Sleep (non-human)
Encyclopedia
Sleep in non-human animals refers to how the behavioral and physiological state of sleep
Sleep
Sleep is a naturally recurring state characterized by reduced or absent consciousness, relatively suspended sensory activity, and inactivity of nearly all voluntary muscles. It is distinguished from quiet wakefulness by a decreased ability to react to stimuli, and is more easily reversible than...

, mainly characterized by reversible unconsciousness, non-responsiveness to external stimuli, and motor passivity, appears in different categories of animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s.
Rats kept from sleeping die within a couple of weeks, but the exact function of sleep is still unknown.

Definition

Sleep can follow a physiological or behavior
Behavior
Behavior or behaviour refers to the actions and mannerisms made by organisms, systems, or artificial entities in conjunction with its environment, which includes the other systems or organisms around as well as the physical environment...

al definition. In the physiological sense, sleep is a state characterized by reversible unconsciousness, special brainwave patterns, sporadic eye movement, loss of muscle tone
Muscle tone
In physiology, medicine, and anatomy, muscle tone is the continuous and passive partial contraction of the muscles, or the muscle’s resistance to passive stretch during resting state. It helps maintain posture, and it declines during REM sleep.-Purpose:Unconscious nerve impulses maintain the...

 (possibly with some exceptions; see below regarding the sleep of birds and of aquatic mammals), and a compensatory increase following deprivation of the state. In the behavioral sense, sleep is characterized by non-responsiveness to external stimuli, the adoption of a typical posture, and the occupation of a sheltered site, all of which is usually repeated on a 24-hour basis. The physiological definition applies well to birds and mammals, but in other animals (whose brain is not as complex), the behavioral definition is more often used. In very simple animals, behavioral definitions of sleep are the only ones possible, and even then the behavioral repertoire of the animal may not be extensive enough to allow distinction between sleep and wakefulness.

Sleep in invertebrates

Sleep as a phenomenon appears to have very old evolutionary roots. The nematode
Nematode
The nematodes or roundworms are the most diverse phylum of pseudocoelomates, and one of the most diverse of all animals. Nematode species are very difficult to distinguish; over 28,000 have been described, of which over 16,000 are parasitic. It has been estimated that the total number of nematode...

 C. elegans
Caenorhabditis elegans
Caenorhabditis elegans is a free-living, transparent nematode , about 1 mm in length, which lives in temperate soil environments. Research into the molecular and developmental biology of C. elegans was begun in 1974 by Sydney Brenner and it has since been used extensively as a model...

 is the most primitive organism in which sleep-like states have been observed. Here, a lethargus phase occurs in short periods preceding each moult
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...

, a fact which may indicate that sleep primitively is connected to developmental processes. Raizen et al.s results furthermore suggest that sleep is necessary for changes in the neural system.

The electrophysiological study of sleep in small invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...

s is complicated. However, even such simple animals as fruit flies
Drosophila melanogaster
Drosophila melanogaster is a species of Diptera, or the order of flies, in the family Drosophilidae. The species is known generally as the common fruit fly or vinegar fly. Starting from Charles W...

 appear to sleep, and systematic disturbance of that state leads to cognitive disabilities
Cognition
In science, cognition refers to mental processes. These processes include attention, remembering, producing and understanding language, solving problems, and making decisions. Cognition is studied in various disciplines such as psychology, philosophy, linguistics, and computer science...

. There are several methods of measuring cognitive functions in fruit flies. A common method is to let the flies choose whether they want to fly through a tunnel that leads to a light source, or through a dark tunnel. Normally, flies are attracted to light. But if sugar
Sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

 is placed in the end of the dark tunnel, and something the flies dislike is placed in the end of the light tunnel, the flies will eventually learn to fly towards darkness rather than light. Flies deprived of sleep require a longer time to learn this and also forget it more quickly. If an arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

 is experimentally kept awake longer than it is used to, then its coming rest period will be prolonged. In cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...

es that rest period is characterized by the antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....

 being folded down and by a decreased sensitivity to external stimuli. Sleep has been described in crayfish
Crayfish
Crayfish, crawfish, or crawdads – members of the superfamilies Astacoidea and Parastacoidea – are freshwater crustaceans resembling small lobsters, to which they are related...

, too, characterized by passivity and increased thresholds for sensory stimuli as well as changes in the EEG
EEG
EEG commonly refers to electroencephalography, a measurement of the electrical activity of the brain.EEG may also refer to:* Emperor Entertainment Group, a Hong Kong-based entertainment company...

 pattern, markedly differing from the patterns found in crayfish when they are awake.

Sleep in fishes and reptiles

Sleep in fish
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...

es is not extensively studied. 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 cave
Cave
A cave or cavern is a natural underground space large enough for a human to enter. The term applies to natural cavities some part of which is in total darkness. The word cave also includes smaller spaces like rock shelters, sea caves, and grottos.Speleology is the science of exploration and study...

s. Other fishes seem to sleep, however. For example, zebrafish, tilapia
Tilapia
Tilapia , is the common name for nearly a hundred species of cichlid fish from the tilapiine cichlid tribe. Tilapia inhabit a variety of fresh water habitats, including shallow streams, ponds, rivers and lakes. Historically, they have been of major importance in artisan fishing in Africa and the...

, tench
Tench
The tench or doctor fish is a freshwater and brackish water fish of the cyprinid family found throughout Eurasia from Western Europe including the British Isles east into Asia as far as the Ob and Yenisei Rivers. It is also found in Lake Baikal...

, brown bullhead
Brown bullhead
The brown bullhead, Ameiurus nebulosus, is a fish of the Ictaluridae family that is widely distributed in North America. It is a species of bullhead catfish and is similar to the black bullhead and yellow bullhead...

, and swell shark become motionless and unresponsive at night (or by day, in the case of the swell shark); Spanish hogfish
Spanish hogfish
The Spanish hogfish is a species of hogfish whose range includes South Florida, the Bahamas, the Caribbean Sea, the Gulf of Mexico, Bermuda, and Brazil.. Typically between 20 cm and 35 cm in size, it can reach 60 cm. It inhabits reefs at depths of 3 m up to 30 m, where it...

 and blue-headed wrasse
Blue-headed wrasse
Thalassoma bifasciatum is a is a species of saltwater fish in the wrasse family of order Perciformes native to the coral reefs of the Caribbean Sea. Like its relative T. amblycephalum it is commonly called "blue-headed wrasse", "blunt-headed wrasse" or "bluehead"...

 can even be lifted by hand all the way to the surface without evoking a response. A 1961 observational study of approximately 200 species in European public aquaria reported many cases of apparent sleep. On the other hand, sleep patterns are easily disrupted and may even disappear during periods of migration, spawning, and parental care.

Reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...

s have been subjected to electrophysiological studies of sleep. That is to say that electrical activity in the brain has been registered when the animals have been asleep. However, the EEG pattern in reptilian sleep differs from what is seen in mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s and other animals. In reptiles, sleep time increases following sleep deprivation
Sleep deprivation
Sleep deprivation is the condition of not having enough sleep; it can be either chronic or acute. A chronic sleep-restricted state can cause fatigue, daytime sleepiness, clumsiness and weight loss or weight gain. It adversely affects the brain and cognitive function. Few studies have compared the...

, and stronger stimuli are needed to awaken the animals when they have been deprived of sleep as compared to when they have slept normally. This suggests that the sleep which follows deprivation is compensatorily deeper.

Sleep in birds

There are significant similarities between sleep in bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s and sleep in mammals, which is one of the reasons for the idea that sleep in higher animals with its division into REM and NREM sleep has evolved together with warm-blooded
Warm-blooded
The term warm-blooded is a colloquial term to describe animal species which have a relatively higher blood temperature, and maintain thermal homeostasis primarily through internal metabolic processes...

ness. Birds compensate for sleep loss in a manner similar to mammals, by deeper or more intense SWS (slow-wave sleep
Slow-wave sleep
Slow-wave sleep , often referred to as deep sleep, consists of stages 3 and 4 of non-rapid eye movement sleep, according to the Rechtschaffen & Kales standard of 1968. As of 2008, the American Academy of Sleep Medicine has discontinued the use of stage 4, such that the previous stages 3 and 4 now...

).

Birds have both REM and NREM sleep, and the EEG patterns of both have similarities to those of mammals. Different birds sleep different amounts, but the associations seen in mammals between sleep and variables such as body mass, brain mass, relative brain mass, basal metabolism
Basal metabolic rate
Basal Metabolic Rate , and the closely related resting metabolic rate , is the amount of daily energy expended by humans and other animals at rest. Rest is defined as existing in a neutrally temperate environment while in the post-absorptive state...

 and other factors (see below) are not found in birds. The only clear explanatory factor for the variations in sleep amounts for birds of different species is that birds who sleep in environments where they are exposed to predators
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...

 have less deep sleep than birds sleeping in more protected environments.

A peculiarity that birds share with aquatic mammals, and possibly also with certain species of lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s (opinions differ about that last point), is the ability for unihemispheric sleep. That is the ability to sleep with one cerebral hemisphere
Cerebral hemisphere
A cerebral hemisphere is one of the two regions of the eutherian brain that are delineated by the median plane, . The brain can thus be described as being divided into left and right cerebral hemispheres. Each of these hemispheres has an outer layer of grey matter called the cerebral cortex that is...

 at a time, while the other hemisphere is awake. When only one hemisphere is sleeping, only the contralateral eye will be shut; that is, when the right hemisphere is asleep the left eye will be shut, and vice versa. The distribution of sleep between the two hemispheres and the amount of unihemispheric sleep are determined both by which part of the brain has been the most active during the previous period of wake—that part will sleep the deepest—and it is also determined by the risk of attacks from predators. Ducks near the perimeter of the flock are likely to be the ones that first will detect predator attacks. These ducks have significantly more unihemispheric sleep than those who sleep in the middle of the flock, and they react to threatening stimuli seen by the open eye.

Opinions partly differ about sleep in migratory birds. The controversy is mainly about whether they can sleep while flying or not. Theoretically, certain types of sleep could be possible while flying, but technical difficulties preclude the recording of brain activity in birds while they are flying.

Sleep duration

Different animals sleep different amounts. Some animals, such as bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...

s, sleep 18–20 hours per day, while others, including giraffe
Giraffe
The giraffe is an African even-toed ungulate mammal, the tallest of all extant land-living animal species, and the largest ruminant...

s, sleep only 3–4 hours per day. There can be big differences even between closely related species. There can also be differences between laboratory and field studies: for example, researchers in 1983 reported that captive sloths slept nearly 16 hours a day, but in 2008, when miniature neurophysiological recorders were developed that could be affixed to wild animals, sloths in nature were found to sleep only 9.6 hours a day.
As for birds, the main rule for mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s (with certain exceptions, see below) is that they have two essentially different stages of sleep: REM and NREM sleep (see above). An animal's feeding habits are associated with its sleep length. The daily need for sleep is highest in carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

s, lower in omnivore
Omnivore
Omnivores are species that eat both plants and animals as their primary food source...

s and lowest in herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s. Human
Human
Humans are the only living species in the Homo genus...

s do not sleep unusually much or unusually little compared to other animals, but we sleep less than many other omnivores. Many herbivores, like Ruminantia
Ruminantia
Ruminantia includes many of the well-known large grazing or browsing mammals: among them cattle, goats, sheep, deer, and antelope. All members of the Ruminantia are ruminants: they digest food in two steps, chewing and swallowing in the normal way to begin with, and then regurgitating the...

 (such as cattle), spend much of their wake time in a state of drowsiness, which perhaps could partly explain their relatively low need for sleep. In herbivores, a direct correlation is apparent between body mass and sleep length; big animals sleep more than smaller ones. This correlation is thought to explain about 25% of the difference in sleep amount between different animals. Also, the length of a particular sleep cycle is associated with the size of the animal; on average, bigger animals will have sleep cycles of longer durations than smaller animals. Sleep amount is also coupled to factors like basal metabolism, brain mass and relative brain mass.

Mammals born with well-developed regulatory systems, such as the horse and giraffe, tend to have less REM sleep than the species which are less developed at birth, such as cats and rats. This appears to echo the greater need for REM sleep among newborns than among adults in most mammal species.

Comparative average sleep periods for various mammals in captivity over 24 hours have been given as: horses - 2.9 hours; elephants - 3 plus; cows - 4.0; giraffes - 4.5 hours; humans - 8.0; rabbits - 8.4; chimpanzees - 9.7; Red foxes - 9.8; dogs - 10.1; House mice - 12.5; cats - 12.5; lions - 13.5; platypuses - 14; chipmunks - 15; Giant armadillos - 18.1; and Little brown bats - 19.9 hours. Reasons given for the wide variations include the fact that mammals "that nap in hiding, like bats or rodents tend to have longer, deeper snoozes than those on constant alert." Lions, which have little fear of predators also have relatively long sleep periods, while elephants have to eat most of the time to support their huge bodies. Little brown bat
Little brown bat
The little brown bat is a species of the genus Myotis , one of the most common bats of North America...

s conserve their energy except for the few hours each night when their insect prey are available and platypus
Platypus
The platypus is a semi-aquatic mammal endemic to eastern Australia, including Tasmania. Together with the four species of echidna, it is one of the five extant species of monotremes, the only mammals that lay eggs instead of giving birth to live young...

es eat a high energy crustacean diet and, therefore, probably don't need to spend as much time awake as many other mammals.

Sleep in monotremes

Since monotreme
Monotreme
Monotremes are mammals that lay eggs instead of giving birth to live young like marsupials and placental mammals...

s, egg-laying mammals, are considered to represent one of the evolutionarily oldest groups of mammals - they have been subject to special interest in the study of mammalian sleep. As early studies of these animals could not find clear evidence for REM sleep, it was initially assumed that such sleep did not exist in monotremes but developed after the monotremes left the rest of the mammals and became a separate, distinct group. However, EEG
EEG
EEG commonly refers to electroencephalography, a measurement of the electrical activity of the brain.EEG may also refer to:* Emperor Entertainment Group, a Hong Kong-based entertainment company...

 registrations of the brain stem
Brain stem
In vertebrate anatomy the brainstem is the posterior part of the brain, adjoining and structurally continuous with the spinal cord. The brain stem provides the main motor and sensory innervation to the face and neck via the cranial nerves...

 in monotremes show a firing pattern that is quite similar to the patterns seen in REM sleep in higher mammals. In fact, the largest amount of REM sleep known in any animal is found in the platypus
Platypus
The platypus is a semi-aquatic mammal endemic to eastern Australia, including Tasmania. Together with the four species of echidna, it is one of the five extant species of monotremes, the only mammals that lay eggs instead of giving birth to live young...

. The average sleep time in a 24-hour period of a platypus is said to be as long as 14 hours. "Why? Maybe because just a little crustacean meal packs a huge caloric punch."

Sleep in aquatic mammals

Among others, seals
Pinniped
Pinnipeds or fin-footed mammals are a widely distributed and diverse group of semiaquatic marine mammals comprising the families Odobenidae , Otariidae , and Phocidae .-Overview: Pinnipeds are typically sleek-bodied and barrel-shaped...

 and whale
Whale
Whale is the common name for various marine mammals of the order Cetacea. The term whale sometimes refers to all cetaceans, but more often it excludes dolphins and porpoises, which belong to suborder Odontoceti . This suborder also includes the sperm whale, killer whale, pilot whale, and beluga...

s belong to the aquatic mammals. Seals are grouped in earless seal
Earless seal
The true seals or earless seals are one of the three main groups of mammals within the seal superfamily, Pinnipedia. All true seals are members of the family Phocidae . They are sometimes called crawling seals to distinguish them from the fur seals and sea lions of the family Otariidae...

s and eared seal
Eared Seal
The eared seals or otariids are marine mammals in the family Otariidae, one of three groupings of Pinnipeds. They comprise 16 species in seven genera commonly known either as sea lions or fur seals, distinct from true seals and the Walrus...

s, which have solved the problem of sleeping in water differently. Eared seals, like whales, show unihemispheric sleep. The sleeping half of the brain does not awaken when they surface to breathe. When one half of a seal's brain shows slow-wave sleep, the flippers and whiskers on its opposite side are immobile. While in the water, these seals have almost no REM sleep and may go a week or two without it. As soon as they move onto land they switch to bilateral REM sleep and NREM sleep comparable to land mammals, surprising researchers with their lack of "recovery sleep" after missing so much REM.
Earless seals sleep bihemispherically like most mammals, under water, hanging at the water surface or on land. They hold their breath while sleeping under water, and wake up regularly to surface and breathe. They can also hang with their nostrils above water and in that position have REM sleep, but they do not have REM sleep underwater.

REM sleep has been observed in the pilot whale
Pilot whale
Pilot whales are cetaceans belonging to the genus Globicephala. There are two extant species, the long-finned pilot whale and the short-finned pilot whale . The two are not readily distinguished at sea and analysis of the skulls is the best way to tell the difference between them...

, a species of dolphin. Whales do not seem to have REM sleep, nor do they seem to have any problems because of this. One reason REM sleep might be difficult in marine settings is the fact that REM sleep causes muscular atony
Atony
Pronounced In medicine, atony refers to a muscle that has lost its strength. It is frequently associated with the conditions Atonic seizure, atonic colon, uterine atony, gastrointestinal atony and choreatic atonia.Atony can also refer to the paralyzed or extremely relaxed state of skeletal...

; that is to say, a functional paralysis of skeletal muscles that can be difficult to combine with the need to breathe regularly.

Unihemispheric sleep

Unihemispheric sleep refers to sleeping with only a single cerebral hemisphere
Cerebral hemisphere
A cerebral hemisphere is one of the two regions of the eutherian brain that are delineated by the median plane, . The brain can thus be described as being divided into left and right cerebral hemispheres. Each of these hemispheres has an outer layer of grey matter called the cerebral cortex that is...

. The phenomenon has been observed in birds and aquatic mammals, as well as in several reptilian species (the latter being disputed: many reptiles behave in a way which could be construed as unihemispheric sleeping, but EEG
EEG
EEG commonly refers to electroencephalography, a measurement of the electrical activity of the brain.EEG may also refer to:* Emperor Entertainment Group, a Hong Kong-based entertainment company...

 studies have given contradictory results). Reasons for the development of unihemispheric sleep are likely that it enables the sleeping animal to receive stimuli - threats, for instance - from its environment, and that it enables the animal to fly or periodically surface to breathe when immersed in water. Only NREM sleep exists unihemispherically, and there seems to exist a continuum in unihemispheric sleep regarding the differences in the hemispheres: in animals exhibiting unihemispheric sleep, conditions range from one hemisphere being in deep sleep with the other hemisphere being awake to one hemisphere sleeping lightly with the other hemisphere being awake. If one hemisphere is selectively deprived of sleep in an animal exhibiting unihemispheric sleep (one hemisphere is allowed to sleep freely but the other is awoken whenever it falls asleep), the amount of deep sleep will selectively increase in the hemisphere that was deprived of sleep when both hemispheres are allowed to sleep freely.

The neurobiological background for unihemispheric sleep is still unclear. In experiments on cats, where the connection between the left and the right halves of the brain stem is severed, the brain hemispheres show a desynchronized EEG where the two hemispheres can sleep independently of each other. In these cats, the state where one hemisphere slept NREM and the other was awake, as well as one hemisphere sleeping NREM with the other state sleeping REM were observed. Interestingly, the cats were never seen to sleep REM sleep with one hemisphere while the other hemisphere was awake. This is in accordance with the fact that REM sleep, as far as is currently known, does not occur unihemispherically.

Sleep in hibernating animals

Animals that hibernate are in a state of torpor
Torpor
Torpor, sometimes called temporary hibernation is a state of decreased physiological activity in an animal, usually characterized by a reduced body temperature and rate of metabolism. Animals that go through torpor include birds and some mammals such as mice and bats...

, differing from sleep. Hibernation markedly reduces the need for sleep, but does not remove it. Hibernating animals end their hibernation a couple of times during the winter so that they can sleep.

External links

  • Sleep in fishes
  • Article originally translated from the Swedish Wikipedia article Sömn hos djur in December 2007
The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
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