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Fish, Shoaling Behavior, and the Role of Pheromones

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Fish, Shoaling Behavior, and the Role of Pheromones
Introduction Shoaling is a widely important mechanism used by fish in freshwater and marine environments. Although used rather loosely in conjunction with the term “schooling,” the two words differ in definition by nature of organization. Where shoaling refers to a congregation of conspecific fish that stay together, schooling indicates that those fish are swimming in a coordinated fashion. Shoaling is not a learned behavior, but one in which a species is born to participate. This conduct evolved in as many as 50% of all fish species for a number of reasons. The incentives for shoaling mostly stem from the simple premise that all animals have three jobs in life: eat, reproduce, and avoid being eaten. From this, we can derive the motives for shoaling behavior. To facilitate these life goals, natural selection has chosen for shoaling behavior to increase foraging success, reproductive convenience, hydrodynamic efficiency, social interaction, and predator avoidance (Moyle and Cech 2003, Brahams and Colgan 1985). Fish that engage in shoaling have developed several methods for facilitating the interaction with others. Individually, shoaling fish have eyes on the sides of their head to better see others in the shoal. Furthermore, it has been demonstrated that vision is important to cohesion because of the inability of most species to maintain schools at night, rather just shoals (Shaw 1961). “Schooling marks” such as prominent stripes or spots help fish recognize others as members of their own species. As a mechanoreceptor, the lateral line system is present in all shoaling fish, assisting in the detection of proximity and movement of other individuals in the shoal. Aside from physiologically based communication adaptations, chemical communication in the aquatic environment plays a large role in fish interaction. There is ample evidence that chemical composition, concentration, flux, and hydrodynamic transport all have profound effects on chemically mediated


Cited: Arnold, Kathryn E. 2000. Kin Recognition in Rainbowfish (Melanotaenia eachamensis): Sex, Sibs and Shoaling. Behavioral Ecology and Sociobiology., 48(5): 385-391. Brahams, M. and Colgan, P. 1985. Risk of predation, hydrodynamic efficiency, and their influence on school structure. Environmental Biology of Fishes., 13(3): 195-202. Chivers, D.P., G.E. Brown, and R.J.F. Smith. 1995. Familiarity and shoal cohesion in fathead minnows (Pimephales promelas)— implications for antipredator behaviour. Canadian Journal of Zoology., 73:955–960. Chivers, Douglas P., Grant E. Brown, and R. Jan F. Smith. 1996. The Evolution of Chemical Alarm Signals: Attracting Predators Benefits Alarm Signal Senders. The American Naturalist., 148(4): 649-659. Dittman, Andrew H. and Thomas Quinn. 1996. Homing in Pacific Salmon: Mechanisms and Ecological Basis. The Journal of Experimental Biology., 199: 83-91. Honda, Haruo. 1980. Female Sex Pheromone of Rainbow Trout, Salmo gairdneri, Involved in Courtship Behaviour. Bulletin of the Japanese Society of Scientific Fisheries., 46(9): 1109-1112. Krause, J., G.D. Ruxton, and D. Rubenstein. 1998. Is there always an influence of shoal size on predator hunting success? Journal of Fish Biology., 52:494–501. Lachlan, R.F., L. Crooks, and K. Laland. 1998. Who follows whom? Shoaling preferences and social learning of foraging information in guppies. Animal Behavior., 56:181–190. Lima, S.L. and L.M. Dill. 1990. Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology., 68: 610-640. Mathis, A., D. P. Chivers, and R. J. F. Smith. 1995. Chemical alarm signals: predator deterrents or predator attractants? American Naturalist., 145: 994–1005. Milinski, H. and R. Heller. 1978. Influence of a predator on the optimal foraging behavior of sticklebacks. Nature., 275: 642-644. Moyle, Peter B Nevitt, Gabrielle A., Andrew H. Dittmano, Thomas P. Quinn, and William J. Moody Jr. 1994. Evidence for a peripheral olfactory memory in imprinted salmon. Proceedings of the National Academy of Sciences., 91: 4288-1292. Olsen, K.H., M. Grahn, and J. Lohm. 2003. The influence of dominance and diet on individual odours in MHC identical juvenile Arctic charr siblings. Journal of Fish Biology., 63: 855–862. Pitcher, T., A. Magurran, and I. Winfield. 1982. Fish in larger shoals find food faster. Behav. Ecology and Sociobiology., 10(2): 149-151. Reiriz, L., A.G. Nicieza, and F. Brana. 1998. Prey selection by experienced and naive juvenile Atlantic salmon. Journal of Fish Biology., 53: 100-114. Roberts, G. 1996. Why individual vigilance increases as group size increases. Animal Behaviour., 51: 1077-1086. Shaw, E Sorensen, P.W., N.E. Stacey. 1998. Evolution and specialization of fish hormonal pheromones, pp. 15-47. In Johnston, R.E., D. Mijller-Schwarze, P.W. Sorensen. Advances in Chemical Signals in Vertebrates. Kluwer Academic/Plenum, New York. 676 p. Stacey, N. E. and P. W. Sorensen. 2005. Reproductive pheromones, pp. 359 - 412. In A.P. Farrell & C. J. Brauner. Fish Physiology, Vol. 24, Behaviour and Physiology of Fish. Academic Press, New York. 480 p. Ward, A.J.W., P.J.B. Hart, and J. Krause. 2004. The effects of habitat- and dietbased cues on association preferences in three-spined sticklebacks. Behavioral Ecology., 15:925–929. Wisenden, Brian D., Keith A. Vollbrecht and Jason L. Brown. 2004. Is there a Fish Alarm Cue? Affirming Evidence from a Wild Study. Animal Behaviour., 67: 59-67. Zimmer, Richard K. and Cheryl Ann Butman. 2000. Chemical Signaling Processes in the Marine Environment. The Biological Bulletin., 198: 168-187.

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