The theory of cellularization, also known as the syncytial theory or ciliate-acoel theory, is a theory to explain the origin of

Metazoa 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 ...

. The idea was proposed by Hadži (1953) and Hanson (1977). The cellularization theory states that metazoans evolved from a unicellular

ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...

with multiple nuclei that went through cellularization. Firstly, the ciliate developed a ventral mouth for feeding and all nuclei moved to one side of the cell. Secondly, an epithelium was created by membranes forming barriers between the nuclei. In this way, a multicellular organism was created from one multinucleate cell (

syncytium A syncytium (; plural syncytia; from Ancient Greek, Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell (biology), cell which can result from multiple cell fusions of uninuclear cells (i.e ...

Arguments

Turbellarian flatworms

By several cellularization processes, the ciliate ancestor evolved into the currently known turbellarian flatworms, which are therefore the most primitive metazoans according to the theory. The theory of cellularization is based on the large similarities between ciliates and flatworms. Both ciliates and flatworms have

cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...

, are

bilaterally symmetric Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...

, and

syncytial A syncytium (; plural syncytia; from Ancient Greek, Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell (biology), cell which can result from multiple cell fusions of uninuclear cells (i.e ...

. Therefore, the theory assumes that bilateral symmetry is more primitive than

radial symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...

. However, current biological evidence shows that the most primitive forms of metazoans show radial symmetry, and thus radially symmetrical animals like

cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latter. Their distinguishing feature is cnidocyt ...

cannot be derived from bilateral flatworms. By concluding that the first multicellular animals were flatworms, it is also suggested that simpler organisms as

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 throug ...

ctenophores Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...

and

ns would have derived from more complex animals. However, most current molecular research has shown that sponges are the most primitive metazoans.

Germ layers are formed simultaneously

The syncytial theory rejects the theory of germ layers. During the development of the turbellaria (

Acoela Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order ...

), three regions are formed without the formation of germ layers. From this, it was concluded that the germ layers are simultaneously formed during the cellularization process. This is in contrast to germ layer theory in which

ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from the o ...

endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gast ...

and

mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical Emb ...

(in more complex animals) build up the embryo.

''Drosophila melanogaster'' development

Evidence for the syncytial theory can also be found in the development of ''

Drosophila melanogaster ''Drosophila melanogaster'' is a species of fly (the taxonomic order Diptera) in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the " vinegar fly" or " pomace fly". Starting with ...

''. First 13 nuclear divisions take place forming a syncytial

blastoderm A blastoderm ( germinal disc, blastodisc) is a single layer of embryonic epithelial tissue that makes up the blastula. It encloses the fluid filled blastocoel. Gastrulation follows blastoderm formation, where the tips of the blastoderm begins the ...

consisting of approximately 6000 nuclei. During the later gastrulation stage, membranes are formed between the nuclei, and cellularization is completed.

Criticism

The macro and micronucleus of Ciliates

There is a lot of evidence against

s being the metazoan ancestor. Ciliates have two types of nuclei: a

micronucleus Micronucleus is the name given to the small nucleus that forms whenever a chromosome or a fragment of a chromosome is not incorporated into one of the daughter nuclei during cell division. It usually is a sign of genotoxic events and chromosomal i ...

which is used as germline nucleus and a

macronucleus A macronucleus (formerly also meganucleus) is the larger type of nucleus in ciliates. Macronuclei are polyploid and undergo direct division without mitosis. It controls the non-reproductive cell functions, such as metabolism. During conjugation, t ...

which regulates the vegetative growth. This division of nuclei is a unique feature of the ciliates and is not found in any other members of the animal kingdom. Therefore, it would be unlikely that ciliates are indeed the ancestors of the metazoans. This is confirmed by molecular

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 ...

research. Ciliates were never found close to animals in any

molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

Flagellated sperm

Furthermore, the syncytial theory cannot explain the

flagellated 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#Cellular level, motility. Many protists with flagella are termed as flagellates. A m ...

sperm of metazoans. Since the ciliate ancestor does not have any

flagella 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 ...

and it is unlikely that the flagella arose as a ''de novo'' trait in metazoans, the syncytial theory makes it almost impossible to explain the origin of flagellated sperm. Due to both the lack of molecular and morphological evidence for this theory, the alternative colonial theory of Haeckel, is currently gaining widespread acceptance.

Related theories

Iron-sulfur world hypothesis

Cellularization occurs in several stages. It begins with the formation of primitive lipids (e.g. fatty acids or isoprenoid acids) in the surface metabolism. These lipids accumulate on or in the mineral base. This lipophilizes the outer or inner surfaces of the mineral base, which promotes condensation reactions over hydrolytic reactions by lowering the activity of water and protons. In the next stage lipid membranes are formed. While still anchored to the mineral base they form a semi-cell bounded partly by the mineral base and partly by the membrane. Further lipid evolution leads to self-supporting lipid membranes and closed cells. The earliest closed cells are pre-cells (''

sensu ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...

'' Kandler) because they allow frequent exchange of genetic material (e.g. by fusions). According to

Woese Carl Richard Woese (; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal RNA, a ...

, this frequent exchange of genetic material is the cause for the existence of the common stem in the tree of life and for a very rapid early evolution.

References

{{Reflist Biological evolution