View source for Cytoplasm

{{Short description|All of the contents of a eukaryotic cell except the nucleus}}
{{Use dmy dates|date=July 2020}}
{{Organelle diagram}}

In [[cell biology]], the '''cytoplasm''' describes all material within a [[eukaryote|eukaryotic]] [[Cell (biology)|cell]], enclosed by the [[cell membrane]], except for the [[cell nucleus]]. The material inside the nucleus and contained within the [[nuclear envelope|nuclear membrane]] is termed the [[nucleoplasm]]. The main components of the cytoplasm are the [[cytosol]] (a gel-like substance), the [[organelle]]s (the cell's internal sub-structures), and various [[cytoplasmic inclusion]]s. The cytoplasm is about 80% water and is usually colorless.<ref>{{cite book | vauthors = Shepherd VA | title = The cytomatrix as a cooperative system of macromolecular and water networks | volume = 75 | pages = 171–223 | date = 2006 | pmid = 16984813 | doi = 10.1016/S0070-2153(06)75006-2 | isbn = 9780121531751 | series = Current Topics in Developmental Biology }}</ref>

The submicroscopic ground cell substance, or cytoplasmic matrix, that remains after the exclusion of the cell [[organelle]]s and particles is [[groundplasm]].  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]]s, [[mitochondria]], plant [[plastid]]s, [[lipid]] droplets, and [[vacuole]]s.

Many cellular activities take place within the cytoplasm, such as many [[metabolic pathway]]s,  including [[glycolysis]], [[photosynthesis]],  and processes such as [[cell division]]. The concentrated inner area is called the [[endoplasm]] and the outer layer is called the [[cell cortex]], or [[ectoplasm (cell biology)|ectoplasm]].

Movement of [[calcium ion]]s in and out of the cytoplasm is a [[Recognition signal|signaling]] activity for [[metabolic]] processes.<ref>{{cite book | last = Hogan | first = C. Michael | date = 2010 | chapter-url = http://www.eoearth.org/article/Calcium?topic=49557 | chapter = Calcium | archive-url = https://web.archive.org/web/20120612123626/http://www.eoearth.org/article/Calcium?topic=49557 | archive-date=12 June 2012 | title = Encyclopedia of Earth | veditors = Jorgensen A, Cleveland C | publisher = National Council for Science and the Environment }}</ref>

In [[plant]]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.<ref>{{cite book | last = von Kölliker | first = Rudolf | name-list-style = vanc  | date = 1863 | chapter-url = https://books.google.com/books?id=5mtARc4NAi0C | title = Handbuch der Gewebelehre des Menschen | chapter = 4. Auflage | location = Leipzig | publisher = Wilhelm Engelmann }}</ref><ref>{{cite book | vauthors = Bynum WF, Browne EJ, Porter R | date = 1981  | url = https://books.google.com/books?id=Ian_AwAAQBAJ | title = Dictionary of the history of science | location = Princeton University Press | isbn = 9781400853410 }}</ref>

There has been certain disagreement on the definition of cytoplasm, as some authors prefer to exclude from it some organelles, especially the [[vacuole]]s<ref>{{cite book |vauthors=Parker J |date=1972 |chapter=Protoplasmic resistance to water deficits |pages=125–176 |veditors=Kozlowski TT |title=Water deficits and plant growth, Vol. III. Plant responses and control of water balance. |publisher=Academic Press |location=New York |isbn=9780323153010 |chapter-url=https://books.google.com/books?id=gOEr2alLRUYC}}</ref> and sometimes the plastids.<ref>{{cite journal| vauthors = Strasburger E |year=1882|title=Ueber den Theilungsvorgang der Zellkerne und das Verhältnis der Kernteilung zur Zellteilung|journal=Arch Mikr Anat |volume=21|pages=476–590|doi=10.1007/BF02952628|hdl=2027/hvd.32044106199177|s2cid=85233009|hdl-access=free |url=https://www.biodiversitylibrary.org/item/49525#page/536/mode/1up|url-status=live|archive-url= https://web.archive.org/web/20170827124018/http://www.biodiversitylibrary.org/item/49525#page/536/mode/1up |archive-date=27 August 2017}}</ref>

==Physical nature==
It remains uncertain how the various components of the cytoplasm interact to allow movement of [[organelle]]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 [[Semipermeable membrane|permeability]] of the cytoplasm.<ref>{{cite book |chapter=Spatial Modeling of Cell Signaling Networks |pmc=3519356 |pmid=22482950 |doi=10.1016/B978-0-12-388403-9.00008-4 |volume=110 |year=2012 |pages=195–221 |vauthors=Cowan AE, Moraru II, Schaff JC, Slepchenko BM, Loew LM |isbn=9780123884039 |title=Computational Methods in Cell Biology}}</ref> An example of such function is [[cell signalling]], a process which is dependent on the manner in which signaling molecules are allowed to [[diffuse]] across the cell.<ref>{{cite journal | vauthors = Holcman D, Korenbrot JI | title = Longitudinal diffusion in retinal rod and cone outer segment cytoplasm: the consequence of cell structure | journal = Biophysical Journal | volume = 86 | issue = 4 | pages = 2566–82 | date = April 2004 | pmid = 15041693 | pmc = 1304104 | doi = 10.1016/S0006-3495(04)74312-X | bibcode = 2004BpJ....86.2566H }}</ref> While small signaling molecules like [[calcium ions]] are able to diffuse with ease, larger molecules and subcellular structures often require aid in moving through the cytoplasm.<ref name="autogenerated1">{{cite journal | vauthors = Parry BR, Surovtsev IV, Cabeen MT, O'Hern CS, Dufresne ER, Jacobs-Wagner C | title = The bacterial cytoplasm has glass-like properties and is fluidized by metabolic activity | journal = Cell | volume = 156 | issue = 1–2 | pages = 183–94 | date = January 2014 | pmid = 24361104 | pmc = 3956598 | doi = 10.1016/j.cell.2013.11.028 | bibcode = 2014APS..MARJ16002P }}</ref> 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]].<ref>{{cite journal|title=The contractile vacuole in Euplotes: An example of the sol-gel reversibility of cytoplasm| doi=10.1002/jez.1400370302 |volume=37 |issue=3 |journal=Journal of Experimental Zoology |pages=259–289 |year=1923 |vauthors=Taylor CV| bibcode=1923JEZ....37..259T }}</ref> 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]] smaller than 100&nbsp;[[nanometer|nm]], the cytoplasm acts like a liquid, while in a larger length scale, it acts like a gel.<ref>{{cite journal |last1=Kwapiszewska |first1=Karina |last2=Szczepański |first2=Krzysztof |title=Nanoscale Viscosity of Cytoplasm Is Conserved in Human Cell Lines |journal=[[The Journal of Physical Chemistry Letters]] |date=31 July 2020 |volume=11 |issue=16 |pages=6914–6920 |doi=10.1021/acs.jpclett.0c01748 |pmid=32787203 |pmc=7450658 |display-authors=1|doi-access=free }}</ref>

===As a glass===

It has been proposed that the cytoplasm behaves like a [[glass]]-forming liquid approaching the [[glass transition]].<ref name="autogenerated1"/> 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).<ref name="autogenerated1"/> 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]].<ref name="autogenerated1"/>

===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.<ref>{{cite journal | vauthors = Guo M, Ehrlicher AJ, Jensen MH, Renz M, Moore JR, Goldman RD, Lippincott-Schwartz J, Mackintosh FC, Weitz DA | title = Probing the stochastic, motor-driven properties of the cytoplasm using force spectrum microscopy | journal = Cell | volume = 158 | issue = 4 | pages = 822–832 | date = August 2014 | pmid = 25126787 | pmc = 4183065 | doi = 10.1016/j.cell.2014.06.051 }}</ref>

==Constituents==
The three major elements of the cytoplasm are the [[cytosol]], [[organelle]]s and [[cytoplasmic inclusions|inclusions]].

===Cytosol===
{{main|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 filament]]s such as [[actin filament]]s and [[microtubule]]s that make up the cytoskeleton, as well as soluble [[protein]]s and small structures such as [[ribosome]]s, [[proteasome]]s, and the mysterious [[Vault (organelle)|vault complexes]].<ref>{{cite journal | vauthors = van Zon A, Mossink MH, Scheper RJ, Sonneveld P, Wiemer EA | s2cid = 21196262 | title = The vault complex | journal = Cellular and Molecular Life Sciences | volume = 60 | issue = 9 | pages = 1828–37 | date = September 2003 | pmid = 14523546 | doi = 10.1007/s00018-003-3030-y | pmc = 11138885 }}</ref> The inner, granular and more fluid portion of the cytoplasm is referred to as endoplasm.[[File:Localisations02eng.jpg|thumb|right|250px|Proteins in different [[cellular compartment]]s and structures [[Protein tag#Protein tags|tagged]] with [[green fluorescent protein]]]]

Due to this network of fibres and high concentrations of dissolved [[macromolecule]]s, such as [[protein]]s, an effect called [[macromolecular crowding]] 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===
{{main|Organelle}}
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]], the [[endoplasmic reticulum]], the [[Golgi apparatus]], [[vacuole]]s, [[lysosome]]s, and in plant cells, [[chloroplast]]s.

===Cytoplasmic inclusions===
{{main|Cytoplasmic inclusion}}
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]] in plants,<ref name=Prychid1999>{{Cite journal |author1=Prychid, Christina J. |author2=Rudall, Paula J. | year = 1999 | title = Calcium Oxalate Crystals in Monocotyledons: A Review of their Structure and Systematics | journal = Annals of Botany | volume = 84 | issue = 6 | pages = 725–739 | doi = 10.1006/anbo.1999.0975 | url = https://academic.oup.com/aob/article-pdf/84/6/725/7983834/840725.pdf | doi-access = free }}</ref><ref name=Prychid2003>{{Cite journal | vauthors = Prychid CJ, Rudall PJ |author3=Gregory, M. | year = 2004 | title = Systematics and Biology of Silica Bodies in Monocotyledons | journal = The Botanical Review | volume = 69 | issue = 4 | pages = 377–440 | doi = 10.1663/0006-8101(2004)069[0377:SABOSB]2.0.CO;2 | jstor = 4354467|s2cid=24520433 }}</ref> to granules of energy-storage materials such as [[starch]],<ref>{{cite journal | vauthors = Ball SG, Morell MK | title = From bacterial glycogen to starch: understanding the biogenesis of the plant starch granule |journal=Annual Review of Plant Biology |volume=54 |pages=207–233 |year=2003 |pmid=14502990 |doi=10.1146/annurev.arplant.54.031902.134927}}</ref> [[glycogen]],<ref>{{cite journal |vauthors=Shearer J, Graham TE |title=New perspectives on the storage and organization of muscle glycogen |journal=Canadian Journal of Applied Physiology |volume=27 |issue=2 |pages=179–203 |date=April 2002 |pmid=12179957 |doi=10.1139/h02-012}}</ref> or [[polyhydroxybutyrate]].<ref>{{cite journal |vauthors=Anderson AJ, Dawes EA |title=Occurrence, metabolism, metabolic role, and industrial uses of bacterial polyhydroxyalkanoates |journal=Microbiological Reviews |volume=54 |issue=4 |pages=450–472 |date=December 1990 |pmid=2087222 |pmc=372789 |doi=10.1128/MMBR.54.4.450-472.1990 }}</ref> A particularly widespread example are [[lipid droplet]]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]]s and [[sterol]]s.<ref>{{cite journal |vauthors=Murphy DJ |title=The biogenesis and functions of lipid bodies in animals, plants and microorganisms |journal=Progress in Lipid Research |volume=40 |issue=5 |pages=325–438 |date=September 2001 |pmid=11470496 |doi=10.1016/S0163-7827(01)00013-3}}</ref> Lipid droplets make up much of the volume of [[adipocyte]]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 [[Viscoelasticity|viscoplastic behavior]] and a measure of the reciprocal rate of bond breakage within the cytoplasmic network.<ref name=Viscoplasticity>{{cite journal |vauthors=Feneberg W, Westphal M, Sackmann E |s2cid=9782043 |title=Dictyostelium cells' cytoplasm as an active viscoplastic body |journal=European Biophysics Journal |volume=30 |issue=4 |pages=284–94 |date=August 2001 |pmid=11548131 |doi=10.1007/s002490100135}}</ref>

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.<ref name="pmid28827333">{{cite journal | vauthors = Hu J, Jafari S, Han Y, Grodzinsky AJ, Cai S, Guo M | title = Size- and speed-dependent mechanical behavior in living mammalian cytoplasm | journal = Proc. Natl. Acad. Sci. U.S.A. | volume = 114 | issue = 36 | pages = 9529–9534 | date = September 2017 | pmid = 28827333 | pmc = 5594647 | doi = 10.1073/pnas.1702488114 | bibcode = 2017PNAS..114.9529H | doi-access = free }}</ref>

== See also ==
* {{annotated link|Amoeboid movement}}
* {{annotated link|Cytoplasmic streaming}}
* {{annotated link|Protoplasm}}
* {{annotated link|Syncytium}}

== References ==
{{Reflist|35em}}

== External links ==
* {{cite book | author = Luby-Phelps K | title = Microcompartmentation and Phase Separation in Cytoplasm | year = 2000 | chapter = Cytoarchitecture and physical properties of cytoplasm: volume, viscosity, diffusion, intracellular surface area | url = http://www.rpgroup.caltech.edu/courses/aph161/Handouts/Luby-Phelps2000.pdf | journal = Int Rev Cytol | volume = 192 | pages = 189–221 | doi = 10.1016/S0074-7696(08)60527-6 | pmid = 10553280 | series = International Review of Cytology | isbn = 9780123645968 | archive-url = https://web.archive.org/web/20080911153732/http://www.rpgroup.caltech.edu/courses/aph161/Handouts/Luby-Phelps2000.pdf | archive-date = 11 September 2008 | url-status = dead }}

{{organelles}}

{{Authority control}}

[[Category:Cell anatomy]]