HOME

TheInfoList



OR:

A multicellular organism is an
organism In biology, an organism () is any life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy (biology), taxonomy into groups such as Multicellular o ...
that consists of more than one cell, in contrast to unicellular organism. 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 and most
fungi A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
are multicellular, as are many
algae Algae ( , ; : alga ) are any of a large and diverse group of photosynthetic, eukaryotic organisms. The name is an informal term for a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from ...
, whereas a few organisms are partially uni- and partially multicellular, like slime molds and social amoebae such as the genus '' Dictyostelium''. Multicellular organisms arise in various ways, for example by cell division or by aggregation of many single cells. Colonial organisms are the result of many identical individuals joining together to form a 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". There are also multinucleate 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 th ...
that can reach 20 cm.


Evolutionary history


Occurrence

Multicellularity has evolved independently at least 25 times in eukaryotes, and also in some prokaryotes, like cyanobacteria, myxobacteria, actinomycetes, '' Magnetoglobus multicellularis'' or '' Methanosarcina''. However, complex multicellular organisms evolved only in six eukaryotic groups:
animals 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 ...
, symbiomycotan fungi, brown algae,
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
,
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 plants. It evolved repeatedly for Chloroplastida (green algae and land plants), once for animals, once for brown algae, three times in the
fungi A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
( chytrids, ascomycetes, and basidiomycetes) and perhaps several times for slime molds 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-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 cells (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 ...
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 ...
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.


Loss of multicellularity

Loss of multicellularity occurred in some groups. 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). It may also have occurred in some
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
(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'' and some Ulvophyceae). In other groups, generally parasites, a reduction of multicellularity occurred, in number or types of cells (e.g., the myxozoans, 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 bl ...
, 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 ( metazoans) 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 galls have been characterized as tumors, 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 A somatic cell (from Ancient Greek σῶμα ''sôma'', meaning "body"), or vegetal cell, is any biological cell forming the body of a multicellular organism other than a gamete, germ cell, gametocyte or undifferentiated stem cell. Such cells compo ...
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).


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, which moved as a multicellular unit. This is essentially what slime molds 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 ''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 s ...
). 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 embryos 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 hard ...
, 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). The Doushantuo Formation has yielded 600 million year old microfossils with evidence of multicellular traits. Until recently, phylogenetic reconstruction has been through anatomical (particularly embryological) 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 plants 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 Alloenzymes (or also called allozymes) are variant forms of an enzyme which differ structurally but not functionally from other allozymes coded for by different alleles at the same locus. These are opposed to isozymes, which are enzymes that perfo ...
, satellite DNA 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, ...
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 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 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'' "withi ...
), 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 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 ''macro''nucleus serves the organism's needs, whereas the ''micro''nucleus is used for sexual reproduction with exchange of genetic material. Slime molds 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.


The colonial theory

The colonial theory of Haeckel, 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 me ...
, though multicellularity is also typically considered to involve cellular differentiation. 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. 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 protists 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's
gastraea theory The Urmetazoan is the hypothetical last common ancestor of all animals or metazoans. It is universally accepted to be a multicellular heterotroph — with the novelties of a germline and oogamy, an extracellular matrix (ECM) and basement membra ...
.


GK-PID

About 800 million years ago, 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.


The role of viruses

Genes borrowed from
viruses A virus is a submicroscopic infectious agent that replicates only inside the living cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room ...
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 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 (anatomy), organ that begins embryonic development, developing from the blastocyst shortly after implantation (embryology), implantation. It plays critical roles in facilitating nutrien ...
and the brain body separation. Two viral components have been identified. The first is syncytin, 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 but before the most recent rise in oxygen. Mills 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.


Snowball Earth hypothesis

A ''
snowball Earth The Snowball Earth hypothesis proposes that, during one or more of Earth's Greenhouse and icehouse Earth, icehouse Climate, climates, the Earth's surface, planet's surface became entirely or nearly entirely Freezing, frozen. It is believed that ...
'' 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 period and consisted of two global glaciation events known as the Sturtian and Marinoan glaciations. Xiao ''et al''. 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 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 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'', 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 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

* Bacterial colony * Embryogenesis * Organogenesis * Unicellular organism


References


External links


Tree of Life Eukaryotes
{{DEFAULTSORT:Multicellular Organism Developmental biology Evolutionary biology Articles containing video clips