Whether fish sleep or not is an open question, to the point of having inspired the title of several popular science books.Feldman, D. (1989) When do fish sleep? And other imponderables of everyday life. Harper and Row, New York.Weis, J.S. (2011) Do fish sleep? Fascinating answers to questions about fishes. Rutgers University Press, New Brunswick. In birds and mammals, sleep is defined by eye closure and the presence of typical patterns of electrical activity in the brain, including the neocortex, but fish lack eyelids and a neocortex. Some species that always live in shoals or that swim continuously (because of a need for ram ventilation of the gills, for example) are suspected never to sleep. There is also doubt about certain blind species that live in caves.Parzefall, J. (1993): Behavioural ecology of cave-dwelling fish; pp. 573–606 in: Pitcher, T.J.(ed.), The Behaviour of Teleost Fish; London: Chapman&Hall.
Other fish do seem to sleep, however, especially when purely behavioral criteria are used to define sleep. For example, zebrafish,Zhdanova, I.V., Wang, S.Y., Leclair, O.U., and Danilova, N.P. (2001) Melatonin promotes sleep-like state in zebrafish, Brain Research 903: 263–268. Yokogawa T, Marin W, Faraco J, Pézeron G, Appelbaum L, et al. (2007) Characterization of Sleep in Zebrafish and Insomnia in Hypocretin Receptor Mutants, PLOS Biology Vol. 5, No. 10, e277 and criticism and rebuttal, at PLoS Biology tilapia,Shapiro, C.M., and Hepburn, H.R. (1976) Sleep in a schooling fish, Tilapia mossambica, Physiology and Behavior 16:613–615 tench,Peyrethon, J., and Dusan-Peyrethon, D. (1967) Étude polygraphique du cycle veille-sommeil d'un téléostéen (Tinca tinca), Compte-Rendus de la Société de Biologie 161: 2533-2537 brown bullhead,Titkov, E.S. (1976) Characteristics of the daily periodicity of wakefulness and rest in the brown bullhead (Ictalurus nebulosus), Journal of Evolutionary Biochemistry and Physiology 12:305–309. and swell sharkNelson, D.R., and Johnson, R.H. (1970) Diel activity rhythms in the nocturnal, bottom-dwelling sharks Heterodontus francisci and Cephaloscyllium ventriosum, Copeia 1970: 732–739. become motionless and unresponsive at night (or by day, in the case of the swell shark); Spanish hogfish and blue-headed wrasse can even be lifted by hand all the way to the surface without evoking a response. On the other hand, sleep patterns are easily disrupted and may even disappear during periods of migration, spawning, and parental care.Reebs, S.G. (2002) Plasticity of diel and circadian activity rhythms in fish, Reviews in Fish Biology and Fisheries 12: 349–371.
Behavioural sleep Instead of examining brain activity for sleep patterns, an alternate approach is to examine any rest/activity cycles that might indicate "behavioural sleep". The following four behavioural criteria are characteristic of sleep in birds and mammals and could be extended to fishes: (1) prolonged inactivity; (2) typical resting posture, often in a typical shelter; (3) alternation with activity in a 24-h cycle; (4) high arousal thresholds. Based on these criteria, many fish species have been observed sleeping. The typical sleep posture of the brown bullhead is with the fins stretched out, the tail lying flat on the bottom, the body inclined to one side at an angle of 10-30 degrees to the vertical, the cardiac and respiratory frequencies much slower than normal, and much less sensitivity to sound and to being touched.Titkov, E.S. (1976) Characteristics of the daily periodicity of wakefulness and rest in the brown bullhead (Ictalurus nebulosus), Journal of Evolutionary Biochemistry and Physiology 12: 305-309.Karmanova, I.G., Belich, A.I., and Lazarev, S.G. (1981) An electrophysiological study of wakefulness and sleeplike states in fish and amphibians, pp. 181-202 In: Brain Mechanisms of Behaviour in Lower Vertebrates (P.R. Laming, ed.). Cambridge University Press, Cambridge. Mozambique tilapia are motionless at the bottom at night, with a lower respiratory rate and no eye movement, and they do not respond as readily as during the day to electrical currents or food delivery.Shapiro, C.M., and Hepburn, H.R. (1976) Sleep in a schooling fish, Tilapia mossambica. Physiology and Behavior 16: 613-615. At night the loach Misgurnus dabryanus "floats on the water" with a reduced respiratory frequency and less sensitivity to external stimuli. Port Jackson sharks and draughtsboard sharks can lay flat on the bottom with their eyes closed for extended periods, with less sensitivity to electrical stimulation and their metabolic rate is reduced during such periods. At night, Spanish hogfish, bluehead wrasse, the wrasse Halichoeres bivittatus, the cunner Tautogolabrus adspersus, and even requiem sharks, can be picked up by hand without eliciting a response.Tauber, E.S., Weitzman, E.D., and Korey, S.R. (1969) Eye movements during behavioral inactivity in certain Bermuda reef fish. Communications in Behavioral Biology A 3: 131-135.Tauber, E.S. (1974) The phylogeny of sleep, pages 133-172 In: Advances in sleep research, vol. 1 (E.D. Weitzman, ed.). Spectrum Publications, New York.Clark, E. (1973) "Sleeping" sharks in Mexico. Underwater Naturalist 8: 4-7.Dew, C.B. (1976) A contribution to the life history of the cunner, Tautogolabrus adspersus, in Fishers Island Sound, Connecticut. Chesapeake Science 17: 101-103. A 1961 observational study of approximately 200 species in European public aquaria reported many cases of apparent sleep.Weber, E. (1961) Über Ruhelagen von Fischen, Zeitschrift für Tierpsychologie 18: 517–533.
Divers can easily see fishes settling down for the night in typical shelters, such as holes and crevices, underneath ledges, amidst vegetation, inside sponges, or buried in sand.Reebs, S.G. (1992) Sleep, inactivity, and circadian rhythms in fish. Pp. 127-135 In Rhythms in Fishes (M.A. Ali, editor). Plenum, New York. Some extra protection can be derived from special secretions, such as the mucous envelope produced by several species of wrasse and parrotfish, either around the fish themselves or at the opening of their shelter. These envelopes screen the sleeping fish from predatorsWinn, H.E., and Bardach, J.E. (1959) Differential food selection by moray eels and a possible role of the mucous envelope of parrot fishes in reduction of predation. Ecology 40: 296-298. and ectoparasites.Grutter, A.S., Rumney, J.G., Sinclair-Taylor, T., Waldie, P., and Franklin, C.E. (2011) Fish mucous cocoons: the "mosquito nets" of the sea. Biology Letters 7: 292-294.
In the laboratory, periods of inactivity often alternate with periods of activity on a 24-h basis, or a near 24-h basis when the lighting conditions are constant. Circadian rhythms of activity have been documented in over 40 different fish species, including hagfish, lamprey, sharks, cyprinids, ictalurids, gymnotids, salmonids, and labrids.Reebs S.G. (2011) Circadian Rhythms in Fish. In: Farrell A.P., (ed.), Encyclopedia of Fish Physiology: From Genome to Environment, volume 1, pp. 736–743. San Diego: Academic Press.Zhdanova, I. and S.G. Reebs. (2006) Circadian rhythms in fish. Pp. 197-238 In Fish Physiology, Vol 24: Behaviour and Physiology of Fishes (K.A. Sloman, R.W. Wilson, and S. Balshine, eds.). Elsevier, New York.
Physiological sleep One physiological characteristic of sleep goes by the name of "homeostatic regulation". This is the notion that animals need a more or less constant amount of sleep every day, so that if a subject is deprived of sleep one day, the amount of sleep tends to "rebound" (increase) the next few days. This has been observed in zebrafish. At night, zebrafish appear to float in the water column, either horizontally or with the head slightly up. The frequency of mouth and gill movement is reduced by almost half and they are twice as hard to arouse as during the day. If they are deprived of this sleep-like behaviour, the sleep bouts thereafter are longer and the arousal threshold is higher than usual, suggesting a rebound effect.Zhdanova, I.V., Wang, S.Y., Leclair, O.U., and Danilova, N.P. (2001) Melatonin promotes sleep-like state in zebrafish. Brain Research 903: 263-268.Yokogawa, T., Marin, W., Faraco, J., Pézeron, G., Appelbaum, L., Zhang, J., Rosa, F., Mourrain, P., and Mignot, E. (2007) Characterization of sleep in zebrafish and insomnia in hypocretin receptor mutants. PLOS Biology 5: 2379-2397. Similarly, in the convict cichlid and brown trout activity decreases on days that follow an experimental disruption of the fish's normal rest behaviour at night.Tobler, I., and Borbély, A.A. (1985) Effet of rest deprivation on motor activity of fish. Journal of Comparative Physiology A 157: 817-822.
Absence of sleep
Many pelagic fish species, such as bluefish, Atlantic mackerel, tuna, bonito, and some sharks, swim continuously and do not show signs, behavioural or otherwise, of sleep.Olla, B.L., and Studholme, A.L. (1978) Comparative aspects of the activity rhythms of tautog, Tautoga onitis, Bluefish, Pomatomus saltatrix, and Atlantic mackerel, Scomber scombrus, as related to their life habits. Pp. 131-151 In: Rhythmic Activity of Fishes (J.E. Thorpe, ed.). Academic Press, London. It has been argued that one function of sleep is to allow the brain to consolidate into memory the things it has learned during the animal's normal period of activity. The brain might not be able to do this while still assailed by new stimuli and new information to process. Therefore, the role of sleep would be to periodically shut down sensory input to allow the brain to form memories. Pelagic species swim in an open-water environment wherein novel stimuli is uncommon. In such species, the sensory input is so low that memory formation could take place even if the fish keeps on moving (a repetitive activity) and does not fall asleep in the traditional sense of the word.Kavanau, J.L. (2010) Schooling by continuously active fishes: clues to sleep's ultimate function, Pp. 57-85 In: Evolution of sleep: phylogenetic and functional perspectives (McNamara, P., Barton, R.A., and Nunn, C.L., eds.). Cambridge University Press, Cambridge.
Some populations of Mexican tetra live in permanently dark caves. They are derived from still extant surface populations. The surface fish exhibit 3–6 hours of continuous inactivity each day, interpreted as sleep, but the cave-dwellers display continuous activity, suggesting that sleep has been lost during evolution in these populations that live in an environment devoid of daily time cues.
Diurnal damselfish normally sleep motionless in crevices within coral reefs at night, but three species (the green chromis, the marginate dascyllus and the whitetail dascyllus) spend the night between coral branches where they beat their fins at a rate about twice that of normal daytime swimming. This creates water currents that keep the inner zone of the coral (and thus the fish themselves) well oxygenated, at levels about four times higher than in the absence of the fish. Though the fish are active (mostly in a repetitive way), they do not respond to light or to the presence of potential predators. The researchers who documented this behaviour called it "sleep-swimming".Goldshmid, R., Holzman, R., Weihs, D., and Genin, A. (2004) Aeration of corals by sleep-swimming fish. Limnology and Oceanography 49: 1832-1839.