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
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 ...
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 ...
, enclosed by 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 ( ...
, 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
The nuclear envelope, also known as the nuclear membrane, is made up of two lipid bilayer membranes that in eukaryotic cells surround the nucleus, which encloses the genetic material.
The nuclear envelope consists of two lipid bilayer membra ...
is termed the
nucleoplasm
The nucleoplasm, also known as karyoplasm, is the type of protoplasm that makes up the cell nucleus, the most prominent organelle of the eukaryotic cell. It is enclosed by the nuclear envelope, also known as the nuclear membrane. The nucleoplasm ...
. The main components of the cytoplasm are
cytosol
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 ...
(a gel-like substance), the
organelles (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
organelles and particles is
groundplasm. It is the
hyaloplasm
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 ...
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
plastids,
lipid
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
droplets, and
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s.
Most cellular activities take place within the cytoplasm, such as many
metabolic pathway
In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell. The reactants, products, and intermediates of an enzymatic reaction are known as metabolites, which are modified by a sequence of chemical reac ...
s including
glycolysis, and processes such as
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 concentrated inner area is called the
endoplasm
Endoplasm generally refers to the inner (often granulated), dense part of a cell's cytoplasm. This is opposed to the ectoplasm which is the outer (non-granulated) layer of the cytoplasm, which is typically watery and immediately adjacent to the ...
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 ion
Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
s in and out of the cytoplasm is a
signaling activity for
metabolic processes.
In
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, movement of the cytoplasm around vacuoles is known as
cytoplasmic streaming.
History
The term was introduced by
Rudolf von Kölliker in 1863, originally as a synonym for
protoplasm, 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
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s and sometimes the plastids.
Physical nature
It remains uncertain how the various components of the cytoplasm interact to allow movement of
organelles 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, a process which is dependent on the manner in which signaling molecules are allowed to
diffuse
Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
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 contractio ...
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
A colloid is a mixture in which one substance consisting of microscopically dispersed insoluble particles is suspended throughout another substance. Some definitions specify that the particles must be dispersed in a liquid, while others extend ...
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 af ...
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
Glass is a non-crystalline, often transparent, amorphous solid that has widespread practical, technological, and decorative use in, for example, window panes, tableware, and optics. Glass is most often formed by rapid cooling (quenching) of ...
-forming liquid approaching the
glass transition.
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.
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
Motor proteins are a class of molecular motors that can move along the cytoplasm of cells. They convert chemical energy into mechanical work by the hydrolysis of ATP. Flagellar rotation, however, is powered by a proton pump.
Cellular function ...
explain the non-
Brownian motion
Brownian motion, or pedesis (from grc, πήδησις "leaping"), is the random motion of particles suspended in a medium (a liquid or a gas).
This pattern of motion typically consists of random fluctuations in a particle's position insi ...
of cytoplasmic constituents.
Constituents
The three major elements of the cytoplasm are the
cytosol
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 ...
,
organelles 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
The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compos ...
filaments, dissolved molecules, and water. The cytosol's filaments include the
protein filament
In biology, a protein filament is a long chain of protein monomers, such as those found in hair, muscle, or in flagella. Protein filaments form together to make the cytoskeleton of the cell. They are often bundled together to provide support, st ...
s such as
actin filament
Microfilaments, also called actin filaments, are protein filaments in the cytoplasm of eukaryotic cells that form part of the cytoskeleton. They are primarily composed of polymers of actin, but are modified by and interact with numerous other pro ...
s and
microtubules that make up the cytoskeleton, as well as soluble
protein
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, respo ...
s 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.
![Localisations02eng](https://upload.wikimedia.org/wikipedia/commons/6/6e/Localisations02eng.jpg)
Due to this network of fibres and high concentrations of dissolved
macromolecules, such as
protein
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, respo ...
s, an effect called
macromolecular crowding
The phenomenon of macromolecular crowding alters the properties of molecules in a solution when high concentrations of macromolecules such as proteins are present. Such conditions occur routinely in living cells; for instance, the cytosol of ''Esch ...
occurs and the cytosol does not act as an
ideal solution. 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
Golgi apparatus
The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic cells. Part of the endomembrane system in the cytoplasm, it packages proteins into membrane-bound vesicles ins ...
,
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s,
lysosome
A lysosome () is a membrane-bound organelle found in many animal cells. They are spherical vesicles that contain hydrolytic enzymes that can break down many kinds of biomolecules. A lysosome has a specific composition, of both its membrane pr ...
s, 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 or
silicon dioxide
Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
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
Polyhydroxybutyrate (PHB) is a polyhydroxyalkanoate (PHA), a polymer belonging to the polyesters class that are of interest as bio-derived and biodegradable plastics. The poly-3-hydroxybutyrate (P3HB) form of PHB is probably the most common typ ...
. A particularly widespread example are
lipid droplet
Lipid droplets, also referred to as lipid bodies, oil bodies or adiposomes, are lipid-rich cellular organelles that regulate the storage and hydrolysis of neutral lipids and are found largely in the adipose tissue. They also serve as a reservoi ...
s, 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 acid
In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
s and
sterols.
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 has been described.
See also
*
*
*
*
References
External links
*
{{Authority control
Cell anatomy