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Horisontaalne geeniülekanne: erinevus redaktsioonide vahel

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2. rida:
'''Horisontaalne geeniülekanne '''ehk '''horisontaalne geenisiire '''on [[geneetiline_info|geneetilise informatsiooni]] kandumine ühest [[organism]]ist teise muul viisil kui läbi traditsioonilise [[paljunemine|paljunemise]]. Nimetatakse ka lateraalseks geeniülekandeks, vastandudes vertikaalsele, kus [[geen]]ide ülekandmine toimub vanempõlvkonnalt järglastele läbi [[seksuaalne paljunemine|seksuaalse]] või [[seksuaalne_paljunemine|aseksuaalse]] [[sigimine|sigimise]]. On näidatud, et horisontaalne geeniülekanne on oluliseks faktoriks paljude organismide [[evolutsioon]]is.
 
Horisontaalne geeniülekanne on peamiseks [[bakter]]ite [[antibiootikumresistentsus]]e põhjuseks <ref name="Transgenic"/><ref name="Kay"/><ref name="Koonin"/><ref name="Nielsen"/> ja mängib olulist rolli nende võimes lagundada uusi ühendeid. Näiteks inimese poolt loodud [[pestitsiid|keemilisi taimekaitsevahendeid]].<ref name="McGowan"/>
Horisontaalne geeniülekanne on peamiseks [[bakter]]ite [[antibiootikumresistentsus]]e põhjuseks<ref>OECD, ''Safety Assessment of Transgenic Organisms, Volume 4: OECD Consensus Documents,'' 2010, pp.171–174</ref><ref>{{cite journal |author=Kay E, Vogel TM, Bertolla F, Nalin R, Simonet P |title=In situ transfer of antibiotic resistance genes from transgenic (transplastomic) tobacco plants to bacteria |journal=Appl. Environ. Microbiol. |volume=68 |issue=7 |pages=3345–51 |year=2002 |month=July |pmid=12089013 |pmc=126776 |url=http://aem.asm.org/cgi/pmidlookup?view=long&pmid=12089013}}</ref><ref>{{cite journal |author=Koonin EV, Makarova KS, Aravind L |title=Horizontal gene transfer in prokaryotes: quantification and classification |journal=Annu. Rev. Microbiol. |volume=55 |pages=709–42 |year=2001 |pmid=11544372 |doi=10.1146/annurev.micro.55.1.709 |url=http://arjournals.annualreviews.org/doi/full/10.1146/annurev.micro.55.1.709?url_ver=Z39.88-2003&rfr_id=ori:rid:crossref.org&rfr_dat=cr_pub%3dpubmed}}</ref><ref>{{cite journal |author=Nielsen KM |title=Barriers to horizontal gene transfer by natural transformation in soil bacteria |journal=APMIS Suppl. |volume=84 |pages=77–84 |year=1998 |pmid=9850687 }}</ref> ja mängib olulist rolli nende võimes lagundada uusi ühendeid. Näiteks inimese poolt loodud [[pestitsiid|keemilisi taimekaitsevahendeid]].<ref>{{cite journal |author=McGowan C, Fulthorpe R, Wright A, Tiedje JM |title=Evidence for interspecies gene transfer in the evolution of 2,4-dichlorophenoxyacetic acid degraders |journal=Appl. Environ. Microbiol. |volume=64 |issue=10 |pages=4089–92 |year=1998 |month=October |pmid=9758850 |pmc=106609 |url=http://aem.asm.org/cgi/pmidlookup?view=long&pmid=9758850}}</ref>
 
Horisontaalses geeniülekandes osalevad tihti mõõdukad [[bakteriofaagid]] ja [[plasmiid]]id.<ref>{{Cite journal |authorname="Naik GA, Bhat LN, Chpoade BA, Lynch JM |title=Transfer of broad-host-range antibiotic resistance plasmids in soil microcosms |journal=Curr. Microbiol. |volume=28 |year=1994 |pages=209–215 |doi=10.1007"/BF01575963 |issue=4}}</ref> Geenid, mis vastutavad antibiootikumresistentsuse eest ühes bakteriliigis, kanduvad üle teistele [[liik]]idele läbi erinevate mehhanismide (näiteks F-pilide kaudu). See on muutumas laialdaseks probleemiks meditsiinis ning üheks oluliseks põhjuseks, miks patsiendid ei tohiks antibiootikume manustada ilma arsti vastava retseptita.<ref>{{cite book |first=Peter J. |lastname="Russell |title=Biology : exploring the diversity of life|year=2009|publisher=Nelson Education|location=Toronto|isbn=0-17-644094-1|url=http:"//www.coursesmart.com/biology-exploring-the-diversity-of-life-1st/russell-wolfe-hertz-starr-fenton-addy-maxwell/dp/9780176440947|edition=1st Canadian}}</ref>
 
Viimastel aastatel on kindlaks tehtud, et bakterite vahel toimub horisontaalne geeniülekanne palju sagedamini kui varem arvati. Seetõttu on [[geneetik]]ud hakanud vertikaalse ülekande kõrvalt sellele ka järjest enam tähelepanu pöörama.<ref name="edwards">{{cite web|url=http://www.physorg.com/news205389256.html|title=Horizontal gene transfer in microbes much more frequent than previously thought|author=Lin Edwards|date= October 4, 2010|publisher=PhysOrg.com|accessdate=2012-01-06}}</ref><ref name="arnold">{{cite web|url=http://www.scientificamerican.com/article.cfm?id=to-share-and-share-alike|title=To Share and Share Alike: Bacteria swap genes with their neighbors more frequently than researchers have realized|author=Carrie Arnold|date= April 18, 2011|publisher=Scientific American|accessdate=2012-01-06}}</ref>
 
Kunstlik horisontaalne geeniülekanne on põhiliseks tööriistaks [[geenitehnoloogia]]s.
 
==Ajalugu==
1951. aastal täheldati horisontaalset geeniülekannet esmakordselt. Artiklis kirjeldati kuidas [[virulentsus]]geeni ülekandumise tagajärjel muutus avirulentne ''Corynebacterium diphtheriae'' tüvi virulentseks.<ref>{{cite journal | author name= Victor J "Freeman | title = Studies on the virulence of bacteriophage-infected strains of ''Corynebacterium Diphtheriae'' | journal = Journal of Bacteriology | year = 1951 | pages = 675–688 | pmid = 14850426 | volume = 61 | issue = 6 | pmc=386063}}<"/ref> Sellega lahendati ka [[difteeria]] mõistatus (bakteriga nakatunud patsientidel ei pidanud esialgu haiguslikke [[sümptom]]eid esinema, need võisid aga hiljem tekkida).<ref>Phillip name="Marguilies "Epidemics: Deadly diseases throughout history". Rosen, New York. 2005.</ref> See fenomen oli ühtlasi esimeseks näiteks [[lüsogeenne_tsükkel|lüsogeense tsükli]] olulisusest.<ref>André name="Lwoff (1965). [http://nobelprize.org/nobel_prizes/medicine/laureates/1965/lwoff-lecture.html "Interaction among Virus, Cell, and Organism"]. Nobel Lecture for the Nobel Prize in Physiology or Medicine.</ref>
 
1959. aastal kirjeldati esimesena bakterite-sisest geeniülekannet. Väljaandes demonstreeriti antibiootikumresistentsuse edastamist ühelt bakteriliigilt teisele.<ref>{{cite journal |authorname="Ochiai K, Yamanaka T, Kimura K, Sawada, O |title=Inheritance of drug resistance (and its transfer) between Shigella strains and Between Shigella and E. coli strains |journal=Hihon Iji Shimpor |volume=1861 |pages=34 |year=1959 |language=Japanese}}<"/ref><ref>{{cite journal |authorname="Akiba T, Koyama K, Ishiki Y, Kimura S, Fukushima T |title=On the mechanism of the development of multiple-drug-resistant clones of Shigella |journal=Jpn. J. Microbiol. |volume=4 |pages=219–27 |year=1960 |month=April |pmid=13681921 }}<"/ref> 1980. aastate keskel ennustas Syvanen <ref>{{cite journal |authorname="Syvanen M |title=Cross-species gene transfer; implications for a new theory of evolution |journal=J. Theor. Biol. |volume=112 |issue=2 |pages=333–43 |year=1985 |month=January |pmid=2984477 |doi=10.1016"/S0022-5193(85)80291-5 |url=http://www.dcn.davis.ca.us/vme/hgt/JTheoBiolvol112pp333-343yr1985.PDF |format=PDF}}</ref>, et on olemas lateraalne geeniülekanne, millel on bioloogiline tähtsus ning mis on osalenud evolutsioonilise ajaloo kujundamises Maal alates elu tekke algusest.
 
1999. aastal märkisid Jain, Rivera ja Lake: "Geenide ja genoomide uuringud näitavad, et [[prokarüoot]]ide vahel on toimunud märkimisväärselt palju horisontaalseid geeniülekandeid <ref>{{cite journal |authorname="Jain R, Rivera MC, Lake JA |title=Horizontal gene transfer among genomes: The complexity hypothesis |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=96 |issue=7 |pages=3801–6 |year=1999 |month=March |pmid=10097118 |pmc=22375 |doi= 10.1073"/pnas.96.7.3801|url=http://www.pnas.org/cgi/pmidlookup?view=long&pmid=10097118|bibcode = 1999PNAS...96.3801J }}</ref> ja sel protsessil on tähtsus ka üherakuliste [[eukarüoodid|eukarüootide]] hulgas ning [[protistid]]e evolutsioonis."<ref>{{cite journal |authorname="Bapteste E, Susko E, Leigh J, MacLeod D, Charlebois RL, Doolittle WF |title=Do orthologous gene phylogenies really support tree-thinking? |journal=BMC Evol. Biol. |volume=5 |pages=33 |year=2005 |pmid=15913459 |pmc=1156881 |doi=10.1186"/1471-2148-5-33 |issue=1}}</ref>
 
Levimus ja olulisus hulkraksete eukarüootide seas on tänapäeval veel ebaselge.<ref>{{cite journal | author name= "Richardson, Aaron O.; Palmer, Jeffrey D. | month = January | year = 2007 | title = Horizontal Gene Transfer in Plants |journal = Journal of Experimental Botany | volume = 58 | pages = 1–9 |url=http://www.sdsc.edu/~shindyal/ejc121304.pdf |format=PDF | doi = 10.1093/jxb/erl148 | pmid = 17030541 | issue = 1}}<"/ref>
 
Järjest kasvava tõendusmaterjali valguses, mis vihjab selle fenomeni evolutsioonilisele olulisusele, on molekulaarbioloogid horisontaalset geeniülekannet kirjeldanud ka kui "uut paradigmat bioloogias".<ref name="gogarten">{{cite journal | author = Gogarten, Peter | year = 2000 | title = Horizontal Gene Transfer: A New Paradigm for Biology | journal = Esalen Center for Theory and Research Conference | url = http://www.esalenctr.org/display/confpage.cfm?confid=10&pageid=105&pgtype=1 | accessdate = 2007-03-18}}</ref>
 
On kardetud, et see protsess kujutab endast varjatud ohtu, kuna võimaldab transgeense [[DNA]] levimist liikide vahel.<ref name="Mae-Wan Ho">{{Cite journal |author=Mae-Wan Ho |year=1999 |title=Cauliflower Mosaic Viral Promoter – A Recipe for Disaster? |journal=Microbial Ecology in Health and Disease |volume=11 |url=http://www.i-sis.org.uk/pdf/CaMV_promoter_recipe_for_disaster.pdf |accessdate=2008-06-09 |pages=194–7}}</ref>
 
==Mehhanismid==
Horisontaalne geeniülekanne võib toimuda mitme erineva mehhanismi kaudu:<ref name="todar"/><ref name="maloy"/>
Horisontaalne geeniülekanne võib toimuda mitme erineva mehhanismi kaudu:<ref name="todar">{{cite web|url=http://textbookofbacteriology.net/themicrobialworld/bactresanti.html|title=Bacterial Resistance to Antibiotics|author=Kenneth Todar |publisher=The Microbial World: Lectures in Microbiology, Department of Bacteriology, University of Wisconsin-Madison |accessdate=January 6, 2012}}</ref><ref name="maloy">{{cite web|url=http://www.sci.sdsu.edu/~smaloy/MicrobialGenetics/topics/genetic-exchange/exchange/exchange.html|title=Horizontal Gene Transfer|author=Stanley Maloy|date= July 15, 2002 |publisher=San Diego State University|accessdate=January 6, 2012}}</ref>▼
* [[Geneetiline_transformatsioon|Transformatsioon]], geneetilise info sisenemine väliskeskkonnast bakterisse läbi [[rakumembraan]]i. Seda protsessi kasutatakse tihti geenitehnoloogias, et sisestada bakteritesse uusi geene, mis võimaldaksid neid kasutada tööstuslikel või meditsiinilistel eesmärkidel.
 
34. rida:
 
==Viirused==
Viirus nimega Sputnik nakatab [[amööb]]e, kuid ei suuda paljuneda kui Mimiviirus pole eelnevalt sama rakku nakatanud.<ref name="pmid18690211">{{cite journal | author = La Scola B, Desnues C, Pagnier I, Robert C, Barrassi L, Fournous G, Merchat M, Suzan-Monti M, Forterre P, Koonin E, Raoult D | title = The virophage as a unique parasite of the giant mimivirus | journal = Nature | volume = 455 | issue = 7209 | pages = 100–4 | year = 2008 | month = September | pmid = 18690211 | doi = 10.1038/nature07218 |bibcode = 2008Natur.455..100L }}</ref> Kuigi Sputniku 13. geenil on vähe ühist ühegi teise teadaoleva geeniga, siis kolm neist on lähedalt suguluses Mimi- ja Mamaviiruse geenidega. See võimaldab oletada, et satelliitviirus saab teostada horisontaalset ülekannet viiruste vahel analoogselt bakteriofaagidele, mis vahendavad geneetilist informatsiooni bakterite vahel.<ref>{{cite journal |authorname="Pearson H |title='Virophage' suggests viruses are alive |journal=Nature |volume=454 |issue=7205 |pages=677 |year=2008 |month=August |pmid=18685665 |doi=10.1038"/454677a |url=http://www.nature.com/news/2008/080806/full/454677a.html |bibcode = 2008Natur.454..677P }}</ref>
 
==Prokarüoodid==
Horisontaalne geeniülekanne on bakterite hulgas väga levinud, seda ka evolutsiooniliselt kaugete liikide vahel. Arvatakse, et see protsess on oluliseks põhjuseks kasvavale ravimresistentsusele. Üks bakterirakk, omandades resistentsuse, kannab vastavad geenid kiiresti üle teistele liikidele.<ref name="pmid19675017">{{cite journal |author=Hawkey PM, Jones AM |title=The changing epidemiology of resistance |journal=[[Journal of Antimicrobial Chemotherapy]] |volume=64 |issue=Suppl 1 |pages=i3–10 |year=2009 |month=September |pmid=19675017 |doi=10.1093/jac/dkp256 |url= }}</ref><ref name= FrancinoMP>{{cite book | author= Francino, MP (editor) | year=2012 | title=Horizontal Gene Transfer in Microorganisms | publisher=[[Caister Academic Press]] | isbn= 978-1-908230-10-2}}</ref> See protsess mängib rolli ka [[virulentsusfaktor]]ite, [[eksotoksiin]]ide ja [[eksoensüüm|-ensüümide]] levikus. On välja pakutud strateegiaid, võitlemaks kindlate [[bakteriaalne infektsioon|bakteriaalsete infektsioonide]] vastu, mille sihtmärgiks oleksid spetsiifilised virulentsusfaktorid ja [[transponeeruvad elemendid|mobiilsed geneetilised elemendid]].
 
==Eukarüoodid==
"Järjestuste võrdlemine viitab paljude geenide hiljutisele horisontaalsele ülekandele erinevate liikide vahel, hõlmates ka [[domeen_(bioloogia)|fülogeneetiliste domeenide]] vahelisi ülekandeid. Mis tähendab, et liikide [[fülogenees|fülogeneetilist]] ajalugu ei saa lõplikult määrata ainult üksikute geenide [[fülogeneesipuu|evolutsioonipuu]] järgi."<ref>[http://opbs.okstate.edu/~melcher/MG/MGW3/MG334.html okstate.edu]</ref>
 
* DNA järjestuste analüüs vihjab, et horisontaalne geeniülekanne on toimunud [[eukarüoot]]ides ka [[kloroplast]]ide ja [[mitokonder|mitokondrite]] [[genoom]]idest tuumagenoomi. [[Endosümbiootiline_teooria|Endosümbioositeooria]] väidab, et kloroplastid ja mitokondrid pärinevad rakkudest, kes on eukarüootide poolt [[fagotsütoos|fagotsüteeritud]] ja jäänud sinna [[endosümbiont]]ideks.<ref>{{cite journal |authorname="Blanchard JL, Lynch M |title=Organellar genes: why do they end up in the nucleus? |journal=Trends Genet. |volume=16 |issue=7 |pages=315–20 |year=2000 |month=July |pmid=10858662 |url=http://linkinghub.elsevier.com/retrieve/pii/S0168-9525(00)02053-9 |doi=10.1016/S0168-9525(00)02053-9 |notes=Discusses theories on how mitochondria and chloroplast genes are transferred into the nucleus, and also what steps a gene needs to go through in order to complete this process.}}<"/ref>
 
* Hästi on dokumenteeritud horisontaalset geeniülekannet bakteritelt [[seened|seentele]], eriti [[pagaripärm]]ile.<ref>{{cite journal |authorname="Hall C, Brachat S, Dietrich FS |title=Contribution of Horizontal Gene Transfer to the Evolution of ''Saccharomyces cerevisiae'' |journal=Eukaryotic Cell |volume=4 |issue=6 |pages=1102–15 |year=2005 |month=June |pmid=15947202 |pmc=1151995 |doi=10.1128/EC.4.6.1102-1115.2005 |url=http://ec.asm.org/cgi/content/full/4/6/1102}} The article argues that horizontal transfer of bacterial DNA to ''Saccharomyces cerevisiae'' has occurred.<"/ref>
 
* Arizona Ülikooli teadlased on leidnud, et herne-lehetäi (''Acyrthosiphon pisum'') genoom sisaldab mitmeid geene, mis on ülekandunud seentelt.<ref>{{cite journal |authorname="Fukatsu T |title=Evolution. A fungal past to insect color |journal=Science |volume=328 |issue=5978 |pages=574–5 |year=2010 |month=April |pmid=20431000 |doi=10.1126/science.1190417 |url=http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=20431000|bibcode = 2010Sci...328..574F }}<"/ref> Taimed, seened ja mikroorganismid suudavad sünteesida [[karotenoidid|karotenoide]], aga toruleeni sünteesivad herne-lehetäid on teadaolevalt ainukesed omataolised loomariigis.
 
* Hiljuti hüpotiseeriti, et [[malaaria]]t põhjustav [[patogeen]] ''[[Plasmodium vivax]]'' on horisontaalse ülekande abil omandanud geneetilist informatsiooni inimestelt, mis võib kaasa aidata tema pikaajalisele vastupidavusele inimkehas.<ref>{{cite web |authorname="Bar D |title=Evidence of Massive Horizontal Gene Transfer Between Humans and ''Plasmodium vivax'' |date=16 February 2011 |work=Nature Precedings |doi=10.1038/npre.2011.5690.1 |url=http://precedings.nature.com/documents/5690/version/1/}}<"/ref>
 
* 2012. aastal kirjeldati uut võimalikku horisontaalse geeniülekande mehhanismi, mida vahendas bakteriofaag ja mis toimus prokarüootide ja eukarüootide vahel.
 
* ''Hypothenemus hampei'' genoomis on geen HhMAN1, mis sarnaneb bakteriaalsete geenidega. Arvatakse, et see on pärit bakteritelt mardika seedekulglast.<ref>{{cite journal|journalname="Nature|year=2012|doi=10.1038"/nature.2012.10116|title=Bacterial gene helps coffee beetle get its fix|last1=Lee Phillips|first1=Melissa}}</ref><ref>{{cite journal|journalname="PNAS|title=Adaptive horizontal transfer of a bacterial gene to an invasive insect pest of coffee|year=2012|doi=10.1073"/pnas.1121190109}}</ref>
 
==Kunstlik horistonaalne geeniülekanne==
Insenergeneetika on põhimõtteliselt geenide ülekandmine horisontaalselt, kuigi selleks kasutatakse kunstlikke [[ekspressioonikassett]]e. "Uinuva kaunitari" transposoonsüsteem <ref>{{cite journal | author name= "Ivics Z., Hackett P.B., Plasterk R.H., Izsvak Z. | year = 1997 | title = Molecular reconstruction of ''Sleeping Beauty'', a Tc1-like transposon from fish, and its transposition in human cells |doi=10.1016"/S0092-8674(00)80436-5 | journal = Cell | volume = 91 | issue = 4| pages = 501–510 | pmid=9390559}}</ref> arendati välja sünteetiliseks geene ülekandvaks vahendajaks, mis baseerus teadaoleval Tc1/meremees transposoonide võimes sisse tungida väga erinevate liikide genoomidesse.<ref>{{cite journal |authorname="Plasterk RH |title=The Tc1"/mariner transposon family |journal=Curr. Top. Microbiol. Immunol. |volume=204 |pages=125–43 |year=1996 |pmid=8556864 }}</ref> Seda süsteemi on kasutatud geenijärjestuste sisendamiseks mitmete loomade genoomidesse.<ref>{{cite journal | author name= "Izsvak Z., Ivics Z., Plasterk R.H. | year = 2000 | title = Sleeping Beauty, a wide host-range transposon vector for genetic transformation in vertebrates | journal = J. Mol. Biol. | volume = 302 | issue = 1| pages = 93–102 | pmid=10964563 | doi=10.1006"/jmbi.2000.4047}}</ref><ref>{{cite journal |authorname="Kurtti TJ, Mattila JT, Herron MJ, ''et al.'' |title=Transgene expression and silencing in a tick cell line: A model system for functional tick genomics |journal=Insect Biochem. Mol. Biol. |volume=38 |issue=10 |pages=963–8 |year=2008 |month=October |pmid=18722527 |pmc=2581827 |doi=10.1016"/j.ibmb.2008.07.008 |url=http://linkinghub.elsevier.com/retrieve/pii/S0965-1748(08)00132-X}}</ref>
 
==Tähtsus evolutsioonis==
[[Image:Elu_puu.jpg|thumb|right|250px|Fülogeneetiline elu puu 16s rRNA subühiku analüüsi alusel.]]
Horisontaalne geeniülekanne on võimalikuks segavaks faktoriks fülogeneesipuude konstrueerimisel ühe geeni järjestuse järgi.<ref name="Lawton">Graham Lawton ''[http://www.newscientist.com/article/mg20126921.600-why-darwin-was-wrong-about-the-tree-of-life.html?page=1 Why Darwin was wrong about the tree of life]'' [[New Scientist]] Magazine issue 2692 21 January 2009 Accessed February 2009</ref> Näiteks kui kaks omavahel kaugelt suguluses olevat bakterit vahetavad geeni, siis neid liike sisaldaval puul märgitakse nad lähedasteks sugulasteks, kuna võrdluse aluseks olev geen on identne. Seda isegi siis, kui ülejäänud geenid on erinevad. Seetõttu on tihti parem kasutada teistsugust informatsiooni fülogeneesipuudelt järelduste tegemiseks, nagu geenide olemasolu või puudumise hindamine ja kaasata fülogeneetilisse analüüsi võimalikult lai valik geene.
 
Enimlevinud geeniks, mille põhjal koostatakse prokarüootide fülogeneetilisi suhteid, on 16S [[Ribosomaalne_RNA|rRNA]] geen. Selle järjestused on küllalt konserveerunud lähedaste sugulaste määramiseks, kuid samas piisavalt muutuvad, et erinevusi mõõta. Viimastel aastatel on arutletud, et ka need geenid võivad horisontaalselt üle kanduda. Ehkki see võib toimuda harva, tuleks siiski 16S rRNA põhjal konstrueeritud fülogeneesipuid uuesti hinnata.<ref>[http://ijs. name="sgmjournals.org/content/55/3/1021.full Genomic analysis of Hyphomonas neptunium contradicts 16S rRNA gene-based phylogenetic analysis: implications for the taxonomy of the orders ‘Rhodobacterales’ and Caulobacteral...]<"/ref>
 
Bioloog Johann Peter Gogarten soovitab, et hiljutiste genoomianalüüside valguses tuleks traditsiooniline "puu" asendada "mosaiigiga", mis kirjeldaks erineva ajalooga geenide kombinatsiooni genoomides. Ning "võrk" oleks sobivaks metafooriks visualiseerimaks tihedat geeniülekannet mikroobide vahel.
70. rida:
Leidub tõendeid, et ajalooliselt on toimunud horisontaalset ülekannet järgmiste geenide puhul:
 
* Lükopeeni tsüklaas, mis on vajalik karotenoidide [[biosüntees]]iks, rohelise [[väävlibakterid|väävlibakteri]] ja [[tsüanobakterid|tsüanobakteri]] vahel.<ref name="Bryant-Frigaard"/>
{{cite journal
|author=D.A. Bryant & N.-U. Frigaard
|month=November
|year=2006
|title=Prokaryotic photosynthesis and phototrophy illuminated
|journal=Trends Microbiol.
|volume=14
|issue=11
|pages=488–96
|doi=10.1016/j.tim.2006.09.001
|pmid=16997562
}}</ref>
 
* TetO geen, mis annab resistentsuse [[tetratsükliin]]ile, ''Campylobacter jejuni'' vahel.<ref>{{cite journal |author=Avrain L, Vernozy-Rozand C, Kempf I |title=Evidence for natural horizontal transfer of tetO gene between ''Campylobacter jejuni'' strains in chickens |journal=J. Appl. Microbiol. |volume=97 |issue=1 |pages=134–40 |year=2004 |pmid=15186450 |doi=10.1111/j.1365-2672.2004.02306.x |url=http://onlinelibrary.wiley.com/resolve/openurl?genre=article&sid=nlm:pubmed&issn=1364-5072&date=2004&volume=97&issue=1&spage=134}}</ref>
91. rida ⟶ 79. rida:
 
==Viited==
{{Viited}}|allikad=
<ref name="Transgenic">OECD, ''Safety Assessment of Transgenic Organisms, Volume 4: OECD Consensus Documents,'' 2010, pp.171–174</ref>
<ref name="Kay">{{cite journal |author=Kay E, Vogel TM, Bertolla F, Nalin R, Simonet P |title=In situ transfer of antibiotic resistance genes from transgenic (transplastomic) tobacco plants to bacteria |journal=Appl. Environ. Microbiol. |volume=68 |issue=7 |pages=3345–51 |year=2002 |month=July |pmid=12089013 |pmc=126776 |url=http://aem.asm.org/cgi/pmidlookup?view=long&pmid=12089013}}</ref>
<ref name="Koonin">{{cite journal |author=Koonin EV, Makarova KS, Aravind L |title=Horizontal gene transfer in prokaryotes: quantification and classification |journal=Annu. Rev. Microbiol. |volume=55 |pages=709–42 |year=2001 |pmid=11544372 |doi=10.1146/annurev.micro.55.1.709 |url=http://arjournals.annualreviews.org/doi/full/10.1146/annurev.micro.55.1.709?url_ver=Z39.88-2003&rfr_id=ori:rid:crossref.org&rfr_dat=cr_pub%3dpubmed}}</ref>
<ref name="Nielsen">{{cite journal |author=Nielsen KM |title=Barriers to horizontal gene transfer by natural transformation in soil bacteria |journal=APMIS Suppl. |volume=84 |pages=77–84 |year=1998 |pmid=9850687 }}</ref>
<ref name="McGowan">{{cite journal |author=McGowan C, Fulthorpe R, Wright A, Tiedje JM |title=Evidence for interspecies gene transfer in the evolution of 2,4-dichlorophenoxyacetic acid degraders |journal=Appl. Environ. Microbiol. |volume=64 |issue=10 |pages=4089–92 |year=1998 |month=October |pmid=9758850 |pmc=106609 |url=http://aem.asm.org/cgi/pmidlookup?view=long&pmid=9758850}}</ref>
<ref name="Blanchard">{{cite journal |author=Blanchard JL, Lynch M |title=Organellar genes: why do they end up in the nucleus? |journal=Trends Genet. |volume=16 |issue=7 |pages=315–20 |year=2000 |month=July |pmid=10858662 |url=http://linkinghub.elsevier.com/retrieve/pii/S0168-9525(00)02053-9 |doi=10.1016/S0168-9525(00)02053-9 |notes=Discusses theories on how mitochondria and chloroplast genes are transferred into the nucleus, and also what steps a gene needs to go through in order to complete this process.}}</ref>
<ref name="Naik">{{Cite journal |author=Naik GA, Bhat LN, Chpoade BA, Lynch JM |title=Transfer of broad-host-range antibiotic resistance plasmids in soil microcosms |journal=Curr. Microbiol. |volume=28 |year=1994 |pages=209–215 |doi=10.1007/BF01575963 |issue=4}}</ref>
<ref name="Russell">{{cite book |first=Peter J. |last=Russell |title=Biology : exploring the diversity of life|year=2009|publisher=Nelson Education|location=Toronto|isbn=0-17-644094-1|url=http://www.coursesmart.com/biology-exploring-the-diversity-of-life-1st/russell-wolfe-hertz-starr-fenton-addy-maxwell/dp/9780176440947|edition=1st Canadian}}</ref>
<ref name="Freeman">{{cite journal | author = Victor J Freeman | title = Studies on the virulence of bacteriophage-infected strains of ''Corynebacterium Diphtheriae'' | journal = Journal of Bacteriology | year = 1951 | pages = 675–688 | pmid = 14850426 | volume = 61 | issue = 6 | pmc=386063}}</ref>
<ref name="Marguilies">Phillip Marguilies "Epidemics: Deadly diseases throughout history". Rosen, New York. 2005.</ref>
<ref name="edwards">{{cite web|url=http://www.physorg.com/news205389256.html|title=Horizontal gene transfer in microbes much more frequent than previously thought|author=Lin Edwards|date= October 4, 2010|publisher=PhysOrg.com|accessdate=2012-01-06}}</ref>
<ref name="arnold">{{cite web|url=http://www.scientificamerican.com/article.cfm?id=to-share-and-share-alike|title=To Share and Share Alike: Bacteria swap genes with their neighbors more frequently than researchers have realized|author=Carrie Arnold|date= April 18, 2011|publisher=Scientific American|accessdate=2012-01-06}}</ref>
<ref name="Lwoff">André Lwoff (1965). [http://nobelprize.org/nobel_prizes/medicine/laureates/1965/lwoff-lecture.html "Interaction among Virus, Cell, and Organism"]. Nobel Lecture for the Nobel Prize in Physiology or Medicine.</ref>
<ref name="Ochiai">{{cite journal |author=Ochiai K, Yamanaka T, Kimura K, Sawada, O |title=Inheritance of drug resistance (and its transfer) between Shigella strains and Between Shigella and E. coli strains |journal=Hihon Iji Shimpor |volume=1861 |pages=34 |year=1959 |language=Japanese}}</ref>
<ref name="Akiba">{{cite journal |author=Akiba T, Koyama K, Ishiki Y, Kimura S, Fukushima T |title=On the mechanism of the development of multiple-drug-resistant clones of Shigella |journal=Jpn. J. Microbiol. |volume=4 |pages=219–27 |year=1960 |month=April |pmid=13681921 }}</ref>
<ref name="Syvanen">{{cite journal |author=Syvanen M |title=Cross-species gene transfer; implications for a new theory of evolution |journal=J. Theor. Biol. |volume=112 |issue=2 |pages=333–43 |year=1985 |month=January |pmid=2984477 |doi=10.1016/S0022-5193(85)80291-5 |url=http://www.dcn.davis.ca.us/vme/hgt/JTheoBiolvol112pp333-343yr1985.PDF |format=PDF}}</ref>
<ref name="Jain">{{cite journal |author=Jain R, Rivera MC, Lake JA |title=Horizontal gene transfer among genomes: The complexity hypothesis |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=96 |issue=7 |pages=3801–6 |year=1999 |month=March |pmid=10097118 |pmc=22375 |doi= 10.1073/pnas.96.7.3801|url=http://www.pnas.org/cgi/pmidlookup?view=long&pmid=10097118|bibcode = 1999PNAS...96.3801J }}</ref>
<ref name="Bapteste">{{cite journal |author=Bapteste E, Susko E, Leigh J, MacLeod D, Charlebois RL, Doolittle WF |title=Do orthologous gene phylogenies really support tree-thinking? |journal=BMC Evol. Biol. |volume=5 |pages=33 |year=2005 |pmid=15913459 |pmc=1156881 |doi=10.1186/1471-2148-5-33 |issue=1}}</ref>
<ref name="Richardson">{{cite journal | author = Richardson, Aaron O.; Palmer, Jeffrey D. | month = January | year = 2007 | title = Horizontal Gene Transfer in Plants |journal = Journal of Experimental Botany | volume = 58 | pages = 1–9 |url=http://www.sdsc.edu/~shindyal/ejc121304.pdf |format=PDF | doi = 10.1093/jxb/erl148 | pmid = 17030541 | issue = 1}}</ref>
<ref name="gogarten">{{cite journal | author = Gogarten, Peter | year = 2000 | title = Horizontal Gene Transfer: A New Paradigm for Biology | journal = Esalen Center for Theory and Research Conference | url = http://www.esalenctr.org/display/confpage.cfm?confid=10&pageid=105&pgtype=1 | accessdate = 2007-03-18}}</ref>
<ref name="Mae-Wan Ho">{{Cite journal |author=Mae-Wan Ho |year=1999 |title=Cauliflower Mosaic Viral Promoter – A Recipe for Disaster? |journal=Microbial Ecology in Health and Disease |volume=11 |url=http://www.i-sis.org.uk/pdf/CaMV_promoter_recipe_for_disaster.pdf |accessdate=2008-06-09 |pages=194–7}}</ref>
▲Horisontaalne geeniülekanne võib toimuda mitme erineva mehhanismi kaudu:<ref name="todar">{{cite web|url=http://textbookofbacteriology.net/themicrobialworld/bactresanti.html|title=Bacterial Resistance to Antibiotics|author=Kenneth Todar |publisher=The Microbial World: Lectures in Microbiology, Department of Bacteriology, University of Wisconsin-Madison |accessdate=January 6, 2012}}</ref><ref name="maloy">{{cite web|url=http://www.sci.sdsu.edu/~smaloy/MicrobialGenetics/topics/genetic-exchange/exchange/exchange.html|title=Horizontal Gene Transfer|author=Stanley Maloy|date= July 15, 2002 |publisher=San Diego State University|accessdate=January 6, 2012}}</ref>
<ref name="maloy">{{cite web|url=http://www.sci.sdsu.edu/~smaloy/MicrobialGenetics/topics/genetic-exchange/exchange/exchange.html|title=Horizontal Gene Transfer|author=Stanley Maloy|date= July 15, 2002 |publisher=San Diego State University|accessdate=January 6, 2012}}</ref>
<ref name="pmid18690211">{{cite journal | author = La Scola B, Desnues C, Pagnier I, Robert C, Barrassi L, Fournous G, Merchat M, Suzan-Monti M, Forterre P, Koonin E, Raoult D | title = The virophage as a unique parasite of the giant mimivirus | journal = Nature | volume = 455 | issue = 7209 | pages = 100–4 | year = 2008 | month = September | pmid = 18690211 | doi = 10.1038/nature07218 |bibcode = 2008Natur.455..100L }}</ref>
<ref name="Pearson">{{cite journal |author=Pearson H |title='Virophage' suggests viruses are alive |journal=Nature |volume=454 |issue=7205 |pages=677 |year=2008 |month=August |pmid=18685665 |doi=10.1038/454677a |url=http://www.nature.com/news/2008/080806/full/454677a.html |bibcode = 2008Natur.454..677P }}</ref>
<ref name="pmid19675017">{{cite journal |author=Hawkey PM, Jones AM |title=The changing epidemiology of resistance |journal=[[Journal of Antimicrobial Chemotherapy]] |volume=64 |issue=Suppl 1 |pages=i3–10 |year=2009 |month=September |pmid=19675017 |doi=10.1093/jac/dkp256 |url= }}</ref>
<ref name= FrancinoMP>{{cite book | author= Francino, MP (editor) | year=2012 | title=Horizontal Gene Transfer in Microorganisms | publisher=[[Caister Academic Press]] | isbn= 978-1-908230-10-2}}</ref>
<ref name="Hall">{{cite journal |author=Hall C, Brachat S, Dietrich FS |title=Contribution of Horizontal Gene Transfer to the Evolution of ''Saccharomyces cerevisiae'' |journal=Eukaryotic Cell |volume=4 |issue=6 |pages=1102–15 |year=2005 |month=June |pmid=15947202 |pmc=1151995 |doi=10.1128/EC.4.6.1102-1115.2005 |url=http://ec.asm.org/cgi/content/full/4/6/1102}} The article argues that horizontal transfer of bacterial DNA to ''Saccharomyces cerevisiae'' has occurred.</ref>
<ref name="Fukatsu">{{cite journal |author=Fukatsu T |title=Evolution. A fungal past to insect color |journal=Science |volume=328 |issue=5978 |pages=574–5 |year=2010 |month=April |pmid=20431000 |doi=10.1126/science.1190417 |url=http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=20431000|bibcode = 2010Sci...328..574F }}</ref>
<ref name="Bar">{{cite web |author=Bar D |title=Evidence of Massive Horizontal Gene Transfer Between Humans and ''Plasmodium vivax'' |date=16 February 2011 |work=Nature Precedings |doi=10.1038/npre.2011.5690.1 |url=http://precedings.nature.com/documents/5690/version/1/}}</ref>
<ref name="Nature">{{cite journal|journal=Nature|year=2012|doi=10.1038/nature.2012.10116|title=Bacterial gene helps coffee beetle get its fix|last1=Lee Phillips|first1=Melissa}}</ref>
<ref name="PNAS">{{cite journal|journal=PNAS|title=Adaptive horizontal transfer of a bacterial gene to an invasive insect pest of coffee|year=2012|doi=10.1073/pnas.1121190109}}</ref>
<ref name="Ivics">{{cite journal | author = Ivics Z., Hackett P.B., Plasterk R.H., Izsvak Z. | year = 1997 | title = Molecular reconstruction of ''Sleeping Beauty'', a Tc1-like transposon from fish, and its transposition in human cells |doi=10.1016/S0092-8674(00)80436-5 | journal = Cell | volume = 91 | issue = 4| pages = 501–510 | pmid=9390559}}</ref>
<ref name="Plasterk">{{cite journal |author=Plasterk RH |title=The Tc1/mariner transposon family |journal=Curr. Top. Microbiol. Immunol. |volume=204 |pages=125–43 |year=1996 |pmid=8556864 }}</ref>
<ref name="Izsvak">{{cite journal | author = Izsvak Z., Ivics Z., Plasterk R.H. | year = 2000 | title = Sleeping Beauty, a wide host-range transposon vector for genetic transformation in vertebrates | journal = J. Mol. Biol. | volume = 302 | issue = 1| pages = 93–102 | pmid=10964563 | doi=10.1006/jmbi.2000.4047}}</ref>
<ref name="Kurtti">{{cite journal |author=Kurtti TJ, Mattila JT, Herron MJ, ''et al.'' |title=Transgene expression and silencing in a tick cell line: A model system for functional tick genomics |journal=Insect Biochem. Mol. Biol. |volume=38 |issue=10 |pages=963–8 |year=2008 |month=October |pmid=18722527 |pmc=2581827 |doi=10.1016/j.ibmb.2008.07.008 |url=http://linkinghub.elsevier.com/retrieve/pii/S0965-1748(08)00132-X}}</ref>
<ref name="Lawton">Graham Lawton ''[http://www.newscientist.com/article/mg20126921.600-why-darwin-was-wrong-about-the-tree-of-life.html?page=1 Why Darwin was wrong about the tree of life]'' [[New Scientist]] Magazine issue 2692 21 January 2009 Accessed February 2009</ref>
<ref name="sgmjournals">[http://ijs.sgmjournals.org/content/55/3/1021.full Genomic analysis of Hyphomonas neptunium contradicts 16S rRNA gene-based phylogenetic analysis: implications for the taxonomy of the orders ‘Rhodobacterales’ and Caulobacteral...]</ref>
<ref name="Bryant-Frigaard">{{cite journal|author=D.A. Bryant & N.-U. Frigaard|month=November|year=2006|title=Prokaryotic photosynthesis and phototrophy illuminated|journal=Trends Microbiol.|volume=14|issue=11|pages=488–96|doi=10.1016/j.tim.2006.09.001|pmid=16997562}}</ref>
 
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