Editing Multicellular organism

{{short description|Organism that consists of more than one cell}}
[[File:C elegans stained.jpg|thumb|The [[nematode]] ''[[Caenorhabditis elegans]]'' stained to highlight the nuclei of its cells]]
A '''multicellular organism''' is an [[organism]] that consists of more than one [[cell (biology)|cell]], unlike [[unicellular organisms]].<ref>{{cite book
  | last = Becker| first = Wayne M.
  | title = The world of the cell
  | publisher = [[Benjamin Cummings|Pearson Benjamin Cummings]]
  | year = 2008
  | isbn = 978-0-321-55418-5
  | page = 480|display-authors=etal}}</ref> All species of [[animal]]s, [[Embryophyte|land plants]] and most [[fungi]] are multicellular, as are many [[algae]], whereas a few organisms are partially uni- and partially multicellular, like [[slime mold]]s and social [[Amoeba|amoebae]] such as the genus ''[[Dictyostelium]]''.<ref>{{Cite book|url=http://www.hup.harvard.edu/catalog.php?isbn=9780674975910|title=Life at the Edge of Sight: A Photographic Exploration of the Microbial World|last1=Chimileski|first1=Scott|last2=Kolter|first2=Roberto|publisher=Harvard University Press|year=2017|isbn=9780674975910}}</ref><ref name="Lyons-2015">{{Cite journal|last1=Lyons|first1=Nicholas A.|last2=Kolter|first2=Roberto|date=April 2015|title=On the evolution of bacterial multicellularity|journal=Current Opinion in Microbiology|volume=24|pages=21–28|doi=10.1016/j.mib.2014.12.007|issn=1879-0364|pmc=4380822|pmid=25597443}}</ref>

Multicellular organisms arise in various ways, for example by [[cell division]] or by aggregation of many single cells.<ref name="Miller">{{cite journal | author=S. M. Miller | year=2010 | title=Volvox, Chlamydomonas, and the evolution of multicellularity | journal=Nature Education | volume=3 | issue=9 | page=65 }}</ref><ref name="Lyons-2015" /> [[Colonial organism]]s are the result of many identical individuals joining together to form a [[colony (biology)|colony]]. However, it can often be hard to separate colonial protists from true multicellular organisms, because the two concepts are not distinct; colonial protists have been dubbed "pluricellular" rather than "multicellular".<ref name="Strickberger">{{cite book
| author1 = Brian Keith Hall
| author2 = Benedikt Hallgrímsson
| author3 = Monroe W. Strickberger
| year = 2008
| title = Strickberger's evolution: the integration of genes, organisms and populations
| edition = 4th
| publisher = Hall/Hallgrímsson
| isbn = 978-0-7637-0066-9
| page = [https://archive.org/details/strickbergersevo0000hall/page/149 149]
| url = https://archive.org/details/strickbergersevo0000hall/page/149
}}</ref><ref>{{cite journal |author=Adl, Sina <!---|last=Adl|first=Sina, MSimpson, Alastair G. B.; Farmer, Mark A.; Andersen, Robert A.; Anderson, O. Roger; Barta, John R.; Bowser, Samuel S.; Brugerolle, Guy; Fensome, Robert A.; Fredericq, Suzanne; James, Timothy Y.; Karpov, Sergei; Kugrens, Paul; Krug, John; Lane, Christopher E.; Lewis, Louise A.; Lodge, Jean; Lynn, Denis H.; Mann, David G.; Mccourt, Richard M.; Mendoza, Leonel; Moestrup,Øjvind; Mozley-Standridge, Sharon E.; Nerad, Thomas A.; Shearer, Carol A.; Smirnov, Alexey V.; Spiegel, Frederick W.;Taylor, Max F.J.R.--->|title=The New Higher Level Classification of Eukaryotes with Emphasis on the Taxonomy of Protists|journal=J. Eukaryot. Microbiol.|date=October 2005|volume=52|issue=5|doi=10.1111/j.1550-7408.2005.00053.x|display-authors=etal|pages=399–451|pmid=16248873|s2cid=8060916|doi-access=free}}</ref> There are also macroscopic organisms that are [[multinucleate]] though technically unicellular, such as the [[Xenophyophorea]] that can reach 20&nbsp;cm.

==Evolutionary history==

=== Occurrence ===

Multicellularity has evolved independently at least 25 times in [[eukaryote]]s,<ref name="Grosberg2007">{{cite journal | last1 = Grosberg | first1 = RK | last2 = Strathmann | first2 = RR | year = 2007 | title = The evolution of multicellularity: A minor major transition? | journal = [[Annu Rev Ecol Evol Syst]] | volume = 38 | pages = 621–654 | doi=10.1146/annurev.ecolsys.36.102403.114735 | url = https://grosberglab.faculty.ucdavis.edu/wp-content/uploads/sites/453/2017/05/2007-Grosberg-R.-K.-and-R.-R.-Strathmann.pdf}}</ref><ref>{{cite journal |last1=Parfrey |first1=L.W. |last2=Lahr |first2=D.J.G. |year=2013 |title=Multicellularity arose several times in the evolution of eukaryotes |url=https://web.archive.org/web/20140725235332/http://www.producao.usp.br/bitstream/handle/BDPI/45022/339_ftp.pdf?sequence=1&isAllowed=y |journal=BioEssays |volume=35 |issue=4 |pages=339–347 |doi=10.1002/bies.201200143 |pmid=23315654 |s2cid=13872783}}</ref> and also in some [[prokaryote]]s, like [[cyanobacteria]], [[myxobacteria]], [[actinomycetes]], ''[[Deltaproteobacteria|Magnetoglobus multicellularis]]'' or ''[[Methanosarcina]]''.<ref name="Lyons-2015"/> However, complex multicellular organisms evolved only in six eukaryotic groups: [[animals]], [[fungi#Phylogeny|symbiomycotan fungi]], [[brown algae]], [[red algae]], [[green algae]], and [[land plant]]s.<ref>{{cite journal |last1=Popper |first1=Zoë A. |last2=Michel |first2=Gurvan |last3=Hervé |first3=Cécile |last4=Domozych |first4=David S. |last5=Willats |first5=William G.T. |last6=Tuohy |first6=Maria G. |last7=Kloareg |first7=Bernard |last8=Stengel |first8=Dagmar B. |year=2011 |title=Evolution and diversity of plant cell walls: From algae to flowering plants |journal=Annual Review of Plant Biology |volume=62 |pages=567–590 |pmid=21351878 |hdl=10379/6762 |hdl-access=free |s2cid=11961888 |doi=10.1146/annurev-arplant-042110-103809}}</ref> It evolved repeatedly for [[Chloroplastida]] (green algae and land plants), once for animals, once for brown algae, three times in the [[fungi]] ([[chytrid]]s, [[ascomycete]]s, and [[basidiomycete]]s)<ref name="Niklas-2014">{{Cite journal |last=Niklas |first=K.J. |year=2014 |title=The evolutionary-developmental origins of multicellularity |journal=American Journal of Botany |volume=101 |issue=1 |pages=6–25 |doi=10.3732/ajb.1300314 |doi-access=free |pmid=24363320}}</ref> and perhaps several times for [[Mycetozoa|slime molds]] and red algae.<ref>{{cite journal |author=Bonner, John Tyler |author-link=John Tyler Bonner |year=1998 |title=The origins of multicellularity |journal=[[Integrative Biology]] |volume=1 |issue=1 |pages=27–36 |issn=1093-4391 |doi=10.1002/(SICI)1520-6602(1998)1:1<27::AID-INBI4>3.0.CO;2-6 |url=http://courses.cit.cornell.edu/biog1101/outlines/Bonner%20-Origin%20of%20Multicellularity.pdf |url-status=unfit |archive-url=https://web.archive.org/web/20120308175112/http://courses.cit.cornell.edu/biog1101/outlines/Bonner%20-Origin%20of%20Multicellularity.pdf |archive-date=March 8, 2012 }}</ref> The first evidence of multicellular organization, which is when unicellular organisms coordinate behaviors and may be an evolutionary precursor to true multicellularity, is from [[cyanobacteria]]-like organisms that lived 3.0–3.5&nbsp;billion years ago.<ref name="Grosberg2007"/> To reproduce, true multicellular organisms must solve the problem of regenerating a whole organism from [[germ cell]]s (i.e., [[sperm]] and [[ovum|egg]] cells), an issue that is studied in [[evolutionary developmental biology]]. Animals have evolved a considerable diversity of [[cell types]] in a multicellular body (100–150 different cell types), compared with 10–20 in plants and fungi.<ref>{{cite book |author1=Margulis, L. |author1-link=Lynn Margulis |author2=Chapman, M.J. |year=2009 |title=Kingdoms and Domains: An illustrated guide to the phyla of life on Earth |edition=4th |place=Amsterdam, NL |publisher=Academic Press / Elsevier |page=116}}</ref>

===Loss of multicellularity===
Loss of multicellularity occurred in some groups.<ref>{{Cite journal |last=Seravin |first=L. N. |year=2001 |title=The principle of counter-directional morphological evolution and its significance for constructing the megasystem of protists and other eukaryotes |url=http://cyberleninka.ru/article/n/the-principle-of-counter-directional-morphological-evolution-and-its-significance-for-construction-the-megasystem-of-protists-and-other |journal=Protistology |volume=2 |pages=6–14}}</ref> Fungi are predominantly multicellular, though early diverging lineages are largely unicellular (e.g., [[Microsporidia]]) and there have been numerous reversions to unicellularity across fungi (e.g., ''[[Saccharomycotina]]'', ''[[Cryptococcus]]'', and other [[yeasts]]).<ref>Parfrey, L.W. & Lahr, D.J.G. (2013), p. 344.</ref><ref>{{cite journal | last1 = Medina | first1 = M. | last2 = Collins | first2 = A. G. | last3 = Taylor | first3 = J. W. | last4 = Valentine | first4 = J. W. | last5 = Lipps | first5 = J. H. | last6 = Zettler | first6 = L. A. Amaral | last7 = Sogin | first7 = M. L. | year = 2003 | title = Phylogeny of Opisthokonta and the evolution of multicellularity and complexity in Fungi and Metazoa | journal = International Journal of Astrobiology | volume = 2 | issue = 3| pages = 203–211 | doi=10.1017/s1473550403001551| bibcode = 2003IJAsB...2..203M | doi-access = free }}</ref> It may also have occurred in some [[red algae]] (e.g., ''[[Porphyridium cruentum|Porphyridium]]''), but they may be primitively unicellular.<ref>Seckbach, Joseph, Chapman, David J. [eds.]. (2010). ''Red algae in the genomic age''. New York, NY, U.S.A.: Springer, p. 252, [https://books.google.com/books?id=fegCa9G-c90C&pg=PA252].</ref> Loss of multicellularity is also considered probable in some [[green algae]] (e.g., ''[[Chlorella|Chlorella vulgaris]]'' and some [[Ulvophyceae]]).<ref name="CocquytVerbruggen2010">{{cite journal|last1=Cocquyt|first1=E.|last2=Verbruggen|first2=H.|last3=Leliaert|first3=F.|last4=De Clerck|first4=O.|title=Evolution and Cytological Diversification of the Green Seaweeds (Ulvophyceae)|journal=Mol. Biol. Evol.|volume=27|issue=9|year=2010|pages=2052–2061|issn=0737-4038|doi=10.1093/molbev/msq091|pmid=20368268|doi-access=free}}</ref><ref>Richter, Daniel Joseph: [http://escholarship.org/uc/item/7xc2p94p The gene content of diverse choanoflagellates illuminates animal origins], 2013.</ref> In other groups, generally parasites, a reduction of multicellularity occurred, in the number or types of cells (e.g., the [[myxozoa]]ns, multicellular organisms, earlier thought to be unicellular, are probably extremely reduced [[cnidarian]]s).<ref>{{cite web|url=http://tolweb.org/Myxozoa/2460|title=Myxozoa|website=tolweb.org|access-date=14 April 2018}}</ref>

===Cancer===
Multicellular organisms, especially long-living animals, face the challenge of [[cancer]], which occurs when cells fail to regulate their growth within the normal program of development. Changes in tissue morphology can be observed during this process. Cancer in animals ([[metazoan]]s) has often been described as a loss of multicellularity and an atavistic reversion towards a unicellular-like state.<ref>{{cite journal | last1 = Davies | first1 = P. C. W. | last2 = Lineweaver | first2 = C. H. | year = 2011 | title = Cancer tumors as Metazoa 1.0: tapping genes of ancient ancestors | journal = Physical Biology | volume = 8 | issue = 1 | page = 015001 | doi=10.1088/1478-3975/8/1/015001 | pmid=21301065 | pmc=3148211| bibcode = 2011PhBio...8a5001D }}</ref> Many genes responsible for the establishment of multicellularity that originated around the appearance of metazoans are deregulated in cancer cells, including genes that control [[cell differentiation]], [[cell adhesion|adhesion]] and [[Cell signaling|cell-to-cell communication]].<ref>{{cite journal | last1 = Domazet-Loso | first1 = T. | last2 = Tautz | first2 = D. | year = 2010 | title = Phylostratigraphic tracking of cancer genes suggests a link to the emergence of multicellularity in metazoa | journal = BMC Biology | volume = 8 | issue = 66 | page = 66 | doi=10.1186/1741-7007-8-66| doi-access = free | pmid=20492640 | pmc=2880965 }}</ref><ref>{{cite journal | first1=F. | last1=Jacques | first2=E. | last2=Baratchart | first3=K. | last3=Pienta | first4=E. | last4=Hammarlund | year = 2022 | title= Origin and evolution of animal multicellularity in the light of phylogenomics and cancer genetics | journal = Medical Oncology | volume=39 | issue =160 | doi=10.1007/s12032-022-01740-w | pmid=35972622  | pmc=9381480 | pages = 1–14}}.</ref> There is a discussion about the possibility of existence of cancer in other multicellular organisms<ref>Richter, D. J. (2013), p. 11.</ref><ref>{{cite journal | last1 = Gaspar | first1 = T. | last2 = Hagege | first2 = D. | last3 = Kevers | first3 = C. | last4 = Penel | first4 = C. | last5 = Crèvecoeur | first5 = M. | last6 = Engelmann | first6 = I. | last7 = Greppin | first7 = H. | last8 = Foidart | first8 = J. M. | year = 1991 | title = When plant teratomas turn into cancers in the absence of pathogens | journal = Physiologia Plantarum | volume = 83 | issue = 4| pages = 696–701 | doi=10.1111/j.1399-3054.1991.tb02489.x}}</ref> or even in [[protozoa]].<ref>Lauckner, G. (1980). Diseases of protozoa. In: ''Diseases of Marine Animals''. Kinne, O. (ed.). [https://www.int-res.com/archive/doma_books/DOMA_Vol_I_(general_aspects,_protozoa_to%20gastropoda).pdf Vol. 1, p. 84], John Wiley & Sons, Chichester, UK.</ref> For example, [[plant gall]]s have been characterized as [[tumors]],<ref>{{cite journal | last1 = Riker | first1 = A. J. | year = 1958 | title = Plant tumors: Introduction | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 44 | issue = 4| pages = 338–9 | doi=10.1073/pnas.44.4.338| pmid = 16590201 | pmc = 335422| bibcode = 1958PNAS...44..338R | doi-access = free }}</ref> but some authors argue that plants do not develop cancer.<ref>{{cite journal | last1 = Doonan | first1 = J. | last2 = Hunt | first2 = T. | year = 1996 | title = Cell cycle. Why don't plants get cancer? | journal = Nature | volume = 380 | issue = 6574| pages = 481–2 | doi=10.1038/380481a0 | pmid=8606760| s2cid = 4318184 | doi-access = free }}</ref>

===Separation of somatic and germ cells===
In some multicellular groups, which are called [[Weismann barrier|Weismannists]], a separation between a sterile [[somatic cell]] line and a [[germ cell]] line evolved. However, Weismannist development is relatively rare (e.g., vertebrates, arthropods, ''[[Volvox]]''), as a great part of species have the capacity for [[somatic embryogenesis]] (e.g., land plants, most algae, many invertebrates).<ref>Ridley M (2004) Evolution, 3rd edition. Blackwell Publishing, p. 295–297.</ref><ref name="Niklas-2014" />

==Origin hypotheses==
[[File:The-Simplest-Integrated-Multicellular-Organism-Unveiled-pone.0081641.s003.ogv|thumb|''[[Tetrabaena socialis]]'' consists of four cells.]]
One hypothesis for the origin of multicellularity is that a group of function-specific cells aggregated into a slug-like mass called a [[Grex (biology)|grex]], which moved as a multicellular unit. This is essentially what [[slime mold]]s do. Another hypothesis is that a primitive cell underwent nucleus division, thereby becoming a [[coenocyte]]. A membrane would then form around each nucleus (and the cellular space and organelles occupied in the space), thereby resulting in a group of connected cells in one organism (this mechanism is observable in [[Drosophila]]). A third hypothesis is that as a unicellular organism divided, the daughter cells failed to separate, resulting in a conglomeration of identical cells in one organism, which could later develop specialized tissues. This is what plant and animal [[embryo]]s do as well as colonial [[choanoflagellate]]s.<ref>{{cite journal|url= |title=Multicellular development in a choanoflagellate|first1=Stephen R.|last1=Fairclough|first2=Mark J.|last2=Dayel|first3=Nicole|last3=King|date=26 October 2010|journal=Current Biology|volume=20|issue=20|pages=R875–R876|doi=10.1016/j.cub.2010.09.014|pmid=20971426|pmc=2978077|bibcode=2010CBio...20.R875F }}</ref><ref>{{Cite news|url=https://www.nytimes.com/2010/12/14/science/14creatures.html|title=In a Single-Cell Predator, Clues to the Animal Kingdom's Birth|first=Sean B.|last=Carroll|newspaper=The New York Times |date=December 14, 2010}}</ref>

Because the first multicellular organisms were simple, soft organisms lacking bone, shell, or other hard body parts, they are not well preserved in the fossil record.<ref>A H Knoll, 2003. ''Life on a Young Planet''. Princeton University Press. {{ISBN|0-691-00978-3}} (hardcover), {{ISBN|0-691-12029-3}} (paperback). An excellent book on the early history of life, very accessible to the non-specialist; includes extensive discussions of early signatures, fossilization, and organization of life.</ref> One exception may be the [[demosponge]], which may have left a chemical signature in ancient rocks. The earliest fossils of multicellular organisms include the contested ''[[Grypania]] spiralis'' and the fossils of the black shales of the [[Palaeoproterozoic]] [[Francevillian Group Fossil]] B Formation in [[Gabon]] ([[Gabonionta]]).<ref>
{{cite journal
| author = El Albani, Abderrazak <!---
| last = El Albani
| first = Abderrazak
| author-link = Abderrazak El Albani
A, Bengtson S, Canfield DE, Bekker A, Macchiarelli R, Mazurier A, Hammarlund EU, Boulvais P, Dupuy JJ, Fontaine C, Fürsich FT, Gauthier-Lafaye F, Janvier P, Javaux E, Ossa FO, Pierson-Wickmann AC, Riboulleau A, Sardini P, Vachard D, Whitehouse M, Meunier A.
--->
| date = 1 July 2010
| title = Large colonial organisms with coordinated growth in oxygenated environments 2.1 Gyr ago
| journal = [[Nature (journal)|Nature]]
| volume = 466
| issue = 7302
| pages = 100–104
| issn = 0028-0836
| doi = 10.1038/nature09166
| pmid = 20596019
|display-authors=etal| bibcode = 2010Natur.466..100A
| s2cid = 4331375
}}</ref> The [[Doushantuo Formation]] has yielded 600 million year old microfossils with evidence of multicellular traits.<ref>{{Cite journal | doi = 10.1038/nature13766| pmid = 25252979| title = Cell differentiation and germ–soma separation in Ediacaran animal embryo-like fossils| journal = Nature| year = 2014| last1 = Chen | first1 = L. | last2 = Xiao | first2 = S. | last3 = Pang | first3 = K. | last4 = Zhou | first4 = C. | last5 = Yuan | first5 = X. | volume=516 | issue = 7530| pages=238–241| bibcode = 2014Natur.516..238C| s2cid = 4448316}}</ref>

Until recently, [[phylogenetic]] reconstruction has been through [[anatomy|anatomical]] (particularly [[embryology|embryological]]) similarities. This is inexact, as living multicellular organisms such as [[animal]]s and [[plant]]s are more than 500 million years removed from their single-cell ancestors. Such a passage of time allows both [[divergent evolution|divergent]] and [[convergent evolution|convergent]] evolution time to mimic similarities and accumulate differences between groups of modern and extinct ancestral species. Modern phylogenetics uses sophisticated techniques such as [[alloenzymes]], [[satellite DNA]] and other molecular markers to describe traits that are shared between distantly related lineages.{{citation needed|date=June 2018}}

The evolution of multicellularity could have occurred in several different ways, some of which are described below:

===The symbiotic theory===
This theory suggests that the first multicellular organisms occurred from [[symbiosis]] (cooperation) of different species of single-cell organisms, each with different roles. Over time these organisms would become so dependent on each other that they would not be able to survive independently, eventually leading to the incorporation of their genomes into one multicellular organism.<ref>{{cite book
| author = Margulis, Lynn
| author-link = Lynn Margulis
| year = 1998
| title = Symbiotic Planet: A New Look at Evolution
| page = 160
| publisher = [[Basic Books]]
| location = New York
| isbn = 978-0-465-07272-9
| url = https://www.questia.com/PM.qst?a=o&d=96971657
| access-date = 2017-09-05
| archive-date = 2010-04-20
| archive-url = https://web.archive.org/web/20100420055859/http://www.questia.com/PM.qst?a=o
| url-status = dead
}}</ref> Each respective organism would become a separate lineage of differentiated cells within the newly created species.{{citation needed|date=November 2023}}

This kind of severely co-dependent symbiosis can be seen frequently, such as in the relationship between [[clown fish]] and [[Heteractis magnifica|Riterri sea anemones]]. In these cases, it is extremely doubtful whether either species would survive very long if the other became extinct. However, the problem with this theory is that it is still not known how each organism's DNA could be incorporated into one single [[genome]] to constitute them as a single species. Although such symbiosis is theorized to have occurred (e.g., [[mitochondria]] and [[chloroplasts]] in animal and plant cells—[[endosymbiosis]]), it has happened only extremely rarely and, even then, the genomes of the endosymbionts have retained an element of distinction, separately replicating their DNA during [[mitosis]] of the host species. For instance, the two or three symbiotic organisms forming the composite [[lichen]], although dependent on each other for survival, have to separately reproduce and then re-form to create one individual organism once more.{{citation needed|date=May 2022}}

===The cellularization (syncytial) theory===
This theory states that a single unicellular organism, with multiple [[Cell nucleus|nuclei]], could have developed [[endomembrane|internal membrane]] partitions around each of its nuclei.<ref>
{{cite book
|vauthors=Hickman CP, Hickman FM | date = 8 July 1974
| title = Integrated Principles of Zoology
| edition = 5th
| publisher = [[Mosby (publisher)|Mosby]]
| isbn = 978-0-8016-2184-0
| page = 112
}}</ref> Many protists such as the [[ciliates]] or [[slime molds]] can have several nuclei, lending support to this [[hypothesis]]. However, the simple presence of multiple nuclei is not enough to support the theory. Multiple nuclei of ciliates are dissimilar and have clear differentiated functions. The [[macronucleus|''macro''nucleus]] serves the organism's needs, whereas the [[micronucleus|''micro''nucleus]] is used for sexual reproduction with exchange of genetic material. Slime molds [[syncytium|syncitia]] form from individual amoeboid cells, like syncitial tissues of some multicellular organisms, not the other way round. To be deemed valid, this theory needs a demonstrable example and mechanism of generation of a multicellular organism from a pre-existing syncytium.{{citation needed|date=May 2022}}

===The colonial theory===
The colonial theory of [[Haeckel]], 1874, proposes that the symbiosis of many organisms of the same species (unlike the [[#The symbiotic theory|symbiotic theory]], which suggests the symbiosis of different species) led to a multicellular organism. At least some - it is presumed land-evolved - multicellularity occurs by cells separating and then rejoining (e.g., [[slime mould|cellular slime molds]]) whereas for the majority of multicellular types (those that evolved within aquatic environments), multicellularity occurs as a consequence of cells failing to separate following division.<ref>{{Cite journal
| doi = 10.1038/420745a
| title = Multicellularity: Evolution and the egg
| year = 2002
| last1 = Wolpert | first1 = L.
| last2 = Szathmáry | first2 = E.
| journal = Nature
| volume = 420
| pages = 745
| pmid = 12490925
| issue = 6917
| bibcode = 2002Natur.420..745W
| s2cid = 4385008
| doi-access = free
}}</ref> The mechanism of this latter colony formation can be as simple as incomplete [[cytokinesis]], though multicellularity is also typically considered to involve [[cellular differentiation]].<ref>{{Cite journal
| doi = 10.1002/bies.20197
| title = A twelve-step program for evolving multicellularity and a division of labor
| year = 2005
| last1 = Kirk | first1 = D. L.
| journal = BioEssays
| volume = 27
| pages = 299–310
| pmid = 15714559
| issue = 3
}}</ref>

[[File:ColonialFlagellateHypothesis.png|thumb|650px]]
The advantage of the Colonial Theory hypothesis is that it has been seen to occur independently in 16 different protoctistan phyla. For instance, during food shortages the amoeba [[Dictyostelium]] groups together in a colony that moves as one to a new location. Some of these amoeba then slightly differentiate from each other. Other examples of colonial organisation in protista are [[Volvocaceae]], such as [[Eudorina]] and [[Volvox]], the latter of which consists of up to 500–50,000 cells (depending on the species), only a fraction of which reproduce.<ref>AlgaeBase. [http://www.algaebase.org/search/genus/detail/?genus_id=43497 Volvox Linnaeus, 1758: 820.]</ref> For example, in one species 25–35 cells reproduce, 8 asexually and around 15–25 sexually. However, it can often be hard to separate colonial [[protist]]s from true multicellular organisms, as the two concepts are not distinct; colonial protists have been dubbed "pluricellular" rather than "multicellular".<ref name="Strickberger"/>

===The synzoospore theory===
Some authors suggest that the origin of multicellularity, at least in Metazoa, occurred due to a transition from temporal to spatial [[cell differentiation]], rather than through a gradual evolution of cell differentiation, as affirmed in [[Haeckel]]'s [[Gastraea|gastraea theory]].<ref>{{Cite journal |url=http://www.kumarlab.net/pdf_new/MikhailovAleoshin09.pdf |doi=10.1002/bies.200800214 |archive-url=https://web.archive.org/web/20160305132401/http://www.kumarlab.net/pdf_new/MikhailovAleoshin09.pdf |archive-date=2016-03-05|title=The origin of Metazoa: A transition from temporal to spatial cell differentiation |year=2009 |last1=Mikhailov |first1=Kirill V. |last2=Konstantinova |first2=Anastasiya V. |last3=Nikitin |first3=Mikhail A. |last4=Troshin |first4=Peter V. |last5=Rusin |first5=Leonid Yu. |last6=Lyubetsky |first6=Vassily A. |last7=Panchin |first7=Yuri V. |last8=Mylnikov |first8=Alexander P. |last9=Moroz |first9=Leonid L. |last10=Kumar |first10=Sudhir |last11=Aleoshin |first11=Vladimir V. |journal=BioEssays |volume=31 |issue=7 |pages=758–768 |pmid=19472368 |s2cid=12795095 }}</ref>

===GK-PID===
About 800 million years ago,<ref name="PTRS-2015">{{cite journal |last1=Erwin |first1=Douglas H. |title=Early metazoan life: divergence, environment and ecology |journal=[[Phil. Trans. R. Soc. B]] |date=9 November 2015 |volume=370 |issue=20150036|pages=20150036 |doi=10.1098/rstb.2015.0036 |pmid=26554036 |pmc=4650120 }}</ref> a minor genetic change in a single molecule called [[guanylate kinase]] protein-interaction domain (GK-PID) may have allowed organisms to go from a single cell organism to one of many cells.<ref name="NYT-20160107">{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |title=Genetic Flip Helped Organisms Go From One Cell to Many |url=https://www.nytimes.com/2016/01/12/science/genetic-flip-helped-organisms-go-from-one-cell-to-many.html |date=7 January 2016 |work=[[New York Times]] |access-date=7 January 2016 }}</ref>

===The role of viruses===
Genes borrowed from [[viruses]] and [[mobile genetic elements]] (MGEs) have recently been identified as playing a crucial role in the differentiation of multicellular tissues and organs and even in sexual reproduction, in the fusion of egg cells and sperm.<ref name="Koonin2016" /><ref>{{Cite web |last=Letzter |first=Rafi |date=2018-02-02 |title=An Ancient Virus May Be Responsible for Human Consciousness |url=https://www.livescience.com/61627-ancient-virus-brain.html |access-date=2022-09-05 |website=[[Live Science]] |language=en}}</ref>
Such fused cells are also involved in metazoan membranes such as those that prevent chemicals from crossing the [[placenta]] and the brain body separation.<ref name="Koonin2016">{{Cite journal |last=Koonin |first=E. V. |year=2016 |title=Viruses and mobile elements as drivers of evolutionary transitions |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=371 |issue=1701 |doi=10.1098/rstb.2015.0442 |pmc=4958936 |pmid=27431520 |doi-access=free}}</ref> Two viral components have been identified. The first is [[syncytin]], which came from a virus.<ref>{{cite journal | url=https://www.ncbi.nlm.nih.gov/pubmed/10693809 | pmid=10693809 | year=2000 | last1=Mi | first1=S. | last2=Lee | first2=X. | last3=Li | first3=X. | last4=Veldman | first4=G. M. | last5=Finnerty | first5=H. | last6=Racie | first6=L. | last7=Lavallie | first7=E. | last8=Tang | first8=X. Y. | last9=Edouard | first9=P. | last10=Howes | first10=S. | last11=Keith Jr | first11=J. C. | last12=McCoy | first12=J. M. | title=Syncytin is a captive retroviral envelope protein involved in human placental morphogenesis | journal=Nature | volume=403 | issue=6771 | pages=785–789 | doi=10.1038/35001608 | bibcode=2000Natur.403..785M | s2cid=4367889 }}</ref>
The second identified in 2002 is called [[EFF1|EFF-1]],<ref>{{Cite journal |last1=Mohler |first1=William A. |last2=Shemer |first2=Gidi |last3=del Campo |first3=Jacob J. |last4=Valansi |first4=Clari |last5=Opoku-Serebuoh |first5=Eugene |last6=Scranton |first6=Victoria |last7=Assaf |first7=Nirit |last8=White |first8=John G. |last9=Podbilewicz |first9=Benjamin |date=March 2002 |title=The Type I Membrane Protein EFF-1 Is Essential for Developmental Cell Fusion |url=https://doi.org/10.1016/S1534-5807(02)00129-6 |journal=Developmental Cell |volume=2 |issue=3 |pages=355–362 |doi=10.1016/s1534-5807(02)00129-6 |pmid=11879640 |issn=1534-5807}}</ref> which helps form the skin of ''[[Caenorhabditis elegans]]'', part of a whole family of FF proteins. Felix Rey, of the Pasteur Institute in Paris, has constructed the 3D structure of the EFF-1 protein<ref>{{Cite journal |doi=10.1016/j.cell.2014.02.020 |doi-access=free|title=Structural Basis of Eukaryotic Cell-Cell Fusion |year=2014 |last1=Pérez-Vargas |first1=Jimena |last2=Krey |first2=Thomas |last3=Valansi |first3=Clari |last4=Avinoam |first4=Ori |last5=Haouz |first5=Ahmed |last6=Jamin |first6=Marc |last7=Raveh-Barak |first7=Hadas |last8=Podbilewicz |first8=Benjamin |last9=Rey |first9=Félix A. |journal=Cell |volume=157 |issue=2 |pages=407–419 |pmid=24725407 }}</ref> and shown it does the work of linking one cell to another, in viral infections.
The fact that all known cell fusion molecules are viral in origin suggests that they have been vitally important to the inter-cellular communication systems that enabled multicellularity. Without the ability of cellular fusion, colonies could have formed, but anything even as complex as a sponge would not have been possible.<ref>Slezak, Michael (2016), "No Viruses? No skin or bones either" (New Scientist, No. 2958, 1 March 2014) p.16</ref>

=== Oxygen availability hypothesis ===
This theory suggests that the oxygen available in the atmosphere of early Earth could have been the limiting factor for the emergence of multicellular life.<ref>{{Cite journal|last=Nursall|first=J. R.|date=April 1959|title=Oxygen as a Prerequisite to the Origin of the Metazoa|url=https://www.nature.com/articles/1831170b0|journal=Nature|language=en|volume=183|issue=4669|pages=1170–1172|doi=10.1038/1831170b0|bibcode=1959Natur.183.1170N|s2cid=4200584|issn=1476-4687}}</ref> This hypothesis is based on the correlation between the emergence of multicellular life and the increase of oxygen levels during this time. This would have taken place after the [[Great Oxidation Event]] but before the most recent rise in oxygen. Mills<ref>{{Cite journal|last1=Mills|first1=D. B.|last2=Ward|first2=L. M.|last3=Jones|first3=C.|last4=Sweeten|first4=B.|last5=Forth|first5=M.|last6=Treusch|first6=A. H.|last7=Canfield|first7=D. E.|date=2014-02-18|title=Oxygen requirements of the earliest animals|journal=Proceedings of the National Academy of Sciences|language=en|volume=111|issue=11|pages=4168–4172|doi=10.1073/pnas.1400547111|pmid=24550467|pmc=3964089|bibcode=2014PNAS..111.4168M|issn=0027-8424|doi-access=free}}</ref> concludes that the amount of oxygen present during the [[Ediacaran]] is not necessary for complex life and therefore is unlikely to have been the driving factor for the origin of multicellularity.{{citation needed|date=November 2023}}

=== Snowball Earth hypothesis ===
A ''[[snowball Earth]]'' is a geological event where the entire surface of the Earth is covered in snow and ice. The term can either refer to individual events (of which there were at least two) or to the [[Cryogenian|larger geologic period during which]] all the known total glaciations occurred.

The most recent snowball Earth took place during the [[Cryogenian]] period and consisted of two global glaciation events known as the [[Sturtian glaciation|Sturtian]] and [[Marinoan glaciation|Marinoan]] glaciations. Xiao ''et al''.<ref>{{cite journal |last1=Lyons |first1=Timothy W. |last2=Droser |first2=Mary L.|last3=Lau |first3=Kimberly V. |last4=Porter|first4=Susannah M. |last5=Xiao |first5=Shuhai |last6=Tang |first6=Qing |date=2018-09-28 |title=After the boring billion and before the freezing millions: Evolutionary patterns and innovations in the Tonian Period |journal=Emerging Topics in Life Sciences |language=en |volume=2 |issue=2 |pages=161–171 |doi=10.1042/ETLS20170165 |pmid=32412616|issn=2397-8554 |hdl=10919/86820 |hdl-access=free |s2cid=90374085 |url=https://portlandpress.com/emergtoplifesci/article/2/2/161/77199/After-the-boring-billion-and-before-the-freezing}}</ref> suggest that between the period of time known as the "[[Boring Billion]]" and the snowball Earth, simple life could have had time to innovate and evolve, which could later lead to the evolution of multicellularity.

The snowball Earth hypothesis in regards to multicellularity proposes that the Cryogenian period in Earth's history could have been the catalyst for the evolution of complex multicellular life. Brocks<ref>{{cite journal |last1=Brocks |first1=Jochen J. |last2=Jarrett |first2=Amber J.M. |last3=Sirantoine |first3=Eva |last4=Hallmann |first4=Christian |last5=Hoshino |first5=Yosuke |last6=Liyanage |first6=Tharika |date=August 2017 |title=The rise of algae in Cryogenian oceans and the emergence of animals |journal=Nature |language=en |volume=548 |issue=7669 |pages=578–581 |doi=10.1038/nature23457 |pmid=28813409 |bibcode=2017Natur.548..578B |s2cid=205258987 |issn=1476-4687 |url=https://www.nature.com/articles/nature23457}}</ref> suggests that the time between the Sturtian Glacian and the more recent Marinoan Glacian allowed for planktonic algae to dominate the seas making way for rapid diversity of life for both plant and animal lineages. Complex life quickly emerged and diversified in what is known as the [[Cambrian explosion]] shortly after the Marinoan.{{citation needed|date=May 2022}}

=== Predation hypothesis ===
The predation hypothesis suggests that to avoid being eaten by predators, simple single-celled organisms evolved multicellularity to make it harder to be consumed as prey. Herron et al.<ref>{{Cite journal|last1=Herron|first1=Matthew D.|last2=Borin|first2=Joshua M.|last3=Boswell|first3=Jacob C.|last4=Walker|first4=Jillian|last5=Chen|first5=I.-Chen Kimberly|last6=Knox|first6=Charles A.|last7=Boyd|first7=Margrethe|last8=Rosenzweig|first8=Frank|last9=Ratcliff|first9=William C.|date=2019-02-20|title=De novo origins of multicellularity in response to predation|journal=Scientific Reports|language=en|volume=9|issue=1|page=2328|doi=10.1038/s41598-019-39558-8|pmid=30787483|pmc=6382799|bibcode=2019NatSR...9.2328H|issn=2045-2322|doi-access=free}}</ref>&nbsp;performed laboratory evolution experiments on the single-celled green alga, ''[[Chlamydomonas reinhardtii]]'', using paramecium as a predator. They found that in the presence of this predator, ''C. reinhardtii'' does indeed evolve simple multicellular features.{{citation needed|date=November 2023}}

== Experimental evolution ==
It is impossible to know what happened when single cells evolved into multicellular organisms hundreds of millions of years ago. However, we can identify [[mutation]]s that can turn single-celled organisms into multicellular ones. This would demonstrate the possibility of such an event. Unicellular species can relatively easily acquire mutations that make them attach to each other—the first step towards multicellularity. Multiple normally unicellular species have been evolved to exhibit such early steps:
* [[Baker's yeast|yeast]] are long known to exhibit ''[[Yeast flocculation|flocculation]]''. One of the first yeast genes found to cause this phenotype is FLO1.<ref>{{Cite journal |last1=Smukalla |first1=Scott |last2=Caldara |first2=Marina |last3=Pochet |first3=Nathalie |last4=Beauvais |first4=Anne |last5=Guadagnini |first5=Stephanie |last6=Yan |first6=Chen |last7=Vinces |first7=Marcelo D. |last8=Jansen |first8=An |last9=Prevost |first9=Marie Christine |last10=Latgé |first10=Jean-Paul |last11=Fink |first11=Gerald R. |last12=Foster |first12=Kevin R. |last13=Verstrepen |first13=Kevin J. |date=2008-11-14 |title=FLO1 is a variable green beard gene that drives biofilm-like cooperation in budding yeast |journal=Cell |volume=135 |issue=4 |pages=726–737 |doi=10.1016/j.cell.2008.09.037 |issn=1097-4172 |pmc=2703716 |pmid=19013280}}</ref> A more strikingly clumped [[phenotype]] is called "snowflake", caused by the loss of a single transcription factor ''Ace2''. "Snowflake" yeast grow into multicellular clusters that sediment quickly; they were identified by directed evolution.<ref>{{cite journal |doi=10.1073/pnas.1305949110 |title=Genome duplication and mutations in ''ACE2'' cause multicellular, fast-sedimenting phenotypes in evolved ''Saccharomyces cerevisiae'' |date=2013 |last1=Oud |first1=Bart |last2=Guadalupe-Medina |first2=Victor |last3=Nijkamp |first3=Jurgen F. |last4=De Ridder |first4=Dick |last5=Pronk |first5=Jack T. |last6=Van Maris |first6=Antonius J. A. |last7=Daran |first7=Jean-Marc |journal=Proceedings of the National Academy of Sciences |volume=110 |issue=45 |doi-access=free |pmid=24145419 |bibcode=2013PNAS..110E4223O |pmc=3831460 }}</ref> More recently (2024), snowflake yeast were subject to over 3,000 generations of further directed evolution, forming macroscopic assemblies on the scale of [[millimeters]]. Changes in multiple genes were identified. In addition, the authors reported that only anaerobic cultures of snowflake yeast evolved this trait, while the aerobic ones did not.<ref>{{Cite journal |last1=Bozdag |first1=G. Ozan |last2=Zamani-Dahaj |first2=Seyed Alireza |last3=Day |first3=Thomas C. |last4=Kahn |first4=Penelope C. |last5=Burnetti |first5=Anthony J. |last6=Lac |first6=Dung T. |last7=Tong |first7=Kai |last8=Conlin |first8=Peter L. |last9=Balwani |first9=Aishwarya H. |last10=Dyer |first10=Eva L. |last11=Yunker |first11=Peter J. |last12=Ratcliff |first12=William C. |date=2023-05-25 |title=De novo evolution of macroscopic multicellularity |url=https://www.nature.com/articles/s41586-023-06052-1 |journal=Nature |language=en |volume=617 |issue=7962 |pages=747–754 |doi=10.1038/s41586-023-06052-1 |pmid=37165189 |pmc=10425966 |bibcode=2023Natur.617..747B |s2cid=236953093 |issn=0028-0836 |pmc-embargo-date=May 10, 2024 }}</ref>
* A range of green algae species have been experimentally evolved to form larger clumps. When ''[[Chlorella vulgaris]]'' is grown with a predator ''[[Ochromonas vallescia]]'', it starts forming small colonies, which are harder to ingest due to the larger size. The same is true for ''[[Chlamydomonas reinhardtii]]'' under predation by ''[[Brachionus calyciflorus]]'' and ''[[Paramecium tetraurelia]]''.

''C. reinhartii'' normally starts as a motile single-celled [[propagule]]; this single cell asexually reproduces by undergoing 2–5 rounds of mitosis as a small clump of non-motile cells, then all cells become single-celled propagules and the clump dissolves. With a few generations under ''Paramecium'' predation, the "clump" becomes a persistent structure: only some cells become propagules. Some populations go further and evolved multi-celled propagules: instead of peeling off single cells from the clump, the clump now reproduces by peeling off smaller clumps.<ref>{{cite journal |doi=10.1038/s41598-019-39558-8|doi-access=free |title=De novo origins of multicellularity in response to predation |date=2019 |last1=Herron |first1=Matthew D. |last2=Borin |first2=Joshua M. |last3=Boswell |first3=Jacob C. |last4=Walker |first4=Jillian |last5=Chen |first5=I-Chen Kimberly |last6=Knox |first6=Charles A. |last7=Boyd |first7=Margrethe |last8=Rosenzweig |first8=Frank |last9=Ratcliff |first9=William C. |journal=Scientific Reports |volume=9 |issue=1 |page=2328 |pmid=30787483 |pmc=6382799 |bibcode=2019NatSR...9.2328H }}</ref>

==Advantages==
<!--The reference Grosberg2007 has a good discussion of the advantages of multicellularity.-->
Multicellularity allows an organism to exceed the size limits normally imposed by [[diffusion]]: single cells with increased size have a decreased surface-to-volume ratio and have difficulty absorbing sufficient nutrients and transporting them throughout the cell. Multicellular organisms thus have the [[Competition (biology)|competitive]] advantages of an increase in size without its limitations. They can have longer lifespans as they can continue living when individual cells die. Multicellularity also permits increasing complexity by allowing [[Cellular differentiation|differentiation]] of cell types within one organism.{{citation needed|date=November 2023}}

Whether all of these can be seen as advantages however is debatable: The vast majority of living organisms are single celled, and even in terms of biomass, single celled organisms are far more successful than animals, although not plants.<ref>
{{cite journal
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</ref>
Rather than seeing traits such as longer lifespans and greater size as an advantage, many biologists see these only as examples of diversity, with associated tradeoffs.{{citation needed|date=November 2023}}

==Gene expression changes in the transition from uni- to multicellularity==

During the evolutionary transition from [[unicellular organism]]s to multicellular organisms, the expression of [[gene]]s associated with reproduction and survival likely changed.<ref name="Grochau-Wright2023">Grochau-Wright ZI, Nedelcu AM, Michod RE. The Genetics of Fitness Reorganization during the Transition to Multicellularity: The Volvocine regA-like Family as a Model. Genes (Basel). 2023 Apr 19;14(4):941. doi: 10.3390/genes14040941. PMID 37107699; PMCID: PMC10137558</ref> In the unicellular state, genes associated with reproduction and survival are expressed in a way that enhances the fitness of individual cells, but after the transition to multicellularity, the pattern of [[gene expression|expression of these genes]] must have substantially changed so that individual cells become more specialized in their function relative to reproduction and survival.<ref name = Grochau-Wright2023/> As the multicellular organism emerged, gene expression patterns became compartmentalized between cells that specialized in reproduction ([[germline]] cells) and those that specialized in survival ([[somatic cell]]s). As the transition progressed, cells that specialized tended to lose their own individuality and would no longer be able to both survive and reproduce outside the context of the group.<ref name = Grochau-Wright2023/>

==See also==
* [[Bacterial colony]]
* [[Embryogenesis]]
* [[Organogenesis]]
* [[Unicellular organism]]

==References==
{{Reflist|30em}}

==External links==
* [http://tolweb.org/Eukaryotes/3 Tree of Life Eukaryotes]. {{Webarchive|url=https://web.archive.org/web/20120129074456/http://tolweb.org/Eukaryotes/3 |date=2012-01-29 }}.

{{Evolution}}

{{DEFAULTSORT:Multicellular Organism}}
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[[Category:Evolutionary biology]]
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