사용자:Aspere/갈라파고스민고삐수염벌레

위키백과, 우리 모두의 백과사전.
갈라파고스민고삐수염벌레

생물 분류ℹ️
역: 진핵생물
계: 동물
문: 환형동물
아강: 정재류
목: 꽃갯지렁이목
과: 시보글리니대과
학명
Riftia pachyptila
M. L. Jones, 1981

갈라파고스민고삐수염벌레(학명: Riftia pachyptila)[1] 또는 관벌레(giant tube worm)[2]는 환형동물문의 해양 무척추동물로, 태평양의 열수분출공 인근에 서식한다. 열수분출공은 주변 온도를 2~30 °C로 유지시켜 주며,[3] 관벌레는 황화 수소의 농도가 높은 환경도 버틸 수 있다. 벌레의 길이는 3 m까지 길어지며,[4] 관 자체의 지름은 4 cm이다.

발견[편집]

연구정 DSV 앨빈.

관벌레는 1977년 지질학자 잭 코리스의 주도로 잠수정 DSV 앨빈을 이용해 갈라파고스 열점을 조사하던 도중 발견하였다.[5] 탐사 중 열수분출공 주변에서 발견한 생물은 이전에 전혀 발견된 적이 없었으며, 조사 팀에 생물학자가 한 명도 없었을 정도로 아무도 예상하지 못했었다. 연구팀은 잠수정을 이용해 조개류, 거대 게, 관벌레 등 여러 표본을 채취하였다.[6][7]

당시 대양저 중앙 열수분출공 주변의 온도는 측정하지 못했으나, 이후 측정 결과 5~30 °C로 비교적 높은 편임이 밝혀졌다. 상대적으로 높은 온도임에도 불구하고 많은 해양 생물이 서식하고 있었다.[8][9]

발달[편집]

관벌레는 비공생적 부유 담륜자 유생으로 태어나며, 성숙하며 고착될 때 공생 박테리아를 얻는다.[10][11] 성체가 생존에 의지하는 공생 박테리아는 배우체에는 들어 있지 않으나, 감염과 비슷한 과정을 통해 주변 환경에서 취득한다. 내측 복강 끝부분에 있는 입에서 시작되는 소화관은 전장, 중장, 후장을 거쳐 항문까지 이어지며, 과거에는 이 소화관을 통해 박테리아를 획득하는 것으로 추정했다. 공생 과정이 생겨나면 중장은 크기가 커지면서 모양이 바뀌어 영양체가 되며, 다른 소화 기관은 성체에서 관찰되지 않는다.[12]

구조[편집]

Isolating the vermiform body from white chitinous tube, a small difference exists from the classic three subdivisions typical of phylum Pogonophora:[13] the prosoma, the mesosoma, and the metasoma.

빨간색 새열을 보여주는 관벌레 집단.

The first body region is the vascularized branchial plume, which is bright red due to the presence of hemoglobin that contain up to 144 globin chains (each presumably including associated heme structures). These tube worm hemoglobins are remarkable for carrying oxygen in the presence of sulfide, without being inhibited by this molecule, as hemoglobins in most other species are.[14][15] The plume provides essential nutrients to bacteria living inside the trophosome. If the animal perceives a threat or is touched, it retracts the plume and the tube is closed due to the obturaculum, a particular operculum that protects and isolates the animal from the external environment.[16]

The second body region is the vestimentum, formed by muscle bands, having a winged shape, and it presents the two genital openings at the end.[17][18] The heart, extended portion of dorsal vessel, enclose the vestimentum.[19]

열수분출공 주변에 서식하는 관벌레는 자신의 영양체에 서식하는 박테리아로부터 유기물을 얻는다.

In the middle part, the trunk or third body region, is full of vascularized solid tissue, and includes body wall, gonads, and the coelomic cavity. Here is located also the trophosome, spongy tissue where a billion symbiotic, thioautotrophic bacteria and sulfur granules are found.[20][21] Since the mouth, digestive system, and anus are missing, the survival of R. pachyptila is dependent on this mutualistic symbiosis.[22] This process, known as chemosynthesis, was recognized within the trophosome by Colleen Cavanaugh.[22]

The soluble hemoglobins, present in the tentacles, are able to bind O2 and H2S, which are necessary for chemosynthetic bacteria. Due to the capillaries, these compounds are absorbed by bacteria.[23] During the chemosynthesis, the mitochondrial enzyme rhodanase catalyzes the disproportionation reaction of the thiosulfate anion S2O32- to sulfur S and sulfite SO32- .[24][25] The R. pachyptila’s bloodstream is responsible for absorption of the O2 and nutrients such as carbohydrates.

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.[15]

In the posterior part, the fourth body region, is the opistosome, which anchors the animal to the tube and is used for the storage of waste from bacterial reactions.[26]

Symbiosis[편집]

The discovery of bacterial invertebrate chemoautotrophic symbiosis, particularly in vestimentiferan tubeworms R. pachyptila[22] and then in vesicomyid clams and mytilid mussels revealed the chemoautotrophic potential of the hydrothermal vent tube worm.[27] Scientists discovered a remarkable source of nutrition that helps to sustain the conspicuous biomass of invertebrates at vents.[27] Many studies focusing on this type of symbiosis revealed the presence of chemoautotrophic, endosymbiotic, sulfur-oxidizing bacteria mainly in R. pachyptila,[28] which inhabits extreme environments and is adapted to the particular composition of the mixed volcanic and sea waters.[29] This special environment is filled with inorganic metabolites, essentially carbon, nitrogen, oxygen, and sulfur. In its adult phase, R. pachyptila lacks a digestive system. To provide its energetic needs, it retains those dissolved inorganic nutrients (sulfide, carbon dioxide, oxygen, nitrogen) into its plume and transports them through a vascular system to the trophosome, which is suspended in paired coelomic cavities and is where the intracellular symbiotic bacteria are found.[21][30][31] The trophosome[32] is a soft tissue that runs through almost the whole length of the tube's coelom. It retains a large number of bacteria on the order of 109 bacteria per gram of fresh weight.[33] Bacteria in the trophosome are retained inside bacteriocytes, thereby having no contact with the external environment. Thus, they rely on R. pachyptila for the assimilation of nutrients needed for the array of metabolic reactions they employ and for the excretion of waste products of carbon fixation pathways. At the same time, the tube worm depends completely on the microorganisms for the byproducts of their carbon fixation cycles that are needed for its growth.

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 called the trophosome.[22] Additional analyses involving stable isotope,[34] enzymatic,[35][27] and physiological[36] characterizations confirmed that the end symbionts of R. pachyptila oxidize reduced-sulfur compounds to synthesize ATP for use in autotrophic carbon fixation through the Calvin cycle. The host tubeworm enables the uptake and transport of the substrates required for thioautotrophy, which are HS−, O2, and CO2, receiving back a portion of the organic matter synthesized by the symbiont population. The adult tubeworm, given its inability to feed on particulate matter and its entire dependency on its symbionts for nutrition, the bacterial population is then the primary source of carbon acquisition for the symbiosis. Discovery of bacterial–invertebrate chemoautotrophic symbioses, initially in vestimentiferan tubeworms[22][27] and then in vesicomyid clams and mytilid mussels,[27] pointed to an even more remarkable source of nutrition sustaining the invertebrates at vents.

Endosymbiosis with chemoautotrophic bacteria[편집]

A wide range of bacterial diversity is associated with symbiotic relationships with R. pachyptila. Many bacteria belong to the phylum Campylobacterota (formerly class Epsilonproteobacteria)[37] as supported by the recent discovery in 2016 of the new species Sulfurovum riftiae belonging to the phylum Campylobacterota, family Helicobacteraceae isolated from R. pachyptila collected from the East Pacific Rise.[38] Other symbionts belong to the class Delta-, Alpha- and Gammaproteobacteria.[37] 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.[32]

Evidence based on 16S rRNA analysis affirms that R. pachyptila chemoautotrophic bacteria belong to two different clades: Gammaproteobacteria[39][21] and Campylobacterota (e.g. Sulfurovum riftiae)[38] 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.[21] 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 lack of a digestive system, R. pachyptila depends entirely on its bacterial symbiont to survive.[40][41]

In the first step of sulfide-oxidation, reduced sulfur (HS−) passes from the external environment into R. pachyptila blood, where, together with O2, it is bound by hemoglobin, forming the complex Hb-O2-HS− and then it is transported to the trophosome, where bacterial symbionts reside. Here, HS− is oxidized to elemental sulfur (S0) or to sulfite (SO32-).[21]

In the second step, the symbionts make sulfite-oxidation by the "APS pathway", to get ATP. In this biochemical pathway, AMP reacts with sulfite in the presence of the enzyme APS reductase, giving APS (adenosine 5'-phosphosulfate). Then, APS reacts with the enzyme ATP sulfurylase in presence of pyrophosphate (PPi) giving ATP (substrate-level phosphorylation) and sulfate (SO42-) as end products.[21] In formulas:

The electrons released during the entire sulfide-oxidation process enter in an electron transport chain, yielding a proton gradient that produces ATP (oxydative phosphorylation). Thus, ATP generated from oxidative phosphorylation and ATP produced by substrate-level phosphorylation become available for CO2 fixation in Calvin cycle, whose presence has been demonstrated by the presence of two key enzymes of this pathway: phosphoribulokinase and RubisCO.[27][42]

To support this unusual metabolism, R. pachyptila has to absorb all the substances necessary for both sulfide-oxidation and carbon fixation, that is: HS−, O2 and CO2 and other fundamental bacterial nutrients such as N and P. 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 phosphorilation. In R. pachyptila the production of hydrogen sulfide starts after 24h of hypoxia. In order 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.

Carbon fixation and organic carbon assimilation[편집]

Unlike metazoans, which respire carbon dioxide as a waste product, R. pachyptila-symbiont association has a demand for a net uptake of CO2 instead, as a cnidarian-symbiont associations.[43] Ambient deep-sea water contains an abundant amount of inorganic carbon in the form of bicarbonate HCO3−, but it is actually the chargeless form of inorganic carbon, CO2, that is easily diffusible across membranes. The low partial pressures of CO2 in the deep-sea environment is due to the seawater alkaline pH and the high solubility of CO2, yet the pCO2 of the blood of R. pachyptila may be as much as two orders of magnitude greater than the pCO2 of deep-sea water.[43]

CO2 partial pressures are transferred to the vicinity of vent fluids due to the enriched inorganic carbon content of vent fluids and their lower pH.[21] 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.[44][21] The facilitation of CO2 uptake by high environmental pCO2 was first inferred based on measures of elevated blood and coelomic fluid pCO2 in tubeworms, and was subsequently demonstrated through incubations of intact animals under various pCO2 conditions.[31]

Once CO2 is fixed by the symbionts, it must be assimilated by the host tissues. The supply of fixed carbon to the host is transported via organic molecules from the trophosome in the hemolymph, but the relative importance of translocation and symbiont digestion is not yet known.[31][45] 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.[45][46]

Sulfide acquisition[편집]

In deep-sea hydrothermal vents, sulfide and oxygen are present in different areas. Indeed, the reducing fluid of hydrothermal vents is rich in sulfide, but poor in oxygen, whereas sea water is richer in dissolved oxygen. Moreover, 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.[47] This causes the substrates to be less available for microbial activity, thus bacteria are constricted to compete with oxygen to get their nutrients. In order to avoid this issue, several microbes have evolved to make symbiosis with eukaryotic hosts.[48][21] In fact, R. pachyptila is able to cover the oxic and anoxic areas to get both sulfide and oxygen[49][50][51] 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.[52][53][54] 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.[55][56][57]

Symbiont acquisition[편집]

The acquisition of a symbiont by a host can occur in these ways:

  • Environmental transfer (symbiont acquired from a free-living population in the environment)
  • Vertical transfer (parents transfer symbiont to offspring via eggs)
  • Horizontal transfer (hosts that share the same environment)

Evidence suggests that R. pachyptila acquires its symbionts through its environment. In fact, 16S rRNA gene analysis showed that vestimentiferan tubeworms belonging to three different genera: Riftia, Oasisia, and Tevnia, share the same bacterial symbiont phylotype.[58][59][60][61][62]

This proves that R. pachyptila takes its symbionts from a free-living bacterial population in the environment. Other studies also support this thesis, because analyzing R. pachyptila eggs, 16S rRNA belonging to the symbiont was not found, showing that the bacterial symbiont is not transmitted by vertical transfer.[63]

Another proof to support the environmental transfer comes from several studies conducted in the late 1990s.[64] PCR was used to detect and identify a R. pachyptila symbiont gene whose sequence was very similar to the fliC gene that encodes some primary protein subunits (flagellin) required for flagellum synthesis. Analysis showed that R. pachyptila symbiont has at least one gene needed for flagellum synthesis. Hence, the question arose as to the purpose of the flagellum. Flagellar motility would be useless for a bacterial symbiont transmitted vertically, but if the symbiont came from the external environment, then a flagellum would be essential to reach the host organism and to colonize it. Indeed, several symbionts use this method to colonize eukaryotic hosts.[65][66][67][68]

Thus, these results confirm the environmental transfer of R. pachyptila symbiont.

번식[편집]

관벌레는 자웅이주 종으로,[69][70] 보통 동태평양이나 갈라파고스 인근 열수분출공 근처에 모여 서식하며, 각 종은 특정 위치에 고착되어 있다.[71] 열수분출공을 두른 집단은 수십 미터까지 펼쳐지기도 한다.[72]

수컷의 정자는 실 모양으로, 첨체(6 μm), 핵(26 μm), 꼬리(98 μm)로 나누어져 있다. 합쳐서 정자의 총 길이는 130 μm이며, 지름은 0.7 μm에서 꼬리 부분으로 갈수록 얇아져 0.2 μm에 이른다. 정액은 정자 약 340~350개가 집합을 이루며, 윗부분은 첨체와 핵, 아랫부분은 꼬리로 이루어져, 횃불과 비슷한 모양을 이룬다. 정자 집합체는 피브릴로 덮여 있어 응집력을 유지한다.

암컷의 난소는 몸통 전체에 이어진 생식선내강을 따라 분포하며, 영양체의 복부 측에 위치한다. 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.[출처 필요]

Males release their sperm into sea water. While the released agglomerations of spermatozoa, referred to as spermatozeugmata, do not remain intact for more than 30 seconds in laboratory conditions, they may maintain integrity for longer periods of time in specific hydrothermal vent conditions. 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. Generally, fertilization in R. pachyptila is considered internal. However, some argue that, as the sperm is released into sea water and only afterwards reaches the eggs in the oviducts, it should be defined as internal-external.[출처 필요]

R. pachyptila is completely dependent on the production of volcanic gases and the presence of sulfide-oxidizing bacteria. Therefore, its metapopulation distribution is profoundly linked to volcanic and tectonic activity that create active hydrothermal vent sites with a patchy and ephemeral distribution. The distance between active sites along a rift or adjacent segments can be very high, reaching hundreds of km.[71] This raises the question regarding larval dispersal. R. pachytpila is capable of larval dispersal across distances of 100 to 200 km[71] and cultured larvae show to be viable for 38 days.[73] 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.[71]

관벌레의 내부에서 공생하는 박테리아는 수정란을 통해 넘어가지 않고, 이후 유생 단계에서 획득한다. R. pachyptila planktonic larvae that are transported through sea-bottom currents until they reach active hydrothermal vents sites, are referred to as trophocores. The trophocore stage lacks endosymbionts, which are acquired once larvae settle in a suitable environment and substrate. 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, namely sulfide- oxidizing strains are paghocytized by epithelial cells found in the midgut are then retained. Bacteria that do not represent possible endosymbionts are digested. This raises questions as to how R. pachyptila manages to discern between essential and nonessential bacterial strains. The worm's ability to recognise a beneficial strain, as well as preferential host-specific infection by bacteria have been both suggested as being the drivers of this phenomenon.[12]

성장[편집]

관벌레는 현재까지 발견된 해양 무척추동물 중 가장 빠르게 성장하는 종으로, 2년 만에 성적 성숙이 이루어지고, 새 거주지를 만들며, 길이 1.5 m 이상으로 성장한다.[74]

관벌레가 사는 환경의 특성상, 관벌레는 다른 심해 생물과 큰 차이를 보이며, 해당과정에 사용하는 효소, 시트르산 회로, 세포 표면에서의 전자전달계는 얕은 물에 사는 생물과 비슷하다. 이는 심해 생물의 기초대사율이 낮은 편이라는 사실과 모순되기 때문에, 심해의 낮은 온도 및 높은 수압으로 인해 기초대사율이 낮아지는 것이 아니며, 열수분출공은 주변 환경을 바꿔 근처에 서식하는 생물에게 생물학적 영향을 준다는 사실을 알 수 있다.[33]


같이 보기[편집]

각주[편집]

  1. ↑ 〈갈라파고스민고삐수염벌레〉. 《두산백과 두피디아》. 2023년 8월 4일에 확인함. 
  2. ↑ 한국미생물학회. 〈관벌레〉. 《미생물학백과》. 2023년 8월 4일에 확인함. 
  3. ↑ Bright M, Lallier FH (2010). 〈The biology of vestimentiferan tubeworms〉. 《Oceanography and Marine Biology》 (PDF). Oceanography and Marine Biology - an Annual Review 48. Taylor & Francis. 213–266쪽. doi:10.1201/ebk1439821169-c4. ISBN 978-1-4398-2116-9. 2013년 10월 31일에 원본 문서 (PDF)에서 보존된 문서. 2013년 10월 30일에 확인함. 
  4. ↑ 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. (2015년 1월 13일). “Sizing ocean giants: patterns of intraspecific size variation in marine megafauna”. 《PeerJ》 (영어) 3: e715. doi:10.7717/peerj.715. PMC 4304853. PMID 25649000. 
  5. ↑ “Giant Tube Worm: Riftia pachyptila”. 《Smithsonian National Museum of Natural History》. 2018년 10월 24일에 원본 문서에서 보존된 문서. 2018년 10월 25일에 확인함. 
  6. ↑ Childress, James J. (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》 (영어) 35 (10–11): 1677–1680. Bibcode:1988DSRA...35.1677C. doi:10.1016/0198-0149(88)90043-X. 
  7. ↑ Lutz, Richard (1991년 1월 1일). “The biology of deep-sea vents and seeps: Alvin's magical mystery tour”. 《Oceanus》 34 (4): 75–83. ISSN 0029-8182. 2019년 12월 22일에 원본 문서에서 보존된 문서. 2019년 12월 22일에 확인함. 
  8. ↑ “Exploring the deep ocean floor: Hot springs and strange creatures”. 
  9. ↑ Koschinsky A, Garbe-Schönberg D, Sander S, Schmidt K, Gennerich HH, Strauss H (2008). “Hydrothermal venting at pressure-temperature conditions above the critical point of seawater, 5°S on the Mid-Atlantic Ridge”. 《Geology》 36 (8): 615. Bibcode:2008Geo....36..615K. doi:10.1130/g24726a.1. 
  10. ↑ Bright M. “Riftia pachyptila”. 2015년 4월 2일에 원본 문서에서 보존된 문서. 2015년 3월 19일에 확인함. 
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