Journal of Comparative and Physiological Psychology 85: 20-28.
  24. ↑ Garland, T., Jr., H. Schutz, M. A. Chappell, B. K. Keeney, T. H. Meek, L. E. Copes, W. Acosta, C. Drenowatz, R. C. Maciel, G. van Dijk, C. M. Kotz, and J. C. Eisenmann. 2011. The biological control of voluntary exercise, spontaneous physical activity and daily energy expenditure in relation to obesity: human and rodent perspectives. Journal of Experimental Biology 214:206-229.
  25. ↑ Coutinho A.E., Fediuc S., Campbell J.E., and Riddell, M.C. 2006. Metabolic effects of voluntary wheel running in young and old Syrian golden hamsters. Physiology and Behavior 87: 360-367.
  26. ↑ Gattermann R., Weinandy R., and Fritzsche P. 2004. Running-wheel activity and body composition in golden hamsters (Mesocricetus auratus). Physiology and Behavior 82: 541-544.
  27. ↑ Kandasamy, R., Calsbeek, J. J., & Morgan, M. M. (2016). Home cage wheel running is an objective and clinically relevant method to assess inflammatory pain in male and female rats. Journal of neuroscience methods, 263, 115-122.
  28. ↑ Cobos, E. J., Ghasemlou, N., Araldi, D., Segal, D., Duong, K., & Woolf, C. J. (2012). Inflammation-induced decrease in voluntary wheel running in mice: a nonreflexive test for evaluating inflammatory pain and analgesia. PAIN®, 153(4), 876-884.
  29. ↑ Rhodes, J.S., Gammie, S.C. and Garland, T., Jr., 2005, Neurobiology of mice selected for high voluntary wheel-running activity, Integrative and Comparative Biology, 45: 438-455.
  30. ↑ Weber, M; Talmon S; Schulze I; Boeddinghaus C; Gross G; Schoemaker H; Wicke KM (May 2009). "Running wheel activity is sensitive to acute treatment with selective inhibitors for either serotonin or norepinephrine reuptake". Psychopharmacology. 203 (4): 753–762. doi:10.1007/s00213-008-1420-4.
  31. ↑ Keeney, B.K., Meek, T.H., Middleton, K.M., Holness, L. F. and Garland, T., Jr., 2012, Sex differences in cannabinoid receptor-1 (CB1) pharmacology in mice selectively bred for high voluntary wheel-running behavior, Pharmacology, Biochemistry and Behavior, 101: 528-537.
  32. ↑ Kolb, E. M., E. L. Rezende, L. Holness, A. Radtke, S. K. Lee, A. Obenaus, and T. Garland, Jr. 2013. Mice selectively bred for high voluntary wheel running have larger midbrains: support for the mosaic model of brain evolution. Journal of Experimental Biology 216:515-523.
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