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{{short description|Evolutionary consequences of transfer of genetic material between organisms of different taxa}}
[[Image:Phylogenetic tree.svg|thumb|right|400px|A three-domain [[phylogenetic tree|tree]] of life showing the separation of [[Bacteria]], [[Archaea]], and [[Eukaryote]] domains. See [[Microorganisms]] article for further explanation.]]{{Essay|date=November 2023}}
'''[[Horizontal gene transfer]]''' ('''HGT''') refers to the transfer of genes between distant branches on the [[tree of life (biology)|tree of life]]. In '''evolution''', it can scramble the information needed to reconstruct the phylogeny of [[Organism|organisms]], how they are related to one another.

HGT can also help scientists to reconstruct and date the tree of life, as a gene transfer can be used as a phylogenetic marker, or as the proof of contemporaneity of the donor and recipient organisms, and as a trace of extinct biodiversity.

HGT happens very infrequently – at the individual organism level, it is highly improbable for any such event to take place. However, on the grander scale of evolutionary history, these events occur with some regularity. On one hand, this forces biologists to abandon the use of individual genes as good markers for the history of life. On the other hand, this provides an almost unexploited large source of information about the past.

== Three domains of life ==

The three main early branches of the [[Tree of life (biology)|tree of life]] have been intensively studied by [[Microorganism|microbiologists]] because the first organisms were microorganisms. Microbiologists (led by [[Carl Woese]]) have introduced the term ''[[Domain (biology)|domain]]'' for the three main branches of this tree, where ''domain'' is a phylogenetic term similar in meaning to biological [[Kingdom (biology)|kingdom]]. To reconstruct this tree of life, the gene sequence encoding the small subunit of [[ribosome|ribosomal]] [[RNA]] (SSU rRNA, [[16s rRNA]]) has proven useful, and the tree (as shown in the picture) relies heavily on information from this single gene.

These three domains of life represent the main evolutionary lineages of early cellular life and currently include [[Bacteria]], [[Archaea]] (single-celled organisms superficially similar to bacteria), and [[Eukaryote|Eukarya]]. Eukarya includes only organisms having a well-defined nucleus, such as [[fungi]], [[protist]]s, and all organisms in the plant and animals kingdoms (see figure).

The gene most commonly used for constructing phylogenetic relationships in [[prokaryote|microorganisms]] is the small subunit ribosomal RNA gene, as its sequences tend to be conserved among members with close phylogenetic distances, yet variable enough that differences can be measured.
<ref>{{cite journal |vauthors=Woese C, etal | title = Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya| journal = PNAS USA| volume = 87 | pages = 4576–9 | year = 1990 | pmid = 2112744 | doi = 10.1073/pnas.87.12.4576 | issue = 12 | pmc = 54159 | bibcode=1990PNAS...87.4576W| doi-access = free}}
*{{cite journal | author = Woese C, Fox G | title = Phylogenetic structure of the prokaryotic domain: the primary kingdoms|journal = PNAS USA |volume = 74 |pages = 5088–90 | year = 1977 | pmid = 270744 | doi = 10.1073/pnas.74.11.5088 | issue = 11 | pmc = 432104|bibcode = 1977PNAS...74.5088W | doi-access = free}}</ref>
The [[SSU rRNA]] as a measure of evolutionary distances was pioneered by [[Carl Woese]] when formulating the first modern "tree of life", and his results led him to propose the [[Archaea]] as a third domain of [[life]]. However, recently it has been argued that SSU rRNA genes can also be horizontally transferred.<ref>{{cite journal | last1 = Yap | first1 = WH | last2 = Zhang | first2 = Z | last3 = Wang | first3 = Y | title = Distinct types of rRNA operons exist in the genome of the actinomycete Thermomonospora chromogena and evidence for horizontal transfer of an entire rRNA operon. | journal = Journal of Bacteriology | volume = 181 | issue = 17 | pages = 5201–9 | year = 1999 | doi = 10.1128/JB.181.17.5201-5209.1999 | pmid = 10464188 | pmc = 94023 }}</ref> Although this may be rare, this possibility is forcing scrutiny of the validity of phylogenetic trees based on SSU rRNAs.

Recent discoveries of "rampant" HGT in microorganisms, and the detection of horizontal movement of even genes for the small subunit of ribosomal RNA, have forced biologists to question the accuracy of at least the early branches in the tree, and even question the validity of trees as useful models of how early evolution occurs.<ref name="Simonson2005">{{cite journal | last1 = Simonson | first1 = AB | last2 = Servin | first2 = JA | last3 = Skophammer | first3 = RG | last4 = Herbold | first4 = CW | last5 = Rivera | first5 = MC | last6 = Lake | first6 = JA | title = Decoding the genomic tree of life. | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 102 | pages = 6608–13 | year = 2005 | issue = Suppl 1 | pmid = 15851667 | pmc = 1131872 | doi = 10.1073/pnas.0501996102 |bibcode = 2005PNAS..102.6608S | doi-access = free }}</ref> In fact, early evolution is considered to have occurred starting from a community of [[progenote]]s, able to exchange large molecules when HGT was the standard. This lateral gene transfer occurred also beyond the [[Darwinian threshold]], after heredity or vertical gene transfer was established.<ref>{{Cite journal|last=Woese|first=C.|date=1998-06-09|title=The universal ancestor|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=95|issue=12|pages=6854–6859|issn=0027-8424|pmid=9618502|pmc=22660|doi=10.1073/pnas.95.12.6854|bibcode=1998PNAS...95.6854W|doi-access=free}}</ref><ref>{{Cite journal|last=Woese|first=Carl R.|date=2002-06-25|title=On the evolution of cells|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=99|issue=13|pages=8742–8747|doi=10.1073/pnas.132266999|issn=0027-8424|pmid=12077305|pmc=124369|bibcode=2002PNAS...99.8742W|doi-access=free}}</ref><br>
"Sequence comparisons suggest recent horizontal transfer of many [[gene]]s among diverse [[species]] including across the boundaries of [[phylogenetic]] "domains". Thus determining the phylogenetic history of a species can not be done conclusively by determining evolutionary trees for single genes."<ref name=":0">[https://web.archive.org/web/19990220101406/http://opbs.okstate.edu/%7Emelcher/MG/MGW3/MG334.html Horizontal Gene Transfer, Oklahoma State]</ref> HGT is thus a potential [[Lurking variable|confounding factor]] in inferring [[phylogenetic tree]]s from the [[sequence]] of one [[gene]]. For example, if two distantly related bacteria have exchanged a gene, a [[phylogenetic tree]] including those species will show them to be closely related even though most other genes have diverged substantially. For this reason it is important to use other information to infer phylogenies, such as the presence or absence of genes, or, more commonly, to include as wide a range of genes for analysis as possible.<ref name=":0" />

Earlier HGTs are thought to have happened. The [[first universal common ancestor]] (FUCA), earliest ancestor of the [[Last universal common ancestor|last common ancestor to all life]] (LUCA), is thought to have had other descendants that had their own lineages.<ref name=":03">{{Citation |last1=Prosdocimi |first1=Francisco |title=The First Universal Common Ancestor (FUCA) as the Earliest Ancestor of LUCA's (Last UCA) Lineage |date=2019 |url=https://doi.org/10.1007/978-3-030-30363-1_3 |work=Evolution, Origin of Life, Concepts and Methods |pages=43–54 |editor-last=Pontarotti |editor-first=Pierre |access-date=2023-11-02 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-30363-1_3 |isbn=978-3-030-30363-1 |s2cid=199534387 |last2=José |first2=Marco V. |last3=de Farias |first3=Sávio Torres|url-access=subscription }}</ref><ref name=":1">{{Cite journal |last1=Harris |first1=Hugh M. B. |last2=Hill |first2=Colin |date=2021 |title=A Place for Viruses on the Tree of Life |journal=Frontiers in Microbiology |volume=11 |doi=10.3389/fmicb.2020.604048 |issn=1664-302X |pmc=7840587 |pmid=33519747 |doi-access=free}}</ref> These now-extinct sister lineages of LUCA descending from FUCA are thought to have horizontally transferred some of their genes into the genome of early descendants of LUCA.<ref name=":1" />

==Choice of metaphor: tree, net, cobweb, or ring==
In his article ''Uprooting the Tree of Life'', W. [[Ford Doolittle]] discusses the [[Last universal ancestor|Last Universal Common Ancestor]] – the root of the Tree of Life – and the problems with that concept posed by HGT.<ref name=":2">{{cite journal | doi = 10.1038/scientificamerican0200-90 | last1 = Doolittle | first1 = WF | year = 2000 | title = Uprooting the tree of life | journal = Sci Am | volume = 282 | issue = 2| pages = 90–5 | pmid = 10710791 | bibcode = 2000SciAm.282b..90D }}</ref> He describes the microorganism ''[[Archaeoglobus fulgidus]]'' as an anomaly with respect to a [[phylogenetic]] tree based upon the code for the [[enzyme]] [[HMGCoA reductase]] – the organism is definitely an archaean, with all the cell lipids and transcription machinery expected of an archaean, but its HMGCoA genes are of bacterial origin. In the article, Doolittle says that while it is now widely accepted that [[mitochondria]] in eukaryotes derived from alpha-proteobacterial cells and that [[chloroplast]]s came from ingested [[cyanobacteria]],<ref name=":2" />

<blockquote>
".. it is no longer safe to assume that those were the only lateral gene transfers that occurred after the first eukaryotes arose. Only in later, multicellular eukaryotes do we know of definite restrictions on horizontal gene exchange, such as the advent of separated (and protected) [[germ cell]]s...

If there had never been any lateral gene transfer, all these individual gene trees would have the same topology (the same branching order), and the ancestral genes at the root of each tree would have all been present in the last universal common ancestor, a single ancient cell. But extensive transfer means that neither is the case: gene trees will differ (although many will have regions of similar topology) ''and'' there would never have been a single cell that could be called the last universal common ancestor..."<ref name=":2" />
</blockquote>

Doolittle suggested that the universal common ancestor cannot have been one particular organism, but must have been a loose, diverse conglomeration of primitive cells that evolved together. These early cells, each with relatively few genes, differed in many ways, and swapped their [[gene]]s freely. Eventually, from these eclectic cells came the three domains of life as we know them today: [[bacteria]], [[archaea]] and [[eukaryote]]. These domains are now recognizably distinct because much of the gene transfer that still occurs is within these domains, rather than between them. Biologist [[Johann Peter Gogarten|Peter Gogarten]] reinforced these arguments, and suggested that the metaphor of a tree does not fit the data from recent genome research, and that biologists should now instead use "the metaphor of a mosaic to describe the different histories combined in individual genomes and use [the] metaphor of a net to visualize the rich exchange and cooperative effects of HGT among microbes."<ref>Gogarten JP 'Horizontal Gene Transfer – A New Paradigm for Biology' [http://www.esalenctr.org/display/confpage.cfm?confid=10&pageid=105&pgtype=1 PhD thesis] {{Webarchive|url=https://web.archive.org/web/20120721232310/http://www.esalenctr.org/display/confpage.cfm?confid=10&pageid=105&pgtype=1 |date=2012-07-21 }}
*{{cite journal |last1=Zhaxybayeva |first1=O |last2=Gogarten |first2=JP |title=Cladogenesis, coalescence and the evolution of the three domains of life. |url=http://web.uconn.edu/gogarten/articles/TIG2004_cladogenesis_paper.pdf |journal=Trends in Genetics |volume=20 |issue=4 |pages=182–7 |year=2004 |pmid=15041172 |doi=10.1016/j.tig.2004.02.004 |access-date=2009-07-10 |archive-date=2009-03-26 |archive-url=https://web.archive.org/web/20090326181517/http://web.uconn.edu/gogarten/articles/TIG2004_cladogenesis_paper.pdf |url-status=live }}</ref>

==Resolution of uncertainty with phylogenomics==
[[Image:Tree microbialgenomes nocobwebs.jpg|none|frame|Tree Based on the Median Tree Algorithm.<ref>{{Cite journal| last1 = Ge | first1 = F.| last2 = Wang | first2 = L.| last3 = Kim | first3 = J.| title = The cobweb of life revealed by genome-scale estimates of horizontal gene transfer| journal = PLOS Biology| volume = 3| issue = 10| pages = e316| year = 2005| pmid = 16122348| pmc = 1233574| doi = 10.1371/journal.pbio.0030316| doi-access = free}} {{open access}}</ref> Branches with [[Bootstrapping (statistics)|bootstrap]] (statistical) scores less than 50% were collapsed. The three domains of life are (A) archaea, (B–J) bacteria, and (K) eukaryote. Species are labeled with different colors based on their inferred HGT rates: red, >4%; yellow, 3–4%; pink, 2–3%; blue, 1–2%; green, <1%. Taxonomy labels are (A) [[Euryarchaeota|Euryarchaea]], (B) [[Pseudomonadota|Proteobacteria]], (C) [[Chlamydiae]], (D) [[Spirochaete|Spirochaetes]], (E) [[Thermotogae]], (F) [[Aquificae]], (G) [[Actinobacteria]], (H) [[Deinococcus]], (I) [[Cyanobacteria]], (J) [[Firmicutes]], and (K) [[Fungus|Fungi]].]]

Despite the uncertainties in reconstructing phylogenies back to the beginnings of life, progress is being made in reconstructing the tree of life in the face of uncertainties raised by HGT. The uncertainty of any inferred phylogenetic tree based on a single gene can be resolved by using several common genes or even evidence from whole genomes.<ref>{{Cite journal
| last1 = Henz | first1 = S.
| last2 = Huson | first2 = D.
| last3 = Auch | first3 = A.
| last4 = Nieselt-Struwe | first4 = K.
| last5 = Schuster | first5 = S.
| title = Whole-genome prokaryotic phylogeny
| journal = Bioinformatics
| volume = 21
| issue = 10
| pages = 2329–2335
| year = 2005
| pmid = 15166018
| doi = 10.1093/bioinformatics/bth324
| doi-access = free
}}
*{{cite journal|doi=10.1186/1471-2148-6-99|last1=Fitzpatrick|first1=DA|last2=Logue|first2=ME|last3=Stajich|first3=JE|last4=Butler|first4=G|title=A fungal phylogeny based on 42 complete genomes derived from supertree and combined gene analysis.|journal=BMC Evolutionary Biology|volume=6|pages=99|year=2006 |pmid = 17121679|pmc=1679813 |doi-access=free }} {{open access}}</ref> One such approach, sometimes called 'multi-locus typing', has been used to deduce phylogenic trees for organisms that exchange genes, such as meningitis bacteria.<ref>{{cite journal | doi = 10.1016/j.tim.2003.08.006 | last1 = Urwin | first1 = R | last2 = Maiden | first2 = MC | year = 2003 | title = Multi-locus sequence typing: a tool for global epidemiology | journal = Trends Microbiol | volume = 11 | issue = 10| pages = 479–87 | pmid = 14557031 }}
*{{cite journal | last1=Yang | first1=Z | title=Likelihood and Bayes estimation of ancestral population sizes in hominoids using data from multiple loci. | url=http://www.genetics.org/cgi/content/full/162/4/1811 | journal=Genetics | volume=162 | issue=4 | pages=1811–23 | year=2002 | doi=10.1093/genetics/162.4.1811 | pmid=12524351 | pmc=1462394 | access-date=2009-07-10 | archive-date=2009-01-06 | archive-url=https://web.archive.org/web/20090106142510/http://www.genetics.org/cgi/content/full/162/4/1811 | url-status=live }}
*{{Cite journal
| last1 = Jennings | first1 = W. B.
| last2 = Edwards | first2 = S. V.
| title = Speciational history of Australian grass finches (Poephila) inferred from thirty gene trees
| journal = Evolution; International Journal of Organic Evolution
| volume = 59
| issue = 9
| pages = 2033–2047
| year = 2005
| pmid = 16261740
| doi=10.1554/05-280.1
| s2cid = 18665436
}}</ref>

[[Jonathan Eisen]] and [[Claire M. Fraser|Claire Fraser]] have pointed out that:
<blockquote>"In building the tree of life, analysis of whole genomes has begun to supplement, and in some cases to improve upon, studies previously done with one or a few genes. For example, recent studies of complete bacterial genomes have suggested that the [[Hyperthermophile|hyperthermophilic]] species are not deeply branching; if this is true, it casts doubt on the idea that the first forms of life were thermophiles. Analysis of the genome of the eukaryotic parasite ''[[Encephalitozoon cuniculi]]'' supports suggestions that the group ''[[Microsporidia]]'' are not deep branching protists but are in fact members of the fungal kingdom. Genome analysis can even help resolve relationships within species, such as by providing new genetic markers for population genetics studies in the bacteria causing anthrax or tuberculosis. In all these studies, it is the additional data provided by a complete genome sequence that allows one to separate the phylogenetic signal from the noise. This is not to say the tree of life is now resolved – we only have sampled a smattering of genomes, and many groups are not yet touched"<ref>{{Cite journal| last1 = Eisen | first1 = J.| last2 = Fraser | first2 = C.| title = Phylogenomics: intersection of evolution and genomics| journal = Science| volume = 300| issue = 5626| pages = 1706–1707| year = 2003| pmid = 12805538| doi = 10.1126/science.1086292|bibcode = 2003Sci...300.1706E | s2cid = 42394233}}</ref></blockquote>

These approaches are enabling estimates of the relative frequency of HGT; the relatively low values that have been observed suggests that the 'tree' is still a valid metaphor for evolution – but the tree is adorned with 'cobwebs' of horizontally transferred genes. This is the main conclusion of a 2005 study of more than 40 complete microbial genomic sequences by Fan Ge, Li-San Wang, and [[Junhyong Kim]]. They estimate the frequency of HGT events at about 2% of core genes per genome.<ref>{{cite journal | last1 = Ge | first1 = Fan | last2 = Wang | first2 = Li-San | last3 = Kim | first3 = Junhyong | title = The Cobweb of Life Revealed by Genome-Scale Estimates of Horizontal Gene Transfer | journal = PLOS Biology | volume = 3 | issue = 10 | pages = e316 | year = 2005 | doi = 10.1371/journal.pbio.0030316 | pmc = 1233574 | pmid=16122348 | doi-access = free }} {{open access}}</ref> Similar whole genome approaches to assessing evolution are also enabling progress in identifying very early events in the tree of life, such as a proposal that eukaryotes arose by fusion of two complete but very diverse prokaryote genomes: one from a bacterium and one from an archaeal cell.<ref name="Simonson2005" />

Such a fusion of organisms [[hypothesis]] for the origin of complex nucleated cells has been put forward by [[Lynn Margulis]] using quite different reasoning about [[symbiosis]] between a bacterium and an archaen arising in an ancient consortium of microbes.<ref>''Acquiring genomes: a theory of the origin of species.'' Margulis L and Sagan D (2002) Basic Books {{ISBN|0-465-04392-5}}</ref>

== Use in evolutionary studies ==
While HGT is often seen as a challenge for the reconstruction of the tree of life, an alternative view is that oppositely it provides additional valuable information for its reconstruction.

First, for the recipient organism, HGT is a [[DNA mutation]] like others, and as such, it can be modeled and used in tree reconstruction and rooting.<ref>{{Cite journal|last1=Abby|first1=Sophie S.|last2=Tannier|first2=Eric|last3=Gouy|first3=Manolo|last4=Daubin|first4=Vincent|date=2012-03-27|title=Lateral gene transfer as a support for the tree of life|journal=Proceedings of the National Academy of Sciences|volume=109|issue=13|pages=4962–4967|doi=10.1073/pnas.1116871109|pmid=22416123|pmc=3323970|bibcode=2012PNAS..109.4962A|doi-access=free}}</ref>

Second, it is necessary that the recipient of a gene acquisition by HGT lives at the same time, or at an ulterior time, as the donor.<ref>{{Cite book|title=Horizontal Gene Transfer|journal= |volume=532|last1=Huang|first1=Jinling|last2=Gogarten|first2=Johann Peter|date=2009|publisher=Humana Press|isbn=9781603278522|series=Methods in Molecular Biology|pages=127–139|language=en|doi=10.1007/978-1-60327-853-9_7|pmid = 19271182}}</ref> In consequence there is an information on the timing of diversification in HGT.<ref>{{Cite journal|last1=Davín|first1=Adrián A.|last2=Tannier|first2=Eric|last3=Williams|first3=Tom A.|last4=Boussau|first4=Bastien|last5=Daubin|first5=Vincent|last6=Szöllősi|first6=Gergely J.|date=2018-04-02|title=Gene transfers can date the tree of life|journal=Nature Ecology & Evolution|volume=2|issue=5|pages=904–909|language=En|doi=10.1038/s41559-018-0525-3|pmid=29610471|pmc=5912509|bibcode=2018NatEE...2..904D |issn=2397-334X}}</ref> This is all the more remarkable since the principal usual source for dating in the living world, the fossil record, is absent precisely where HGT is abundant, in the microbial world.

Third, it provides information about the extinct biodiversity, because transfers are likely from extinct species.<ref>{{Cite journal|last1=Szöllősi|first1=Gergely J.|last2=Tannier|first2=Eric|last3=Lartillot|first3=Nicolas|last4=Daubin|first4=Vincent|date=2013-05-01|title=Lateral Gene Transfer from the Dead|journal=Systematic Biology|language=en|volume=62|issue=3|pages=386–397|doi=10.1093/sysbio/syt003|pmid=23355531|pmc=3622898|issn=1063-5157|arxiv=1211.4606}}</ref>

==See also==
*[[Evolutionary developmental biology]]
*[[Horizontal gene transfer]]
*[[Inferring horizontal gene transfer]]
*[[Tree of life (biology)]]

==References==
{{Reflist|30em}}
*{{citizendium|title=Prokaryote phylogeny and evolution}}

==Further reading==
* {{cite journal | last1= Gogarten | first1= JP | last2= Townsend | first2= JP | title= Horizontal gene transfer, genome innovation and evolution. | journal= Nature Reviews. Microbiology | volume= 3 | issue= 9 | pages= 679–87 | year= 2005 | pmid= 16138096 | doi= 10.1038/nrmicro1204 | s2cid= 8092665 }}. One article in a whole issue of the journal ''Nature Reviews Microbiology'' largely devoted to HGT.
* {{cite journal | doi = 10.1126/science.1086568 | last1 = Daubin | first1 = V | last2 = Moran | year = 2003 | first2 = NA | last3 = Ochman | first3 = H | title = Phylogenetics and the cohesion of bacterial genomes | journal = Science | volume = 301 | issue = 5634| pages = 829–32 | pmid = 12907801 |bibcode = 2003Sci...301..829D | s2cid = 11268678 }}
* {{cite journal | doi = 10.1126/science.1086292 | last1 = Eisen | first1 = JA | last2 = Fraser | first2 = CM | year = 2003 | title = Viewpoint phylogenomics: intersection of evolution and genomics | journal = Science | volume = 300 | issue = 5626| pages = 1706–7 | pmid = 12805538 | bibcode=2003Sci...300.1706E| s2cid = 42394233 }}
* {{cite journal | last1 = Ge | first1 = F | year = 2005 | last2 = Wang | first2 = LS | last3 = Kim | first3 = J | title = The Cobweb of Life revealed by genome-scale estimates of horizontal gene transfer | journal = PLOS Biology | volume = 3 | issue = 10| page = e316 | pmid = 16122348 | pmc = 1233574 | doi = 10.1371/journal.pbio.0030316 | doi-access = free }} {{open access}}
* {{cite journal | doi = 10.1093/bioinformatics/bth324 | last1 = Henz | first1 = SR | year = 2005 | last2 = Huson | first2 = DH | last3 = Auch | first3 = AF | last4 = Nieselt-Struwe | first4 = K | last5 = Schuster | first5 = SC | title = Whole-genome prokaryotic phylogeny | journal = Bioinformatics | volume = 21 | issue = 10| pages = 2329–35 | pmid = 15166018 | doi-access = free }}
* {{cite journal | last1 = Lerat | first1 = E | last2 = Daubin | year = 2005 | first2 = V | last3 = Ochman | first3 = H | last4 = Moran | first4 = NA | title = Evolutionary origins of genomic repertoires in bacteria | journal = PLOS Biology | volume = 3 | issue = 5| page = e130 | pmid = 15799709 | pmc = 1073693 | doi = 10.1371/journal.pbio.0030130 | doi-access = free }} {{open access}}
* {{cite journal | doi = 10.1016/j.tim.2003.08.006 | last1 = Urwin | first1 = R | last2 = Maiden | first2 = MC | year = 2003 | title = Multi-locus sequence typing: a tool for global epidemiology | journal = Trends Microbiol | volume = 11 | issue = 10| pages = 479–87 | pmid = 14557031 }}
* {{cite journal | doi = 10.1073/pnas.97.9.4453 | last1 = Adoutte | first1 = A | last2 = Balavoine | year = 2000 | first2 = G | last3 = Lartillot | first3 = N | last4 = Lespinet | first4 = O | last5 = Prud'homme | first5 = B | last6 = De Rosa | first6 = R | title = The new animal phylogeny: reliability and implications | journal = Proc Natl Acad Sci USA | volume = 97 | issue = 9| pages = 4453–6 | pmid = 10781043 | pmc = 34321 |bibcode = 2000PNAS...97.4453A | doi-access = free }}
* {{cite journal | doi = 10.1038/nrg1603 | last1 = Delsuc | first1 = F | last2 = Brinkmann | year = 2005 | first2 = H | last3 = Philippe | first3 = H| title = Phylogenomics and the reconstruction of the tree of life | journal = Nat Rev Genet | volume = 6 | issue = 5| pages = 361–75 | pmid = 15861208 | citeseerx = 10.1.1.333.1615 | s2cid = 16379422 }}
* {{cite journal | doi = 10.1093/molbev/msj011 | last1 = Steenkamp | first1 = ET | last2 = Wright | year = 2006 | first2 = J | last3 = Baldauf | first3 = SL | title = The protistan origins of animals and fungi | journal = Mol Biol Evol | volume = 23 | issue = 1| pages = 93–106 | pmid = 16151185 | doi-access = free }}
* {{cite journal | doi = 10.1038/nrmicro1234 | last1 = Thomas | first1 = CM | last2 = Nielsen | first2 = KM. | year = 2005 | title = Mechanisms of, and barriers to, horizontal gene transfer between bacteria | journal = Nat Rev Microbiol | volume = 3 | issue = 9| pages = 711–21 | pmid = 16138099 | s2cid = 1231127 }}

{{DEFAULTSORT:Horizontal Gene Transfer In Evolution}}
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[[Category:Evolutionary biology]]

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