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Aestivation ( la, aestas (summer); also spelled estivation in
American English American English, sometimes called United States English or U.S. English, is the set of varieties of the English language native to the United States. English is the most widely spoken language in the United States and in most circumstances ...
) is a state of animal
dormancy Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be c ...
, similar to
hibernation Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It mos ...
, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a lowered
metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
rate, that is entered in response to high temperatures and arid conditions. It takes place during times of heat and dryness, the hot dry season, which are often the summer months. Invertebrate and vertebrate animals are known to enter this state to avoid damage from high temperatures and the risk of
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
. Both terrestrial and aquatic animals undergo aestivation. Fossil records suggest that aestivation may have evolved several hundred million years ago.


Physiology

Organisms that aestivate appear to be in a fairly "light" state of dormancy, as their physiological state can be rapidly reversed, and the organism can quickly return to a normal state. A study done on '' Otala lactea'', a snail native to parts of Europe and Northern Africa, shows that they can wake from their dormant state within ten minutes of being introduced to a wetter environment. The primary physiological and biochemical concerns for an aestivating animal are to conserve energy, retain water in the body, ration the use of stored energy, handle the nitrogenous end products, and stabilize bodily organs, cells, and
macromolecule A macromolecule is a very large molecule important to biophysical processes, such as a protein or nucleic acid. It is composed of thousands of covalently bonded atoms. Many macromolecules are polymers of smaller molecules called monomers. The ...
s. This can be quite a task as hot temperatures and arid conditions may last for months, in some cases for years. The depression of metabolic rate during aestivation causes a reduction in macromolecule synthesis and degradation. To stabilize the macromolecules, aestivators will enhance antioxidant defenses and elevate
chaperone proteins In molecular biology, molecular chaperones are proteins that assist the conformational folding or unfolding of large proteins or macromolecular protein complexes. There are a number of classes of molecular chaperones, all of which function to ass ...
. This is a widely used strategy across all forms of hypometabolism. These physiological and biochemical concerns appear to be the core elements of hypometabolism throughout the animal kingdom. In other words, animals which aestivate appear to go through nearly the same physiological processes as animals that hibernate.


Invertebrates


Mollusca

Gastropoda The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. The ...
: some air-breathing
land snail A land snail is any of the numerous species of snail that live on land, as opposed to the sea snails and freshwater snails. ''Land snail'' is the common name for terrestrial gastropod mollusks that have shells (those without shells are known ...
s, including species in the genera ''
Helix A helix () is a shape like a corkscrew or spiral staircase. It is a type of smooth space curve with tangent lines at a constant angle to a fixed axis. Helices are important in biology, as the DNA molecule is formed as two intertwined helic ...
, Cernuella, Theba, Helicella, Achatina'' and '' Otala'', commonly aestivate during periods of heat. Some species move into shaded vegetation or rubble. Others climb up tall plants, including crop species as well as bushes and trees, and will also climb man-made structures such as posts, fences, etc. Their habit of climbing vegetation to aestivate has caused more than one introduced snail species to be declared an agricultural nuisance. To seal the opening to their shell to prevent water loss, pulmonate land snails secrete a membrane of dried mucus called an
epiphragm An epiphragm (from the Ancient Greek ἐπί, '' epi '' " upon, on, over " and φράγμα, '' -phrágma '' "fence") is a temporary structure which can be created by many species of shelled, air-breathing land snails, terrestrial pulmonate gast ...
. In certain species, such as ''
Helix pomatia ''Helix pomatia'', common names the Roman snail, Burgundy snail, or escargot, is a species of large, edible, air-breathing land snail, a pulmonate gastropod terrestrial mollusc in the family Helicidae.MolluscaBase eds. (2021). MolluscaBase. ...
'', this barrier is reinforced with calcium carbonate, and thus it superficially resembles an operculum, except that it has a tiny hole to allow some oxygen exchange. There is a decrease in
metabolic rate Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
and reduced rate of water loss in aestivating snails like ''
Rhagada tescorum ''Rhagada'' is a genus of air-breathing land snails, terrestrial pulmonate gastropod mollusks in the subfamily Hadrinae of the family Camaenidae.MolluscaBase eds. (2021). MolluscaBase. Rhagada Albers, 1860. Accessed through: World Register of Mar ...
'', ''
Sphincterochila boissieri ''Sphincterochila zonata zonata'' is a subspecies of air-breathing land snail, a terrestrial molluscs, terrestrial pulmonate gastropod mollusk in the family Sphincterochilidae. MolluscaBase eds. (2022). MolluscaBase. Sphincterochila zonata zonata ...
'' and others.


Arthropoda

Insecta Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs o ...
: Lady beetles (''
Coccinellidae Coccinellidae () is a widespread family of small beetles ranging in size from . They are commonly known as ladybugs in North America and ladybirds in Great Britain. Some entomologists prefer the names ladybird beetles or lady beetles as they ...
'') have been reported to aestivate. Mosquitoes also are reported to undergo aestivation. False honey ants are well known for being winter active and aestivate in temperate climates.
Bogong moth The bogong moth (''Agrotis infusa'') is a temperate species of night-flying moth, notable for its biannual long-distance seasonal migrations towards and from the Australian Alps, similar to the diurnal monarch butterfly. During the autumn an ...
s will aestivate over the summer to avoid the heat and lack of food sources. Adult alfalfa weevils (''
Hypera postica ''Hypera postica'', commonly known as the alfalfa weevil, is a species of beetle in the superfamily Curculionoidea; it can be found in alfalfa fields throughout Europe. Considered a destructive threat to alfalfa production in North America, se ...
'') aestivate during the summer in the southeastern United States, during which their metabolism, respiration, and nervous systems show a dampening of activity.
Crustacea Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group c ...
: An example of a crustacean undergoing aestivation is with the Australian crab ''Austrothelphusa transversa ,'' which undergoes aestivation underground during the dry season''.''


Vertebrates


Reptiles and amphibians

Non-mammalian animals that aestivate include North American desert tortoises,
crocodile Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant me ...
s, and
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s. Some amphibians (e.g. the
cane toad The cane toad (''Rhinella marina''), also known as the giant neotropical toad or marine toad, is a large, terrestrial true toad native to South and mainland Central America, but which has been introduced to various islands throughout Oceania ...
and
greater siren The greater siren (''Siren lacertina'') is an eel-like amphibian and one of the three members of the genus '' Siren''. The largest of the sirens and one of the largest amphibians in North America, the greater siren resides in the coastal plains o ...
) aestivate during the hot dry season by moving underground where it is cooler and more humid. The
California red-legged frog The California red-legged frog (''Rana draytonii'') is a species of frog found in California (USA) and northern Baja California (Mexico). It was formerly considered a subspecies of the northern red-legged frog (''Rana aurora''). The frog is an IU ...
may aestivate to conserve energy when its food and water supply is low. The water-holding frog has an aestivation cycle. It buries itself in sandy ground in a secreted, water-tight mucus cocoon during periods of hot, dry weather.
Australian Aboriginals Aboriginal Australians are the various Indigenous peoples of the Australian mainland and many of its islands, such as Tasmania, Fraser Island, Hinchinbrook Island, the Tiwi Islands, and Groote Eylandt, but excluding the Torres Strait Isla ...
discovered a means to take advantage of this by digging up one of these frogs and squeezing it, causing the frog to empty its bladder. This dilute urine—up to half a glassful—can be drunk. However, this will cause the death of the frog which will be unable to survive until the next rainy season without the water it had stored. The
western swamp turtle The western swamp turtle or western swamp tortoise (''Pseudemydura umbrina'') is a critically endangered species of freshwater turtle endemic to a small portion of Western Australia. It is the only member of the genus ''Pseudemydura'' in the mon ...
aestivates to survive hot summers in the ephemeral swamps it lives in. It buries itself in various media which change depending on location and available substrates. Because the species is critically endangered, the
Perth Zoo Perth Zoo is a zoological park in South Perth, Western Australia. The zoo first opened in 1898 and by 2011 housed 1258 animals of 164 species and an extensive botanical collection. It is a full institutional member of the Zoo and Aquarium Ass ...
began a conservation and breeding program for it. However, zookeepers were unaware of the importance of their aestivation cycle and during the first summer period would perform weekly checks on the animals. This repeated disturbance was detrimental to the health of the animals, with many losing significant weight and some dying. The zookeepers quickly changed their procedures and now leave their captive turtles undisturbed during their aestivation period.


Fish

African lungfish ''Protopterus'' is the genus of four species of lungfish found in Africa. ''Protopterus'' was formerly thought to be the sole genus in the family Protopteridae, but more recent studies have classified it with ''Lepidosiren'' in the family Lepi ...
also aestivate as can salamanderfish.


Mammals

Although relatively uncommon, a small number of mammals aestivate. Animal physiologist Kathrin Dausmann of
Philipps University of Marburg The Philipps University of Marburg (german: Philipps-Universität Marburg) was founded in 1527 by Philip I, Landgrave of Hesse, which makes it one of Germany's oldest universities and the oldest still operating Protestant university in the worl ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
, and coworkers presented evidence in a 2004 edition of ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...
'' that the Malagasy
fat-tailed dwarf lemur The fat-tailed dwarf lemur (''Cheirogaleus medius''), also known as the lesser dwarf lemur, western fat-tailed dwarf lemur, or spiny forest dwarf lemur, is endemic to Madagascar. Hibernation Recent research has shown that ''C. medius'' hibernat ...
hibernates or aestivates in a small tree hole for seven months of the year. According to the Oakland Zoo in California, four-toed hedgehogs are thought to aestivate during the dry season.


See also

* Critical thermal maximum * Hibernation induction trigger *
Torpor Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature and metabolic rate. Torpor enables animals to survive periods of reduced food availability. The term "torpor" can refer to the time ...


References


Further reading

*


External links


Abstract of an Australian paper on aestivation in snails






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