In evolutionary biology, the term cellularization (cellularisation) has been used in theories to explain the evolution of cells, for instance in the pre-cell theory, [1] [2] [3] dealing with the evolution of the first cells on this planet, and in the syncytial theory [4] attempting to explain the origin of Metazoa from unicellular organisms.
Processes of cell development in multinucleate cells (syncytium, plural syncytia) of animals and plants are also termed cellularization, often called syncytium cellularization.
According to Otto Kandler's pre-cell theory, [1] [2] [3] early evolution of life and primordial metabolism (see Iron-Sulfur world hypothesis - metabolism first scenario, according to Wächtershäuser [5] [6] ) led to the early diversification of life through the evolution of a multiphenotypical population of pre-cells, [1] [2] [3] from which the three founder groups A, B, C and then, from them, the precursor cells (here named proto-cells) of the three domains of life [7] emerged successively. [1] [2] [3] [8]
In this scenario the three domains of life did not originate from an ancestral nearly complete “first cell“ nor a cellular organism often defined as the last universal common ancestor (LUCA [9] [10] [11] ), but from a population of evolving pre-cells. Kandler introduced the term cellularization for his concept of a successive evolution of cells by a process of evolutionary improvements. [1] [2] [3]
His concept may explain the quasi-random distribution of evolutionarily important features among the three domains and, at the same time, the existence of the most basic biochemical features (genetic code, set of protein amino acids etc.) in all three domains (unity of life), as well as the close relationship between the Archaea and the Eucarya. Kandler’s pre-cell theory is supported by Wächtershäuser. [8]
According to Kandler, the protection of fragile primordial life forms from their environment by the invention of envelopes (i.e. membranes, walls) was an essential improvement. For instance, the emergence of rigid cell walls by the invention and elaboration of peptidoglycan [12] in bacteria (domain Bacteria) may have been a prerequisite for their successful survival, radiation and colonisation of virtually all habitats of the geosphere and hydrosphere. [3]
A coevolution of the biosphere and the geosphere is suggested: “The evolving life could venture into a larger variety of habitats, even into microaerobic habitats in shallow, illuminated surface waters. The continuous changes in the physical environment on the aging and cooling Earth led to further diversification of habitats and favored opportunistic radiation of primitive life into numerous phenotypes on the basis of each of the different chemolithoautotrophies. Concomitantly, with the accumulation of organic matter derived from chemolithoautotrophic life, opportunistic and obligate heterotrophic life may also have developed”. [1] : 155f
The details of Kandler's proposal for the early diversification of life are represented in a scheme, where numbers indicate evolutionary improvements. [3]
This theory is also known as a theory of cellularization. It is a theory to explain the origin of the Metazoa. The idea was proposed by Hadži (1953) [4] and Hanson (1977). [13]
This cellularization (syncytial) theory states that metazoans evolved from a unicellular ciliate 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). [14]
According to the syncytial theory, the ciliate ancestor, by several cellularization processes, evolved into the currently known turbellarian flatworms, which are therefore the most primitive metazoans. The theory of cellularization is based on the large similarities between ciliates and flatworms. Both ciliates and flatworms have cilia, are bilaterally symmetric, and syncytial. Therefore, the theory assumes that bilateral symmetry is more primitive than radial symmetry. However, current biological evidence shows that the most primitive forms of metazoans show radial symmetry, and thus radially symmetrical animals like cnidaria cannot be derived from bilateral flatworms. [15]
By concluding that the first multicellular animals were flatworms, it is also suggested that simpler organisms as sponges, ctenophores and cnidarians would have derived from more complex animals. [16] However, most current molecular research has shown that sponges are the most primitive metazoans. [17] [18]
The syncytial theory rejects the theory of germ layers. During the development of the turbellaria (Acoela), 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, endoderm and mesoderm (in more complex animals) build up the embryo. [19]
There is a lot of evidence against ciliates being the metazoan ancestor. Ciliates have two types of nuclei: a micronucleus which is used as germline nucleus and a macronucleus which regulates the vegetative growth. [20] This division of nuclei is a unique feature of the ciliates and is not found in any other members of the animal kingdom. [21] Therefore, it would be unlikely that ciliates are indeed the ancestors of the metazoans. This is confirmed by molecular phylogenetic research. Ciliates were never found close to animals in any molecular phylogeny. [22]
Furthermore, the syncytial theory cannot explain the flagellated sperm of metazoans. Since the ciliate ancestor does not have any flagella 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. [19]
Due to both the lack of molecular and morphological evidence for this theory, the alternative colonial theory of Haeckel, is currently gaining widespread acceptance.
For more theories see main article Multicellular organisms.
The development of cells in a syncytium (multinucleate cells) is termed syncytium cellularization. Syncytia are quite frequent in animals and plants. Syncytium cellularization occurs for instance in the embryonic development of animals and in endosperm development of plants. Here two examples:
In the embryonic development of Drosophila melanogaster , first 13 nuclear divisions take place forming a syncytial blastoderm consisting of approximately 6000 nuclei. During the later gastrulation stage, membranes are formed between the nuclei, and cellularization is completed. [23]
The term syncytium cellularization is used for instance for a process of cell development in the endosperm of the Poaceae , e.g. barley (Hordeum vulgare), [24] rice (Oryza sativa). [25]
The cell is the basic structural and functional unit of all forms of life. Every cell consists of cytoplasm enclosed within a membrane; many cells contain organelles, each with a specific function. The term comes from the Latin word cellula meaning 'small room'. Most cells are only visible under a microscope. Cells emerged on Earth about 4 billion years ago. All cells are capable of replication, protein synthesis, and motility.
Zoology is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one of the primary branches of biology. The term is derived from Ancient Greek ζῷον, zōion ('animal'), and λόγος, logos.
Placozoa is a phylum of marine and free-living (non-parasitic) animals. They are blob-like animals composed of aggregations of cells. Moving in water by ciliary motion, eating food by engulfment, reproducing by fission or budding, placozoans are described as "the simplest animals on Earth." Structural and molecular analyses have supported them as among the most basal animals, thus, constituting a primitive metazoan phylum.
The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm of ciliated cells surrounding one or more reproductive cells.
A multicellular organism is an organism that consists of more than one cell, unlike unicellular organisms. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- and partially multicellular, like slime molds and social amoebae such as the genus Dictyostelium.
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 inside the mass. The word syncytium in animal embryology is used to refer to the coenocytic blastoderm of invertebrates. A coenocytic colony is referred to as a coenobium, and most coenobia are composed of a distinct number of cells, often as a multiple of two.
A syncytium or symplasm is a multinucleate cell that can result from multiple cell fusions of uninuclear cells, in contrast to a coenocyte, which can result from multiple nuclear divisions without accompanying cytokinesis. The muscle cell that makes up animal skeletal muscle is a classic example of a syncytium cell. The term may also refer to cells interconnected by specialized membranes with gap junctions, as seen in the heart muscle cells and certain smooth muscle cells, which are synchronized electrically in an action potential.
The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid in most species, which may be auxin-driven. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread, while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as certain orchids, lack endosperm in their seeds.
Trichoplax adhaerens is one of the four named species in the phylum Placozoa. The others are Hoilungia hongkongensis, Polyplacotoma mediterranea and Cladtertia collaboinventa. Placozoa is a basal group of multicellular animals, possible relatives of Cnidaria. Trichoplax are very flat organisms commonly less than 4 mm in diameter, lacking any organs or internal structures. They have two cellular layers: the top epitheloid layer is made of ciliated "cover cells" flattened toward the outside of the organism, and the bottom layer is made up of cylinder cells that possess cilia used in locomotion, and gland cells that lack cilia. Between these layers is the fibre syncytium, a liquid-filled cavity strutted open by star-like fibres.
Stuart Alan Newman is a professor of cell biology and anatomy at New York Medical College in Valhalla, NY, United States. His research centers around three program areas: cellular and molecular mechanisms of vertebrate limb development, physical mechanisms of morphogenesis, and mechanisms of morphological evolution. He also writes about social and cultural aspects of biological research and technology.
Multinucleate cells are eukaryotic cells that have more than one nucleus, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinated, synchronous manner where all nuclei divide simultaneously or asynchronously where individual nuclei divide independently in time and space. Certain organisms may have a multinuclear stage of their life cycle. For example, slime molds have a vegetative, multinucleate life stage called a plasmodium.
The terms pre-cell (precell), proto-cell (protocell), etc. are frequently used to designate hypothetical ancestral entities precursing complete cells. The meanings of these terms vary with the different hypotheses for the early evolution of life and, accordingly, with the corresponding publications.
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.
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 membrane, cell-cell and cell-ECM adhesions and signaling pathways, collagen IV and fibrillar collagen, different cell types, spatial regulation and a complex developmental plan, and relegated unicellular stages.
Holozoa is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa and the protists Choanoflagellata, Filasterea, Pluriformea and Ichthyosporea. Along with fungi and some other groups, Holozoa is part of the Opisthokonta, a supergroup of eukaryotes. Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic.
The eukaryotes constitute the domain of Eukarya or Eukaryota, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, and many unicellular organisms are eukaryotes. They constitute a major group of life forms alongside the two groups of prokaryotes: the Bacteria and the Archaea. Eukaryotes represent a small minority of the number of organisms, but given their generally much larger size, their collective global biomass is much larger than that of prokaryotes.
Otto Kandler was a German botanist and microbiologist. Until his retirement in 1986 he was professor of botany at the Ludwig Maximilian University of Munich.
Eukaryogenesis, the process which created the eukaryotic cell and lineage, is a milestone in the evolution of life, since eukaryotes include all complex cells and almost all multicellular organisms. The process is widely agreed to have involved symbiogenesis, in which an archeon and a bacterium came together to create the first eukaryotic common ancestor (FECA). This cell had a new level of complexity and capability, with a nucleus, at least one centriole and cilium, facultatively aerobic mitochondria, sex, a dormant cyst with a cell wall of chitin and/or cellulose and peroxisomes. It evolved into a population of single-celled organisms that included the last eukaryotic common ancestor (LECA), gaining capabilities along the way, though the sequence of the steps involved has been disputed, and may not have started with symbiogenesis. In turn, the LECA gave rise to the eukaryotes' crown group, containing the ancestors of animals, fungi, plants, and a diverse range of single-celled organisms.
Sphaeroforma arctica, is a unicellular eukaryote with a pivotal position in the tree of life. It was first isolated from the arctic marine amphipod Gammarus setosus. Like other Ichthyosporeans such as Creolimax and Abeoforma, Sphaeroforma arctica are spherical cells characterized with their capacity to grow into multi-nucleated coenocytes. However, a unique feature of S. arctica, is that no obvious budding, hyphal, amoeboid, sporal or flagellated growth stages have been observed in laboratory growth conditions.
Syssomonas is a monotypic genus of unicellular flagellated protists containing the species Syssomonas multiformis. It is a member of Pluriformea inside the lineage of Holozoa, a clade containing animals and their closest protistan relatives. It lives in freshwater habitats. It has a complex life cycle that includes unicellular amoeboid and flagellated phases, as well as multicellular aggregates, depending on the growth medium and nutritional state.
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