Avian sleep

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In birds, sleep consists of "periods of eye closure interrupted by short periods of eye-opening."[ This quote needs a citation ] During the short periods of eye-opening, electroencephalographic (EEG) studies indicate that the birds are still sleeping; the voltage level in the brain is identical. [1] Birds restore their arousal thresholds during sleep. During their short eye-open periods, sleeping birds can mobilize almost instantaneously when threatened by a predator. Avian species have been found to rely on flock size and height for predatory precautions. [1] Between the eye-opening and group sleep, these precautions allow sleep to be beneficial and safe.

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The amount of sleep necessary to function can vary by species. Pectoral sandpipers migrate from the Southern Hemisphere to the Arctic Circle, their mating ground (where they breed during daylight. Since the sandpipers are polygamous, they mate (or search for a mate) for the duration of daylight. Males do not require as much sleep during this time; some have been observed to give up 95 percent of their sleep time during the nineteen mating days. [2] Most act similarly to humans when sleep-deprived, getting them into potentially life-threatening situations or slowing their migration speed. [3]

Comparative anatomy of avian brain and nervous system

The typical avian nervous system is similar to that of mammals. The central nervous system includes the brain and spinal cord, and a peripheral nervous system consists of nerves and sensory organs. Key attributes have evolved compared to other species, especially vision; avian visual capabilities are believed to be more advanced than any other group of vertebrates. In addition to larger eyes, birds have larger-than-average optic lobes. With a larger, more intricate optic lobe, some bird species can view the ultraviolet (UV) spectrum (beyond the visual range of the human eye). This UV visual capability facilitates hunting, as seen in nighthawks. UV-sensitive cone opsin is typically responsible for avian ability to see UV, but some species have circumvented this; owls can see UV light, but lack opsins. They compensate for this with essential enzymes which allow heightened rod sensitivity. [4] UV is seen by several other animal groups, including cats and insects (where it has appeared to evolve in response to predator-prey relationships).

Trade-offs in anatomy and physiology are common, and this is seen in the olfactory lobes of most avian species. Possibly due to the larger-than-average optic lobes, avian olfactory lobes are relatively small; few bird species use smell to find food. Falcons and eagles do not tend to have larger cerebellums for flying. According to comparative neuroanatomy researcher Ludwig Edinger, avian brains consist mostly of basal ganglia (responsible for instinctive behavior, rather than behavioral plasticity). Scientists have challenged some of Edinger's findings, and called for the renaming of avian nervous-system organs to reflect their similarity to those of mammals.

REM and slow-wave sleep

Avian sleep shares two similarities with that of mammals: rapid eye movement (REM) and slow-wave sleep (SWS). REM sleep is believed to have an important effect on motor functions and memory storage. EEGs show high-amplitude and low-frequency waves during REM sleep; SWS tends toward lower-amplitude, higher-frequency waves, and is believed to be a form of deep sleep. During SWS, membrane potentials in the neurons of the neocortex oscillate slowly.

A number of avian species exhibit unihemispheric slow-wave sleep: the ability to rest one half of the brain in SWS, while the other half appears to be awake. This type of sleep has also been seen in dolphins and whales. The organism is typically able to keep one eye open during this process, which allows added vigilance in high-predation environments. [5] The evolution of this trait in birds and aquatic mammals is of interest to researchers because of the pressures involved. Unihemispheric SWS is thought to have evolved in aquatic mammals because they must return to the surface for oxygen; it is believed to help birds avoid predation, demonstrating homoplasy in the two groups.

Dove experiment

In a study of how the Barbary dove's sleep patterns are affected by flock size, D. W. Lendrum intended to prove that larger flocks reduced overall vigilance, and the apparent increase in predation risk of a smaller flock would harm the doves' sleep cycle. [2] At the beginning of the study, the doves were caged alone or in pairs of cages containing two, three or six. They were then placed in one of two environments. In the calm environment, Lendrum walked alone past the cage between 10 am and noon; in the aggressive environment, Lendrum walked past the cage with a domesticated ferret at the same time of day. Lendrum discovered that the birds in the calm environment spent substantially more time with their eyes closed than those in the aggressive environment. [2]

Lendrum collected data on the doves' opened- and closed-eye sleep; flocking was associated with an increase in a bird's overall eye-closure time and a decrease in its amount of eye-opening. [2] In the presence of a predator, Lendrum found that the doves exhibited higher levels of individual vigilance and increase in open-eye sleep; this reduced the active-sleep component of their total sleep time. [2]

Perch height

Predators are believed to play a large role in an organism's sleeping patterns. To adapt to predation, two common techniques have evolved: positioning oneself out of harm's way while sleeping, and sleeping more lightly (such as unihemispheric sleep). In birds, perch height is believed to play a significant role in sleep; lower perch height has been shown to reduce the number and length of REM sleep episodes in pigeons, and a higher perch increases REM sleep and decreases slow-wave sleep. [6] Findings also suggest that the time spent awake by pigeons increases when nesting on lower perches. Lower perch height correlates to a higher risk of predation; REM sleep would place the pigeon in more danger, since it is a less reactive form of sleep. [6]

Light pollution

Light is one of the more common threats to sufficient sleep for birds living in anthropogenic environments, known as "artificial light pollution at night" (ALAN). ALAN eliminates darkness, a necessity for rest. Disrupting the birds' light and dark cycles can impact circadian rhythms, eventually harming sleep patterns. Biologist Thomas Raap conducted a study which suggested that exposure to ALAN affected the sleep behavior of Eurasian blue tits (Cyanistes caeruleus). [7] In this study, birds woke up earlier due to ALAN factors such as seasonal timekeeping. Because light usually indicates a day's passage to birds, exposure to light pollution disrupts their ability to measure the length of a day. Outside densely-populated areas, there is normally about a five-percent drop in sleep duration for blue-tit females during their nesting period. The researchers found a 50-percent reduction in the females' sleep duration during this period in urban centers, and suggested that the effects of ALAN were responsible. [7]

Related Research Articles

Nocturnality animal behavior characterized by activity during the night and sleeping during the day

Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite.

Rapid eye movement sleep Unique phase of sleep in mammals and birds, characterized by random/rapid movement of the eyes

Rapid eye movement sleep is a unique phase of sleep in mammals and birds, distinguishable by random/rapid movement of the eyes, accompanied with low muscle tone throughout the body, and the propensity of the sleeper to dream vividly.

Rock dove Species of bird

The rock dove, rock pigeon, or common pigeon is a member of the bird family Columbidae. In common usage, this bird is often simply referred to as the "pigeon".

Sleep cycle

The sleep cycle is an oscillation between the slow-wave and REM (paradoxical) phases of sleep. It is sometimes called the ultradian sleep cycle, sleep–dream cycle, or REM-NREM cycle, to distinguish it from the circadian alternation between sleep and wakefulness. In humans this cycle takes 1–2 hours. Humans usually grow from time period of 1 AM to 3 PM during night.

Delta wave A high amplitude low frequency brain wave

A delta wave is a high amplitude brain wave with a frequency of oscillation between 0.5 and 4 hertz. Delta waves, like other brain waves, are recorded with an electroencephalogram (EEG) and are usually associated with the deep stage 3 of NREM sleep, also known as slow-wave sleep (SWS), and aid in characterizing the depth of sleep.

Non-rapid eye movement sleep(NREM), also known as quiescent sleep, is, collectively, sleep stages 1–3, previously known as stages 1–4. Rapid eye movement sleep (REM) is not included. There are distinct electroencephalographic and other characteristics seen in each stage. Unlike REM sleep, there is usually little or no eye movement during these stages. Dreaming is rare during NREM sleep, and muscles are not paralyzed as in REM sleep. People who do not go through the sleeping stages properly get stuck in NREM sleep, and because muscles are not paralyzed a person may be able to sleepwalk. According to studies, the mental activity that takes place during NREM sleep is believed to be thought-like, whereas REM sleep includes hallucinatory and bizarre content. The mental activity that occurs in NREM and REM sleep is a result of two different mind generators, which also explains the difference in mental activity. In addition, there is a parasympathetic dominance during NREM. During the period of Non-REM sleep, the mindset of a person is more organized. The reported differences between the REM and NREM activity are believed to arise from differences in the memory stages that occur during the two types of sleep. It has been found through several experiments that low levels of stage 3 sleep are observed in about 40-50% of people with acute and chronic schizophrenia.

Herd group of animals

A herd is a social group of certain animals of the same species, either wild or domestic. The form of collective animal behavior associated with this is referred to as herding.

Sleep inertia, most-known as Hicham's syndrome, is a physiological state of impaired cognitive and sensory-motor performance that is present immediately after awakening. It persists during the transition of sleep to wakefulness, where an individual will experience feelings of drowsiness, disorientation and a decline in motor dexterity. Impairment from sleep inertia may take several hours to dissipate. In the majority of cases, morning sleep inertia is experienced for 15 to 30 minutes after waking.

Anti-predator adaptation Defensive feature of prey for selective advantage

Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avoiding detection, warding off attack, fighting back, or escaping when caught.

Slow-wave sleep

Slow-wave sleep (SWS), often referred to as deep sleep, consists of stage three of non-rapid eye movement sleep. Initially, SWS consisted of both Stage 3, which has 20–50 percent delta wave activity, and Stage 4, which has more than 50 percent delta wave activity.

Unihemispheric slow-wave sleep

Unihemispheric slow-wave sleep (USWS) is sleep where one half of the brain rests while the other half remains alert. This is in contrast to normal sleep where both eyes are shut and both halves of the brain show unconsciousness. In USWS, also known as asymmetric slow-wave sleep, one half of the brain is in deep sleep, a form of non-rapid eye movement sleep and the eye corresponding to this half is closed while the other eye remains open. When examined by low-voltage electroencephalography (EEG), the characteristic slow-wave sleep tracings are seen from one side while the other side shows a characteristic tracing of wakefulness. The phenomenon has been observed in a number of terrestrial, aquatic and avian species.

Flock (birds) group of birds conducting flocking behavior in the midst of flight, or while foraging or roosting

A flock is a gathering of a group of same species animals in order to forage or travel with one another. In avians flocks are typically seen in association with migration. While this is true it can also be seen that flocking is important in safety from predation and foraging benefits. Living in a flock can also come at a cost to the birds living within it.

Bird colony large congregation of birds at a particular location

A bird colony is a large congregation of individuals of one or more species of bird that nest or roost in proximity at a particular location. Many kinds of birds are known to congregate in groups of varying size; a congregation of nesting birds is called a breeding colony. Colonial nesting birds include seabirds such as auks and albatrosses; wetland species such as herons; and a few passerines such as weaverbirds, certain blackbirds, and some swallows. A group of birds congregating for rest is called a communal roost. Evidence of colonial nesting has been found in non-neornithine birds (Enantiornithes), in sediments from the Late Cretaceous (Maastrichtian) of Romania.

Communal roosting

Communal roosting is an animal behavior where a group of individuals, typically of the same species, congregate in an area for a few hours based on an external signal and will return to the same site with the reappearance of the signal. Environmental signals are often responsible for this grouping, including nightfall, high tide, or rainfall. The distinction between communal roosting and cooperative breeding is the absence of chicks in communal roosts. While communal roosting is generally observed in birds, the behavior has also been seen in bats, primates, and insects. The size of these roosts can measure in the thousands to millions of individuals, especially among avian species.

Sleep in non-human animals sleep in non-human beings

Sleep in non-human animals refers to a behavioral and physiological state characterized by altered consciousness, reduced responsiveness to external stimuli, and homeostatic regulation. Sleep is observed in mammals, birds, reptiles, amphibians, and some fish, and, in some form, in insects and even in simpler animals such as nematodes. The internal circadian clock promotes sleep at night for diurnal organisms and in the day for nocturnal organisms. Sleep patterns vary widely among species. It appears to be a requirement for all mammals and most other animals.

Bird vision senses for birds

Vision is the most important sense for birds, since good eyesight is essential for safe flight, and this group has a number of adaptations which give visual acuity superior to that of other vertebrate groups; a pigeon has been described as "two eyes with wings". The avian eye resembles that of a reptile, with ciliary muscles that can change the shape of the lens rapidly and to a greater extent than in the mammals. Birds have the largest eyes relative to their size in the animal kingdom, and movement is consequently limited within the eye's bony socket. In addition to the two eyelids usually found in vertebrates, it is protected by a third transparent movable membrane. The eye's internal anatomy is similar to that of other vertebrates, but has a structure, the pecten oculi, unique to birds.

Ponto-geniculo-occipital waves or PGO waves are distinctive wave forms of propagating activity between three key brain regions: the pons, lateral geniculate nucleus, and occipital lobe; specifically, they are phasic field potentials. These waves can be recorded from any of these three structures during and immediately before REM sleep. The waves begin as electrical pulses from the pons, then move to the lateral geniculate nucleus residing in the thalamus, and then finally end up in the primary visual cortex of the occipital lobe. The appearances of these waves are most prominent in the period right before REM sleep, albeit they have been recorded during wakefulness as well. They are theorized to be intricately involved with eye movement of both wake and sleep cycles in many different animals.

Sleep and memory

The relationship between sleep and memory has been studied since at least,the early 19th century. Memory, the cognitive process of storing and retrieving past experiences, learning and recognition, is a product of brain plasticity, the structural changes within synapses that create associations between stimuli. Stimuli are encoded within milliseconds; however, the long-term maintenance of memories can take additional minutes, days, or even years to fully consolidate and become a stable memory that is accessible. Therefore, the formation of a specific memory occurs rapidly, but the evolution of a memory is often an ongoing process.

Neuroscience of sleep Study of the neuroscientific and physiological basis of the nature of sleep

The neuroscience of sleep is the study of the neuroscientific and physiological basis of the nature of sleep and its functions. Traditionally, sleep has been studied as part of psychology and medicine. The study of sleep from a neuroscience perspective grew to prominence with advances in technology and proliferation of neuroscience research from the second half of the twentieth century.

Vigilance, in the field of behavioural ecology, refers to an animal's examination of its surroundings in order to heighten awareness of predator presence. Vigilance is an important behaviour during foraging as animals must often venture away from the safety of shelter to find food. However being vigilant comes at the expense of time spent feeding so there is a trade-off between the two. The length of time animals devote to vigilance is dependent on many factors including predation risk and hunger.

References

  1. 1 2 Lesku, J. & Rattenborg, N. (2014). "Avian sleep"Current Biology24 (1): R12–R14.
  2. 1 2 3 4 5 Lendrem, D. W. (June 3, 2006). "Sleeping and vigilance in birds, II. An experimental study of the Barbary dove (Streptopelia risoria)".
  3. Jiang, Y.; Chai, Y.; Yang, F.; Xu, S.; Basner, M.; Detre, J. A.; Dinges, D. F. & Rao, H. (2018). "Effects Of Sleep Deprivation And Recovery Sleep On Human Brain Network Organization". Sleep41 (suppll): A85–A86.
  4. Rattenborg, N. (2006). "Evolution of slow-wave sleep and palliopallial connectivity in mammals and birds: A hypothesis"Brain Research Bulletin69 (1): 20–29.
  5. Rattenborg, Niels C.; Lima, Steven L. & Amlaner, Charles J. (1999). "Facultative control of avian unihemispheric sleep under the risk of predation". Behavioural Brain Research105 (2): 163-172. doi : 10.1016/S0166-4328(99)00070-4
  6. 1 2 Tisdale, R. K.; Lesku, J. A.; Beckers, G. J. L.; Vyssotski, A. L. & Rattenborg, N. C. (October 1, 2018). "The low-down on sleeping down low: pigeons shift to lighter forms of sleep when sleeping near the ground".
  7. 1 2 Raap, T.; Sun, J.; Pinxten, R. & Eens, M. (2017). "Disruptive effects of light pollution on sleep in free-living birds: Season and/or light intensity-dependent?". Behavioural Processes: 144. doi : 10.1016/j.beproc.2017.08.011