Tardigrade

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Tardigrade
Fossil range: Early Cambrian to Recent[1]
The tardigrade Hypsibius dujardini
The tardigrade Hypsibius dujardini
Scientific classification
Kingdom: Animalia
Subkingdom: Ecdysozoa
Phylum: Tardigrada
Spallanzani, 1777
Classes [2]

Heterotardigrada
Mesotardigrada
Eutardigrada

Tardigrades (commonly known as water bears) form the phylum Tardigrada, part of the superphylum Ecdysozoa. They are microscopic, water-dwelling, segmented animals 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 in 1777. The name water bear comes from the way they walk, because it walks the same way. The biggest adults may reach a body length of 1.5 mm, the smallest below 0.1 mm. Freshly hatched larvae 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 (above 6,000 m), to the deep sea (below 4,000 m) and from the polar regions to the equator.

The most convenient place to find tardigrades is on lichens and mosses. Other environments are dunes, beaches, soil and marine or freshwater sediments, where they may occur quite frequently (up to 25,000 animals per litre). Tardigrades often can be found by soaking a piece of moss in spring water.[3]

Tardigrades are polyextremophiles and are able to survive in extreme environments that would kill almost any other animal. Some can survive temperatures close to absolute zero[4], temperatures as high as 151 °C (303 °F), 1,000 times more radiation than any other animal[5], nearly a decade without water, and even the vacuum of space.[6]

Contents

[edit] Anatomy and morphology

Tardigrades have a body with four segments (not counting the head), four pairs of legs without joints, and feet with claws or toes. The cuticle contains chitin and is moulted. They have a ventral nervous system with one ganglion per segment, and a multilobed brain. Instead of a coelom they have a haemocoel. The only place where a true coelom can be found is around the gonad (coelomic pouch). The pharynx is of a triradiate, muscular, sucking kind, armed with stylets. Although some species are parthenogenetic, males and females are usually present, each with a single gonad. 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. [7][8]

[edit] Ecology and life history

[edit] Feeding ecology

Most tardigrades are phytophagous (plant eaters) or bacteriophagous (bacteria eaters), but some are predatory[9] (e.g. Milnesium tardigradum).[10]

[edit] Physiology

[edit] Extreme environments

Tardigrades are polyextremophiles; scientists have reported their existence in hot springs, on top of the Himalayas, under layers of solid ice and in ocean sediments. Many species can be found in a milder environment like lakes, ponds and meadows, 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.

Modern tardigrades

Tardigrades are one of the few groups of species that are capable of reversibly suspending their metabolism and going into a state of cryptobiosis. 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 trehalose, which protects their membranes.

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

  • 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 is −273.15 °C).[11]
  • Pressure — they can withstand the extremely low pressure of a vacuum and also very high pressures, more than 1200 times atmospheric pressure. It has recently been demonstrated that tardigrades can survive the vacuum of open space and solar radiation combined for at least 10 days.[12] 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. [13]
  • Dehydration — tardigrades have been shown to survive nearly one decade in a dry state.[14] Another researcher reported that a tardigrade survived over a period of 120 years in a dehydrated state, but soon died after 2 to 3 minutes.[15] Subsequent research has cast doubt on its accuracy since only small movements in the water bear's leg were detected.[16]
  • Radiation — as shown by Raul M. May from the University of Paris, tardigrades can withstand 5,700 grays or 570,000 rads 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 hydration state provides fewer reactants for the ionizing radiation.

Recent experiments conducted by Cai and Zabder have also shown that these tardigrades can undergo chemobiosis — a cryptobiotic response to high levels of environmental toxins. However, their results have yet to be verified.[17][18] In September 2008, a space launch showed that tardigrades can survive the extreme environment of outer space 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.[19]

[edit] Evolutionary relationships and history

Recent DNA and RNA sequencing data[citation needed] indicate that tardigrades are the sister group to the arthropods and Onychophora. These groups have been traditionally thought[citation needed] of as close relatives of the annelids, but newer schemes consider them Ecdysozoa, together with the roundworms (Nematoda) and several smaller phyla. The Ecdysozoa-concept resolves the problem of the nematode-like pharynx as well as some data from 18S-rRNA and HOX (homeobox) 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 deposits in Siberia and a few rare specimens from Cretaceous amber.[20]

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.[20]

The rare specimens in Cretaceous amber comprise Milnesium swolenskyi, from New Jersey, the oldest, whose claws and mouthparts are indistinguishable from the living M. tartigradum; and two specimens from western Canada, 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 from The Hobbit by J. R. R. Tolkien and the species named after his student William M. Legg); however, it bears a strong resemblance to many living specimens in the family Hipsiblidae.[20][21]

Aysheaia from the middle Cambrian Burgess shale might be related to tardigrades.

[edit] Genomes and genome sequencing

Tardigrade genomes vary in size, from about 75 to 800 megabase pairs of DNA. [22] 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. [23]

[edit] References

  1. ^ Budd, G.E. (2001). "Tardigrades as ‘stem-group arthropods’: the evidence from the Cambrian fauna". Zool. Anz 240: 265–279. doi:10.1078/0044-5231-00034. 
  2. ^ Tardigrada (TSN 155166). Integrated Taxonomic Information System.
  3. ^ Goldstein, B. and Blaxter, M. (2002). "Quick Guide: Tardigrades". Current Biology 12: R475. doi:10.1016/S0960-9822(02)00959-4. 
  4. ^ Bertolani, R. et al (2004). "Experiences with dormancy in tardigrades". Journal of Limnology 63(Suppl 1): 16–25. 
  5. ^ Radiation tolerance in the tardigrade Milnesium tardigradum
  6. ^ Rachel Courtland (2008-09-08). "'Water bears' are first animal to survive space vacuum".
  7. ^ Seki, K & Toyoshima, M. (1998). Preserving tardigrades under pressure. Nature 395: 853–854.
  8. ^ Ian M. Kinchin (1994) The Biology of Tardigrades, Ashgate Publishing
  9. ^ Lindahl, K. (2008-03-15). "Tardigrade Facts".
  10. ^ Morgan, Clive I. (1977). "Population Dynamics of two Species of Tardigrada, Macrobiotus hufelandii (Schultze) and Echiniscus (Echiniscus) testudo (Doyere), in Roof Moss from Swansea". The Journal of Animal Ecology 46 (1): 263–279. doi:10.2307/3960. 
  11. ^ Ramel, G. (2005-11-11). "The Water Bears (Phylum Tardigrada)".
  12. ^ K. Ingemar Jönsson, Elke Rabbow, Ralph O. Schill, Mats Harms-Ringdahl and Petra Rettberg (2008). "Tardigrades survive exposure to space in low Earth orbit". Current Biology 18 (17): R729-R731. doi:10.1016/j.cub.2008.06.048. http://space.newscientist.com/article/dn14690-water-bears-are-first-animal-to-survive-in-space.html. 
  13. ^ Seki, K & Toyoshima, M. (1998). "Preserving tardigrades under pressure". Nature 395: 853–854. doi:10.1038/27576. 
  14. ^ Guidetti, R. & Jönsson, K.I. (2002). "Long-term anhydrobiotic survival in semi-terrestrial micrometazoans". Journal of Zoology 257: 181–187. doi:10.1017/S095283690200078X. 
  15. ^ Manga Science Volume VI by Yoshitoh Asari, ISBN-05-202039-1, 1998
  16. ^ Guidetti, R. & Jönsson, K.I. (2002). "Long-term anhydrobiotic survival in semi-terrestrial micrometazoans". Journal of Zoology 257: 181. doi:10.1017/S095283690200078X. 
  17. ^ Franceschi, T. (1948). "Anabiosi nei tardigradi". Bolletino dei Musei e degli Istituti Biologici dell'Università di Genova 22: 47–49. 
  18. ^ Jönsson, K. I. & R. Bertolani (2001). "Facts and fiction about long-term survival in tardigrades". Journal of Zoology 255: 121–123. doi:10.1017/S0952836901001169. 
  19. ^ 'Water bears' can survive in the vacuum of space
  20. ^ a b c David A. Grimaldi and Michael S. Engel (2005). Evolution of the Insects. Cambridge University Press. pp. 96–97. ISBN 0521821495. 
  21. ^ Kenneth W. Cooper (1964). "The first fossil tardigrade: Beorn leggi, from Cretaceous Amber". Psyche – Journal of Entomology 71 (2): 41. 
  22. ^ [1]
  23. ^ Gabriel, W et al. (2007). "The tardigrade Hypsibius dujardini, a new model for studying the evolution of development". Developmental Biology 312: 545–559. doi:10.1016/j.ydbio.2007.09.055. 

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