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The Urmetazoan is the hypothetical
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of all
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 or metazoans. It is universally accepted to be a multicellular heterotroph — with the novelties of a
germline In biology and genetics, the germline is the population of a multicellular organism's cells that pass on their genetic material to the progeny (offspring). In other words, they are the cells that form the egg, sperm and the fertilised egg. They ...
and
oogamy Oogamy is an extreme form of anisogamy where the gametes differ in both size and form. In oogamy the large female gamete (also known as ovum) is immobile, while the small male gamete (also known as sperm) is mobile. Oogamy is a common form of a ...
, an
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide stru ...
(ECM) and
basement membrane The basement membrane is a thin, pliable sheet-like type of extracellular matrix that provides cell and tissue support and acts as a platform for complex signalling. The basement membrane sits between Epithelium, epithelial tissues including mesot ...
, cell-cell and cell-ECM adhesions and signaling pathways,
collagen IV Collagen IV (ColIV or Col4) is a type of collagen found primarily in the basal lamina. The collagen IV C4 domain at the C-terminus is not removed in post-translational processing, and the fibers link head-to-head, rather than in parallel. Also, ...
and fibrillar collagen, different cell types (as well as expanded gene and protein families), spatial regulation and a complex developmental plan, and relegated unicellular stages.


Choanoflagellates

All animals are posited to have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
from a
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
d eukaryote. Their closest known
living Living or The Living may refer to: Common meanings *Life, a condition that distinguishes organisms from inorganic objects and dead organisms ** Living species, one that is not extinct *Personal life, the course of an individual human's life * Hu ...
relatives are the
choanoflagellate The choanoflagellates are a group of free-living unicellular and colonial flagellate eukaryotes considered to be the closest living relatives of the animals. Choanoflagellates are collared flagellates, having a funnel shaped collar of interconne ...
s, collared
flagellates A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
whose cell morphology is similar to the choanocyte cells of certain
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
s.
Molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...
studies place animals in a supergroup called the
opisthokont The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
s, which also includes the choanoflagellates,
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 ...
, and a few small parasitic
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. The name comes from the posterior location of the
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
in motile cells, such as most animal spermatozoa, whereas other
eukaryotes 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 ...
tend to have anterior flagella as well.


Five hypotheses

Five different hypotheses for the animals' last common ancestor have been put forward. The relationships between the different animal phyla themselves are not well-resolved, so discriminating between the hypotheses is difficult.


Placula hypothesis

Under the placula hypothesis, the last common ancestor of animals was a somewhat amorphous blob, lacking any form of symmetry or axis. The centre of this blob became raised slightly from the sediment, creating a cavity that assisted feeding on the sea floor below. As this cavity developed, it became deeper and deeper, until the organisms resembled a thimble, with an inside and outside. Some sponges and cnidaria have this kind of body form. The development of the bilaterian body plan follows from this hypothesis; the
urbilaterian The urbilaterian (from German ur- 'original') is the hypothetical last common ancestor of the bilaterian clade, i.e., all animals having a bilateral symmetry. Appearance Its appearance is a matter of debate, for no representative has been (or ...
would develop its symmetry as one end of the placula became adapted for moving forward, which would result in a left-right symmetry.


Planula hypothesis

This idea, proposed by
Otto Bütschli Johann Adam Otto Bütschli (3 May 1848 – 2 February 1920) was a German zoologist and professor at the University of Heidelberg. He specialized in invertebrates and insect development. Many of the groups of protists were first recognized by him. ...
, suggests that metazoa are derived from
planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores. Some groups of Nemerteans also produce larvae that are very similar to the planula, which ...
; that is, the larva of certain
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
, or the adult form of the
placozoa The Placozoa are a basal form of marine free-living (non-parasitic) multicellular organism. They are the simplest in structure of all animals. Three genera have been found: the classical ''Trichoplax adhaerens'', ''Hoilungia hongkongensis'', an ...
ns. Under this hypothesis, the larva became sexually mature through
paedomorphosis Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
, and could reproduce without passing through a
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
phase.


Gastraea hypothesis

The Gastraea hypothesis was proposed by
Ernst 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 ...
,Haeckel, E. 1874. ''Die Gastraea-Theorie, die phylogenetische Classification des Thierreichs und die Homologie der Keimblätter''. Jenaische Zeitschr. Naturwiss. 8:1-55. shortly after his work on the
calcareous sponge The calcareous sponges of class Calcarea are members of the animal phylum Porifera, the cellular sponges. They are characterized by spicules made of calcium carbonate in the form of calcite or aragonite. While the spicules in most species hav ...
s. He proposed that this group of sponges is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
with all
eumetazoans Eumetazoa (), also known as diploblasts, Epitheliozoa, or Histozoa, are a proposed basal animal clade as a sister group of the Porifera (sponges). The basal eumetazoan clades are the Ctenophora and the ParaHoxozoa. Placozoa is now also seen as a ...
, including the
bilaterians The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly an ...
. This suggests that the
gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrula. Be ...
and the
gastrula Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of Cell (biology), cells), or in mammals the blastocyst is reorganized into a multilayered structure known as ...
stage are universal for eumetazoans. It has been perceived as problematic that gastrulation by invagination is by no means universal among eumetazoans. Only recently has an invagination been confirmed in a Calcarea sponge, albeit too early to form a remaining inner space (
archenteron The primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesen ...
).


Bilaterogastraea hypothesis

This hypothesis was developed by Gösta Jägersten as an adaptation of
Ernst 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 ...
's Gastraea hypothesis. He proposed that the Bilaterogastraea have a two-stage life cycle, with a
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
juvenile and a
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
adult stage. The invagination of the original gastrula stage he saw as bilaterally symmetric rather than radially symmetric.


Phagocytella hypothesis

Proposed by
Élie Metchnikoff Ilya Ilyich Mechnikov (russian: Илья Ильич Мечников; – 15 July 1916), also spelled Élie Metchnikoff, was a Russian zoologist best known for his pioneering research in immunology. Belkin, a Russian science historian, explains ...
.


See also

* * ' * * * * * * *


References


External links


Biota
(Taxonomicon)

(Systema Naturae 2000)
Vitae
(BioLib) *
Wikispecies Wikispecies is a wiki-based online project supported by the Wikimedia Foundation. Its aim is to create a comprehensive open content catalogue of all species; the project is directed at scientists, rather than at the general public. Jimmy Wales ...
 – a free directory of life {{portal bar, Biology, Evolutionary biology, Marine life, Paleontology Evolutionary biology Evolution of animals Last common ancestors