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In
cell biology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...
, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the
cell nucleus The cell nucleus (pl. nuclei; from Latin or , meaning ''kernel'' or ''seed'') is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, h ...
. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The main components of the cytoplasm are cytosol (a gel-like substance), the
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s (the cell's internal sub-structures), and various
cytoplasmic inclusion In cellular biology, inclusions are diverse intracellularShively, J. M. (ed.). (2006). ''Microbiology Monographs Vol. 1: Inclusions in Prokaryotes''. Berlin, Heidelberg: Springer-Verlaglink non-living substances ( ergastic substances) that are no ...
s. The cytoplasm is about 80% water and is usually colorless. The submicroscopic ground cell substance or cytoplasmic matrix which remains after exclusion of the cell
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s and particles is
groundplasm The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells (intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondri ...
. It is the hyaloplasm of light microscopy, a highly complex, polyphasic system in which all resolvable cytoplasmic elements are suspended, including the larger organelles such as the
ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...
s,
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
, the plant
plastid The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosy ...
s, lipid droplets, and vacuoles. Most cellular activities take place within the cytoplasm, such as many metabolic pathways including
glycolysis Glycolysis is the metabolic pathway that converts glucose () into pyruvate (). The free energy released in this process is used to form the high-energy molecules adenosine triphosphate (ATP) and reduced nicotinamide adenine dinucleotide (NADH ...
, and processes such as cell division. The concentrated inner area is called the endoplasm and the outer layer is called the
cell cortex The cell cortex, also known as the actin cortex, cortical cytoskeleton or actomyosin cortex, is a specialized layer of cytoplasmic proteins on the inner face of the cell membrane. It functions as a modulator of membrane behavior and cell surface p ...
or the ectoplasm. Movement of calcium ions in and out of the cytoplasm is a
signaling In signal processing, a signal is a function that conveys information about a phenomenon. Any quantity that can vary over space or time can be used as a signal to share messages between observers. The ''IEEE Transactions on Signal Processing'' ...
activity for
metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
processes. In plants, movement of the cytoplasm around vacuoles is known as
cytoplasmic streaming Cytoplasmic streaming, also called protoplasmic streaming and cyclosis, is the flow of the cytoplasm inside the cell, driven by forces from the cytoskeleton. It is likely that its function is, at least in part, to speed up the transport of mol ...
.


History

The term was introduced by Rudolf von Kölliker in 1863, originally as a synonym for
protoplasm Protoplasm (; ) is the living part of a cell that is surrounded by a plasma membrane. It is a mixture of small molecules such as ions, monosaccharides, amino acid, and macromolecules such as proteins, polysaccharides, lipids, etc. In some defini ...
, but later it has come to mean the cell substance and organelles outside the nucleus. There has been certain disagreement on the definition of cytoplasm, as some authors prefer to exclude from it some organelles, especially the vacuoles and sometimes the plastids.


Physical nature

It remains uncertain how the various components of the cytoplasm interact to allow movement of
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s while maintaining the cell's structure. The flow of cytoplasmic components plays an important role in many cellular functions which are dependent on the permeability of the cytoplasm. An example of such function is
cell signalling In biology, cell signaling (cell signalling in British English) or cell communication is the ability of a cell to receive, process, and transmit signals with its environment and with itself. Cell signaling is a fundamental property of all cellula ...
, a process which is dependent on the manner in which signaling molecules are allowed to diffuse across the cell. While small signaling molecules like
calcium ions Calcium ions (Ca2+) contribute to the physiology and biochemistry of organisms' cells. They play an important role in signal transduction pathways, where they act as a second messenger, in neurotransmitter release from neurons, in contraction of ...
are able to diffuse with ease, larger molecules and subcellular structures often require aid in moving through the cytoplasm. The irregular dynamics of such particles have given rise to various theories on the nature of the cytoplasm.


As a sol-gel

There has long been evidence that the cytoplasm behaves like a sol-gel. It is thought that the component molecules and structures of the cytoplasm behave at times like a disordered colloidal solution (sol) and at other times like an integrated network, forming a solid mass (gel). This theory thus proposes that the cytoplasm exists in distinct fluid and solid phases depending on the level of interaction between cytoplasmic components, which may explain the differential dynamics of different particles observed moving through the cytoplasm. A papers suggested that at
length scale In physics, length scale is a particular length or distance determined with the precision of at most a few orders of magnitude. The concept of length scale is particularly important because physical phenomena of different length scales cannot aff ...
smaller than 100  nm, the cytoplasm acts like a liquid, while in a larger length scale, it acts like a gel.


As a glass

Recently it has been proposed that the cytoplasm behaves like a glass-forming liquid approaching the
glass transition The glass–liquid transition, or glass transition, is the gradual and reversible transition in amorphous materials (or in amorphous regions within semicrystalline materials) from a hard and relatively brittle "glassy" state into a viscous or rubb ...
. In this theory, the greater the concentration of cytoplasmic components, the less the cytoplasm behaves like a liquid and the more it behaves as a solid glass, freezing more significant cytoplasmic components in place (it is thought that the cell's metabolic activity can fluidize the cytoplasm to allow the movement of such more significant cytoplasmic components). A cell's ability to vitrify in the absence of metabolic activity, as in dormant periods, may be beneficial as a defense strategy. A solid glass cytoplasm would freeze subcellular structures in place, preventing damage, while allowing the transmission of tiny proteins and metabolites, helping to kickstart growth upon the cell's revival from
dormancy Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be clo ...
.


Other perspectives

Research has examined the motion of cytoplasmic particles independent of the nature of the cytoplasm. In such an alternative approach, the aggregate random forces within the cell caused by motor proteins explain the non- Brownian motion of cytoplasmic constituents.


Constituents

The three major elements of the cytoplasm are the cytosol,
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s and inclusions.


Cytosol

The cytosol is the portion of the cytoplasm not contained within membrane-bound organelles. Cytosol makes up about 70% of the cell volume and is a complex mixture of cytoskeleton filaments, dissolved molecules, and water. The cytosol's filaments include the protein filaments such as actin filaments and
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
s that make up the cytoskeleton, as well as soluble proteins and small structures such as
ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...
s,
proteasome Proteasomes are protein complexes which degrade unneeded or damaged proteins by proteolysis, a chemical reaction that breaks peptide bonds. Enzymes that help such reactions are called proteases. Proteasomes are part of a major mechanism by w ...
s, and the mysterious vault complexes. The inner, granular and more fluid portion of the cytoplasm is referred to as endoplasm. Due to this network of fibres and high concentrations of dissolved
macromolecule A macromolecule is a very large molecule important to biophysical processes, such as a protein or nucleic acid. It is composed of thousands of covalently bonded atoms. Many macromolecules are polymers of smaller molecules called monomers. The ...
s, such as proteins, an effect called macromolecular crowding occurs and the cytosol does not act as an
ideal solution In chemistry, an ideal solution or ideal mixture is a solution that exhibits thermodynamic properties analogous to those of a mixture of ideal gases. The enthalpy of mixing is zero as is the volume change on mixing by definition; the closer to zero ...
. This crowding effect alters how the components of the cytosol interact with each other.


Organelles

Organelles (literally "little organs") are usually membrane-bound structures inside the cell that have specific functions. Some major organelles that are suspended in the cytosol are the
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
, the
endoplasmic reticulum The endoplasmic reticulum (ER) is, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – rough endoplasmic reticulum ( ...
, the Golgi apparatus, vacuoles, lysosomes, and in plant cells,
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s.


Cytoplasmic inclusions

The inclusions are small particles of insoluble substances suspended in the cytosol. A huge range of inclusions exist in different cell types, and range from crystals of
calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrate ...
or silicon dioxide in plants, to granules of energy-storage materials such as
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
,
glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one o ...
, or polyhydroxybutyrate. A particularly widespread example are lipid droplets, which are spherical droplets composed of lipids and proteins that are used in both prokaryotes and eukaryotes as a way of storing lipids such as fatty acids and
sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the go ...
s. Lipid droplets make up much of the volume of
adipocyte Adipocytes, also known as lipocytes and fat cells, are the cells that primarily compose adipose tissue, specialized in storing energy as fat. Adipocytes are derived from mesenchymal stem cells which give rise to adipocytes through adipogenesis. I ...
s, which are specialized lipid-storage cells, but they are also found in a range of other cell types.


Controversy and research

The cytoplasm, mitochondria, and most organelles are contributions to the cell from the maternal gamete. Contrary to the older information that disregards any notion of the cytoplasm being active, new research has shown it to be in control of movement and flow of nutrients in and out of the cell by viscoplastic behavior and a measure of the reciprocal rate of bond breakage within the cytoplasmic network. The material properties of the cytoplasm remain an ongoing investigation. A method of determining the mechanical behaviour of living cell mammalian cytoplasm with the aid of
optical tweezers Optical tweezers (originally called single-beam gradient force trap) are scientific instruments that use a highly focused laser beam to hold and move microscopic and sub-microscopic objects like atoms, nanoparticles and droplets, in a manner simila ...
has been described.


See also

* * * *


References


External links

* {{Authority control Cell anatomy