A coenocyte () is a

multinucleate Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinat ...

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 ...

which can result from multiple nuclear divisions without their accompanying

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 ...

, in contrast to a

syncytium 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) ...

, which results from cellular aggregation followed by dissolution of the

cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...

s inside the mass. The word syncytium 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 ...

embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, '' -logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos and ...

is used to refer to the coenocytic

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 for ...

invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...

s. A coenocytic colony is referred to as a coenobium (plural coenobia), and most coenobia are composed of a distinct number of cells, often as a multiple of two (4, 8, etc.). Research suggests that coenobium formation may be a defense against grazing in some species.

Physiological examples

Protists

Diplomonad The diplomonads (Greek for "two units") are a group of flagellates, most of which are parasitic. They include ''Giardia duodenalis'', which causes giardiasis in humans. They are placed among the metamonads, and appear to be particularly close r ...

s, like ''

Giardia ''Giardia'' ( or ) is a genus of anaerobic flagellated protozoan parasites of the phylum Metamonada that colonise and reproduce in the small intestines of several vertebrates, causing the disease giardiasis. Their life cycle alternates between ...

'', have two nuclei.

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

s have cells that contain two nuclei: 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 ...

and 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 ...

. 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 ...

apicomplexa The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. T ...

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 ...

s is a form of a coenocyte (i.e. a plasmodium in the general sense) as well as the plasmodia of

microsporidia Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or prot ...

n (

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

myxosporidia Myxosporea is a class of microscopic parasites, belonging to the Myxozoa clade within Cnidaria. They have a complex life cycle which comprises vegetative forms in two hosts, an aquatic invertebrate (generally an annelid but sometimes a bryozoan) ...

n (

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 in ...

) parasites. The trophont of

syndinea The Syndiniales are an order of early branching dinoflagellates (also known as Marine Alveolates, "MALVs"), found as parasites of crustaceans, fish, algae, cnidarians, and protists (ciliates, radiolarians, other dinoflagellates). The troph ...

n (Dinoflagellata) parasites.

Xenophyophorea Xenophyophorea is a clade of foraminiferans. Members of this class are multinucleate unicellular organisms found on the ocean floor throughout the world's oceans, at depths of . They are a kind of foraminiferan that extract minerals from their s ...

are giant cells with numerous nuclei, and is common on the

abyssal plain An abyssal plain is an underwater plain on the deep ocean floor, usually found at depths between and . Lying generally between the foot of a continental rise and a mid-ocean ridge, abyssal plains cover more than 50% of the Earth's surface. T ...

Algae

Coenocytic cells are present in diverse and unrelated groups of algae, including

Xanthophyceae Yellow-green algae or the Xanthophyceae (xanthophytes) are an important group of heterokont algae. Most live in fresh water, but some are found in marine and soil habitats. They vary from single-celled flagellates to simple colonial and filamento ...

(e.g., ''Vaucheria''),

red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...

(e.g., ''Griffithsia'') and

green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...

(e.g., the internodal cells of '' Chara''). In the siphonous

green alga The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...

Bryopsidales Bryopsidales is an order of green algae, in the class Ulvophyceae. Characteristics The thallus is filamentous and much branched and may be packed into a mass. It is coenocytic, having multi-nucleate cells consisting of cytoplasm contained withi ...

and some

Dasycladales Dasycladales is an order of large unicellular green algae in the class Ulvophyceae. It contains two families, the Dasycladaceae and the Polyphysaceae. These single celled algae are from 2 mm to 200 mm long. They live on substrates ...

the entire thallus is a single multinucleate cell, which can be many meters across (e.g. ''

Caulerpa ''Caulerpa'' is a genus of seaweeds in the family Caulerpaceae (among the green algae). They are unusual because they consist of only one cell with many nuclei, making them among the biggest single cells in the world. A species in the Mediterran ...

''). However, in some cases, crosswalls may occur during reproduction. The green algal order

Cladophorales Cladophorales are an order of green algae, in the class Ulvophyceae.See the NCBI The National Center for Biotechnology Information (NCBI) is part of the United States National Library of Medicine (NLM), a branch of the National Institutes o ...

is characterized by siphonocladous organization, i.e., the

thalli Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms were ...

are composed of many coenocytic cells. In contrast to the Cladophorales where nuclei are organized in regularly spaced cytoplasmic domains, the cytoplasm of

exhibits streaming, enabling transportation of organelles, transcripts and nutrients across the plant. The

Sphaeropleales Sphaeropleales is an order of green algae that used to be called Chlorococcales. The order includes some of the most common freshwater planktonic algae such as ''Scenedesmus'' and ''Pediastrum''. The Spaeropleales includes vegetatively non-motile ...

also contain several common freshwater species that are coenocytic, namely

Scenedesmus ''Scenedesmus'' is a genus of green algae, in the class Chlorophyceae. They are colonial and non-motile. Taxonomy Currently, there are 74 taxonomically accepted species of ''Scenedesmus''. Additionally, several subgenera have been identified, ...

Hydrodictyon The water net (genus ''Hydrodictyon'') is a taxon of freshwater green algae in the family Hydrodictyaceae. ''Hydrodictyon'' does well in clean, eutrophic water, and has become a nuisance in New Zealand, where it has been recently introduced. Th ...

, and

Pediastrum ''Pediastrum'' is a genus of green algae, in the family Hydrodictyaceae. It is a photoautotrophic, nonmotile coenobial (fixed number of cells) green algae that inhabits freshwater environments. Morphology ''Pediastrum'' is a genus usually loo ...

Myxogastrids (slime molds)

:''See

Plasmodium (life cycle) A plasmodium is a living structure of cytoplasm that contains many nuclei, rather than being divided into individual cells each with a single nucleus. Plasmodia are best known from slime molds, but are also found in parasitic Myxosporea, and so ...

.''

Plants

The

endosperm The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the embryo and ...

plants Plants are predominantly Photosynthesis, photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all curr ...

begins to grow when one fertilized cell (the primary endosperm cell) becomes a coenocyte. Different species produce coenocytes with different numbers of nuclei before the PEC eventually begins to subdivide, with some growing to contain thousands of nuclei.

Fungi

Some

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 ...

(Such as Glomeromycota, Chytridiomycota and Neocalligomastigomycota) may contain multiple nuclei in a coenocytic

mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrate (biology), substrates. A typical single ...

. A coenocyte functions as a single coordinated unit composed of multiple cells linked structurally and functionally, i.e. through gap junctions. Fungal mycelia in which

hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...

e lack

septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five coun ...

are known as "aseptate" or "coenocytic".

Metazoans: invertebrates

Many insects, such as the model organism ''

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 Ch ...

'', lay eggs that initially develop as "syncytial" blastoderms, i.e. early on the

embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...

s exhibit incomplete

cell division Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...

. The nuclei undergo

S-phase S phase (Synthesis Phase) is the phase of the cell cycle in which DNA is replicated, occurring between G1 phase and G2 phase. Since accurate duplication of the genome is critical to successful cell division, the processes that occur during ...

(DNA replication) and sister chromatids get pulled apart and re-assembled into nuclei containing full sets of homologous chromosomes, but

does not occur. Thus, the nuclei multiply in a common cytoplasmic space. The early embryo "syncytium" of invertebrates such as ''Drosophila'' is important for "syncytial" specification of cell differentiation. The egg cell cytoplasm contains localized

mRNA In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of Protein biosynthesis, synthesizing a protein. mRNA is ...

molecules such as those that encode the

transcription factors In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The func ...

Bicoid Homeotic protein bicoid is encoded by the ''bcd'' maternal effect gene in ''Drosophilia''. Homeotic protein bicoid concentration gradient patterns the anterior-posterior (A-P) axis during ''Drosophila'' embryogenesis. Bicoid was the first prot ...

and Nanos. Bicoid protein is expressed in a gradient that extends from the anterior end of the early embryo, whereas Nanos protein is concentrated at the posterior end. At first, the nuclei of the early embryo rapidly and synchronously divide in the "syncytial" blastoderm and then migrate through the cytoplasm and position themselves in a monolayer around the periphery, leaving only a small number of nuclei in the center of the egg, which will become yolk nuclei. The position of the nuclei along the embryonic axes determines the relative exposure of different amounts of Bicoid, Nanos, and other

morphogens A morphogen is a substance whose non-uniform distribution governs the pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positions of the various ...

. Those nuclei with more Bicoid will activate genes that promote differentiation of cells into head and thorax structures. Nuclei exposed to more Nanos will activate genes responsible for differentiation of posterior regions, such as the abdomen and

germ cells Germ or germs may refer to: Science * Germ (microorganism), an informal word for a pathogen * Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually * Germ layer, a primary layer of cells that forms during embr ...

. The same principles hold true for the specification of the dorso-ventral axis – higher concentration of nuclear Dorsal protein on the ventral side of the egg specify the ventral fate, whereas absence thereof allows dorsal fates. After the nuclei are positioned in a monolayer underneath the egg membrane, the membrane begins to slowly invaginate, thus separating the nuclei into cellular compartments; during this period, the egg is called a cellular blastoderm. The

pole cell In early ''Drosophila'' development, the first 13 cells pass through mitosis are nuclear divisions (karyokinesis) without cytokinesis, resulting in a multinucleate cell (generally referred to as a syncytium, but strictly a coenocyte). Pole cells a ...

s – the germline anlage – are the first cells to separate fully.

Pathological examples

Certain mutations and the activation of certain

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 genes can lead to bacteria forming "filament-like" cells with multiple chromosomes but without cellular division. These mechanisms or mistakes may lead to a similar structure to a coenocyte, though bacteria do not possess nuclei. This fact has been used in certain synthetic biology applications, for example, to create cell-derived fibers for an organically grown concrete.

Etymology

As with much

international scientific vocabulary International scientific vocabulary (ISV) comprises scientific and specialized words whose language of origin may or may not be certain, but which are in current use in several modern languages (that is, translingually, whether in naturalized, loa ...

, English got the word ''coenocyte'' (''cœnocyte'') from

New Latin New Latin (also called Neo-Latin or Modern Latin) is the revival of Literary Latin used in original, scholarly, and scientific works since about 1500. Modern scholarly and technical nomenclature, such as in zoological and botanical taxonomy ...

, in which its combining forms, '' coeno-'' + '' -cyte'', are based on ancient

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

: κοινός (''koinós'') = "common" + κύτος (''kýtos'') = "box, i.e. cell"). The stressed vowel is œ, which in scientific English usually sounds like

long e In linguistics, vowel length is the perceived length of a vowel sound: the corresponding physical measurement is duration. In some languages vowel length is an important phonemic factor, meaning vowel length can change the meaning of the word, ...

and usually shifts a preceding ''c'' to be soft; this explains how there is a degree of regularity in "how one gets a "see-no" sound from ''coeno-''," which might seem irregular at first glance.

References

{{reflist

External links

* http://www.ige.tohoku.ac.jp/outou/outou-e/ceonocytes-e.html Cells Fungal morphology and anatomy