organism
In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
that consists of more than one
cell
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
, in contrast to
unicellular organism
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
.
All species of
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s,
land plants
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sist ...
and most
fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
are multicellular, as are many
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, whereas a few organisms are partially uni- and partially multicellular, like
slime mold
Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mul ...
s and social amoebae such as the genus ''
Dictyostelium
''Dictyostelium'' is a genus of single- and multi-celled eukaryotic, phagotrophic bacterivores. Though they are Protista and in no way fungal, they traditionally are known as "slime molds". They are present in most terrestrial ecosystems as ...
''.
Multicellular organisms arise in various ways, for example by
cell division
Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...
or by aggregation of many single cells.
Colonial organism
In biology, a colony is composed of two or more conspecific individuals living in close association with, or connected to, one another. This association is usually for mutual benefit such as stronger defense or the ability to attack bigger prey.
...
s are the result of many identical individuals joining together to form a
colony
In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the ''metropole, metropolit ...
. 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".
There are also
multinucleate
Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinat ...
though technically unicellular organisms that are macroscopic, such as the
xenophyophorea
Xenophyophorea is a clade of foraminiferans. Members of this class are multinucleate unicellular organisms found on the ocean floor throughout the world's oceans, at depths of . They are a kind of foraminiferan that extract minerals from their s ...
that can reach 20 cm.
Evolutionary history
Occurrence
Multicellularity has evolved independently at least 25 times in
eukaryote
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s, and also in some
prokaryote
A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...
s, like
cyanobacteria
Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
,
myxobacteria
The myxobacteria ("slime bacteria") are a group of bacteria that predominantly live in the soil and feed on insoluble organic substances. The myxobacteria have very large genomes relative to other bacteria, e.g. 9–10 million nucleotides except ...
,
actinomycetes
The Actinomycetales is an order of Actinomycetota. A member of the order is often called an actinomycete. Actinomycetales are generally gram-positive and anaerobic and have mycelia in a filamentous and branching growth pattern. Some actinomycete ...
Methanosarcina
''Methanosarcina'' is a genus of euryarchaeote archaea
Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as ...
''. However, complex multicellular organisms evolved only in six
eukaryotic
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
groups:
animals
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
brown algae
Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and po ...
green algae
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
, and
land plant
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sist ...
s. It evolved repeatedly for
Chloroplastida
Viridiplantae (literally "green plants") are a clade of eukaryotic organisms that comprise approximately 450,000–500,000 species and play important roles in both terrestrial and aquatic ecosystems. They are made up of the green algae, which ar ...
(green algae and land plants), once for animals, once for brown algae, three times in the
fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
(
chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
basidiomycete
Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basi ...
s) and perhaps several times for
slime molds
Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
and red algae. 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
Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
-like organisms that lived 3.0–3.5 billion years ago. To reproduce, true multicellular organisms must solve the problem of regenerating a whole organism from
germ cell
Germ or germs may refer to:
Science
* Germ (microorganism), an informal word for a pathogen
* Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually
* Germ layer, a primary layer of cells that forms during emb ...
s (i.e.,
sperm
Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
and
egg
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
cell types
A cell type is a classification used to identify cells that share morphological or phenotypical features. A multicellular organism may contain cells of a number of widely differing and specialized cell types, such as muscle cells and skin cells, ...
in a multicellular body (100–150 different cell types), compared with 10–20 in plants and fungi.
Loss of multicellularity
Loss of multicellularity occurred in some groups. Fungi are predominantly multicellular, though early diverging lineages are largely unicellular (e.g.,
Microsporidia
Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or pr ...
) and there have been numerous reversions to unicellularity across fungi (e.g., ''
Saccharomycotina
Saccharomycotina is a subdivision (subphylum) of the division (phylum) Ascomycota in the kingdom Fungi. It comprises most of the ascomycete yeasts. The members of Saccharomycotina reproduce by budding and they do not produce ascocarps (fruitin ...
'', ''
Cryptococcus
''Cryptococcus'' is a genus of fungi in the family Cryptococcaceae that includes both yeasts and filamentous species. The filamentous, sexual forms or teleomorphs were formerly classified in the genus ''Filobasidiella'', while ''Cryptococcus'' ...
'', and other
yeasts
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
). It may also have occurred in some red algae (e.g., '' Porphyridium''), but it is possible that they are primitively unicellular. Loss of multicellularity is also considered probable in some
green algae
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
(e.g., ''
Chlorella vulgaris
''Chlorella vulgaris'' is a species of green microalga in the division Chlorophyta. It is mainly used as a dietary supplement or protein-rich food additive in Japan.
Description
''C. vulgaris'' is a green eukaryotic microalga in the genus ...
'' and some
Ulvophyceae
The Ulvophyceae or ulvophytes are a class of green algae, distinguished mainly on the basis of ultrastructural morphology, life cycle and molecular phylogenetic data.Graham LE, Graham JM, Wilcox LW (2009) ''Algae''. 2nd Edition. Benjamin Cummin ...
). In other groups, generally parasites, a reduction of multicellularity occurred, in number or types of cells (e.g., the
myxozoa
Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. O ...
ns, multicellular organisms, earlier thought to be unicellular, are probably extremely reduced cnidarians).
Cancer
Multicellular organisms, especially long-living animals, face the challenge of
cancer
Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal b ...
, 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
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s) has often been described as a loss of multicellularity. There is a discussion about the possibility of existence of cancer in other multicellular organisms or even in
protozoa
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
. For example,
plant gall
Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
s have been characterized as
tumors
A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
, but some authors argue that plants do not develop cancer.
Separation of somatic and germ cells
In some multicellular groups, which are called Weismannists, a separation between a sterile somatic cell line and a
germ cell
Germ or germs may refer to:
Science
* Germ (microorganism), an informal word for a pathogen
* Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually
* Germ layer, a primary layer of cells that forms during emb ...
line evolved. However, Weismannist development is relatively rare (e.g., vertebrates, arthropods, ''
Volvox
''Volvox'' is a polyphyletic genus of chlorophyte green algae in the family Volvocaceae. It forms spherical colonies of up to 50,000 cells. They live in a variety of freshwater habitats, and were first reported by Antonie van Leeuwenhoek in 170 ...
''), as a great part of species have the capacity for
somatic embryogenesis
Somatic embryogenesis is an artificial process in which a plant or embryo is derived from a single somatic cell. Somatic embryos are formed from plant cells that are not normally involved in the development of embryos, i.e. ordinary plant tissue. N ...
(e.g., land plants, most algae, many invertebrates).
Origin hypotheses
One hypothesis for the origin of multicellularity is that a group of function-specific cells aggregated into a slug-like mass called a
grex
Grex or GREX may refer to:
* Grex (biology), a multicellular aggregate of amoeba of the phyla Acrasiomycota or Dictyosteliomycota
* Grex (horticulture), (pl. greges) a kind of group used in horticultural nomenclature applied to the progeny of an a ...
, which moved as a multicellular unit. This is essentially what
slime mold
Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mul ...
s do. Another hypothesis is that a primitive cell underwent nucleus division, thereby becoming a
coenocyte
A coenocyte () is a multinucleate cell which can result from multiple nuclear divisions without their accompanying cytokinesis, in contrast to a syncytium, which results from cellular aggregation followed by dissolution of the cell membranes insid ...
. 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
''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species ...
). 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
An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
s do as well as colonial choanoflagellates.
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. One exception may be the
demosponge
Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a har ...
, which may have left a chemical signature in ancient rocks. The earliest fossils of multicellular organisms include the contested ''
Grypania
''Grypania'' is an early, tube-shaped fossil from the Proterozoic eon. The organism, with a size over one centimeter and consistent form, could have been a giant bacterium, a bacterial colony, or a eukaryotic alga. The oldest probable ''Grypania' ...
spiralis'' and the fossils of the black shales of the
Palaeoproterozoic
The Paleoproterozoic Era (;, also spelled Palaeoproterozoic), spanning the time period from (2.5–1.6 Ga), is the first of the three sub-divisions ( eras) of the Proterozoic Eon. The Paleoproterozoic is also the longest era of the Earth's ...
Francevillian Group Fossil
The Francevillian biota (also known as Gabon macrofossils or Gabonionta) is a group of 2.1-billion-year-old Palaeoproterozoic, macroscopic organisms known from fossils found in Gabon in the Palaeoproterozoic Francevillian B Formation, a black shal ...
B Formation in
Gabon
Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
(
Gabonionta
The Francevillian biota (also known as Gabon macrofossils or Gabonionta) is a group of 2.1-billion-year-old Palaeoproterozoic, macroscopic Organism, organisms known from Fossil, fossils found in Gabon in the Palaeoproterozoic Francevillian B Forma ...
). The
Doushantuo Formation
The Doushantuo Formation (formerly transcribed as Toushantuo or Toushantou, from ) is a geological formation in western Hubei, eastern Guizhou, southern Shaanxi, central Jiangxi, and other localities in China. It is known for the fossil Lagerst ...
has yielded 600 million year old microfossils with evidence of multicellular traits.
Until recently,
phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
reconstruction has been through
anatomical
Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
(particularly
embryological
Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos and ...
) similarities. This is inexact, as living multicellular organisms such as
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s and
plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
s are more than 500 million years removed from their single-cell ancestors. Such a passage of time allows both divergent and 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 Satellite DNA consists of very large arrays of tandemly repeating, non-coding DNA. Satellite DNA is the main component of functional centromeres, and form the main structural constituent of heterochromatin.
The name "satellite DNA" refers to the ...
and other molecular markers to describe traits that are shared between distantly related lineages.
The evolution of multicellularity could have occurred in a number of different ways, some of which are described below:
The symbiotic theory
This theory suggests that the first multicellular organisms occurred from
symbiosis
Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
(cooperation) of different species of single-cell organisms, each with different roles. Over time these organisms would become so dependent on each other they would not be able to survive independently, eventually leading to the incorporation of their genomes into one multicellular organism. Each respective organism would become a separate lineage of differentiated cells within the newly created species.
This kind of severely co-dependent symbiosis can be seen frequently, such as in the relationship between
clown fish
Clownfish or anemonefish are fishes from the subfamily Amphiprioninae in the family Pomacentridae. Thirty species of clownfish are recognized: one in the genus '' Premnas'', while the remaining are in the genus ''Amphiprion''. In the wild, t ...
and 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
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
to constitute them as a single species. Although such symbiosis is theorized to have occurred (e.g.,
mitochondria
A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
and
chloroplasts
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, ...
in animal and plant cells—
endosymbiosis
An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship.
(The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
), 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
In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
of the host species. For instance, the two or three symbiotic organisms forming the composite
lichen
A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.nuclei, could have developed internal membrane partitions around each of its nuclei. Many protists such as the
ciliates
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
or
slime molds
Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
can have several nuclei, lending support to this
hypothesis
A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
. 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 ''macro''nucleus serves the organism's needs, whereas the ''micro''nucleus is used for sexual reproduction with exchange of genetic material. Slime molds
syncitia
A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus) ...
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.
The colonial theory
The colonial theory of
Haeckel
Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...
, 1874, proposes that the symbiosis of many organisms of the same species (unlike the 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., 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. The mechanism of this latter colony formation can be as simple as incomplete
cytokinesis
Cytokinesis () is the part of the cell division process during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division in mitosis and mei ...
, though multicellularity is also typically considered to involve
cellular differentiation
Cellular differentiation is the process in which a stem cell alters from one type to a differentiated one. Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a multicellular ...
.
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
''Dictyostelium'' is a genus of single- and multi-celled eukaryotic, phagotrophic bacterivores. Though they are Protista and in no way fungal, they traditionally are known as "slime molds". They are present in most terrestrial ecosystems as ...
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
The Volvocaceae are a family of unicellular or colonial biflagellates, including the typical genus ''Volvox''. The family was named by Ehrenberg in 1834,From p. 281: ''"VOLVOCINA Nova Familia."'' (Volvocina New Family.) ote: According to p. ...
Volvox
''Volvox'' is a polyphyletic genus of chlorophyte green algae in the family Volvocaceae. It forms spherical colonies of up to 50,000 cells. They live in a variety of freshwater habitats, and were first reported by Antonie van Leeuwenhoek in 170 ...
, the latter of which consists of up to 500–50,000 cells (depending on the species), only a fraction of which reproduce. 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
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s from true multicellular organisms, as the two concepts are not distinct; colonial protists have been dubbed "pluricellular" rather than "multicellular".
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
Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...
About 800 million years ago, a minor genetic change in a single molecule called
guanylate kinase
In enzymology, a guanylate kinase () is an enzyme that catalyzes the chemical reaction
: ATP + GMP \rightleftharpoons ADP + GDP
Thus, the two substrates of this enzyme are ATP and GMP, whereas its two products are ADP and GDP.
This enzyme b ...
protein-interaction domain (GK-PID) may have allowed organisms to go from a single cell organism to one of many cells.
The role of viruses
Genes borrowed from
viruses
A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea.
Since Dmitri Ivanovsky's 1 ...
and
mobile genetic elements
Mobile genetic elements (MGEs) sometimes called selfish genetic elements are a type of genetic material that can move around within a genome, or that can be transferred from one species or replicon to another. MGEs are found in all organisms. In ...
(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 cell and sperm.
Such fused cells are also involved in metazoan membranes such as those that prevent chemicals crossing the
placenta
The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate mater ...
and the brain body separation. Two viral components have been identified. The first is
syncytin
Syncytin-1 also known as enverin is a protein found in humans and other primates that is encoded by the ERVW-1 gene ( endogenous retrovirus group W envelope member 1). Syncytin-1 is a cell-cell fusion protein whose function is best characterized ...
, which came from a virus.
The second identified in 2007 is called EFF1, 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 EFF1 protein 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.
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. 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
The Great Oxidation Event (GOE), also called the Great Oxygenation Event, the Oxygen Catastrophe, the Oxygen Revolution, the Oxygen Crisis, or the Oxygen Holocaust, was a time interval during the Paleoproterozoic era when the Earth's atmosphere ...
but before the most recent rise in oxygen. Mills concludes that the amount of oxygen present during the
Ediacaran
The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
is not necessary for complex life and therefore is unlikely to have been the driving factor for the origin of multicellularity.
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 larger geologic period during which all the known total glaciations occurred.
The most recent snowball Earth took place during the
Cryogenian
The Cryogenian (from grc, κρύος, krýos, meaning "cold" and , romanized: , meaning "birth") is a geologic period that lasted from . It forms the second geologic period of the Neoproterozoic Era, preceded by the Tonian Period and followed by ...
period and consisted of two global glaciation events known as the
Sturtian
The Sturtian glaciation was a Snowball Earth glaciation, or perhaps multiple glaciations, during the Cryogenian Period when the Earth experienced repeated large-scale glaciations. The duration of the Sturtian glaciation has been variously defined, ...
and
Marinoan
The Marinoan glaciation, sometimes also known as the Varanger glaciation, was a period of worldwide glaciation that lasted from approximately 650 to 632.3 ± 5.9 Ma (million years ago) during the Cryogenian period. This glaciation possibly cove ...
glaciations. Xiao ''et al''. suggest that between the period of time known as the "
Boring Billion
The Boring Billion, otherwise known as the Mid Proterozoic and Earth's Middle Ages, is the time period between 1.8 and 0.8 billion years ago (Ga) spanning the middle Proterozoic eon, characterized by more or less tectonic stability, climatic st ...
" 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 history could have been the catalyst for the evolution of complex multicellular life. Brocks 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
The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil recor ...
shortly after the Marinoan.
Predation hypothesis
The predation hypothesis suggests that in order to avoid being eaten by predators, simple single-celled organisms evolved multicellularity to make it harder to be consumed as prey. Herron et al. performed laboratory evolution experiments on the single-celled green alga, ''
Chlamydomonas reinhardtii
''Chlamydomonas reinhardtii'' is a single-cell green alga about 10 micrometres in diameter that swims with two flagella. It has a cell wall made of hydroxyproline-rich glycoproteins, a large cup-shaped chloroplast, a large pyrenoid, and an ...
'', using paramecium as a predator. They found that in the presence of this predator, ''C. reinhardtii'' does indeed evolve simple multicellular features.
Advantages
Multicellularity allows an organism to exceed the size limits normally imposed by
diffusion
Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
: 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
competitive
Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...
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 differentiation of cell types within one organism.
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.
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.
See also
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Bacterial colony
In biology, a colony is composed of two or more conspecific individuals living in close association with, or connected to, one another. This association is usually for mutual benefit such as stronger defense or the ability to attack bigger prey. ...
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Embryogenesis
An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
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Organogenesis
Organogenesis is the phase of embryonic development that starts at the end of gastrulation and continues until birth. During organogenesis, the three germ layers formed from gastrulation (the ectoderm, endoderm, and mesoderm) form the internal org ...
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Unicellular organism
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...