Edge Rewrite
everything Source code View original
// HTMLRewriter · presentation

This page was redesigned at the edge.

Cloudflare fetched the original article and streamed it through HTMLRewriter to apply an entirely new visual system without rebuilding the source page.

// request.cf · coarse context

A page that knows where it met you.

Only coarse request metadata is shown. This demo does not display or persist visitor IP addresses.

Country
US
Cloudflare location
CMH
Connection
HTTP/2
Language
Not provided

Ray ID: a23540b99f075751

Jump to content

Riftia

From Wikipedia, the free encyclopedia

Giant tubeworm
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Annelida
Clade: Pleistoannelida
Clade: Sedentaria
Order: Sabellida
Family: Siboglinidae
Genus: Riftia
M. L. Jones, 1981
Species:
R. pachyptila
Binomial name
Riftia pachyptila
M. L. Jones, 1981

Riftia is a monotypic genus of polychaete annelid worms in the Siboglinidae family,[1] which contains the sole species Riftia pachyptila, commonly known as the giant tubeworm or giant beardworm.

R. pachyptila is found in association with geologically active regions of the deep sea. Riftia worms are sessile and are found clustered together in large colonies around deep-sea hydrothermal vents in the East Pacific Rise and the Galapagos Rift.[2] The size of a patch of tube worms surrounding a vent can span tens of metres, and individuals can reach a length of 3 m (9 ft 10 in).[3][4] R. pachyptila has the fastest growth rate of any known marine invertebrate, and have been known to colonize a new site, grow to sexual maturity, and increase in length to 4.9 ft (1.5 m) in less than two years.[5]

Adult Riftia worms lack a digestive system and rely on their symbiotic relationship with chemotrophic sulfur-oxidising bacteria to provide them with energy. The hydrothermal vent habitat in which Riftia lives provides a natural ambient temperature ranging from 2–30 degrees Celsius (36–86 °F)[6] and emits large amounts of chemicals such as hydrogen sulfide that are utilised by the tube worm for bacterial chemosynthesis.

Historically, the genus Riftia was placed within the obsolete phyla Pogonophora. They are now understood to be annelids, with their closest relatives being other deep sea tube worms such as Escarpia and Lamellibrachia.

Description

[edit]

The generic name Riftia alludes to the rift that formed the geothermal vents where the species inhabits, while pachyptila (pachy; thick + ptilon; feather) refers to the anterior plume of the worm.[7] The holotype, USNM 59951, is held by the National Museum of Natural History (USNM).[8]

Anatomy

[edit]

Like other siboglinade tube worms, R. pachyptila has a vermiform body that secretes a chitinous tube for protection and support.[9] Riftia worms can reach a length of 3 m (9 ft 10 in),[3] and their bodies have a diameter of 4 cm (1.6 in).

R. pachyptila community with red branchial plumes visible.

Riftia have a red feather-like structure called a branchial plume at the anterior end which is used to acquire chemicals for chemosynthesis. The plume is highly vascularised, and its red colour is due to the presence of hemoglobin. The structure of the plume maximises the surface area needed to absorb chemicals from the water efficently. If the tubeworm perceives a threat or is touched, it retracts the plume into its tube and seals the tube with its obturaculum, an operculum made of collegen.[10] The respiratory lamellae are supported by the obturaculum.[11]

Beneath the plume is the vestimentum, a muscular body region that contains the heart and the two genital openings.[12][13][14]

Anatomy of Riftia pachyptila, including microstructure and metabolic process

The middle region of the body houses the coelomic cavity, gonads, and the trophosome, a spongy tissue where symbiotic bacteria and sulfur granules are found.[15][16] Adult Riftia have no mouth, digestive system or anus as they do not eat. Instead, they rely on bacteria in their trophosome to provide them with nutrition.[17]

The posterior part of the worm is the opisthosome, which anchors the animal to the tube and is used for the storage of waste from bacterial reactions.[18]

Tubes

[edit]
Riftia pachyptila being extracted out of its tube

Riftia's tubes are composed of chitin,[19] secreted out from cup-microvilli-like structures within glands which form crystallite chitin layers over time.[20] The tube is cylindrical, flexible, and closed at the prosterier end.[21] Riftia tubes can reach 2 m in length and 5 cm in diameter and are very thick compared to those of other tubeworms, especially at the base.[22][6] These tubes are very resistant to enzymatic attack by bacteria, and can take years to decompose.[23] The worms are able to adjust both the top and the base of their tubes, which allows for some adaptability in the highly competitive and crowded spaces they grow in- a worm may need to adjust its position to maximise its access to vent fluid as more worms settle in the colony.[24] Riftia tubes grow at a rate of 10 to 85 cm (3.9 to 33.5 in) per year.[25][26]

Physiology

[edit]

Metabolism

[edit]

R. pachyptila differs significantly in its physiology from other deep-sea species that do not inhabit hydrothermal vent sites. While deep sea species typically have low metabolic rates, the enzyme activity relating to glycolysis, the citric acid cycle and electron transport in the tissues of R. pachyptila is similar to that of shallow-living animals, suggesting a significantly faster metabolism than is typical of species found in the deep sea.[27]

Hydrothermal vent tubeworms get organic compounds from bacteria that live in their trophosome

Relationship with endosymbiotic bacteria

[edit]

In its adult phase, R. pachyptila lacks a digestive system and cannot eat. Rather than consuming organic matter, Riftia gains nutrients from the environment by utilising a process known as chemoautotrophic symbiosis.

In this process, nutrition is gained via sulfur-oxidizing endosymbiotic bacteria that live in a specialised organ inside the worm called the trophosome, a soft tissue that runs almost the whole length of the tube worm's body cavity.[17] These bacteria are able to convert common chemicals in the hydrothermal vent environment into carbohydrates that can be consumed by the tube worm.[16][28][29] These chemicals typically include carbon, nitrogen, oxygen, and sulfur, which are transported by the tube worm into its trophosome to provide the bacteria with nutrition.

Riftia hemoglobins are atypical in that they are able to carry oxygen in the presence of sulfide without being inhibited by it.[30][31] Hemoglobins in the branchial plume are able to bind O2 and H2S, which are necessary for bacterial chemosynthesis. Capillaries in the branchial plume move these compounds to the trophosome, where they can be absorbed by bacteria.[32] Bacteria in the trophosome live inside specialised cells (bacteriocytes) and have no contact with the external environment, relying entirely on the tube worm for nutrients as well as for the removal of its waste products (carbohydrates).

Life history

[edit]
Riftia tubes covered in epibionts
Alongside mussels and sea anemones

Ecology

[edit]

Riftia is able to coexist with other large sessile animals through niche partitioning.[33] Riftia requires areas of high water flow, temperature, and sulfide concentrations, in contrast to Calyptogena magnifica's need of low flow and temperature, and Bathymodiolus thermophilus' flexibility; B. thermophilus mussels may grow in places ranging from the periphery of the vent field, to directly on Riftia's tubes.[33]

It is suggested that the smaller-bodied tubeworm Tevnia may facilitate later vent colonization by Riftia.[34][35]

Reproduction

[edit]

R. pachyptila is dioecious.[36][37] The male's spermatozoa are thread-shaped and are about 130 μm long overall. The sperm is arranged into an agglomeration of around 340–350 individual spermatozoa that form a torch-like shape, referred to as spermatozeugmata. [citation needed]

The ovaries of females run within the gonocoel along the entire length of the trunk, ventral to the trophosome. Eggs at different maturation stages can be found in the middle area of the ovaries, and depending on their developmental stage, are referred to as: oogonia, oocytes, and follicular cells. When the oocytes mature, they acquire protein and lipid yolk granules.[citation needed]

Males release their sperm into seawater. Usually, the spermatozeugmata swim into the female's tube. Movement of the cluster is conferred by the collective action of each spermatozoon moving independently. Reproduction has also been observed involving only a single spermatozoon reaching the female's tube. [citation needed]

Larval stage

[edit]

R. pachyptila develops from a free-swimming, pelagic, non-symbiotic trochophore larva, which enters juvenile (metatrochophore) development, becoming sessile, and subsequently acquiring symbiotic bacteria.[38][39] R. pachyptila larvae are planktonic and move through sea-bottom currents until they reach active hydrothermal vent sites.

The digestive tract is present in the larval stage and consists of a mouth foregut, midgut, hindgut, and anus. After symbionts are established in the midgut, they undergo substantial remodeling and enlargement to become the trophosome, while the remainder of the digestive tract has not been detected in adult specimens.[40]

R. pachyptila is dependent on volcanic gases and the presence of sulfide-oxidizing bacteria. It acquires these from hydrothermal vent sites that are patchy and ephemeral. The distance between active sites along a rift can be hundreds of km.[2] R. pachytpila is capable of larval dispersal across distances of 100 to 200 km[2] and cultured larvae have been shown to be viable for 38 days.[41][42] Though dispersal is considered to be effective, the genetic variability observed in R. pachyptila metapopulation is low compared to other vent species. This may be due to high extinction events and colonization events, as R. pachyptila is one of the first species to colonize a new active site.[2]

Symbiont acquisition

[edit]

The symbiotic bacteria, on which adult worms depend for sustenance, are not present in the larvae. They are acquired from the environment through the skin in a process similar to an infection.

Free-living bacteria found in the water column are ingested randomly and enter the worm through a ciliated opening of the branchial plume. This opening is connected to the trophosome through a duct that passes through the brain. Once the bacteria are in the gut, the ones that are beneficial to the individual, are phaghocytized by epithelial cells found in the midgut are then retained. Bacteria that do not represent possible endosymbionts are digested.[40]

Lifespan

[edit]

Some sources list Riftia pachyptila as one of the world's longest living organisms,[43] but the unstable vent environment and high ecological turnover makes this claim unlikely.[44] Tube worms inhabiting stable cold seep environments such as Lamellibranchia and Escarpia have better claims to longevity, with age measurements of both species exceeding a century.[45][46][47][48]

Hydrogen sulfide chemosynthesis

[edit]
Diagram of Photosynthesis
Sunlight is necessary for the process of photosynthesis.

Background

[edit]

Most ecosystems on earth rely on photosynthetic organisms, such as plants to form the base of their food web. Photosynthetic organisms use light energy from sunlight to convert carbon dioxide and water into carbohydrates and oxygen. The carbohydrates produced provide energy to the organisms, as well as to other organisms higher up in the food chain that consume them.

Chemosynthesis
A chart depicting Riftia chemosynthesis: 1: Hydrothermal Vents 2: Hydrogen Sulfide 3: Oxygen 4: Sulfur-oxidising bacteria 5: Tubeworms 6: Energy 7: Carbohydrates

However, in deep sea environments no sunlight is able to reach the sea floor, and photosynthesis becomes impossible. Hydrothermal vent ecosystems are instead based on chemosynthetic microbes such as sulfur-oxidising bacteria. These microbes are able to generate carbohydrates utilising the energy released from chemical reactions in the same way that photosynthetic organisms utilise light energy. The carbohydrates produced by chemosynthetic microbes can then be consumed by other organisms, forming the basis of the hydrothermal vent food chain.

While there is no one chemical pathway that is utilised for chemosynthesis, different kinds of chemotropic organisms specialise in consuming particular chemicals. The bacteria that live in symbiosis with Riftia are sulfur-oxidising chemotrophs, meaning that they utilise sulfides in their chemosynthesis. This form of chemosynthesis is referred to as thioautotrophy.[18][better source needed]

Bacterial symbiosis

[edit]

Riftia worms live in symbiosis with the sulfur-oxidising bacteria that live inside their trophosome. The tube worm provides the bacteria with the chemicals required for chemosynthesis, and the bacteria produce carbohydrates that can be consumed by the tube worm. Because the tube worm has no digestive system, it is entirely reliant on carbohydrates produced by their symbiotic bacteria.[49]

The host tubeworm intakes the chemicals required for thioautotrophy from the mineral-rich vent environment and transports them to the bacteria within it's trophosome, which houses a large quantity of bacteria (approximately 109 bacteria per gram of fresh weight).

Riftia worms take in hydrogen sulfide, carbon dioxide, and oxygen from the mineral-rich vent environment through their plume. These chemicals pass into the tube worm's blood, where they are bound by hemoglobin and transported to the trophosome. The bacterial symbionts within the trophosome utilise these chemicals to perform thioautotrophic chemosynthesis.[16] The simplified chemical equation for this process is as follows:

CO2carbon
dioxide + 4H2Shydrogen
sulfide +O2oxygen→CH2Ocarbohydrate+ 4Ssulfur + 3H2Owater[49]

In order to support their metabolism, R. pachyptila has to absorb all the substances necessary for both sulfide-oxidation and carbon fixation as well as other fundamental bacterial nutrients such as nitrogen and phosphorus. This means that the tubeworm must be able to access both oxic and anoxic areas.

Oxidation of reduced sulfur compounds requires the presence of oxidized reagents such as oxygen and nitrate. Hydrothermal vents are characterized by conditions of high hypoxia. In hypoxic conditions, sulfur-storing organisms start producing hydrogen sulfide. Therefore, the production of in H2S in anaerobic conditions is common among thiotrophic symbiosis. H2S can be damaging for some physiological processes as it inhibits the activity of cytochrome c oxidase, consequentially impairing oxidative phosphorylation. In R. pachyptila the production of hydrogen sulfide starts after 24h of hypoxia. To avoid physiological damage some animals, including Riftia pachyptila are able to bind H2S to haemoglobin in the blood to eventually expel it in the surrounding environment.

Nitrate and nitrite are toxic, but are required for biosynthetic processes. The chemosynthetic bacteria within the trophosome convert nitrate to ammonium ions, which then are available for production of amino acids in the bacteria, which are in turn released to the tube worm. To transport nitrate to the bacteria, R. pachyptila concentrates nitrate in its blood, to a concentration 100 times more concentrated than the surrounding water. The exact mechanism of R. pachyptila's ability to withstand and concentrate nitrate is still unknown.[31]

Carbon fixation and organic carbon assimilation

[edit]

R. pachyptila must uptake CO2. for its metabolism.[50] CO2 uptake in the worm is enhanced by the higher pH of its blood (7.3–7.4), which favors the bicarbonate ion and thus promotes a steep gradient across which CO2 diffuses into the vascular blood of the plume.[51][16]

Once the CO2 is fixed by the symbionts, it can be consumed by the host tissues. The fixed carbon to is transported via organic molecules from the trophosome in the hemolymph.[29][52]

Studies proved that within 15 min, the label first appears in symbiont-free host tissues, and that indicates a significant amount of release of organic carbon immediately after fixation. After 24 h, labeled carbon is clearly evident in the epidermal tissues of the body wall. Results of the pulse-chase autoradiographic experiments were also evident with ultrastructural evidence for digestion of symbionts in the peripheral regions of the trophosome lobules.[52][53]

Sulfide acquisition

[edit]

In deep-sea hydrothermal vents, sulfide and oxygen are present in different areas. The reducing fluid of hydrothermal vents is rich in sulfide, but poor in oxygen, whereas seawater is richer in dissolved oxygen. Sulfide is immediately oxidized by dissolved oxygen to form partly, or totally, oxidized sulfur compounds like thiosulfate (S2O32-) and ultimately sulfate (SO42-), respectively less, or no longer, usable for microbial oxidation metabolism.[54] This causes the substrates to be less available for microbial activity, thus bacteria are constricted to compete with oxygen to get their nutrients.

To avoid this issue, several microbes have evolved to make symbiosis with eukaryotic hosts.[55][16] In fact, R. pachyptila is able to cover the oxic and anoxic areas to get both sulfide and oxygen[56][57][58] thanks to its hemoglobin that can bind sulfide reversibly and apart from oxygen by functional binding sites determined to be zinc ions embedded in the A2 chains of the hemoglobins.[59][60][61] and then transport it to the trophosome, where bacterial metabolism can occur. It has also been suggested that cysteine residues are involved in this process.[62][63][64]

During the chemosynthesis, the mitochondrial enzyme rhodanase catalyzes the disproportionation reaction of the thiosulfate anion S2O32- to sulfur S and sulfite SO32- .[65][66] The R. pachyptila's bloodstream is responsible for absorption of the O2 and nutrients such as carbohydrates.

Endosymbiont species
[edit]

A wide range of bacterial diversity is associated with symbiotic relationships with R. pachyptila. Many of these bacteria belong to the phylum Campylobacterota[67] as supported by the recent discovery in 2016 of the new species Sulfurovum riftiae collected from the East Pacific Rise.[68] Other symbionts belong to the classes Delta-, Alpha- and Gammaproteobacteria.[67]

The Candidatus Endoriftia persephone (Gammaproteobacteria) is a facultative R. pachyptila symbiont and has been shown to be a mixotroph, thereby exploiting both Calvin Benson cycle and reverse TCA cycle (with an unusual ATP citrate lyase) according to availability of carbon resources and whether it is free living in the environment or inside a eukaryotic host. The bacteria apparently prefer a heterotrophic lifestyle when carbon sources are available.[69]

Evidence based on 16S rRNA analysis affirms that R. pachyptila chemoautotrophic bacteria belong to two different clades: Gammaproteobacteria[70][16] and Campylobacterota (e.g. Sulfurovum riftiae)[68] that get energy from the oxidation of inorganic sulfur compounds such as hydrogen sulfide (H2S, HS−, S2-) to synthesize ATP for carbon fixation via the Calvin cycle.[16] Unfortunately, most of these bacteria are still uncultivable. Symbiosis works so that R. pachyptila provides nutrients such as HS−, O2, CO2 to bacteria, and in turn it receives organic matter from them. Thus, because of a lack of a digestive system, R. pachyptila depends entirely on its bacterial symbiont to survive.[71][72]

History & Discovery

[edit]
DSV Alvin, the US Navy research submarine used for the expedition.

R. pachyptila was first encountered in 1977 during a geological expedition by Jack Corliss to the Galápagos Rift. The initial purpose of the expedition was to prove the existence of hot springs (as hydrothermal vents) on the sea floor.[8] These vents were assumed to be devoid of life, as the heat emitted from such vents averages around 350–380 °C (662–716 °F)).[73][74]

However, the expedition observed a multitude of species found exclusively near hydrothermal vents, including Riftia, crabs, Bathymodiolus thermophilus and Calyptogena magnifica.[75] The vent ecosystem at 00°48′15″N 86°13′29″W had such an abundance of giant tube worms that it was dubbed the "Rose garden" site, the "roses" referring to the red-plumed Riftia worms.[8] While initially thought to be an isolated phenomenon, many such ecosystems have since been discovered on geologically active sections of the sea floor.

An 1985 expedition to the Rose garden site found that the Riftia were fewer in number, having been displaced by clams and mussels.[33] Another expedition in 2002 found that it had been destroyed by a lava flow sometime in the prior decade, but a second vent ecosystem was found near the original site, dubbed "Rosebud".[8]

Initial evidence for a chemoautotrophic symbiosis in R. pachyptila came from microscopic and biochemical analyses showing Gram-negative bacteria packed within a highly vascularized organ in the tubeworm trunk.[17] Chemosynthesis was first recognised within the trophosome by Colleen Cavanaugh.[17]

See also

[edit]

References

[edit]
  1. ↑ Ruppert E, Fox R, Barnes R (2007). Invertebrate Zoology: A functional Evolutionary Approach (7th ed.). Belmont: Thomson Learning. ISBN 978-0-03-025982-1.
  2. 1 2 3 4 Coykendall DK, Johnson SB, Karl SA, Lutz RA, Vrijenhoek RC (April 2011). "Genetic diversity and demographic instability in Riftia pachyptila tubeworms from eastern Pacific hydrothermal vents". BMC Evolutionary Biology. 11 (1) 96. Bibcode:2011BMCEE..11...96C. doi:10.1186/1471-2148-11-96. PMC 3100261. PMID 21489281.
  3. 1 2 McClain, Craig R.; Balk, Meghan A.; Benfield, Mark C.; Branch, Trevor A.; Chen, Catherine; Cosgrove, James; Dove, Alistair D.M.; Gaskins, Lindsay C.; Helm, Rebecca R. (13 January 2015). "Sizing ocean giants: patterns of intraspecific size variation in marine megafauna". PeerJ. 3 e715. Bibcode:2015PeerJ...3.e715M. doi:10.7717/peerj.715. PMC 4304853. PMID 25649000.
  4. ↑ Cary SC, Felbeck H, Holland ND (1989). "Observations on the reproductive biology of the hydrothermal vent tube worm Riftia pachyptila". Marine Ecology Progress Series. 52: 89–94. Bibcode:1989MEPS...52...89C. doi:10.3354/meps052089.
  5. ↑ Lutz RA, Shank TM, Fornari DJ, Haymon RM, Lilley MD, Von Damm KL, Desbruyeres D (1994). "Rapid growth at deep-sea vents". Nature. 371 (6499): 663–664. Bibcode:1994Natur.371..663L. doi:10.1038/371663a0. S2CID 4357672.
  6. 1 2 Bright M, Lallier FH (2010). "The biology of vestimentiferan tubeworms". Oceanography and Marine Biology (PDF). Oceanography and Marine Biology - an Annual Review. Vol. 48. Taylor & Francis. pp. 213–266. doi:10.1201/ebk1439821169-c4 (inactive 16 July 2026). ISBN 978-1-4398-2116-9. Archived from the original (PDF) on 31 October 2013. Retrieved 30 October 2013.{{cite book}}: CS1 maint: DOI inactive as of July 2026 (link)
  7. ↑ Jones, Meredith L (1981). "Riftia Pachyptila, New Genus, New Species, The Vestimentiferan Worm From The Galapagos Rift Geothermal Vents (Pogonophora)". Proceedings of the Biological Society of Washington. 93: 1295–1313. Retrieved 13 August 2025.
  8. 1 2 3 4 "Giant Tube Worm: Riftia pachyptila". Smithsonian National Museum of Natural History. Archived from the original on 24 October 2018. Retrieved 25 October 2018.
  9. ↑ Hilário, Ana; Capa, María; Dahlgren, Thomas G.; Halanych, Kenneth M.; Little, Crispin T. S.; Thornhill, Daniel J.; Verna, Caroline; Glover, Adrian G. (14 February 2011). "New Perspectives on the Ecology and Evolution of Siboglinid Tubeworms". PLOS ONE. 6 (2) e16309. Bibcode:2011PLoSO...616309H. doi:10.1371/journal.pone.0016309. ISSN 1932-6203. PMC 3038861. PMID 21339826.
  10. ↑ Monaco A, Prouzet P (2 October 2015). Marine Ecosystems: Diversity and Functions. John Wiley & Sons. ISBN 978-1-119-23246-9.
  11. ↑ Andersen, Ann C.; Hamraoui, L.; Zaoui, D. (January 2001). "The obturaculum of Riftia pachyptila (Annelida, Vestimentifera): Ultrastructure and function of the obturacular muscles and extracellular matrix". Cahiers de Biologie Marine. 42 (3): 219–237. Retrieved 25 August 2025.
  12. ↑ Desbruyères D, Segonzac M (1997). Handbook of Deep-sea Hydrothermal Vent Fauna. Editions Quae. ISBN 978-2-905434-78-4.
  13. ↑ Gibson RN, Atkinson RJ, Gordon JD (12 May 2010). Oceanography and Marine Biology: An Annual Review. CRC Press. ISBN 978-1-4398-5925-4.
  14. ↑ Bartolomaeus T, Purschke G (30 March 2006). Morphology, Molecules, Evolution and Phylogeny in Polychaeta and Related Taxa. Dordrecht: Springer Science & Business Media. ISBN 978-1-4020-3240-0.
  15. ↑ "Hydrothermal vents Terza parte". www.biologiamarina.eu. Retrieved 10 December 2019.
  16. 1 2 3 4 5 6 7 Stewart FJ, Cavanaugh CM (2006). Overmann J (ed.). "Symbiosis of thioautotrophic bacteria with Riftia pachyptila". Progress in Molecular and Subcellular Biology. 41. Springer-Verlag: 197–225. doi:10.1007/3-540-28221-1_10. ISBN 978-3-540-28210-5. PMID 16623395.
  17. 1 2 3 4 Cavanaugh CM, Gardiner SL, Jones ML, Jannasch HW, Waterbury JB (July 1981). "Prokaryotic cells in the hydrothermal vent tube worm Riftia pachyptila Jones: possible chemoautotrophic symbionts". Science. 213 (4505): 340–342. Bibcode:1981Sci...213..340C. doi:10.1126/science.213.4505.340. PMID 17819907.
  18. 1 2 "Riftia pachyptila – Wikipedia – Symbiotic relationship between chemosynthetic bacteria and riftia tube worms". Archived from the original on 10 December 2019. Retrieved 10 December 2019.
  19. ↑ Gaill, F.; Hunt, S. (1986). "Tubes of deep sea hydrothermal vent worms Riftia pachyptila (Vestimentifera) and Alvinella pompejana (Annelida)". Marine Ecology Progress Series. 34: 267–274. Bibcode:1986MEPS...34..267G. doi:10.3354/meps034267.
  20. ↑ Ravaux, Juliette; Shillitto, Bruce; Gaill, Françoise; Gay, Lucien; Voss-Foucart, Marie-Françoise; Childress, James J. (1998). "Tube synthesis and growth processes in the hydrothermal vent tube-worm Riftia pachyptila" (PDF). Cahiers de Biologie Marine. 39: 325–326. doi:10.21411/CBM.A.6D61BBD1. Retrieved 13 August 2025.
  21. ↑ Bright, Monika; Lallie, François (12 May 2010), "The Biology of Vestimentiferan Tubeworms", in Gibson, R; Atkinson, R; Gordon, J (eds.), Oceanography and Marine Biology, vol. 20103650, CRC Press, pp. 213–265, doi:10.1201/ebk1439821169-c4 (inactive 16 July 2026), ISBN 978-1-4398-2116-9, retrieved 16 July 2026{{citation}}: CS1 maint: DOI inactive as of July 2026 (link)
  22. ↑ Ann C. Andersen; Sylvie Jolivet; Stéphanie Claudinot; François H Lallier (February 2011). "Biometry of the branchial plume in the hydrothermal vent tubeworm Riftia pachyptila (Vestimentifera; Annelida)". Canadian Journal of Zoology. 80 (2): 320–332. doi:10.1139/z02-005.
  23. ↑ Ravaux, J.; Zbinden, M.; Voss-Foucart, M. F.; Compère, P.; Goffinet, G.; Gaill, F. (10 May 2003). "Comparative degradation rates of chitinous exoskeletons from deep-sea environments". Marine Biology. 143 (2): 405–412. Bibcode:2003MarBi.143..405R. doi:10.1007/s00227-003-1086-8.
  24. ↑ Gaill, F.; Shillito, B.; Ménard, F.; Goffinet, G.; Childress, JJ (1997). "Rate and process of tube production by the deep-sea hydrothermal vent tubeworm Riftia pachyptila". Marine Ecology Progress Series. 148: 135–143. Bibcode:1997MEPS..148..135G. doi:10.3354/meps148135.
  25. ↑ Fustec, A.; Desbruyères, D.; Laubier, L. (1988). "Estimation de la biomasse des peuplements associés aux sources hydrothermales profondes de la dorsale du Pacifique oriental, à 13°N" (PDF). Oceanologica Acta. 8: 15–22.
  26. ↑ Lutz, Richard A.; Shank, Timothy M.; Fornari, Daniel J.; Haymon, Rachel M.; Lilley, Marvin D.; von Damm, Karen L.; Desbruyeres, Daniel (1994). "Rapid growth at deep-sea vents". Nature. 371 (6499): 663–664. Bibcode:1994Natur.371..663L. doi:10.1038/371663a0.
  27. ↑ Hand SC, Somero GN (August 1983). "Energy Metabolism Pathways of Hydrothermal Vent Animals: Adaptations to a Food-Rich and Sulfide-Rich Deep-Sea Environment". The Biological Bulletin. 165 (1): 167–181. Bibcode:1983BiolB.165..167H. doi:10.2307/1541362. JSTOR 1541362.
  28. ↑ Childress JJ, Lee RW, Sanders NK, Felbeck H, Oros DR, Toulmond A, Desbruyeres D, Kennicutt II MC, Brooks J (1993). "Inorganic carbon uptake in hydrothermal vent tubeworms facilitated by high environmental pCO2". Nature. 362 (6416): 147–149. Bibcode:1993Natur.362..147C. doi:10.1038/362147a0. S2CID 4341949.
  29. 1 2 Childress JJ, Arp AJ, Fisher CR (1984). "Metabolic and blood characteristics of the hydrothermal vent tube-worm Riftia pachyptila". Marine Biology. 83 (2): 109–124. Bibcode:1984MarBi..83..109C. doi:10.1007/bf00394718. S2CID 84806980.
  30. ↑ Zal F, Lallier FH, Green BN, Vinogradov SN, Toulmond A (April 1996). "The multi-hemoglobin system of the hydrothermal vent tube worm Riftia pachyptila. II. Complete polypeptide chain composition investigated by maximum entropy analysis of mass spectra". The Journal of Biological Chemistry. 271 (15): 8875–81. doi:10.1074/jbc.271.15.8875. PMID 8621529.
  31. 1 2 Hahlbeck E, Pospesel MA, Zal F, Childress JJ, Felbeck H (July 2005). "Proposed nitrate binding by hemoglobin in Riftia pachyptila" (Free full text). Deep-Sea Research Part I: Oceanographic Research Papers. 52 (10): 1885–1895. doi:10.1016/j.dsr.2004.12.011.
  32. ↑ Childress J, Fisher CR (1992). "The biology of hydrothermal vent animals: physiology, biochemistry, and autotrophic symbioses". Oceanography and Marine Biology: An Annual Review.
  33. 1 2 3 Childress JJ (October 1988). "Biology and chemistry of a deep-sea hydrothermal vent on the Galapagos Rift; the Rose Garden in 1985. Introduction". Deep Sea Research Part A. Oceanographic Research Papers. 35 (10–11): 1677–1680. Bibcode:1988DSRA...35.1677C. doi:10.1016/0198-0149(88)90043-X.
  34. ↑ Mullineaux, L. S.; Fisher, C. R.; Peterson, C. H.; Schaeffer, S. W. (2000). "Tubeworm succession at hydrothermal vents: Use of biogenic cues to reduce habitat selection error?". Oecologia. 123 (2): 275–284. Bibcode:2000Oecol.123..275M. doi:10.1007/s004420051014. PMID 28308732.
  35. ↑ Mullineaux, Lauren S.; Metaxas, Anna; Beaulieu, Stace E.; Bright, Monika; Gollner, Sabine; Grupe, Benjamin M.; Herrera, Santiago; Kellner, Julie B.; Levin, Lisa A.; Mitarai, Satoshi; Neubert, Michael G.; Thurnherr, Andreas M.; Tunnicliffe, Verena; Watanabe, Hiromi K.; Won, Yong-Jin (21 February 2018). "Exploring the Ecology of Deep-Sea Hydrothermal Vents in a Metacommunity Framework". Frontiers in Marine Science. 5 49. Bibcode:2018FrMaS...5...49M. doi:10.3389/fmars.2018.00049.
  36. ↑ Jones ML (July 1981). "Riftia pachyptila Jones: Observations on the Vestimentiferan Worm from the Galapagos Rift". Science. 213 (4505): 333–6. Bibcode:1981Sci...213..333J. doi:10.1126/science.213.4505.333. PMID 17819904.
  37. ↑ Karaseva NP, Rimskaya-Korsakova NN, Galkin SV, Malakhov VV (December 2016). "Taxonomy, geographical and bathymetric distribution of vestimentiferan tubeworms (Annelida, Siboglinidae)". Biology Bulletin. 43 (9): 937–969. Bibcode:2016BioBu..43..937K. doi:10.1134/S1062359016090132. S2CID 16005133.
  38. ↑ Bright M. "Riftia pachyptila". Archived from the original on 2 April 2015. Retrieved 19 March 2015.
  39. ↑ Adams DK, Arellano SM, Govenar B (March 2012). "Larval dispersal: Vent life in the ocean column" (PDF). Oceanography.
  40. 1 2 Jones ML, Gardiner SL (October 1989). "On the early development of the vestimentiferan tube worm Ridgeia sp. and observations on the nervous system and trophosome of Ridgeia sp. and Riftia pachyptila" (PDF). Biological Bulletin. 177 (2): 254–276. Bibcode:1989BiolB.177..254J. doi:10.2307/1541941. JSTOR 1541941. Archived from the original (PDF) on 23 September 2015. Retrieved 19 March 2015.
  41. ↑ Marsh AG, Mullineaux LS, Young CM, Manahan DT (May 2001). "Larval dispersal potential of the tubeworm Riftia pachyptila at deep-sea hydrothermal vents". Nature. 411 (6833): 77–80. Bibcode:2001Natur.411...77M. doi:10.1038/35075063. PMID 11333980. S2CID 411076.
  42. ↑ Abe, Shige. "Raising Baby Tubeworms". astrobiology.nasa.gov. NASA. Retrieved 13 August 2025.
  43. ↑ Murad (17 April 2023). "The World's Longest-Living Animals – Unraveling the Secrets of Longevity". sciquest.org. Retrieved 13 August 2025.
  44. ↑ "Giant Riftia Tubeworms – 2015". youtube.com. EVNautilus. 3 March 2023. Retrieved 14 August 2025.
  45. ↑ Cordes, Erik E.; Arthur, Michael A.; Shea, Katriona; Arvidson, Rolf S.; Fisher, Charles R. (22 February 2005). "Modeling the Mutualistic Interactions between Tubeworms and Microbial Consortia". PLOS Biology. 3 (3) e77. doi:10.1371/journal.pbio.0030077. PMC 1044833. PMID 15736979.
  46. ↑ Sharmishtha, D.; Miles, L. L.; Barnabei, M.S.; Fisher, C. R. (2006). "The hydrocarbon seep tubeworm Lamellibrachia luymesi primarily eliminates sulfate and hydrogen ions across its roots to conserve energy and ensure sulfide supply". Journal of Experimental Biology. 209 (19): 3795–3805. Bibcode:2006JExpB.209.3795D. doi:10.1242/jeb.02413. PMID 16985196. S2CID 45656093.
  47. ↑ Durkin, Alanna; Fisher, Charles R.; Cordes, Erik E. (8 July 2017). "Extreme longevity in a deep-sea vestimentiferan tubeworm and its implications for the evolution of life history strategies". The Science of Nature. 104 (7–8): 63. Bibcode:2017SciNa.104...63D. doi:10.1007/s00114-017-1479-z. PMID 28689349. S2CID 11287549.
  48. ↑ Gruber, Karl (20 July 2017). "Giant deep-sea worms may live to be 1000 years old or more". New Scientist.
  49. 1 2 "Chemosynthesis Fact Sheet". Deep Ocean Education Project. Retrieved 18 July 2026.
  50. ↑ Van Dover CL, Lutz RA (2004). "Experimental ecology at deep-sea hydrothermal vents: a perspective". Journal of Experimental Marine Biology and Ecology. 300 (1–2): 273–307. Bibcode:2004JEMBE.300..273V. doi:10.1016/j.jembe.2003.12.024.
  51. ↑ Goffredi SK, Childress JJ (2001). "Activity and inhibitor sensitivity of ATPases in the hydrothermal vent tubeworm Riftia pachyptila: a comparative approach". Marine Biology. 138 (2): 259–265. Bibcode:2001MarBi.138..259G. doi:10.1007/s002270000462. S2CID 83539580.
  52. 1 2 Bright M, Keckeis H, Fisher CR (19 May 2000). "An autoradiographic examination of carbon fixation, transfer and utilization in the Riftia pachyptila symbiosis". Marine Biology. 136 (4): 621–632. Bibcode:2000MarBi.136..621B. doi:10.1007/s002270050722. S2CID 84235858.
  53. ↑ Bright M, Sorgo A (11 May 2005). "Ultrastructural reinvestigation of the trophosome in adults of Riftia pachyptila (Annelida, Siboglinidae)". Invertebrate Biology. 122 (4): 347–368. doi:10.1111/j.1744-7410.2003.tb00099.x.
  54. ↑ Zhang JZ, Millero FJ (1993). "The products from the oxidation of H2S in seawater". Geochimica et Cosmochimica Acta. 57 (8): 1705–1718. Bibcode:1993GeCoA..57.1705Z. doi:10.1016/0016-7037(93)90108-9.
  55. ↑ Cavanaugh CM (1985). "Symbioses of chemoautotrophic bacteria and marine invertebrates from hydrothermal vents and reducing sediments". Bulletin of the Biological Society of Washington. 6: 373–388.
  56. ↑ Cavanaugh CM (February 1994). "Microbial symbiosis: patterns of diversity in the marine environment". American Zoologist. 34 (1): 79–89. doi:10.1093/icb/34.1.79.
  57. ↑ Fisher CR (1996). Ecophysiology of primary production at deep-sea vents and seeps (PDF).
  58. ↑ Polz MF, Ott JA, Bregut M, Cavanaugh CM (2000). "When Bacteria Hitch a Ride". ASM News-American Society for Microbiology. 66 (9): 531–539.
  59. ↑ Flores, Jason F.; Fisher, Charles R.; Carney, Susan L.; Green, Brian N.; Freytag, John K.; Schaeffer, Stephen W.; Royer, William E. (22 February 2005). "Sulfide binding is mediated by zinc ions discovered in the crystal structure of a hydrothermal vent tubeworm hemoglobin". Proceedings of the National Academy of Sciences. 102 (8): 2713–2718. Bibcode:2005PNAS..102.2713F. doi:10.1073/pnas.0407455102. ISSN 0027-8424. PMC 549462. PMID 15710902.
  60. ↑ Flores, Jason F.; Hourdez, téphane M. (2006). "The zinc-mediated sulfide-binding mechanism of hydrothermal vent tubeworm 400-kDa hemoglobin". Cahiers de Biologie Marine. 47: 371–377. doi:10.21411/cbm.a.11ae33dc.
  61. ↑ Royer, William E.; Zhu, Hao; Gorr, Thomas A.; Flores, Jason F.; Knapp, James E. (2005). "Allosteric Hemoglobin Assembly: Diversity and Similarity". Journal of Biological Chemistry. 280 (30): 27477–27480. doi:10.1074/jbc.r500006200. ISSN 0021-9258. PMID 15932877.
  62. ↑ Zal F, Suzuki T, Kawasaki Y, Childress JJ, Lallier FH, Toulmond A (December 1997). "Primary structure of the common polypeptide chain b from the multi-hemoglobin system of the hydrothermal vent tube worm Riftia pachyptila: an insight on the sulfide binding-site". Proteins. 29 (4): 562–574. doi:10.1002/(SICI)1097-0134(199712)29:4<562::AID-PROT15>3.0.CO;2-K. PMID 9408952. S2CID 25706128.
  63. ↑ Zal F, Leize E, Lallier FH, Toulmond A, Van Dorsselaer A, Childress JJ (July 1998). "S-Sulfohemoglobin and disulfide exchange: the mechanisms of sulfide binding by Riftia pachyptila hemoglobins". Proceedings of the National Academy of Sciences of the United States of America. 95 (15): 8997–9002. Bibcode:1998PNAS...95.8997Z. doi:10.1073/pnas.95.15.8997. PMC 21191. PMID 9671793.
  64. ↑ Bailly X, Jollivet D, Vanin S, Deutsch J, Zal F, Lallier F, Toulmond A (September 2002). "Evolution of the sulfide-binding function within the globin multigenic family of the deep-sea hydrothermal vent tubeworm Riftia pachyptila". Molecular Biology and Evolution. 19 (9): 1421–133. doi:10.1093/oxfordjournals.molbev.a004205. PMID 12200470.
  65. ↑ Corbera J (21 April 2017). "La vida que brolla a la foscor: les fumaroles hidrotermals submarines". L'Atzavara. 27: 39–53. ISSN 2339-9791.
  66. ↑ Simon V, Purcarea C, Sun K, Joseph J, Frebourg G, Lechaire JP, Gaill F, Hervé G (18 January 2000). "The enzymes involved in synthesis and utilization of carbamylphosphate in the deep-sea tube worm Riftia pachyptila". Marine Biology. 136 (1): 115–127. Bibcode:2000MarBi.136..115S. doi:10.1007/s002270050014. S2CID 85309709.
  67. 1 2 López-García P, Gaill F, Moreira D (April 2002). "Wide bacterial diversity associated with tubes of the vent worm Riftia pachyptila". Environmental Microbiology. 4 (4): 204–15. Bibcode:2002EnvMi...4..204L. doi:10.1046/j.1462-2920.2002.00286.x. PMID 12010127. S2CID 3940740.
  68. 1 2 Giovannelli D, Chung M, Staley J, Starovoytov V, Le Bris N, Vetriani C (July 2016). "Sulfurovum riftiae sp. nov., a mesophilic, thiosulfate-oxidizing, nitrate-reducing chemolithoautotrophic epsilonproteobacterium isolated from the tube of the deep-sea hydrothermal vent polychaete Riftia pachyptila". International Journal of Systematic and Evolutionary Microbiology. 66 (7): 2697–2701. doi:10.1099/ijsem.0.001106. PMID 27116914.
  69. ↑ Robidart JC, Bench SR, Feldman RA, Novoradovsky A, Podell SB, Gaasterland T, et al. (March 2008). "Metabolic versatility of the Riftia pachyptila endosymbiont revealed through metagenomics". Environmental Microbiology. 10 (3): 727–37. Bibcode:2008EnvMi..10..727R. doi:10.1111/j.1462-2920.2007.01496.x. PMID 18237306.
  70. ↑ Distel DL, Lane DJ, Olsen GJ, Giovannoni SJ, Pace B, Pace NR, et al. (June 1988). "Sulfur-oxidizing bacterial endosymbionts: Analysis of phylogeny and specificity by 16S rRNA sequences". Journal of Bacteriology. 170 (6): 2506–10. doi:10.1128/jb.170.6.2506-2510.1988. PMC 211163. PMID 3286609.
  71. ↑ Fisher CR (1995). "Toward an appreciation of hydrothennal-vent animals: Their environment, physiological ecology, and tissue stable isotope values". Washington DC American Geophysical Union Geophysical Monograph Series. Geophysical Monograph Series. 91: 297–316. Bibcode:1995GMS....91..297F. doi:10.1029/GM091p0297. ISBN 978-1-118-66399-8.
  72. ↑ Nelson DC (1995). "Chemoautotrophic and methanotrophic endosymbiotic bacteria at deep-sea vents and seeps". The Microbiology of Deep-Sea Hydrothermal Vents.
  73. ↑ "Exploring the deep ocean floor [This Dynamic Earth, USGS]". pubs.usgs.gov. Retrieved 5 July 2026.
  74. ↑ Koschinsky, Andrea; Garbe-Schönberg, Dieter; Sander, Sylvia; Schmidt, Katja; Gennerich, Hans-Hermann; Strauss, Harald (1 August 2008). "Hydrothermal venting at pressure-temperature conditions above the critical point of seawater, 5°S on the Mid-Atlantic Ridge". Geology. 36 (8): 615–618. Bibcode:2008Geo....36..615K. doi:10.1130/G24726A.1. ISSN 0091-7613.
  75. ↑ Corliss, John B.; Dymond, Jack; Gordon, Louis I.; Edmond, John M.; von Herzen, Richard P.; Ballard, Robert D.; Green, Kenneth; Williams, David; Bainbridge, Arnold; Crane, Kathy; van Andel, Tjeerd H. (16 March 1979). "Submarine Thermal Springs on the Galápagos Rift". Science. 203 (4385): 1073–1083. Bibcode:1979Sci...203.1073C. doi:10.1126/science.203.4385.1073. ISSN 0036-8075. PMID 17776033.
[edit]
Retrieved from "https://en.wikipedia.org/w/index.php?title=Riftia&oldid=1366783942"