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Multinucleate cells (also known as multinucleated cells or polynuclear cells) are
eukaryotic The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
cells that have more than one nucleus, i.e., multiple nuclei share one common
cytoplasm The cytoplasm describes all the material within a eukaryotic or prokaryotic cell, enclosed by the cell membrane, including the organelles and excluding the nucleus in eukaryotic cells. The material inside the nucleus of a eukaryotic cell a ...
.
Mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...
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 mold Slime mold or slime mould is an informal name given to a polyphyletic assemblage of unrelated eukaryotic organisms in the Stramenopiles, Rhizaria, Discoba, Amoebozoa and Holomycota clades. Most are near-microscopic; those in the Myxogastria ...
s have a vegetative, multinucleate life stage called a
plasmodium ''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a Hematophagy, blood-feeding insect host (biology), host which then inj ...
. Multinucleate cells, depending on the mechanism by which they are formed, can be divided into "
syncytia A syncytium (; : syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell that can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus), ...
" (formed by
cell fusion Cell fusion is an important cellular process in which several uninucleate cells (cells with a single nucleus) combine to form a multinucleate cell, known as a syncytium. Cell fusion occurs during differentiation of myoblasts, osteoclasts and ...
) or " coenocytes" (formed by nuclear division not being followed by
cytokinesis Cytokinesis () is the part of the cell division process and part of mitosis 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 ...
). Some
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
, such as '' Mycoplasma pneumoniae'', a
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
of the
respiratory tract The respiratory tract is the subdivision of the respiratory system involved with the process of conducting air to the alveoli for the purposes of gas exchange in mammals. The respiratory tract is lined with respiratory epithelium as respirato ...
, may display multinuclear filaments as a result of a delay between genome replication and cellular division.


Terminology

Some biologists use the term "acellular" to refer to multinucleate cell forms ( syncitia and plasmodia), such as to differentiate "acellular" slime molds from the purely "cellular" ones (which do not form such structures). This usage is incorrect and highly misleading to
laymen In religious organizations, the laity () — individually a layperson, layman or laywoman — consists of all members who are not part of the clergy, usually including any non-ordained members of religious orders, e.g. a nun or a lay brother. ...
, and as such it is discouraged. Some use the term "syncytium" in a wide sense, to mean any type of multinucleate cell, while others differentiate the terms for each type.


Physiological examples


Syncytia

Syncytia A syncytium (; : syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell that can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus), ...
are multinuclear cells that can form either through normal biological processes, such as the mammalian
placenta The placenta (: placentas or placentae) 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 ...
, or under the influence of certain pathogens, such as HIV, via fusion of the plasma membrane. Other examples include the
skeletal muscle Skeletal muscle (commonly referred to as muscle) is one of the three types of vertebrate muscle tissue, the others being cardiac muscle and smooth muscle. They are part of the somatic nervous system, voluntary muscular system and typically are a ...
cells of
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
, the tapetal cells of
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s, and the storage cells of
Douglas-fir The Douglas fir (''Pseudotsuga menziesii'') is an evergreen conifer species in the pine family, Pinaceae. It is the tallest tree in the Pinaceae family. It is native to western North America and is also known as Douglas-fir, Douglas spruce, Or ...
seeds. The polymorphonuclear leukocytes of mammals are not polynuclear cells, although the lobes of their nuclei are so deeply bifurcated that they can appear so under non-optimal microscopy.
Osteoclast An osteoclast () is a type of bone cell that breaks down bone tissue. This function is critical in the maintenance, repair, and bone remodeling, remodeling of bones of the vertebrate, vertebral skeleton. The osteoclast disassembles and digests th ...
s are multinuclear cells that are found commonly in the human body that aid in the maintenance and repair of the bones by secreting acid that dissolves bone matter. They are typically found to have 5 nuclei per cell, due to the fusion of preosteoclasts. The chlorarachniophytes form multinucleate cells by fusion, being syncytia and not coenocytes. This syncytia is called
plasmodium ''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a Hematophagy, blood-feeding insect host (biology), host which then inj ...
, in the sense of a multinucleate
protoplast Protoplast (), is a biology, biological term coined by Johannes von Hanstein, Hanstein in 1880 to refer to the entire cell, excluding the cell wall. Protoplasts can be generated by stripping the cell wall from plant, bacterium, bacterial, or f ...
without a
cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...
which exhibits amoeboid movement. Other examples include some plasmodiophorids, some haplosporidians, and the grex of cellular slime moulds ( dictyostelids and acrasids).


Placenta

The
placenta The placenta (: placentas or placentae) 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 ...
, a temporary organ that transports nutrients, oxygen, waste, and other materials between a mother and a developing fetus, is partially composed of a syncytial layer that forms the interface between the foetus and the mother. In addition to performing simple interface duties, the placental syncytia also acts as a barrier to infection 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. Viruses are found in almo ...
,
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
, and
protozoa Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
, which is likely due to unique
cytoskeletal The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all Cell (biology), cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane ...
properties of these cells.


Coenocytes

Furthermore, multinucleate cells are produced from specialized
cell cycle The cell cycle, or cell-division cycle, is the sequential series of events that take place in a cell (biology), cell that causes it to divide into two daughter cells. These events include the growth of the cell, duplication of its DNA (DNA re ...
s in which nuclear division occurs without cytokinesis, thus leading to large coenocytes or plasmodia. In filamentous fungi, multinucleate cells may extend over hundreds of meters so that different regions of a single cell experience dramatically different microenvironments. Other examples include, the plasmodia of plasmodial slime molds ( myxogastrids) and the schizont of the ''
Plasmodium ''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a Hematophagy, blood-feeding insect host (biology), host which then inj ...
''
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
which causes
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
.


Pathological examples

Multinucleated cells can also occur under pathological conditions as the consequence of a disturbed
cell cycle The cell cycle, or cell-division cycle, is the sequential series of events that take place in a cell (biology), cell that causes it to divide into two daughter cells. These events include the growth of the cell, duplication of its DNA (DNA re ...
control (e.g., some binucleated cells and metastasizing
tumor 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 ...
cells).


Human Immunodeficiency Virus

As previously mentioned, syncytia may be induced through the actions of HIV, where
T-cell T cells (also known as T lymphocytes) are an important part of the immune system and play a central role in the adaptive immune response. T cells can be distinguished from other lymphocytes by the presence of a T-cell receptor (TCR) on their ce ...
s are fused by the action of virus-derived
proteins Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, re ...
on the
cell membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of a cell from the outside environment (the extr ...
. During viral replication in T
lymphoid cells A lymphocyte is a type of white blood cell (leukocyte) in the immune system of most vertebrates. Lymphocytes include T cells (for cell-mediated and cytotoxic adaptive immunity), B cells (for humoral, antibody-driven adaptive immunity), and i ...
, large amounts of
viral envelope A viral envelope is the outermost layer of many types of viruses. It protects the genetic material in their life cycle when traveling between host cells. Not all viruses have envelopes. A viral envelope protein or E protein is a protein in the en ...
Glycoprotein Glycoproteins are proteins which contain oligosaccharide (sugar) chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known a ...
( Env) are synthesized and trafficked to the cell membrane where they can be incorporated into new virus particles. However, some of the Env molecules interact with neighboring T-cell receptors, which brings the cells into close enough proximity to enable trigger events culminating in the fusion of two host cells, likely due to the close contact of the two plasma membranes. This interaction is likely specific to CD4+ T-cells, as cells lacking this receptor were unable to form syncytia in laboratory conditions.


Not multinucleate

Although not normally viewed as a case of multinucleation,
plant cell Plant cells are the cells present in Viridiplantae, green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids ...
s share a common cytoplasm by
plasmodesma Plasmodesmata (singular: plasmodesma) are microscopic channels which traverse the cell walls of plant cells and some algal cells, enabling transport and communication between them. Plasmodesmata evolved independently in several lineages, and spec ...
ta, and most cells in
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
tissues are in communication with their neighbors via
gap junction Gap junctions are membrane channels between adjacent cells that allow the direct exchange of cytoplasmic substances, such small molecules, substrates, and metabolites. Gap junctions were first described as ''close appositions'' alongside tight ...
s. Although
endosymbiosis An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
among eukaryotes can result in multiple nuclei in a cell, these are not considered "multinucleate" because the nuclei are not surrounded by the same cytoplasm. These extra nuclei are called
nucleomorph Nucleomorphs are small, vestigial eukaryotic nuclei found between the inner and outer pairs of membranes in certain plastids. They are thought to be vestiges of red and green algal nuclei that were engulfed by a larger eukaryote. Because the nucl ...
s.


References

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