Learning About Danger: Chemical Alarm Cues and the Assessment of Predation Risk by Fishes

Fish Cognition and Behavior

Volumn , Issue , 2007, Pages 49-69

  Brown, Grant E ^a   Chivers, Douglas P ^b  

^a Gina Cody School of Engineering and Computer Science   (Canada)

^b UNIVERSITY OF SASKATCHEWAN   (Canada)

Author keywords

Acquired predator recognition; Chemical alarm cues; Fish cognition; Predator diet cues; Temporal variability

Indexed keywords

EID: 84948170014     PISSN: None     EISSN: None     Source Type: Book    
DOI: 10.1002/9780470996058.ch4     Document Type: Chapter

References (118)

1. Alvarez, D. & Nicieza, A.G. (2003) Predator avoidance behaviour in wild and hatcheryreared brown trout: the role of experience and domestication. Journal of Fish Biology, 63, 1565-1577.
2. Berejikian, B.A., Smith, R.J.F., Tezak, E.P., Schroder, S.L. & Knudsen, C.M. (1999) Chemical alarm signals and complex hatchery rearing habitats affect antipredator behaviour and survival of Chinook salmon (Oncorhynchus tshawytscha) juveniles. Canadian Journal of Fisheries and Aquatic Sciences, 56, 830-838.
3. Berejikian, B.A., Tezak, E.P. & LaRae, A.L. (2003) Innate and enhanced predator recognition in hatchery-reared Chinook salmon. Environmental Biology of Fishes, 67, 241-251.
4. Brönmark, C. & Miner, J.G. (1992) Predator-induced phenotypic change in body morphology in crucian carp. Science, 258, 1348-1350.
5. Brown, C. & Day, R. (2002) The future of stock enhancements: bridging the gap between hatchery practice and conservation biology. Fish and Fisheries, 3, 79-94.
6. Brown, C. & Laland, K.N. (2003) Social learning in fishes: a review. Fish and Fisheries, 4, 280-288.
7. Brown, C. & Warburton, K. (1997) Predator recognition and anti-predator responses in the rainbowfish, Melanotaenia eachamensis. Behavioral Ecology and Sociobiology, 41, 61-68.
8. Brown, C., Davidson, T. & Laland, K. (2003) Environmental enrichment and prior experience of live prey improve foraging behaviour in hatchery-reared Atlantic salmon. Journal of Fish Biology, 63(A), 187-196.
9. Brown, G.E. & Chivers, D.P. (2005) Learning as an adaptive response to predation. In: P. Barbosa & I. Castellanos (eds), Ecology of Predator-Prey Interactions, pp. 34-54. Oxford University Press, Oxford.
10. Brown, G.E. & Dreier, V.M. (2002) Predator inspection behaviour and attack cone avoidance in a characin fish: the effects of predator diet and prey experience. Animal Behaviour, 63, 1175-1181.
11. Brown, G.E. & Godin, J.-G.J. (1999) Who dares, learns: chemical inspection behaviour and acquired predator recognition in a characin fish. Animal Behaviour, 57, 475-481.
12. Brown, G.E. & Magnavacca, G. (2003) Predator inspection behaviour in a characin fish: an interaction between chemical and visual information? Ethology, 109, 739-750.
13. Brown, G.E. & Schwarzbauer, E.M. (2001) Chemical predator inspection and attack cone avoidance in a characin fish: the effects of predator diet. Behaviour, 138, 727-739.
14. Brown, G.E. & Smith, R.J.F. (1996) Foraging trade-offs in fathead minnows (Pimephales promelas): acquired predator recognition in the absence of an alarm response. Ethology, 102, 776-785.
15. Brown, G.E. & Smith, R.J.F. (1998) Acquired predator recognition in juvenile rainbow trout (Oncorhynchus mykiss): conditioning hatchery-reared fish to recognize chemical cues of a predator. Canadian Journal of Fisheries and Aquatic Sciences, 55, 611-617.
16. Brown, G.E., Chivers, D.P. & Smith, R.J.F. (1995a) Localized defecation by pike: a response to labeling by cyprinid alarm pheromone? Behavioral Ecology and Sociobiology, 36, 105-110.
17. Brown, G.E., Chivers, D.P. & Smith, R.J.F. (1995b) Fathead minnows avoid conspecific and heterospecific alarm pheromone in the faeces of northern pike. Journal of Fish Biology, 47, 387-393.
18. Brown, G.E., Chivers, D.P. & Smith, R.J.F. (1997) Differential learning rates of chemical versus visual cues of a northern pike by fathead minnows in a natural habitat. Environmental Biology of Fishes, 49, 89-96.
19. Brown, G.E., Paige, J.A. & Godin, J.-G.J. (2000a) Chemically-mediated predator inspection behaviour in the absence of predator visual cues by a characin fish, Hemigrammus erythrozonus. Animal Behaviour, 60, 315-321.
20. Brown, G.E., Adrian, J.C., Jr., Smyth, E., Leet, H. & Brennan, S. (2000b) Ostariophysan alarm pheromones: laboratory and field tests of the functional significance of nitrogen oxides. Journal of Chemical Ecology, 26, 139-154.
21. Brown, G.E., LeBlanc, V.J. & Porter, L.E. (2001a) Ontogenetic changes in the response of largemouth bass (Micropterus salmoides, Centrarchidae, Perciformes) to heterospecific alarm pheromones. Ethology, 107, 401-414.
22. Brown, G.E., Adrian, J.C., Jr. & Shih, M.L. (2001b) Behavioural responses of fathead minnows to hypoxanthine-3-N-oxide at varying concentrations. Journal of Fish Biology, 58, 1465-1470.
23. Brown, G.E., Adrian, J.C., Jr., Patton, T. & Chivers, D.P. (2001c) Fathead minnows learn to recognize predator odour when exposed to concentrations of artificial alarm pheromone below their behavioural response thresholds. Canadian Journal of Zoology, 79, 2239-2245.
24. Brown, G.E., Golub, J.L. & Plata, D.L. (2001d) Attack cone avoidance during predator inspection visits by wild finescale dace (Phoxinus neogaeus): the effects of predator diet. Journal of Chemical Ecology, 27, 1657-1666.
25. Brown, G.E., Adrian, J.C., Jr., Kaufman, I.H., Erickson, J.L. & Gershaneck, D. (2001e) Responses to nitrogen-oxides by Characiforme fishes suggests evolutionary conservation in ostariophysan alarm pheromones. In: Marchlewska-Koj, J.J. Lepri & D. Müller-Schwarze (eds), Chemical Signals in Vertebrates, Vol. 9, pp. 305-312. Plenum Press, New York.
26. Brown, G.E., Adrian, J.C. Jr., Lewis, M.G. & Tower, J.M. (2002a) The effects of reduced pH on chemical alarm signaling in ostariophysan fishes. Canadian Journal of Fisheries and Aquatic Sciences, 59, 1331-1338.
27. Brown, G.E., Gershaneck, D.L., Plata, D.L. & Golub, J.L. (2002b) Ontogenetic changes in response to heterospecific alarm cues by juvenile largemouth bass are phenotypically plastic. Behaviour, 139, 913-927.
28. Brown, G.E., Adrian, J.C., Jr., Naderi, N.T., Harvey, M.C. & Kelly, J.M. (2003) Nitrogen oxides elicit antipredator responses in juvenile channel catfish, but not in convict cichlids or rainbow trout: Conservation of the Ostariophysan alarm pheromone. Journal of Chemical Ecology, 29, 1781-1796.
29. Brown, G.E., Poirier, J.-F. & Adrian, J.C., Jr. (2004) Assessment of local predation risk: the role of subthreshold concentrations of chemical alarm cues. Behavioral Ecology, 15, 810-815.
30. Brown, G.E., Bongiorno, T., DiCapua, D.M., Ivan, L.I. & Roh, E. (2006) Effects of group size on the threat-sensitive response to varying concentrations of chemical alarm cues by juvenile convict cichlids. Canadian Journal of Zoology, 84, 1-8.
31. Chivers, D.P. & Mirza, R.S. (2001) Predator diet cues and the assessment of predation risk by aquatic vertebrates: a review and prospectus. In: A. Marchlewska-Koj, J.J. Lepri & D. Müller-Schwarze (eds), Chemical Signals in Vertebrates, Vol. 9, pp. 227-284. Kluwer Academic, New York.
32. Chivers, D.P. & Smith, R.J.F. (1994a) Fathead minnows, Pimephales promelas, acquire predator recognition when alarm substance is associated with the sight of unfamiliar fish. Animal Behaviour, 48, 597-605.
33. Chivers, D.P. & Smith, R.J.F. (1994b) The role of experience and chemical alarm signaling in predator recognition by fathead minnows, Pimephales promelas. Journal of Fish Biology, 44, 273-285.
34. Chivers, D.P. & Smith, R.J.F. (1995) Free-living fathead minnows rapidly learn to recognize pike as predators. Journal of Fish Biology, 46, 949-954.
35. Chivers, D.P. & Smith, R.J.F. (1998) Chemical alarm signalling in aquatic predator-prey systems: a review and prospectus. Écoscience, 5, 338-352.
36. Chivers, D.P., Brown, G.E. & Smith, R.J.F. (1995) Acquired recognition of chemical stimuli from pike, Esox lucius, by brook sticklebacks, Culaea inconstans (Osteichthyes, Gasterosteidae). Ethology, 99, 234-242.
37. Chivers, D.P., Brown, G.E. & Smith, R.J.F. (1996) The evolution of chemical alarm signals: attracting predators benefits alarm signal senders. American Naturalist, 148, 649-659.
38. Chivers, D.P., Mirza, R.S. & Johnston, J.G. (2002) Learned recognition of heterospecific alarm cues enhances survival during encounters with predators. Behaviour, 139, 929-938.
39. Darwish, T.L., Mirza, R.S., Leduc, A.O.H.C. & Brown, G.E. (2005) Acquired recognition of novel predator odour cocktails by juvenile glowlight tetras. Animal Behaviour, 70, 83-89.
40. Dugatkin, L.A. & Godin, J.-G.J. (1992a) Prey approaching predators: a cost-benefit perspective. Annals Zoologici Fennici, 29, 233-252.
41. Dugatkin, L.A. & Godin, J.-G.J. (1992b) Predator inspection, shoaling and foraging under predation hazard in the Trinidadian guppy, Poecilia reticulata. Environmental Biology of Fishes, 34, 265-276.
42. Dupuch, A., Magnan, P. & Dill, L.M. (2004) Sensitivity of northern redbelly dace, Phoxinus eos, to chemical alarm cues. Canadian Journal of Zoology, 82, 407-415.
43. Ferrari, M.C.O. & Chivers, D.P. (2006a) The role of learning in the development of threatsensitive predator avoidance: how do fathead minnows incorporate conflicting information? Animal Behaviour, 71, 19-26.
44. Ferrari, M.C.O. & Chivers, D.P. (2006b) The role of latent inhibition in acquired predator recognition by fathead minnows. Canadian Journal of Zoology, 84, 505-509.
45. Ferrari, M.C.O., Trowell, J.J., Brown, G.E. & Chivers, D.P. (2005) The role of learning in the development of threat-sensitive predator avoidance by fathead minnows. Animal Behaviour, 70, 777-784.
46. Foam, P.E., Harvey, M.C., Mirza, R.S. & Brown, G.E. (2005a) Heads up: juvenile convict cichlids rely on chemosensory information to make threat-sensitive foraging decisions. Animal Behaviour, 70, 601-607.
47. Foam, P.E., Mirza, R.S., Chivers, D.P. & Brown, G.E. (2005b) Juvenile convict cichlids (Archocentrus nigrofasciatus) allocate foraging in response to temporal variation in predation risk. Behaviour, 142, 129-144.
48. Gazdewich, K.J. & Chivers, D.P. (2002) Acquired predator recognition by fathead minnows: influence of habitat characteristics on survival. Journal of Chemical Ecology, 28, 439-445.
49. Gilliam, J.F. & Fraser, D.F. (2001) Movement in corridors: enhancement by predation threat, disturbance, and habitat structure. Ecology, 82, 124-135.
50. Godin, J.-G.J. & Davis, S.A. (1995) Who dares, benefits: predator approach behaviour in the guppy (Poecilia reticulata) deters predator pursuit. Proceedings of the Royal Society of London, Series B, 259, 193-200.
51. Godin, J.-G.J. & Dugatkin, L.A. (1996) Female mating preferences for bold males in the guppy, Poecilia reticulata. Proceedings of the National Academy of Sciences, 93, 10262-10267.
52. Godin, J.-G.J. & Smith, S.A. (1988) A fitness cost of foraging in the guppy. Nature, 333, 69-71.
53. Golub, J.L., Vermette, V. & Brown, G.E. (2005) The response of pumpkinseed sunfish to conspecific and heterospecific chemical alarm cues under natural conditions: the effects of stimulus type, habitat and ontogeny. Journal of Fish Biology, 66, 1073-1081.
54. Gotceitas, V. & Brown, J.A. (1993) Substrate selection by juvenile Atlantic cod (Gadus morhua): effects of predation risk. Oecologia, 93, 31-37.
55. Griffin, A.S. (2004) Social learning about predators: A review and prospectus. Learning and Behavior, 32, 131-140.
56. Hartman, E.J. & Abrahams, M.V. (2000) Sensory compensation and the detection of predators: the interaction between chemical and visual information. Proceedings of the Royal Society of London, Series B - Biological Sciences, 267, 571-575.
57. Hawkins, L.A., Magurran, A.E. & Armstrong, J.D. (2004) Innate predator recognition in newly-hatched Atlantic salmon. Behaviour, 141, 1249-1262.
58. Jachner, A. & Rydz, M.A. (2002) Behavioural response of roach (Cyprinidae) to different doses of chemical alarm cues (Schreckstoff). Archieves Hydrobiologica, 155, 369-381.
59. Kats, L.B. & Dill, L.M. (1998) The scent of death: chemosensory assessment of predation risk by prey animals. Écoscience, 5, 361-394.
60. Kelley, J.L. & Magurran, A.E. (2003) Learned predator recognition and antipredator responses in fishes. Fish and Fisheries, 4, 216-226.
61. Korpi, N.L. & Wisenden, B.D. (2001) Learned recognition of novel predator odour by zebra danios, Danio rerio, following time-shifted presentation of alarm cues and predator odour. Environmental Biology of Fishes, 61, 205-211.
62. Kristensen, E.A. & Closs, G.P. (2004) Anti-predator response of naive and experienced common bully to chemical alarm cues. Journal of Fish Biology, 64, 643-652.
63. Kusch, R.C., Mirza, R.S. & Chivers, D.P. (2004) Making sense of predator scents: investigating the sophistication of predator assessment abilities of fathead minnows. Behavioural Ecology and Sociobiology, 55, 551-555.
64. Lachlan, R.F., Crooks, L. & Laland, K.N. (1998) Who follows whom? Shoaling preferences and social learning of foraging information in guppies. Animal Behaviour, 56, 181-190.
65. Laland, K.N. & Williams, K. (1997) Shoaling generates social learning of foraging information in guppies. Animal Behaviour, 53, 1161-1169.
66. Lawrence, B.J. & Smith, R.J.F. (1989) Behavioral response of solitary fathead minnows, Pimephales promelas, to alarm substance. Journal of Chemical Ecology, 15, 209-219.
67. Leduc, A.O.H.C., Noseworthy, M.K., Adrian, J.C., Jr & Brown, G.E. (2003) Detection of conspecific and heterospecific alarm signals by juvenile pumpkinseeds under weakly acidic conditions. Journal of Fish Biology, 63, 1331-1336.
68. Leduc, A.O.H.C., Kelly, J.M. & Brown, G.E. (2004a) Detection of conspecific chemical alarm cues by juvenile salmonids under neutral and weakly acidic conditions: laboratory and field tests. Oecologia, 139, 318-324.
69. Leduc, A.O.H.C., Ferrari, M.C.O., Kelly, J.M. & Brown, G.E. (2004b) Learning to recognize novel predators under weakly acidic conditions: the effects of reduced pH on acquired predator recognition by juvenile rainbow trout (Oncorhynchus mykiss). Chemoecology, 14, 107-112.
70. Licht, T. (1989) Discriminating between hungry and satiated predators: the response of guppies (Poecilia reticulata) from high and low predation sites. Ethology, 82, 238-243.
71. Lima, S.L. & Bednekoff, P.A. (1999) Temporal variation in danger drives antipredator behavior: the predation risk allocation hypothesis. American Naturalist, 153, 649-659.
72. Lima, S.L. & Dill, L.M. (1990) Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology, 68, 619-640.
73. Magurran, A.E. (1989) Acquired recognition of predator odour in the European minnow (Phoxinus phoxinus). Ethology, 82, 216-233.
74. Magurran, A.E. & Seghers, B.H. (1990) Population differences in predator recognition and attack cone avoidance in the guppy Poecilia reticulata. Animal Behaviour, 40, 443-452.
75. Marcus, J.P. & Brown, G.E. (2003) Response of pumpkinseed sunfish to conspecific chemical alarm cues: an interaction between ontogeny and stimulus concentration. Canadian Journal of Zoology, 81, 1671-1677.
76. Mathis, A. & Smith, R.J.F. (1993a) Chemical alarm signals increase the survival time of fathead minnows (Pimephales promelas) during encounters with northern pike (Esox lucius). Behavioral Ecology, 4, 260-265.
77. Mathis, A. & Smith, R.J.F. (1993b) Chemical labelling of northern pike, Esox lucius, by the alarm pheromone of fathead minnows, Pimephales promelas. Journal of Chemical Ecology, 19, 1967-1979.
78. Mathis, A. & Smith, R.J.F. (1993c) Fathead minnows (Pimephales promelas) learn to recognize pike (Esox lucius) as predators on the basis of chemical stimuli in the pike's diet. Animal Behaviour, 46, 645-656.
79. Mathis, A., Chivers, D.P. & Smith, R.J.F. (1996) Cultural transmission of predator recognition in fishes: intraspecific and interspecific learning. Animal Behaviour, 51, 185-201.
80. McPherson, T.D., Mirza, R.S. & Pyle, G.G. (2004) Responses of wild fishes to alarm chemicals in pristine and metal-contaminated lakes. Canadian Journal of Zoology, 82, 694-700.
81. Mirza, R.S. & Chivers, D.P. (2000) Predator-recognition training enhances survival of brook trout: evidence from laboratory and field-enclosure studies. Canadian Journal of Zoology, 78, 2198-2208.
82. Mirza, R.S. & Chivers, D.P. (2001a) Chemical alarm signals enhance survival of brook charr, Salvelinus fontinalis, during encounters with predatory chain pickerel, Esox niger. Ethology, 107, 989-1006.
83. Mirza, R.S. & Chivers, D.P. (2001b) Do juvenile yellow perch use diet cues to assess the level of threat posed by intraspecific predators? Behaviour, 138, 1249-1258.
84. Mirza, R.S. & Chivers, D.P. (2001c) Do chemical alarm signals enhance survival of aquatic vertebrates?: an analysis of the current research paradigm. In: A. Marchlewska-Koj, J.J. Lepri, & D. Müller-Schwarze (eds), Chemical Signals in Vertebrates, Vol. 9, pp. 19-26. Kluwer Academic, New York.
85. Mirza, R.S. & Chivers, D.P. (2001d) Are chemical alarm signals conserved within salmonid fishes? Journal of Chemical Ecology, 27, 1641-1655.
86. Mirza, R.S. & Chivers, D.P. (2001e) Learned recognition of heterospecific alarm signals: the importance of mixed predator diet. Ethology, 107, 1007-1018.
87. Mirza, R.S. & Chivers, D.P. (2003a) Response of juvenile rainbow trout to varying concentrations of chemical alarm cue: response thresholds and survival during encounters with predators. Canadian Journal of Zoology, 81, 88-95.
88. Mirza, R.S. & Chivers, D.P. (2003b) Influence of body size on the responses of fathead minnows, Pimephales promelas, to damselfly alarm cues. Ethology, 109, 691-699.
89. Mirza, R.S. & Chivers, D.P. (2003c) Fathead minnows learn to recognize heterospecific alarm cues they detect in the diet of a known predator. Behaviour, 140, 1359-1369.
90. Mirza, R.S., Scott, J.J. & Chivers, D.P. (2001) Differential responses of male and female red swordtails to chemical alarm cues. Journal of Fish Biology, 59, 716-728.
91. Mirza, R.S., Mathis, A. & Chivers, D.P. (2006) Does temporal variation in predation risk influence the intensity of anti-predator responses? A test of the risk allocation hypothesis. Ethology, 112, 44-51.
92. Murphy, K.E. & Pitcher, T.J. (1997) Predator attack motivation influences the inspection behaviour of European minnows. Journal of Fish Biology, 50, 407-417.
93. Olson, M.H. (1996) Ontogenetic niche shifts in largemouth bass: variability and consequences for first-year growth. Ecology, 77, 179-190.
94. Olson, M.H., Mittelback, G.G. & Osenberg, C.W. (1995) Competition between predator and prey: resource-based mechanisms and implication for stage-structured dynamics. Ecology, 76, 1758-1771.
95. Pollock, M.S. & Chivers, D.P. (2003) Does habitat complexity influence the ability of fathead minnows to learn heterospecific chemical alarm cues? Canadian Journal of Zoology, 81, 923-927.
96. Pollock, M.S. & Chivers, D.P. (2004) The effects of density on the learned recognition of heterospecific alarm cues. Ethology, 110, 341-349.
97. Pollock, M.S., Chivers, D.P., Mirza, R.S. & Wisenden, B.D. (2003) Fathead minnows, Pimephales promelas, learn to recognize chemical alarm cues of introduced brook stickleback, Culaea inconstans. Environmental Biology of Fishes, 66, 313-319.
98. Roh, E., Mirza, R.S. & Brown, G.E. (2004) Quality or quantity?: the role of donor condition in the production of chemical alarm cues in juvenile convict cichlids. Behaviour, 141, 1235-1248.
99. Schotz, N.L., Truelove, N.K., French, B.L., Berejikian, B.A., Quinn, T.P., Casillas, E. & Collier, T.K. (2000) Diazinon disrupts antipredator and homing behaviors in Chinook salmon (Oncorhynchus tshawytscha). Canadian Journal of Fisheries and Aquatic Sciences, 57, 1911-1918.
100. Schutz, F. (1956) A comparative study about the fright reaction of fish and their distribution. Zeitschrift für Vergleichende Physiologie, 38, 84-135. (In German.)
101. Schwartz, B. (1984) Psychology of Learning and Behavior, 2nd edn. W.W. Norton, New York.
102. Scott, G.R., Sloman, K.A., Rouleau, C. & Wood, C.M. (2003) Cadmium disrupts behavioural and physiological responses to alarm substance in juvenile rainbow trout (Oncorhynchus mykiss). Journal of Experimental Biology, 206, 1779-1790.
103. Shively, R.S., Poe, T.P. & Sauter, S.T. (1996) Feeding response by northern squawfish to a hatchery release of juvenile salmonids in the Clearwater River, Idaho. Transactions of the American Fisheries Society, 125, 230-236.
104. Sih, A. (1992) Prey uncertainty and the balancing of antipredator and foraging needs. American Naturalist, 139, 1052-1069.
105. Sih, A., Ziemba, R. & Harding, K.C. (2000) New insights on how temporal variation in predation risk shapes prey behavior. Trends in Ecology and Evolution, 15, 3-4.
106. Smith, M.E. & Belk, M.C. (2001) Risk assessment in western mosquitofish (Gambusia affinis): do multiple cues have additive effects? Behavioral Ecology and Sociobiology, 51, 101-107.
107. Smith, R.J.F. (1999) What good is smelly stuff in the skin? Cross-function and cross-taxa effects in fish 'alarm substances'. In: R.E. Johnston, D. Müller-Schwarze & P.W. Sorenesen (eds), Advances in Chemical Signals in Vertebrates, pp 475-488. Kluwer Academic, New York.
108. Smith, R.J.F. & Smith, M.J. (1989) Predator-recognition behaviour in two species of gobies, Asterropteryx semipunctatus and Gnatholepis anjerensis. Ethology, 81, 279-290.
109. Suboski, M.D. (1990) Releaser-induced recognition learning. Psychological Reviews, 97, 271-284.
110. Thorpe, W.H. (1963) Learning and Instinct in Animals, 2nd edn. Methuen, London.
111. Vilhunen, S. (2006) Repeated antipredator conditioning: a pathway to habituation or to better avoidance? Journal of Fish Biology, 68, 25-43.
112. Vilhunen, S. & Hirvonen, H. (2003) Innate antipredator response of Arctic charr (Salvelinus alpinus) depend on predator species and their diet. Behavioral Ecology and Sociobiology, 55, 1-10.
113. Vilhunen, S., Hirvonen, H. & Laakkonen, M.V.-M. (2005) Less is more: social learning of predator recognition requires a low demonstrator to observer ratio in Arctic charr (Salvelinus alpinus). Behavioral Ecology and Sociobiology, 57, 275-282.
114. Werner, E.E. & Gilliam, J.F. (1984) The ontogenetic niche and species interactions in sizestructured populations. Annual Reviews in Ecology and Systematics, 15, 393-425.
115. Wisenden, B.D. (2000) Olfactory assessment of predation risk. Philosophical Transactions of the Royal Society, 355, 1205-1208.
116. Wisenden, B.D. & Harter, K.R. (2001) Motion, not shape, facilitates association of predation risk with novel objects by fathead minnows (Pimephales promelas). Ethology, 107, 357-364.
117. Yunker, W.K., Wein, D.E. & Wisenden, B.D. (1999) Conditioned alarm behavior in fathead minnows (Pimephales promelas) resulting from association of chemical alarm pheromone with a non-biological visual stimulus. Journal of Chemical Ecology, 25, 2677-2686.
118. Zhao, X. & Chivers, D.P. (2005) Response of juvenile goldfish (Carassius auratus) to chemical cues: relationship between response intensity, response duration and the level of predation risk. In: R.T. Mason, M. LeMaster & D. Müller-Schwarze (eds), Chemical Signals in Vertebrates, Vol. 10, pp. 334-341. Springer, New York.