Alvinella pompejana

Alvinella pompejana
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Annelida
Clade:	Pleistoannelida
Clade:	Sedentaria
Order:	Terebellida
Family:	Alvinellidae
Genus:	Alvinella
Species:	A. pompejana
Binomial name
	Alvinella pompejana; Desbruyères and Laubier, 1980

Last updated April 19, 2024

Introduction

In 1980 Daniel Desbruyères and Lucien Laubier, just few years after the discovery of the first hydrothermal vent system, identified one of the most heat-tolerant animals on Earth — Alvinella pompejana, the Pompeii worm.^[1] It was described as a deep-sea polychaete that resides in tubes near hydrothermal vents, along the seafloor. In 1997, marine biologist Craig Cary and colleagues found the same worms in a new section of Pacific Ocean, near Costa Rica, also attached to hydrothermal vents. The new discovery and subsequent work led to important progress in the scientific knowledge of these special worms.^[2]

They can reach up to 13 cm (5.1 in) in length and are pale gray, with red tentacle-like gills on their heads. Perhaps most fascinating, their tail ends are often resting in temperatures as high as 80 °C (176 °F), while their feather-like heads stick out of the tubes into water that is much cooler, 22 °C (72 °F).^[3] Scientists are attempting to understand how Pompeii worms can withstand such extreme temperatures by studying the bacteria that form a "fleece-like" covering on their backs. The bacteria have also been discovered to be chemolithotrophic, contributing to the ecology of the vent community. Recent research suggests the bacteria might play an important role in the feeding of the worms.^[4]

Attaching themselves to black smokers, the worms have been found to thrive at sustained temperatures of from 45 to 60 °C (113 to 140 °F) and even 105 °C (221 °F) for a short time,^[3] making the Pompeii worm the most heat-tolerant complex animal known to science after the tardigrades (or water bears), which are able to survive temperatures over 150 °C.

Biology

Reaching a length of up to 13 cm (5.1 in), Pompeii worms have "hairy" backs; these "hairs" are actually colonies of bacteria such as Nautilia profundicola , which are thought to afford the worm some degree of insulation. Glands on the worm's back secrete a mucus on which the bacteria feed, a form of symbiosis. The Pompeii worms form large, aggregate colonies enclosed in long tubes.

Alvinella pompejana has relatively simple organ systems centering around its rod-like heart. Its outermost organ is the gills along its feather-shaped head, four external gills present as leaf-like structures^[5] with a red colour due to their hemoglobin. The heart provides blood to these organs using contractions, pushing blood along the dorsal and ventral vessels. Beneath the heart lies the animal's stomach which connects to an oesophagus that is used to consume food. Finally, surrounding the organs is a coelom filled with coelomocytes, a type of phagocyte that acts as an immune system for the animal.^[5]

The Pompeii Worm feeds upon chemosynthetic bacteria, using retractable, ciliated feeding tentacles to scoop them up or absorbing their organic matter within the hydrothermal vent fluid.^[6] These tentacles are coated with 4 types of secretory cells along the entire epidermis. The tentacle's muscle cells seem to be filled with homogenous blood cells and individual hat-shaped cells, the hat-shaped cells have a condensed nucleus and it is hypothesised that these are heavily modified sperm cells.^[7]

The gills of A. pompejana are pinnate with many thin outgrowths. This organ is ultrastructurally similar to the gills of terebellidae and the epidermis is irregularly folded inwards. This gives the blood access to a space very close to the skin of A. pompejana, thus allowing more effective oxygen diffusion. Secretory cells of the goblet type have also been observed, along with hairlike receptor cells known as bipolar ciliary receptor cells. It is important to note that the Pompeii worm's epibiotic bacteria are absent from both the gills and tentacles.^[7]

Physiology

Alvinella pompejana's blood is abnormally cool at 20-30 °C. This is due to their blood's high positive cooperativity at these temperatures, thus haemoglobin is more likely to gain oxygen, an essential feature in an environment with sparse oxygen levels. Additionally, A. pompejana's gills have the highest specific surface area of any polychaete and small diffusion distances between the circulatory system and external seawater further assist in oxygenation.^[8]

Alvinellidae have an exceptionally high congenital oxygen affinity, in addition to a high heat requirement for oxygenation (𝚫H). Therefore, to allow adequate release of oxygen within the body, the Pompeii worm has acidic blood with a pH range of 6.6-6.9. A lower pH decreases the energy required to unbind oxygen from haemoglobin by utilising the bohr effect. The effect gradually compounds as the haemoglobin's 4 O₂ binding sites are emptied, it is maximal when the haemoglobin is fully deoxygenated. As a result, they can release all possible oxygen without expending excessive energy. To further compound this, A. pompejana's vascular haemoglobin has a lower oxygen affinity when compared to coelomic haemoglobin. When it is time to rebind oxygen to the haemoglobin, the blood must be cooled to 20-30 °C to regain optimal cooperativity. However, hydrothermal vents reach much higher temperatures. This makes the low blood temperature strange and seemingly hard to maintain.^[8]

The current hypothesis is that the polychaete worm maintains an “external microenvironment” that does not exceed this 20-30 °C limit, possibly through cold water being pulled into the tube during the worms repeated exit and entry in addition to the bacteria circulating this water around the worms body.^[9] This is supported by evidence that A. pompejana's mitochondrion break down when 30 °C is exceeded. Despite this the worm still manages to live in vents that exceed 50 °C. Furthermore, an inverse relationship is drawn between 𝚫H value and the temperature range of an environment; a higher 𝚫H means a habitat has a smaller range of temperatures with fewer fluctuations. As A. pompejana has a high 𝚫H it is reasonable to theorise that their habitable temperature range is quite small with few fluctuations.^[8]

Reproduction

Alvinella pompejana are a gonochoric species with distinct differences between the male and female genital pores (located at the base of the gills).^[5] Males have a pair of tentacles near the mouth that are absent in females. However, females possess a genital tract consisting of oviducts and spermathecae. The exact method of reproduction is unknown but it is believed that it is a complex multi-step process. However, it is known that the Pompeii worm's habitat rapidly changes and is extremely unstable, and thus their reproduction has adapted accordingly.

The size of the female oocytes suggests that the embryo is lecithotrophic meaning the only nutrition is within the yolk of the egg. Prior to fertilisation these oocytes are flattened spheres with an undulating membrane and a slightly off centre germinal vesicle that is less dense than the surrounding cytoplasm. Upon dilution in seawater, they become spherical and the GV disappears, at this point the whole oocyte appears homogeneous. This process does not require sperm.^[10]

The current hypothesis for egg synthesis and spawning is as follows: first the yolk is formed through a long process within the animal's coelom, next, the mature eggs are stored and finally, the eggs are spawned when either an environmental or biological change occurs (eg. sperm transfer).^[11] The method for transfer of spermatozoa it is likely achieved through pseudo-copulatory behaviour as the worms have been observed diving head-first into tubes in a display that may be mating.^[12]

When it is deemed appropriate to fertilise the eggs they are seemingly selected based upon size and then individually passed through the spermathecae. This method of fertilisation is more efficient than having all eggs be passed through at once.^[11]

Between 30 minutes and several hours after fertilisation has been achieved, the fertilisation envelope progressively elevates. This begins at a single point along the periphery of the oocyte. Then, prior to the first cleavage, a polar lobe forms thus resulting in asymmetrical cleavages. The diameter ratio between the asymmetrical blastomeres remains fairly constant (1:1.5) with "4 cell" embryos typically possessing 1 cell bigger than the other 3. This is a similar to the pattern observed in other polychaetes.^[10]

Early Development

Although in situ observations are yet to be made of Alvinella pompejana's early development, experiments have been conducted in order to hypothesise the embryonic and early stages of this polychaete's life.^[10]

Embryos of the Pompeii Worm are unable to tolerate both low and high temperatures, only developing in temperatures higher than 2 °C and lower than 20 °C. At 2 °C, the temperature of the abyssal sea, embryos enter a state of arrested development and at 20 °C the embryos die. Thus, they must not develop in hydrothermal vent colonies as temperatures much higher than 20 °C are achieved in this habitat.^[10]

Both in vitro and in situ incubation supported the hypothesis that embryonic development within an adult colony is impossible. To test their theory, researchers placed embryos in 3 areas, (I1) at the base of the chimney, (I2) in a Riftia pachyptila colony and (I3) in an adult colony. Results are presented in the below table with the highest survival and development rate coming from the (I1) and (I2) placements. It is also important to note that although the max temperature directly next to the (I3) incubator was 17 °C, temperatures as high as 27 °C were recorded 20 cm away.^[10]


	Survival (%)	Embryos with cleavage after 5 days (%)	Mean temp (°C)	Max temp (°C)	Min pH	Mean pH
I1	≤100%	70%	4±2 °C	9 °C	7.3	7.7
I2	≤100%	70%	6±2 °C	11 °C	7.3	7.5
I3	10%	0%	13±4 °C	17 °C	7.1	7.2

After considering all results, as well as prior studies, it was concluded that the 3 most likely hypotheses are as follows.

Embryos develop within a Riftia pachyptila colony (often near hydrothermal vents)
Embryos develop at the base of hydrothermal chimneys
Embryos enter a state of arrest and float in the abyssal column until a warm environment is found before resuming development (this would allow the dispersal of the species)

However, new evidence of an ecosystem underneath hydrothermal vents has suggested that the worm larvae may instead travel underground to colonise new vent systems^[13]

Tubes

The Pompeii worm is known for the tubes that it inhabits. Further research has uncovered that these tubes are unusually stable glycoprotein structures with high sulfur levels. Their construction was originally unknown but it is now hypothesised that the structure is made from a secretion from the epidermis of Alvinella pompejana. The fibril layers of the tubing are layered in a similar manner to plywood with each layer being slightly random. Within these tubes a layer of filamentous and rod-shaped bacteria are present, strangely these same bacteria can be found embedded within the walls of the tubes. It has been assumed that these are the bacteria that live in the worm's tubing, they seem to become trapped when the mucous is secreted. Additionally bacteria have been found to be a source of elemental sulfur which explains the sulfur detected within their tube walls.^[14]

Name

Pompeii worms get their name from the Roman city of Pompeii that was destroyed during an eruption of Mount Vesuvius in AD 79. Its family name Alvinellidae and genus name Alvinella both derive from DSV Alvin , the three-person submersible vehicle used during the discovery of hydrothermal vents and their fauna during the late 1970s. The family Alvinellidae contains eight other species, but none matches the Pompeii worm's heat tolerance.

Symbiotic Bacteria

While it is not yet known precisely how the Pompeii worm survives these severe vent conditions, scientists suspect the answer lies in the fleece-like bacteria on the worm's back; this layer may be up to 1 cm thick. The bacteria may possess special proteins, "eurythermal enzymes", providing the bacteria—and by extension the worms—protection from a wide range of temperatures.^[3] Studies are hampered by the difficulties of sampling; It is currently quite difficult for Pompeii Worms to survive decompression.

The bacteria are known to live in a mutualistic relationship with A. pompejana, making them both symbiotrophs. The main nutrition for the Pompeii worm is derived from chemosynthetic bacteria, this is why it chooses to live in such intense environments. This is due to the toxic metal levels of hydrothermal vent fluid, a factor chemosynthetic bacteria require. Unfortunately, the low pH, low oxygen levels and aforementioned metals within the fluid create an environment only fit for extremophiles; A. pompejana has physiological traits to assist in combating this but they are not enough. As a result, they have developed a strong relationship with the bacteria. The bacteria detoxify the fluid, this allows the worm to feed and live. Then, as the worm lives it respires, thus giving the bacteria a carbon source to feed upon. As an additional benefit, the bacteria gain shelter and surfaces to multiply upon due to the worm's tubes.^[1]

Study of the Pompeii worm's seemingly life-sustaining bacteria could lead to significant advances in the biochemical, pharmaceutical, textile, paper, and detergent industries.

Behavior

Pompeii worms simultaneously keep their heads (including the gills) in much cooler water while their tails are exposed to hot water. Since their internal temperature has yet to be measured, a Pompeii worm may survive exposure to hot water by dissipating heat through its head to keep its internal temperature within the realm previously known to be compatible with animal survival.^{[ citation needed ]}

Related Research Articles

Siboglinidae is a family of polychaete annelid worms whose members made up the former phyla Pogonophora and Vestimentifera. The family is composed of around 100 species of vermiform creatures which live in thin tubes buried in sediment (Pogonophora) or in tubes attached to hard substratum (Vestimentifera) at ocean depths ranging from 100 to 10,000 m. They can also be found in association with hydrothermal vents, methane seeps, sunken plant material, and whale carcasses.

Any worm that lives in a marine environment is considered a water worm. Marine worms are found in several different phyla, including the Platyhelminthes, Nematoda, Annelida, Chaetognatha, Hemichordata, and Phoronida. For a list of marine animals that have been called "sea worms", see sea worm.

<span class="mw-page-title-main">Polychaete</span> Class of annelid worms

Polychaeta is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes. Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made of chitin. More than 10,000 species are described in this class. Common representatives include the lugworm and the sandworm or clam worm Alitta.

<span class="mw-page-title-main">Chemosynthesis</span> Biological process building organic matter using inorganic compounds as the energy source

In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules and nutrients into organic matter using the oxidation of inorganic compounds or ferrous ions as a source of energy, rather than sunlight, as in photosynthesis. Chemoautotrophs, organisms that obtain carbon from carbon dioxide through chemosynthesis, are phylogenetically diverse. Groups that include conspicuous or biogeochemically important taxa include the sulfur-oxidizing Gammaproteobacteria, the Campylobacterota, the Aquificota, the methanogenic archaea, and the neutrophilic iron-oxidizing bacteria.

Hydrothermal vents are fissures on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspots. The dispersal of hydrothermal fluids throughout the global ocean at active vent sites creates hydrothermal plumes. Hydrothermal deposits are rocks and mineral ore deposits formed by the action of hydrothermal vents.

The Alvinellidae are a family of small, deep-sea polychaete worms endemic to hydrothermal vents in the Pacific Ocean. Belonging to the order Terebellida, the family contains two genera, Alvinella and Paralvinella; the former genus contains two valid species and the latter eight. Members of the family are termed alvinellids.

Riftia pachyptila, commonly known as the giant tube worm and less commonly known as the giant beardworm, is a marine invertebrate in the phylum Annelida related to tube worms commonly found in the intertidal and pelagic zones. R. pachyptila lives on the floor of the Pacific Ocean near hydrothermal vents. The vents provide a natural ambient temperature in their environment ranging from 2 to 30 °C, and this organism can tolerate extremely high hydrogen sulfide levels. These worms can reach a length of 3 m, and their tubular bodies have a diameter of 4 cm (1.6 in).

Mirocaris is a genus of shrimp associated with hydrothermal vents. Sometimes considered the only genus of the family Mirocarididae, Mirocaris is usually placed in the broader family Alvinocarididae. Mirocaris is characterized by a dorsoventrally flattened, non-dentate rostrum, as well as the possession of episodes on the third maxilliped through to the fourth pteropod. The genus contains two species, M. fortunata and M. indica. The two species are found in different oceans, and can be distinguished by the pattern of setation on the claw of the first pereiopod.

Paralvinella sulfincola, also known as the Sulfide worm, is a species of polychaete worm of the Alvinellidae family that thrives on undersea hot-water vents. It dwells within tubes in waters surrounding hydrothermal vents, in close proximity to super-heated fluids reaching over 300 °C (572 °F). The upper thermal limit for this polychaete is unknown; however, it is unlikely they can survive in constant temperatures over 50 °C (122 °F). It may tentatively be named a metazoan extremophile or, more specifically, a thermophile.

Lamellibrachia is a genus of tube worms related to the giant tube worm, Riftia pachyptila. They live at deep-sea cold seeps where hydrocarbons leak out of the seafloor, and are entirely reliant on internal, sulfide-oxidizing bacterial symbionts for their nutrition. The symbionts, gammaproteobacteria, require sulfide and inorganic carbon. The tube worms extract dissolved oxygen and hydrogen sulfide from the sea water with the crown of plumes. Species living near seeps can also obtain sulfide through their "roots", posterior extensions of their body and tube. Several sorts of hemoglobin are present in the blood and coelomic fluid to bind to the different components and transport them to the symbionts.

Terebellida make up an order of the Polychaeta class, commonly referred to as "bristle worms". Together with the Sabellida, the Spionida and some enigmatic families of unclear taxonomic relationship, they make up the subclass Canalipalpata, one of the three main clades of polychaetes. Like most polychaetes, almost all members of the Terebellida are marine organisms. Most are small, sessile detritivores which live in small tubes they build from mud or similar substrate, or burrow in the sand. Their central nervous system displays characteristic apomorphies.

Chrysomallon squamiferum, commonly known as the scaly-foot gastropod, scaly-foot snail, sea pangolin, or volcano snail is a species of deep-sea hydrothermal-vent snail, a marine gastropod mollusc in the family Peltospiridae. This vent-endemic gastropod is known only from deep-sea hydrothermal vents in the Indian Ocean, where it has been found at depths of about 2,400–2,900 m (1.5–1.8 mi). C. squamiferum differs greatly from other deep-sea gastropods, even the closely related neomphalines. In 2019, it was declared endangered on the IUCN Red List, the first species to be listed as such due to risks from deep-sea mining of its vent habitat.

Colleen Marie Cavanaugh is an American academic microbiologist best known for her studies of hydrothermal vent ecosystems. As of 2002, she is the Edward C. Jeffrey Professor of Biology in the Department of Organismic and Evolutionary Biology at Harvard University and is affiliated with the Marine Biological Laboratory and the Woods Hole Oceanographic Institution. Cavanaugh was the first to propose that the deep-sea giant tube worm, Riftia pachyptila, obtains its food from bacteria living within its cells, an insight which she had as a graduate student at Harvard. Significantly, she made the connection that these chemoautotrophic bacteria were able to play this role through their use of chemosynthesis, the biological oxidation of inorganic compounds to synthesize organic matter from very simple carbon-containing molecules, thus allowing organisms such as the bacteria to exist in deep ocean without sunlight.

Serpula is a genus of sessile, marine annelid tube worms that belongs to the family Serpulidae. Serpulid worms are very similar to tube worms of the closely related sabellid family, except that the former possess a cartilaginous operculum that occludes the entrance to their protective tube after the animal has withdrawn into it. The most distinctive feature of worms of the genus Serpula is their colorful fan-shaped "crown". The crown, used by these animals for respiration and alimentation, is the structure that is most commonly seen by scuba divers and other casual observers.

A trophosome is a highly vascularised organ found in some animals that houses symbiotic bacteria that provide food for their host. Trophosomes are contained by the coelom of the vestimentiferan tube worms and in the body of symbiotic flatworms of the genus Paracatenula.

Nautilia profundicola is a Gram-negative chemolithoautotrophic bacterium found around hydrothermal vents in the deep ocean. It was first discovered in 1999 on the East Pacific Rise at depth of 2,500 metres (8,200 ft), on the surface of the polychaete worm Alvinella pompejana. Nautilia profundicola lives symbiotically on the dorsal hairs of A. pompejana but they may also form biofilms and live independently on the walls of hydrothermal vents. The ability of N. profundicola to survive in an anaerobic environment rich in sulfur, H₂ and CO₂ of varying temperature makes it a useful organism to study, as these are the conditions that are theorized to have prevailed around the time of the earliest life on earth.

Vibrio diabolicus is a polysaccharide-secreting bacterium isolated from a deep-sea hydrothermal vent polychaete annelid, Alvinella pompejana. It is facultatively anaerobic, heterotrophic, and mesophilic.

Microbial symbiosis in marine animals was not discovered until 1981. In the time following, symbiotic relationships between marine invertebrates and chemoautotrophic bacteria have been found in a variety of ecosystems, ranging from shallow coastal waters to deep-sea hydrothermal vents. Symbiosis is a way for marine organisms to find creative ways to survive in a very dynamic environment. They are different in relation to how dependent the organisms are on each other or how they are associated. It is also considered a selective force behind evolution in some scientific aspects. The symbiotic relationships of organisms has the ability to change behavior, morphology and metabolic pathways. With increased recognition and research, new terminology also arises, such as holobiont, which the relationship between a host and its symbionts as one grouping. Many scientists will look at the hologenome, which is the combined genetic information of the host and its symbionts. These terms are more commonly used to describe microbial symbionts.

Tevnia is a genus of giant tube worm in the family Siboglinidae, with only one species, Tevnia jerichonana, living in a unique deep-sea environment. These deep sea marine species survive in environments like hydrothermal vents. These vents give off gas and toxic chemicals with the addition of having superheated temperatures. The giant tube worm prefers environments such as these despite the harsh temperature and toxic sea water.

The hydrothermal vent microbial community includes all unicellular organisms that live and reproduce in a chemically distinct area around hydrothermal vents. These include organisms in the microbial mat, free floating cells, or bacteria in an endosymbiotic relationship with animals. Chemolithoautotrophic bacteria derive nutrients and energy from the geological activity at Hydrothermal vents to fix carbon into organic forms. Viruses are also a part of the hydrothermal vent microbial community and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.

References

1 2 Desbruyères, D.; Laubier, L. (1980). "Alvinella pompejana gen.sp. nov., Ampharetidae aberrant des sources hydrothermales de la ride Est-Pacifique". Oceanologica Acta. 3 (3): 267–274.
↑ Scientists Succeed at First-Ever Attempt to Sequence DNA at Sea. NSF-OLPA. 1 November 2001
1 2 3 Grime, J. Philip; Pierce, Simon (2012). The Evolutionary Strategies that Shape Ecosystems. John Wiley & Sons. pp. 121–123. ISBN 978-1-118-22327-7.
↑ Grzymski, J. J.; et al. (2008). "Metagenome analysis of an extreme microbial symbiosis reveals eurythermal adaptation and metabolic flexibility". PNAS . 105 (45): 17516–17521. Bibcode:2008PNAS..10517516G. doi: 10.1073/pnas.0802782105 . PMC 2579889 . PMID 18987310.
1 2 3 Jouin-Toulmond, C.; Zal, F.; Hourdez, S. (1997). "Genital apparatus and ultrastructure of the spermatozoa in Alvinella pompejana (Annelida: Polychaeta)". Cahiers de Biologie Marine (2).
↑ Gaill, Francoise; Desbruyeres, Daniel; Laubier, Lucien. "Relationships between the "Pompeii worms" and their epibiotic bacteria" (PDF). Oceanologica.
1 2 Storch, V.; Gaill, F. (1 September 1986). "Ultrastructural observations on feeding appendages and gills ofAlvinella pompejana (Annelida, Polychaeta)". Helgoländer Meeresuntersuchungen. 40 (3): 309–319. doi: 10.1007/BF01983738 . ISSN 1438-3888. S2CID 32663219.
1 2 3 Toulmond, A.; Slitine, F. E. I.; De Frescheville, J.; Jouin, C. (December 1990). "Extracellular Hemoglobins of Hydrothermal Vent Annelids: Structural and Functional Characteristics in Three Alvinellid Species". The Biological Bulletin. 179 (3): 366–373. doi:10.2307/1542329. ISSN 1939-8697. JSTOR 1542329. PMID 29314955.
↑ Yazıcı, Ayşe Meriç; Persson, Erik (January 2023). "A New Model Inspired by the Pompeii Worm to Reverse Overheating in Nanosatellites". Space: Science & Technology. 3. doi: 10.34133/space.0017 . ISSN 2692-7659.
1 2 3 4 5 "Influence of environmental conditions on early development of the hydrothermal vent polychaete Alvinella pompejana". journals.biologists.com. Retrieved 8 November 2022.
1 2 PRADILLON, FLORENCE; GAILL, FRANÇOISE (1 July 2003). "Oogenesis characteristics in the hydrothermal vent polychaete Alvinella pompejana". Invertebrate Reproduction & Development. 43 (3): 223–235. doi:10.1080/07924259.2003.9652541. ISSN 0792-4259. S2CID 83704748.
↑ Desbruyères, D.; Chevaldonné, P.; Alayse, A. -M.; Jollivet, D.; Lallier, F. H.; Jouin-Toulmond, C.; Zal, F.; Sarradin, P. -M.; Cosson, R.; Caprais, J. -C.; Arndt, C.; O’Brien, J.; Guezennec, J.; Hourdez, S.; Riso, R. (1 January 1998). "Biology and ecology of the "Pompeii worm" (Alvinella pompejana Desbruyères and Laubier), a normal dweller of an extreme deep-sea environment: A synthesis of current knowledge and recent developments". Deep Sea Research Part II: Topical Studies in Oceanography. 45 (1): 383–422. doi:10.1016/S0967-0645(97)00083-0. ISSN 0967-0645.
↑ Mock-Bunting, Logan (8 August 2023). "Scientists Discover New Ecosystem Underneath Hydrothermal Vents". Schmidt Ocean Institute. Retrieved 17 August 2023.
↑ Gaill, F; Hunt, S (19 December 1986). "Tubes of deep sea hydrothermal vent worms Riftia pachyptila (Vestimentif era) and Alvinella pompejana (Annelida)" (PDF). Marine Ecology Progress Series . 34: 267–274. doi:10.3354/meps034267.

External links

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.

[:2-1] 1 2 Desbruyères, D.; Laubier, L. (1980). "Alvinella pompejana gen.sp. nov., Ampharetidae aberrant des sources hydrothermales de la ride Est-Pacifique". Oceanologica Acta. 3 (3): 267–274.

[2] Scientists Succeed at First-Ever Attempt to Sequence DNA at Sea. NSF-OLPA. 1 November 2001

[Grime-3] 1 2 3 Grime, J. Philip; Pierce, Simon (2012). The Evolutionary Strategies that Shape Ecosystems. John Wiley & Sons. pp. 121–123. ISBN 978-1-118-22327-7.

[4] Grzymski, J. J.; et al. (2008). "Metagenome analysis of an extreme microbial symbiosis reveals eurythermal adaptation and metabolic flexibility". PNAS . 105 (45): 17516–17521. Bibcode:2008PNAS..10517516G. doi: 10.1073/pnas.0802782105 . PMC 2579889 . PMID 18987310.

[:3-5] 1 2 3 Jouin-Toulmond, C.; Zal, F.; Hourdez, S. (1997). "Genital apparatus and ultrastructure of the spermatozoa in Alvinella pompejana (Annelida: Polychaeta)". Cahiers de Biologie Marine (2).

[6] Gaill, Francoise; Desbruyeres, Daniel; Laubier, Lucien. "Relationships between the "Pompeii worms" and their epibiotic bacteria" (PDF). Oceanologica.

[:4-7] 1 2 Storch, V.; Gaill, F. (1 September 1986). "Ultrastructural observations on feeding appendages and gills ofAlvinella pompejana (Annelida, Polychaeta)". Helgoländer Meeresuntersuchungen. 40 (3): 309–319. doi: 10.1007/BF01983738 . ISSN 1438-3888. S2CID 32663219.

[:5-8] 1 2 3 Toulmond, A.; Slitine, F. E. I.; De Frescheville, J.; Jouin, C. (December 1990). "Extracellular Hemoglobins of Hydrothermal Vent Annelids: Structural and Functional Characteristics in Three Alvinellid Species". The Biological Bulletin. 179 (3): 366–373. doi:10.2307/1542329. ISSN 1939-8697. JSTOR 1542329. PMID 29314955.

[9] Yazıcı, Ayşe Meriç; Persson, Erik (January 2023). "A New Model Inspired by the Pompeii Worm to Reverse Overheating in Nanosatellites". Space: Science & Technology. 3. doi: 10.34133/space.0017 . ISSN 2692-7659.

[:6-10] 1 2 3 4 5 "Influence of environmental conditions on early development of the hydrothermal vent polychaete Alvinella pompejana". journals.biologists.com. Retrieved 8 November 2022.

[:1-11] 1 2 PRADILLON, FLORENCE; GAILL, FRANÇOISE (1 July 2003). "Oogenesis characteristics in the hydrothermal vent polychaete Alvinella pompejana". Invertebrate Reproduction & Development. 43 (3): 223–235. doi:10.1080/07924259.2003.9652541. ISSN 0792-4259. S2CID 83704748.

[:0-12] Desbruyères, D.; Chevaldonné, P.; Alayse, A. -M.; Jollivet, D.; Lallier, F. H.; Jouin-Toulmond, C.; Zal, F.; Sarradin, P. -M.; Cosson, R.; Caprais, J. -C.; Arndt, C.; O’Brien, J.; Guezennec, J.; Hourdez, S.; Riso, R. (1 January 1998). "Biology and ecology of the "Pompeii worm" (Alvinella pompejana Desbruyères and Laubier), a normal dweller of an extreme deep-sea environment: A synthesis of current knowledge and recent developments". Deep Sea Research Part II: Topical Studies in Oceanography. 45 (1): 383–422. doi:10.1016/S0967-0645(97)00083-0. ISSN 0967-0645.

[13] Mock-Bunting, Logan (8 August 2023). "Scientists Discover New Ecosystem Underneath Hydrothermal Vents". Schmidt Ocean Institute. Retrieved 17 August 2023.

[14] Gaill, F; Hunt, S (19 December 1986). "Tubes of deep sea hydrothermal vent worms Riftia pachyptila (Vestimentif era) and Alvinella pompejana (Annelida)" (PDF). Marine Ecology Progress Series . 34: 267–274. doi:10.3354/meps034267.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]