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Multinucleate
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
s (also known as multinucleated or polynuclear cells) are
eukaryotic 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 ...
cells that have more than one
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...
per cell, i.e., multiple nuclei share one common
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
.
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 ...
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 ''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 blood-feeding insect host which then injects parasites into a vert ...
. Although not normally viewed as a case of multinucleation,
plant cell Plant cells are the cells present in green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids with the capabi ...
s share a common cytoplasm by
plasmodesmata 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 ...
, and most cells in
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 ...
tissues are in communication with their neighbors via
gap junctions Gap junctions are specialized intercellular connections between a multitude of animal cell-types. They directly connect the cytoplasm of two cells, which allows various molecules, ions and electrical impulses to directly pass through a regulate ...
. Multinucleate cells, depending on the mechanism by which they are formed, can be divided into "
syncytia A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus) ...
" (formed by cell fusion) or "
coenocyte 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 insid ...
s" (formed by nuclear division not being followed by
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 meios ...
). A number of
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s are known to have two nuclei. Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA: one from the dinoflagellate and the other from a symbiotic
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
. Some bacteria, such as ''
Mycoplasma pneumoniae ''Mycoplasma pneumoniae'' is a very small bacterium in the class Mollicutes. It is a human pathogen that causes the disease mycoplasma pneumonia, a form of atypical bacterial pneumonia related to cold agglutinin disease. ''M. pneumoniae'' is char ...
'', a pathogen of the respiratory tract, 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 A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus) ...
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, and as such it is strongly 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 are multinuclear cells that can form either through normal biological processes, such as the mammalian placenta, or under the influence of certain pathogens, such as
HIV The human immunodeficiency viruses (HIV) are two species of ''Lentivirus'' (a subgroup of retrovirus) that infect humans. Over time, they cause acquired immunodeficiency syndrome (AIDS), a condition in which progressive failure of the immune ...
, via fusion of the plasma membrane. Other examples include the
skeletal muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
cells of
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
, the
tapetal cell The tapetum is a specialised layer of nutritive cells found within the anther, of flowering plants, where it is located between the sporangenous tissue and the anther wall. Tapetum is important for the nutrition and development of pollen grains ...
s of
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
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 native to western North America and is also known as Douglas-fir, Douglas spruce, Oregon pine, and Columbian pine. There are three va ...
seeds. The
polymorphonuclear leukocytes Granulocytes are cells in the innate immune system characterized by the presence of specific granules in their cytoplasm. Such granules distinguish them from the various agranulocytes. All myeloblastic granulocytes are polymorphonuclear. They hav ...
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 remodeling of bones of the vertebral skeleton. The osteoclast disassembles and digests the composite of hydrated pro ...
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
chlorarachniophyte The chlorarachniophytes are a small group of exclusively marine algae widely distributed in tropical and temperate waters. They are typically mixotrophic, ingesting bacteria and smaller protists as well as conducting photosynthesis. Normally the ...
s 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 blood-feeding insect host which then injects parasites into a vert ...
, in the sense of a multinucleate protoplast without a cell wall which exhibits amoeboid movement. Other examples include some
plasmodiophorid The Phytomyxea are a class of parasites that are cosmopolitan, obligate biotrophic protist parasites of plants, diatoms, oomycetes and brown algae. They are divided into the orders Plasmodiophorida ( ICZN, or Plasmodiophoromycota, ICBN) and Ph ...
s, some
haplosporid The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecul ...
ians, and the
grex Grex or GREX may refer to: * Grex (biology), a multicellular aggregate of amoeba of the phyla Acrasiomycota or Dictyosteliomycota * Grex (horticulture), (pl. greges) a kind of group used in horticultural nomenclature applied to the progeny of an a ...
of
cellular slime moulds Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
(
dictyostelid The dictyostelids (Dictyostelia/Dictyostelea, International Code of Zoological Nomenclature, ICZN, or Dictyosteliomycetes, ICBN) are a group of cellular slime molds, or social amoebae. Multicellular behavior When food (normally bacteria) is rea ...
s and
acrasid The family Acrasidae (ICZN, or Acrasiomycota, ICBN) is a family of slime molds which belongs to the excavate group Percolozoa. The name element - comes from the Greek ''akrasia'', meaning "acting against one's judgement". This group consists ...
s).


Placenta

The
placenta The placenta 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 the physically separate mater ...
, 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, bacteria, and protozoa, which is likely due to unique cytoskeletal properties of these cells.


Coenocytes

Furthermore, multinucleate cells are produced from specialized
cell cycle The cell cycle, or cell-division cycle, is the series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and subs ...
s in which nuclear division occurs without cytokinesis, thus leading to large coenocytes or plasmodia. In
filamentous fungi A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. Not ...
, 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 (
myxogastrid Myxogastria/Myxogastrea (myxogastrids, ICZN) or Myxomycetes ( ICN), is a class of slime molds that contains 5 orders, 14 families, 62 genera, and 888 species. They are colloquially known as the ''plasmodial'' or ''acellular' ...
s) and the
schizont Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism is ...
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 blood-feeding insect host which then injects parasites into a vert ...
''
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
which causes
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...
.


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 series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and subs ...
control (e.g., some
binucleated cells Binucleated cells are cells that contain two nuclei. This type of cell is most commonly found in cancer cells and may arise from a variety of causes. Binucleation can be easily visualized through staining and microscopy. In general, binucleation ...
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 Human Immunodeficiency Virus, where
T-cell A T cell is a type of lymphocyte. T cells are one of the important white blood cells 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 rec ...
s are fused by the action of virus-derived proteins on the cell membrane. During viral replication in T lymphoid cells, 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. Numerous human pathogenic viruses in circulation are encase ...
Glycoprotein Glycoproteins are proteins which contain oligosaccharide 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 as glycos ...
(
Env env is a shell command for Unix and Unix-like operating systems. It is used to either print a list of environment variables or run another utility in an altered environment without having to modify the currently existing environment. Using 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.


References

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