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Tardigrade



 
 
Tardigrades (commonly known as water bears) form the phylum
Phylum

A phylum "Phylum" is adopted from the Greek phylai, the clan-based voting groups in Greek city-states. is a taxonomic rank below Kingdom and above Class ....
 Tardigrada, part of the superphylum Ecdysozoa
Ecdysozoa

The Ecdysozoa are a grouping of protostome animals, including the Arthropoda , roundworm, and several smaller phylum . They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes....
. They are microscopic, water-dwelling, segmented 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....
s with eight legs. Tardigrades were first described by Johann August Ephraim Goeze in 1773 (kleiner Wasserbär = little water bear). The name Tardigrada means "slow walker" and was given by Spallanzani
Lazzaro Spallanzani

Lazzaro Spallanzani was an Italian biologist and physiologist who made important contributions to the experimental study of bodily functions and animal reproduction, and whose research of biogenesis paved the way for the investigations of Louis Pasteur....
 in 1777. The name water bear comes from the way they walk, reminiscent of a bear's.






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Tardigrades (commonly known as water bears) form the phylum
Phylum

A phylum "Phylum" is adopted from the Greek phylai, the clan-based voting groups in Greek city-states. is a taxonomic rank below Kingdom and above Class ....
 Tardigrada, part of the superphylum Ecdysozoa
Ecdysozoa

The Ecdysozoa are a grouping of protostome animals, including the Arthropoda , roundworm, and several smaller phylum . They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes....
. They are microscopic, water-dwelling, segmented 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....
s with eight legs. Tardigrades were first described by Johann August Ephraim Goeze in 1773 (kleiner Wasserbär = little water bear). The name Tardigrada means "slow walker" and was given by Spallanzani
Lazzaro Spallanzani

Lazzaro Spallanzani was an Italian biologist and physiologist who made important contributions to the experimental study of bodily functions and animal reproduction, and whose research of biogenesis paved the way for the investigations of Louis Pasteur....
 in 1777. The name water bear comes from the way they walk, reminiscent of a bear's. The biggest adults may reach a body length of 1.5 mm
Millimetre

The millimetre is a Units of measurement of length in the metric system, equal to one thousandth of a metre, which is the current International System of Units SI base unit of length....
, the smallest below 0.1 mm. Freshly hatched larva
Larva

A larva is a young form of animal with indirect developmental biology, going through or undergoing metamorphosis .The larva can look completely different from the adult form, for example, a caterpillar differs from a butterfly....
e may be smaller than 0.05 mm.

More than 1000 species of tardigrades have been described. Tardigrades occur over the entire world, from the high Himalayas
Himalayas

The Himalaya Range or Himalayas for short , meaning "abode of snow" ), is a mountain range in Asia, separating the Indian subcontinent from the Tibetan Plateau....
 (above 6,000 m
Metre

The metre or meter is a Unit of measurement of length. It is the SI base unit of length in the metric system and in the International System of Units , used around the world for general and scientific purposes....
), to the deep sea
Deep sea

File:Nur04506.jpgThe deep sea, or deep layer, is the lowest layer in the ocean, existing below the thermocline, at a depth of 1000 fathoms or more....
 (below 4,000 m) and from the polar region
Polar region

Earth polar regions are the areas of the globe surrounding the geographical pole also known as Geographical zone. The North Pole and South Pole being the centers, these regions are dominated by the polar ice caps, resting respectively on the Arctic Ocean and the continent of Antarctica....
s to the equator
Equator

The equator is the intersection of the Earth's surface with the Plane perpendicular to the Earth's rotation and containing the Earth's center of mass....
.

The most convenient place to find tardigrades is on lichen
Lichen

Lichens are composite organisms consisting of a symbiosis association of a fungus with a Photosynthesis partner , usually either a green algae or Cyanobacteria ....
s and moss
Moss

Mosses are small, soft plants that are typically 1?10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations....
es. Other environments are dune
Dune

In physical geography, a dune is a hill of sand built by aeolian processes. Dunes are subject to different forms and sizes based on their interaction with the wind....
s, beach
Beach

File:MiamiSouthBeachPanoramaEdit.jpgA beach is a geology landform along the shoreline of a body of water. It usually consists of loose particles which are often composed of Rock , such as sand, gravel, shingle beach, pebbles, or cobble....
es, soil
Soil

Soil is the naturally occurring, unconsolidated or loose covering on the Earth's surface. Soil is composed of particles of broken rock that have been altered by chemical and environmental processes including weathering and erosion....
 and marine
Ocean

An ocean is a major body of Seawater, and a principal component of the hydrosphere. Approximately 71% of the Earth's surface is covered by ocean, a World Ocean that is customarily divided into several principal oceans and smaller seas....
 or freshwater
Freshwater

Freshwater is a word that refers to bodies of water such as ponds, lakes, rivers and streams containing low concentrations of dissolved salts and other total dissolved solids....
 sediments, where they may occur quite frequently (up to 25,000 animals per litre
Litre

The litre or liter is a unit of volume. There are two official symbols: the Latin letter L in lower and upper case . The lower case L is often written as a cursive l to avoid confusion with the number 1 in antiqua fonts....
). Tardigrades often can be found by soaking a piece of moss
Moss

Mosses are small, soft plants that are typically 1?10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations....
 in spring water.

Tardigrades are polyextremophile
Polyextremophile

A polyextremophile is an organism which combines several extremophilic features. For example, a polyextremophile living at the summit of a mountain in the Atacama Desert might be a radioresistant xerophile, a psychrophile, and an oligotroph....
s and are able to survive in extreme environments that would kill almost any other animal. Some can survive temperatures close to absolute zero
Absolute zero

Absolute zero is a temperature marked by a 0 entropy configuration. It is the coldest temperature theoretically possible, and cannot be reached, by artificial or natural means....
, temperatures as high as 151 °C (303 °F), 1,000 times more radiation than any other animal, nearly a decade without water, and even the vacuum of space.

Anatomy and morphology

Tardigrades have a body with four segments (not counting the head), four pairs of legs without joints, and feet with claw
Claw

A claw is a curved, pointed appendage, found at the end of a toe or finger in most mammals, birds, and some reptiles. Somewhat similar fine hooked structures are found in arthropods such as beetles and spiders, at the end of the leg or Arthropod leg for gripping a surface as the creature walks....
s or toe
Toe

Toes are the Digit s of the foot of an animal. Many animal species such as cats walk on their toes, and are described as being digitigrade....
s. The cuticle
Cuticle

In biology, a cuticle or cuticula is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or part of an organism, that provide protection....
 contains chitin
Chitin

Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world....
 and is moulted
Ecdysis

Ecdysis is the molting of the cuticula in arthropods and related groups . Since the cuticula of these animals is also the skeletal support of the body and is inelastic, it is shed during growth and a new, larger covering is formed....
. They have a ventral nervous system with one ganglion
Ganglion

In anatomy, a ganglion is a biological tissue.Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to Retinal ganglion cells....
 per segment, and a multilobed brain
Brain

The brain is the center of the nervous system in all vertebrate, and most invertebrate, animals. Some primitive animals such as cnidarian and echinoderm have a decentralized nervous system without a brain, while sponges lack any nervous system at all....
. Their pigment-cup eyes are rhabdomeric. Instead of a coelom
Coelom

The coelom is a fluid filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages....
 they have a haemocoel. The only place where a true coelom can be found is around the gonad (coelomic pouch). The pharynx
Pharynx

FunctionsThe pharynx is part of the digestive system and respiratory system of many organisms.Because both food and Earth's atmosphere pass through the pharynx, a flap of connective tissue called the epiglottis closes over the trachea when food is swallowed to prevent choking or Pulmonary aspiration....
 is of a triradiate, muscular, sucking kind, armed with stylet
Stylet

A stylet is a hard, sharp, anatomical structure found in some invertebrates.For example, the word stylet or stomatostyle, is used for the primitive piercing mouthparts of some nematodes and some nemerteans....
s. Although some species are parthenogenetic
Parthenogenesis

Parthenogenesis is an asexual form of reproduction found in females where growth and development of embryos or seeds occurs without fertilization by a male....
, males and females are usually present, each with a single gonad
Gonad

The gonad is the organ that makes gametes. The gonads in males are the testes and the gonads in females are the ovaries. The product, gametes, are haploid germ cells....
. Tardigrades are eutelic (all adult tardigrades of the same species are believed to have the same number of cells) and oviparous. Some tardigrade species have as many as about 40,000 cells in each adult's body, others have far fewer.

Ecology and life history

Most tardigrades are phytophagous (plant eaters) or bacteriophagous (bacteria eaters), but some are predatory (e.g. Milnesium tardigradum).

Physiology


Extreme environments

Tardigrades are polyextremophile
Polyextremophile

A polyextremophile is an organism which combines several extremophilic features. For example, a polyextremophile living at the summit of a mountain in the Atacama Desert might be a radioresistant xerophile, a psychrophile, and an oligotroph....
s; scientists have reported their existence in hot spring
Hot spring

A hot spring is a Spring that is produced by the emergence of Geothermal groundwater from the earth's crust . There are hot springs all over the earth, on every continent and even under the oceans and seas....
s, on top of the Himalayas
Himalayas

The Himalaya Range or Himalayas for short , meaning "abode of snow" ), is a mountain range in Asia, separating the Indian subcontinent from the Tibetan Plateau....
, under layers of solid ice
Ice

Ice is a solid phases of matter, usually crystalline solid, of a non-metallic substance that is liquid or gas at room temperature, such as ammonia ice or methane ice....
 and in ocean sediments. Many species can be found in a milder environment like lake
Lake

A lake is a terrain feature , a body of liquid on the surface of a world that is localized to the bottom of basin and moves slowly if it moves at all....
s, pond
Pond

A pond is a body of water smaller than a lake, both being examples of terrain feature. Although the term pond is universally used to describe waterbodies that are smaller than lakes, an internationally recognised size cutoff has not yet been agreed, with values ranging from 2 hectares to 8 hectares used to distinguish the smaller from...
s and meadow
Meadow

A meadow is a field vegetated primarily by grass and other non-woody plants . It may be cut for hay or grazing by livestock such as cattle, sheep or goats....
s, while others can be found in stone walls and roofs. Tardigrades are most common in moist environments, but can stay active wherever they can retain at least some moisture.

Tardigrades are one of the few groups of species that are capable of reversibly suspending their metabolism
Metabolism

Metabolism is the set of chemical reactions that occur in living organisms in order to maintain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments....
 and going into a state of cryptobiosis
Cryptobiosis

Cryptobiosis is an ametabolic state of life entered by an organism in response to adverse environmental conditions such as desiccation, freezing, and oxygen deficiency....
. Several species regularly survive in a dehydrated state for nearly ten years. Depending on the environment they may enter this state via anhydrobiosis, cryobiosis, osmobiosis or anoxybiosis. While in this state their metabolism lowers to less than 0.01% of normal and their water content can drop to 1% of normal. Their ability to remain desiccated for such a long period is largely dependent on the high levels of the non-reducing sugar
Sugar

Sugar is a class of edible crystalline substances, mainly sucrose, lactose, and fructose. Human taste buds interpret its flavor as sweet. Sugar as a basic food carbohydrate primarily comes from sugar cane and from sugar beet, but also appears in fruit, honey, sorghum, sugar maple , and in many other sources....
 trehalose
Trehalose

Trehalose, also known as mycose, is a natural alpha-linked disaccharide formed by an a, a-1, 1-glucoside bond between two a-glucose units....
, which protects their membrane
Cell membrane

The cell membrane is the interface between the cellular machinery inside the cell and the fluid outside.It is a semipermeable lipid bilayer found in all cell ....
s. In this cryptobiotic state the tardigrade is known as a tun

Tardigrades have been known to withstand the following extremes while in this state:
  • Temperature
    Temperature

    In physics, temperature is a physical property of a Physical system that underlies the common notions of hot and cold; something that feels hotter generally has the greater temperature....
     — tardigrades can survive being heated for a few minutes to 151 °C, or being chilled for days at –200 °C, or for a few minutes at –272 °C. (absolute zero
    Absolute zero

    Absolute zero is a temperature marked by a 0 entropy configuration. It is the coldest temperature theoretically possible, and cannot be reached, by artificial or natural means....
     is -273.15 °C).
  • Pressure
    Pressure

    Pressure is the force per unit area applied to an object in a direction surface normal to the surface. Gauge pressure is the pressure relative to the local atmospheric or ambient pressure....
     — they can withstand the extremely low pressure of a vacuum
    Vacuum

    A vacuum is a volume of space that is essentially empty of matter, such that its gaseous pressure is much less than atmospheric pressure. The word comes from the Latin term for "empty," but in reality, no volume of space can ever be perfectly empty....
     and also very high pressures, more than 1200 times atmospheric pressure
    Atmospheric pressure

    Atmospheric pressure is sometimes defined as the force per unit area exerted against a surface by the weight of air above that surface at any given point in the Earth's atmosphere....
    . It has recently been demonstrated that tardigrades can survive the vacuum of open space and solar radiation combined for at least 10 days. Recent research has notched up another feat of endurance: they can withstand 6,000 atmospheres, which is nearly six times the pressure of water in the deepest ocean trench.
  • Dehydration
    Dehydration

    Dehydration is the removal of water from an object. In Physiology terms, it entails a relative deficiency of water molecules in relation to other dissolved solutes....
     — tardigrades have been shown to survive nearly one decade in a dry state.
  • Radiation
    Radiation

    In physics, radiation describes any process in which energy emitted by one body travels through a medium or through space, ultimately to be absorbed by another body....
     — as shown by Raul M. May from the University of Paris, tardigrades can withstand 5,700 gray
    Gray (unit)

    The gray is the SI unit of absorbed dose due to ionizing radiation ....
    s or 570,000 rad
    Rad (unit)

    The rad is a unit of absorbed radiation dose, with symbol rad. The rad was first proposed in 1918 as "that quantity of X rays which when absorbed will cause the destruction of the [malignant mammalian] cells in question..." It was defined in Centimetre gram second system of units in 1953 as the dose causing 100 ergs of energy to be absorb...
    s of x-ray radiation. (Ten to twenty grays or 1,000–2,000 rads could be fatal to a human). The only explanation thus far for this ability is that their lowered water state provides fewer reactants for the ionizing radiation
    Ionizing radiation

    Ionizing radiation consists of subatomic particle radiation or electromagnetic radiation that are energetic enough to detach electrons from atoms or molecules, ionize them....
    .


Recent experiments conducted by Cai and Zabder have also shown that these tardigrades can undergo chemobiosis — a cryptobiotic
Cryptobiosis

Cryptobiosis is an ametabolic state of life entered by an organism in response to adverse environmental conditions such as desiccation, freezing, and oxygen deficiency....
 response to high levels of environmental toxins. However, their results have yet to be verified. In September 2008, a space launch showed that tardigrades can survive the extreme environment of outer space
Outer space

Outer space comprises the relatively empty regions of the universe outside the atmospheres of celestial bodies. Outer space is used to distinguish it from airspace and terrestrial locations....
 for 10 days. After being rehydrated back on earth, over 68% of the subjects protected from high-energy UV radiation survived and many of these produced viable embryos, and a handful survived full exposure to the sun.

Evolutionary relationships and history

Recent DNA
DNA

Deoxyribonucleic acid is a nucleic acid that contains the genetics instructions used in the development and functioning of all known living organisms and some viruses....
 and RNA
RNA

Ribonucleic acid is a type of molecule that consists of a long chain of nucleotide units. Each nucleotide consists of a nucleobase, a ribose sugar, and a phosphate....
 sequencing data indicate that tardigrades are the sister group to the arthropod
Arthropod

Arthropods are animals belonging to the Scientific classification Arthropoda , and include the insects, arachnids, crustaceans, and others....
s and Onychophora
Velvet worm

The velvet worms form a clade within the Ecdysozoa and can be simply described as "worms with legs". Most common in the Southern Hemisphere, they prey on smaller animals such as insects, which they catch by squirting a sticky slime....
. These groups have been traditionally thought of as close relatives of the annelid
Annelid

The annelids, collectively called Annelida , are a large Scientific classification of animals comprising the segmented worms, with about 15,000 modern species including the well-known earthworms and leeches....
s, but newer schemes consider them Ecdysozoa
Ecdysozoa

The Ecdysozoa are a grouping of protostome animals, including the Arthropoda , roundworm, and several smaller phylum . They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes....
, together with the roundworms (Nematoda) and several smaller phyla. The Ecdysozoa
Ecdysozoa

The Ecdysozoa are a grouping of protostome animals, including the Arthropoda , roundworm, and several smaller phylum . They were first defined by Aguinaldo et al. in 1997, based mainly on trees constructed using 18S ribosomal RNA genes....
-concept resolves the problem of the nematode-like pharynx
Pharynx

FunctionsThe pharynx is part of the digestive system and respiratory system of many organisms.Because both food and Earth's atmosphere pass through the pharynx, a flap of connective tissue called the epiglottis closes over the trachea when food is swallowed to prevent choking or Pulmonary aspiration....
 as well as some data from 18S-rRNA and HOX (homeobox
Homeobox

A homeobox is a DNA sequence found within genes that are involved in the regulation of patterns of development in animals, fungus and plants. Genes that have a homeobox are called homeobox genes and form the homeobox gene family....
) gene data, which indicate a relation to roundworms.

The minute sizes of tardigrades and their membranous integuments make their fossilization both difficult to detect and highly unlikely. The only known fossil specimens comprise some from mid-Cambrian
Cambrian

The Cambrian is a geologic period that began about Mya at the end of the Proterozoic eon and ended about Ma with the beginning of the Ordovician period ....
 deposits in Siberia
Siberia

Siberia , is the name given to the vast region constituting almost all of North Asia and for the most part currently serving as the massive central and eastern portion of the Russian Federation, having served in the same capacity previously for the Soviet Union from its beginning, and the Russian Empire beginning in the 16th century....
 and a few rare specimens from Cretaceous
Cretaceous

The Cretaceous , usually abbreviated K for its German translation Kreide, is a geologic period from circa to million years ago . In the geologic timescale, the Cretaceous follows on the Jurassic period and is followed by the Paleogene period....
 amber
Amber

Amber is fossil tree resin, which is appreciated for its color and beauty. Good quality amber is used for the manufacture of ornamental objects and jewelry....
.

The Siberian tardigrades differ from living tardigrades in several ways. They have three pairs of legs rather than four; they have a simplified head morphology; and they have no posterior head appendages. It is considered that they probably represent a stem group of living tardigrades.

The rare specimens in Cretaceous amber comprise Milnesium swolenskyi, from New Jersey
New Jersey

New Jersey is a state in the Mid-Atlantic States and Northeastern United States regions of the United States. It is bordered on the north by New York, on the east by the Hudson River and the Atlantic Ocean, on the southwest by Delaware, and on the west by Pennsylvania....
, the oldest, whose claws and mouthparts are indistinguishable from the living M. tartigradum; and two specimens from western Canada
Canada

Canada is a country occupying most of northern North America, extending from the Atlantic Ocean in the east to the Pacific Ocean in the west and northward into the Arctic Ocean....
, some 15–20 million years younger than M. swolenskyi. Of the two latter, one has been given its own genus and family, Beorn leggi (the genus named by Cooper after the character Beorn
Beorn

In J. R. R. Tolkien's Middle-earth Tolkien's legendarium, Beorn was a shapeshifting , a man who could assume the appearance of a great black bear....
 from The Hobbit
The Hobbit

The Hobbit, or There and Back Again is an award-winning Juvenile fantasy and children's book by J. R. R. Tolkien, written in the tradition of the fairy tale....
 by J. R. R. Tolkien
J. R. R. Tolkien

John Ronald Reuel Tolkien, Order of the British Empire was an English people English literature, poetry, Philology, and university professor, best known as the author of the classic high fantasy works The Hobbit, The Lord of the Rings and The Silmarillion....
 and the species named after his student William M. Legg); however, it bears a strong resemblance to many living specimens in the family Hipsiblidae.

Aysheaia
Aysheaia

Aysheaia was a genus of soft-bodied, caterpillar-shaped organisms average body length of 1?6 cm. The genus name commemorates a mountain peak named "Ashea" due north of the Waptia Glacier....
 from the middle Cambrian
Cambrian

The Cambrian is a geologic period that began about Mya at the end of the Proterozoic eon and ended about Ma with the beginning of the Ordovician period ....
 Burgess shale
Burgess Shale

The Burgess Shale Formation is one of the world's most celebrated fossil localities, and is famous for the exceptional preservation of the fossils found within it, in which the soft parts are preserved....
 has been proposed as a sister-taxon to an arthropod-tardigrade clade.

Genomes and genome sequencing

Tardigrade genomes vary in size, from about 75 to 800 megabase pairs of DNA. The genome of a tardigrade species, Hypsibius dujardini, is being sequenced. Hypsibius dujardini has a compact genome and a generation time of about two weeks, and it can be cultured indefinitely and cryopreserved.

External links