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KLITE789 (bincang | sumb.)
Teg: Suntingan sumber 2017
KLITE789 (bincang | sumb.)
Teg: Suntingan sumber 2017
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</div>Salasilah dipermudah menunjukkan [[Homoscleromorpha]]<br/>paling rapat dengan haiwan kompleks<ref name="SperlingPeterson2007PoriferanParaphyly"/></div>
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In the 1990s sponges were widely regarded as a [[monophyletic]] group, all of them having descended from a [[last universal common ancestor|common ancestor]] that was itself a sponge, and as the "sister-group" to all other [[animals|metazoans]] (multi-celled animals), which themselves form a monophyletic group. On the other hand, some 1990s analyses also revived the idea that animals' nearest evolutionary relatives are [[choanoflagellate]]s, single-celled organisms very similar to sponges' [[choanocytes]] – which would imply that most Metazoa evolved from very sponge-like ancestors and therefore that sponges may not be monophyletic, as the same sponge-like ancestors may have given rise both to modern sponges and to non-sponge members of Metazoa.<ref name="BorchielliniManuelEtAl2001SpongeParaphylyAndOriginOfMetazoa">{{cite journal | vauthors = Borchiellini C, Manuel M, Alivon E, Boury-Esnault N, Vacelet J, Le Parco Y | title = Sponge paraphyly and the origin of Metazoa | journal = Journal of Evolutionary Biology | volume = 14 | issue = 1 | pages = 171–179 | date = January 2001 | pmid = 29280585 | doi = 10.1046/j.1420-9101.2001.00244.x }}</ref>
 
InPada the1990-an, 1990sspan spongesdianggap wereumum widelysebagai regarded as akumpulan [[monophyleticmonofiletik]] group, all of them havingsemuanya descendedketurunan fromdari asatu [[lastleluhur universal common ancestorterakhir|commonleluhur ancestorbersama]] thatyang wasjuga itselfmerupakan a spongespan, anddan assebagai the"kumpulan "sister-groupsaudara" tokepada all othersegala [[animalshaiwan|metazoansmetazoa]] (multi-celledhaiwan animalsmultisel), whichlain themselvesyang formsendirinya ajuga monophyleticmembentuk group.satu Onkumpulan themonofiletik. otherAkan handtetapi, someada 1990sjuga analisis dari dekad 1990-an analysesyang alsojuga revivedmembangkitkan thesemula idea thatbahawa animals'saudara nearestevolusi evolutionaryterdekat relativesbagi arehaiwan ialah [[choanoflagellatekoanoflagelat]]s, single-cellediaitu organismsorganisme veryunisel similaryang toamat sponges'serupa dengan sel [[choanocyteskoanosit]] pada span, dan whichini wouldakan implymembayangkan thatbahawa mostkebanyakan Metazoa evolvedberevolusi fromdari veryleluhur sponge-likeyang ancestorsamat andserupa thereforedengan thatspan, spongesdan mayoleh notitu bespan monophyletic,mungkin asbukan themonofiletik samekerana sponge-likeleluhur ancestorsseakan mayspan haveyang givensama risemungkin bothmenurunkan tokedua-dua modernspan spongesmoden anddan to nonahli-spongeahli membersMetazoa ofyang Metazoabukan span.<ref name="BorchielliniManuelEtAl2001SpongeParaphylyAndOriginOfMetazoa">{{cite journal | vauthors = Borchiellini C, Manuel M, Alivon E, Boury-Esnault N, Vacelet J, Le Parco Y | title = Sponge paraphyly and the origin of Metazoa | journal = Journal of Evolutionary Biology | volume = 14 | issue = 1 | pages = 171–179 | date = January 2001 | pmid = 29280585 | doi = 10.1046/j.1420-9101.2001.00244.x }}</ref>
Analyses since 2001 have concluded that [[Eumetazoa]] (more complex than sponges) are more closely related to particular groups of sponges than to the rest of the sponges. Such conclusions imply that sponges are not monophyletic, because the [[most recent common ancestor|last common ancestor]] of all sponges would also be a direct ancestor of the Eumetazoa, which are not sponges. A study in 2001 based on comparisons of [[ribosome]] [[DNA]] concluded that the most fundamental division within sponges was between [[glass sponge]]s and the rest, and that Eumetazoa are more closely related to [[Calcarea|calcareous sponges]], those with calcium carbonate spicules, than to other types of sponge.<ref name="BorchielliniManuelEtAl2001SpongeParaphylyAndOriginOfMetazoa"/> In 2007 one analysis based on comparisons of [[RNA]] and another based mainly on comparison of spicules concluded that demosponges and glass sponges are more closely related to each other than either is to calcareous sponges, which in turn are more closely related to Eumetazoa.<ref name="MüllerLiEtAl2007UniqueSkeletonOfSiliceousSponges"/><ref>{{cite journal | vauthors = Medina M, Collins AG, Silberman JD, Sogin ML | title = Evaluating hypotheses of basal animal phylogeny using complete sequences of large and small subunit rRNA | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 98 | issue = 17 | pages = 9707–12 | date = August 2001 | pmid = 11504944 | pmc = 55517 | doi = 10.1073/pnas.171316998 | bibcode = 2001PNAS...98.9707M }}</ref>
 
AnalysesAnalisis sincesejak 2001 havetelah concludedmencapai thatkesimpulan bahawa [[Eumetazoa]] (morelebih complexkompleks thandaripada spongesspan) aremempunyai morepertalian closelylebih relatedrapat todengan particularkumpulan groupsspan oftertentu spongesdaripada thanspan-span toyang the rest of the spongeslain. SuchKesimpulan conclusionssedemikian implymembayangkan thatbahawa spongesspan arebukanlah notmonofiletik monophyletic, because thekerana [[most recentLeluhur commonuniversal ancestorterakhir|lastleluhur commonbersama ancestorterakhir]] ofkepada allsegala spongesspan wouldjuga alsoadalah beleluhur alangsung direct ancestor of thebagi Eumetazoa, whichyang arebukan not spongesspan. A study inKajian 2001 based on comparisonsberasaskan ofperbandingan [[ribosomeDNA]] [[DNAribosom]] concludedmencapai kesimpulan bahawa pembahagian thatyang thepaling mostasas fundamentaldi divisionkalangan withinspan spongesadalah wasdi betweenantara [[glassspan spongekaca]]s anddan thespan-span restlain, and thatmalah Eumetazoa are more closelylebih relatedrapat todengan [[Calcarea|calcareousspan spongesberkalsium]], thoseyang withmempunyai calciumspikul carbonatekalsium spiculeskarbonat, than to otherberbanding typeslain-lain ofjenis spongespan.<ref name="BorchielliniManuelEtAl2001SpongeParaphylyAndOriginOfMetazoa"/> InPada 2007, onesatu analysisanalisis basedberdasarkan on comparisons ofperbandingan [[RNA]], anddan anothersatu basedlagi mainlyberasaskan onperbandingan comparisonspikul, ofmencapai spiculeskesimpulan concludedbahawa thatdemospan demospongesdan andspan glasskaca spongeslebih arerapat moredengan closelysatu relatedsama tolain eachberbanding othersetiap thansatu eitherdengan isspan to calcareous spongesberkalsium, which in turn are moreyang closelylebih relatedrapat todengan Eumetazoa.<ref name="MüllerLiEtAl2007UniqueSkeletonOfSiliceousSponges"/><ref>{{cite journal | vauthors = Medina M, Collins AG, Silberman JD, Sogin ML | title = Evaluating hypotheses of basal animal phylogeny using complete sequences of large and small subunit rRNA | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 98 | issue = 17 | pages = 9707–12 | date = August 2001 | pmid = 11504944 | pmc = 55517 | doi = 10.1073/pnas.171316998 | bibcode = 2001PNAS...98.9707M }}</ref>
Other anatomical and biochemical evidence links the Eumetazoa with [[Homoscleromorpha]], a sub-group of demosponges. A comparison in 2007 of [[cell nucleus|nuclear]] [[DNA]], excluding glass sponges and [[Ctenophora|comb jellies]], concluded that: [[Homoscleromorpha]] are most closely related to Eumetazoa; calcareous sponges are the next closest; the other demosponges are evolutionary "aunts" of these groups; and the [[chancelloriidae|chancelloriids]], bag-like animals whose fossils are found in [[Cambrian]] rocks, may be sponges.<ref name="SperlingPeterson2007PoriferanParaphyly">{{cite journal| vauthors = Sperling EA, Pisani D, Peterson KJ |year=2007 |access-date=2008-11-04 |title=Poriferan paraphyly and its implications for Precambrian paleobiology |journal=Journal of the Geological Society of London |volume=286 |pages=355–368 |doi=10.1144/SP286.25 |url=http://www.dartmouth.edu/~peterson/Sperling,%20Pisani%20and%20Peterson.pdf |url-status=dead |archive-date=May 9, 2009 |archive-url=https://web.archive.org/web/20090509061759/http://www.dartmouth.edu/~peterson/Sperling,%20Pisani%20and%20Peterson.pdf |bibcode=2007GSLSP.286..355S }}</ref> The [[sperm]] of Homoscleromorpha share with those of Eumetazoa features that those of other sponges lack. In both Homoscleromorpha and Eumetazoa layers of cells are bound together by attachment to a carpet-like basal membrane composed mainly of "type IV" [[collagen]], a form of collagen not found in other sponges – although the spongin fibers that reinforce the mesohyl of all demosponges is similar to "type IV" collagen.<ref name="ExpositoCluzelEtAl2002EvolutionOfCollagens">{{cite journal | vauthors = Exposito JY, Cluzel C, Garrone R, Lethias C | title = Evolution of collagens | journal = The Anatomical Record | volume = 268 | issue = 3 | pages = 302–16 | date = November 2002 | pmid = 12382326 | doi = 10.1002/ar.10162 }}</ref>
 
OtherBukti-bukti anatomicalanatomi anddan biochemicalbiokimia evidenceselanjutnya linksmenghubungkan theEumetazoa Eumetazoadengan withsatu subkumpulan demospan [[Homoscleromorpha]], a sub-group of demosponges. AKajian comparison2007 inberdasarkan 2007 ofperbandingan [[cell nucleus|nuclearDNA]] [[DNAnukleus sel|nukleus]], excludingtidak glasstermasuk spongesspan andkaca dan [[Ctenophora|combampai jelliesbulat]], concludedmenyimpulkan thatbahawa: [[Homoscleromorpha]] arepaling mostrapat closelypertalian related todengan Eumetazoa; calcareousspan spongesberkalsium arekedua the next closestterdekat; thelain-lain otherdemospan demospongesmerupakan are evolutionary"ibu "auntssaudara" ofevolusi thesebagi groupskumpulan-kumpulan tersebut; and thedan [[chancelloriidae|chancelloriidsChancelloriidae]], bag-likehaiwan seakan animalskarung whoseyang fossilsfosilnya areditemui founddalam inbatuan [[CambrianKambria]] rocks, maymungkin beadalah spongesspan.<ref name="SperlingPeterson2007PoriferanParaphyly">{{cite journal| vauthors = Sperling EA, Pisani D, Peterson KJ |year=2007 |access-date=2008-11-04 |title=Poriferan paraphyly and its implications for Precambrian paleobiology |journal=Journal of the Geological Society of London |volume=286 |pages=355–368 |doi=10.1144/SP286.25 |url=http://www.dartmouth.edu/~peterson/Sperling,%20Pisani%20and%20Peterson.pdf |url-status=dead |archive-date=May 9, 2009 |archive-url=https://web.archive.org/web/20090509061759/http://www.dartmouth.edu/~peterson/Sperling,%20Pisani%20and%20Peterson.pdf |bibcode=2007GSLSP.286..355S }}</ref> The [[spermSperma]] of Homoscleromorpha sharemempunyai withpersamaan thoseciri-ciri ofdengan sperma Eumetazoa featuresyang thattiada thosepada of other spongesspan lacklain. InBagi bothkedua-dua Homoscleromorpha anddan Eumetazoa, layerslapisan-lapisan ofsel cellsdiikat aredengan boundterlekat togetherpada by attachment to a carpet-likemembran basal membraneseakan composedhamparan mainlyyang ofdiperbuat "typekhususnya IV"daripada [[collagenkolagen]], a"jenis formIV", ofsejenis collagenkolagen notyang foundtidak interdapat otherpada spongesspan-span lain, althoughnamun theserat-serat spongin fibersyang thatmemperkukuh reinforcemesohil thebagi mesohylsemua ofdemospan alladalah demospongesserupa isdengan similar tokolagen "typejenis IV" collagen.<ref name="ExpositoCluzelEtAl2002EvolutionOfCollagens">{{cite journal | vauthors = Exposito JY, Cluzel C, Garrone R, Lethias C | title = Evolution of collagens | journal = The Anatomical Record | volume = 268 | issue = 3 | pages = 302–16 | date = November 2002 | pmid = 12382326 | doi = 10.1002/ar.10162 }}</ref>
 
[[File:Bathocyroe fosteri.jpg|thumb|left|Seekor [[ctenophora|ampai bulat]].]]
TheAnalisis-analisis analysesyang describeddihuraikan abovedi concludedatas thattelah spongesmencapai arekesimpulan closestbahawa tospan theadalah ancestorspaling ofrapat alldengan leluhur bagi segala Metazoa, ofbagi allsemua multi-celledhaiwan animalsmultisel includingtermasuk bothspan spongesdan andhaiwan-haiwan moreyang complexlebih groupskompleks. HoweverAkan tetapi, anothersatu comparisonlagi inkajian 2008 ofyang membandingkan 150 genesgem indalam eachsetiap ofsatu daripada 21 genera,genus ranginghidupan fromdari fungikulat tohingga humansmanusia buttetapi includingtermasuk onlyhanya twodua speciesspesies of spongespan, suggestedmembayangkan thatbahawa [[ctenophora|combmakhluk jellies]]ampai bulat ([[ctenophoraCtenophora]]) areadalah theketurunan mostMetazoa basalpaling lineageasas ofdi thekalangan Metazoasampel. includedJika inini thebenar, sample.maka Ifsama thisada isampai correct,bulat eithermoden moderntumbuh combstruktur jellieskompleks developedsecara theirberasingan complexdaripada structuresMetazoa independentlylain, ofataupun otherleluhur Metazoa,span or sponges'lebih ancestorskompleks weresedangkan moresemua complexspan andyang alldiketahui knownkini spongessekadar arejelmaan drasticallyyang simplifiedteramat formsdipermudah. TheKajian studyini recommendedmengesyorkan furtheragar analysesdiadakan usinganalisis alanjutan widermenggunakan rangebanyak oflagi spongesjenis andspan other simpledan Metazoa suchringkas yang lain asseperti [[Placozoa]].<ref name="Dunn_2008" /> TheHasil resultsanalisis ofini suchditerbitkan an analysis, published inpada 2009, suggestmencadangkan thatbahawa akita returnmungkin toharus thekembali previouske viewpandangan may be warrantedlama. 'Family treesSalasilah' constructed using ayang combinationdibina ofmenggunakan allkombinasi availablesegala data – morphologicalmorfologi, developmentalperkembangan anddan molecularmolekul yang concludedsedia thatada, themencapai spongeskesimpulan arebahawa inspan factsememangnya asatu monophyletickumpulan groupmonofiletik, anddan withbersama thedengan [[cnidarianCnidaria]]s form theia sistermembentuk groupkumpulan tosaudara thekepada bilateriansbilateria.<ref name=plos2009>{{cite journal | vauthors = Schierwater B, Eitel M, Jakob W, Osigus HJ, Hadrys H, Dellaporta SL, Kolokotronis SO, Desalle R | display-authors = 6 | title = Concatenated analysis sheds light on early metazoan evolution and fuels a modern "urmetazoon" hypothesis | journal = PLOS Biology | volume = 7 | issue = 1 | pages = e20 | date = January 2009 | pmid = 19175291 | pmc = 2631068 | doi = 10.1371/journal.pbio.1000020 }}</ref>
 
ASatu verykajian largeyang andmelibatkan internally consistent alignment ofpenjajaran 1,719 proteinsprotein yang sangat besar dan attekal thedalam metazoanskala scalemetazoa, publishedterbitan intahun 2017, showedmenunjukkan thatbahawa (i) spongesspanrepresenteddiwakili byoleh Homoscleromorpha, Calcarea, Hexactinellida, anddan Demospongiae – areadalah monophyleticmonofiletik, (ii) spongesspan areadalah sister-groupkumpulan tosaudara allkepada othersemua multicellularhaiwan animalsmultisel lain, (iii) ctenophoresCtenophora emergemuncul assebagai theketurunan second-earliesthaiwan branchingyang animalkedua lineageterawal bercabang, anddan (iv) [[placozoansPlacozoa]] emergemuncul assebagai theketurunan thirdhaiwan animal lineageketiga, followeddiikuti byCnidaria, cnidarianskumpulan sister-groupsaudara tokepada bilateriansbilateria.<ref name = "Simion_2017" />
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