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Cell (biology)

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From Wikipedia, the free encyclopedia
Basic unit of life forms
This article is about the basic unit of lifeforms. For the branch of biology that studies them, seeCell biology.

Cell
Structure of an animal cell (eukaryotic)
Aeukaryotic cell as inanimals (left) and aprokaryotic cell as inbacteria (right)
Identifiers
MeSHD002477
THH1.00.01.0.00001
FMA686465
Anatomical terminology

Thecell is the basic structural and functional unit of allforms of life ororganisms. The term comes from theLatin wordcellula meaning 'small room'. A biological cell basically consists of asemipermeablecell membrane enclosingcytoplasm that contains genetic material. Most cells are only visible under amicroscope. Except for highly-differentiated cell types (examples includered blood cells andgametes) most cells are capable ofreplication, andprotein synthesis. Some types of cell aremotile. Cellsemerged on Earth about four billion years ago.

All organisms are grouped intoprokaryotes, andeukaryotes. Prokaryotes aresingle-celled, and includearchaea, andbacteria. Eukaryotes can be single-celled ormulticellular, and includeprotists,plants,animals, most types offungi, and some species ofalgae. All multicellular organisms are made up of many differenttypes of cell. Thediploid cells that make up the body of aplant oranimal are known assomatic cells, and in animals excludes thehaploidgametes.

Prokaryotic cells lack themembrane-boundnucleus present in eukaryotic cells, and instead have anucleoid region. In eukaryotic cells the nucleus is enclosed in thenuclear membrane. Eukaryotic cells contain othermembrane-bound organelles such asmitochondria, which provide energy for cell functions, andchloroplasts, in plants that create sugars byphotosynthesis. Othernon-membrane-bound organelles may beproteinaceous such as theribosomes present (though different) in both groups. A unique membrane-boundprokaryotic organelle themagnetosome has been discovered inmagnetotactic bacteria.

Cells were discovered byRobert Hooke in 1665, who named them after their resemblance tocells in amonastery.Cell theory, developed in 1839 byMatthias Jakob Schleiden andTheodor Schwann, states that all organisms are composed of one or more cells, that cells are the fundamental unit of structure and function in all organisms, and that all cells come from pre-existing cells.

Types

Organisms are broadly grouped intoeukaryotes, andprokaryotes. Eukaryotic cells possess amembrane-boundnucleus, andprokaryotic cells lack a nucleus but have anucleoid region.[1] Prokaryotes aresingle-celled organisms, whereas eukaryotes can be either single-celled ormulticellular. Single-celled eukaryotes includemicroalgae such asdiatoms. Multicellular eukaryotes include allanimals, andplants, mostfungi, and some species ofalgae.[2][3][4]

PropertyArchaeaBacteriaEukaryota
Cell membraneEther-linkedlipidsEster-linked lipidsEster-linked lipids
Cell wallGlycoprotein, orS-layer; rarelypseudopeptidoglycanPeptidoglycan, S-layer, or no cell wallVarious structures
Gene structureCircular chromosomes, similartranslation andtranscription to EukaryotaCircular chromosomes, unique translation and transcriptionMultiple, linear chromosomes, but translation and transcription similar to Archaea
Internalcell structureNo membrane-boundorganelles (?[5]) ornucleusNo membrane-bound organelles or nucleusMembrane-bound organelles and nucleus
Metabolism[6]Various, includingdiazotrophy, withmethanogenesis unique to ArchaeaVarious, includingphotosynthesis,aerobic andanaerobic respiration,fermentation, diazotrophy, andautotrophyPhotosynthesis, cellular respiration, and fermentation; no diazotrophy
ReproductionAsexual reproduction,horizontal gene transferAsexual reproduction, horizontal gene transferSexual and asexual reproduction
Protein synthesis initiationMethionineFormylmethionineMethionine
RNA polymeraseOneOneMany
EF-2/EF-GSensitive todiphtheria toxinResistant to diphtheria toxinSensitive to diphtheria toxin

Prokaryotes

Main article:Prokaryote
Structure of a typicalbacterial cell, generally similar with thearchaeal cell structure. Thebacterial flagellum shown, differs from thearchaellum in archaea
Bacteria andarchaea divide bybinary fission

Allprokaryotes are single-celled and includebacteria andarchaea, two of thethreedomains of life.[7] Prokaryotic cells were likely the first form oflife on Earth,[8][9] characterized by having vitalbiological processes includingcell signaling. They are simpler and smaller than eukaryotic cells, lack anucleus, and the other usually presentmembrane-bound organelles.[10]Prokaryotic organelles are less complex, and are typically non-membrane-bound.[11] All prokaryotic cellssecrete different substances from their membranes, includingexoenzymes, andextracellular polymeric substances.

Most prokaryotes are the smallest of all organisms, ranging from 0.5 to 2.0 μm in diameter.[12] The largest bacterium known,Thiomargarita magnifica, is visible to the naked eye with an average length of1 cm, but can be as much as2 cm[13][14]

Bacteria

Main article:Bacterial cell structure

Bacteria are enclosed in acell envelope, that protects the interior from the exterior.[15] It generally consists of aplasma membrane covered by acell wall which, for some bacteria, is covered by a third gelatinous layer called abacterial capsule. The capsule may bepolysaccharide as inpneumococci,meningococci orpolypeptide asBacillus anthracis orhyaluronic acid as instreptococci.Mycoplasma only possess the cell membrane.[16] The cell envelope gives rigidity to the cell and separates the interior of the cell from its environment, serving as a protective mechanical and chemical filter.[17] The cell wall consists ofpeptidoglycan and acts as an additional barrier against exterior forces.[18][17] The cell wall acts to protect the cell mechanically and chemically from its environment, and is an additional layer of protection to the cell membrane. It also prevents the cell from expanding and bursting (cytolysis) fromosmotic pressure due to ahypotonic environment.[19]

TheDNA of a bacterium typically consists of a singlecircular chromosome that is in direct contact with thecytoplasm in a region called thenucleoid. Some bacteria contain multiple circular or evenlinear chromosomes.[20][21][22] The cytoplasm also containsribosomes and various inclusions wheretranscription takes place alongsidetranslation.[23][24]Extrachromosomal DNA asplasmids, are usually circular and encode additional genes, such as those ofantibiotic resistance.[25] Linear bacterial plasmids have been identified in several species ofspirochete bacteria, including species ofBorrelia which causes Lyme disease.[26] Theprokaryotic cytoskeleton in bacteria is involved in the maintenance of cell shape,polarity and cytokinesis.[27]

Compartmentalization is a feature of eukaryotic cells but some species of bacteria, have protein-based organelle-likemicrocompartments such asgas vesicles, andcarboxysomes, andencapsulin nanocompartments.[28][29][30][31] Certain membrane-boundprokaryotic organelles have also been discovered. They include themagnetosome ofmagnetotactic bacteria,[29] and theanammoxosome ofanammox bacteria.[32][33]

Cell-surface appendages can includeflagella, andpili, protein structures that facilitate movement and communication between cells.[34] The flagellum stretches from the cytoplasm through the cell membrane and extrudes through the cell wall.[35]Fimbriae are short attachment pili, the other type of pilus is the longerconjugative type.[36] Fimbriae are formed of anantigenic protein calledpilin, and are responsible for the attachment of bacteria to specific receptors on host cells.[37]

Archaea

Main article:Archaea

Archaea are enclosed in a cell envelope consisting of a plasma membrane and a cell wall.An exception to this is theThermoplasma that only has the cell membrane.[16] The cell membranes of archaea are unique, consisting ofether-linked lipids. Theprokaryotic cytoskeleton has homologues of eukaryoticactin andtubulin.[27] A unique form of metabolism in the archaean ismethanogenesis. Theircell-surface appendage equivalent of the flagella is the differently structured and uniquearchaellum.[38][36] The DNA is contained in a circular chromosome in direct contact with the cytoplasm, in a region known as the nucleoid. Ribosomes are also found freely in the cytoplasm, or attached to the cell membrane where DNA processing takes place.[23][39]

The archaea are noted for theirextremophile species, and many are selectively evolved to thrive in extreme heat, cold, acidic, alkaline, or high salt conditions.[40] There are no known archaean pathogens.[41]

Eukaryotes

Main article:Eukaryote

Eukaryotes can be single-celled, as indiatoms (microscopic algae), or multicellular, as inanimals,plants, mostfungi, and somealgae.[42]Multicellular organisms are made up of many differenttypes of cell known overall assomatic cells.[43] Eukaryotes are distinguished by the presence of a membrane-boundnucleus.[44] The nucleus gives the eukaryote its name, which means "true nut" or "true kernel", where "nut" means the nucleus.[45] A eukaryotic cell can be 2 to 1000 times larger in diameter than a typical prokaryotic cell.[46]

Eukaryotic cells have acell membrane that surrounds a gel-likecytoplasm. The cytoplasm contains thecytoskeleton, the cell nucleus, the endoplasmic reticulum, ribosomes, the Golgi apparatus, mitochondria,lysosomes,peroxisomes,endosomes, vacuoles and vesicles, and may have a cell wall, chloroplasts,vaults, andcell-surface appendages. There are many cell variations among the different eukaryote groups.

The membranes of most of the organelles including the cell membrane are sometimes referred to as theendomembrane system.[47] All of these membranes are involved in thesecretory andendocytic pathways, modifying, packaging, and transporting proteins and lipids to and from the trans-Golgi network.[48] Inmammalian cells, endocytosis includes early, late, and recyclingendosomes.[48]

Most distinct cell types arise from a singletotipotent cell, called azygote, thatdifferentiates into hundreds of different cell types during the course ofdevelopment. Differentiation of cells is driven by different environmental cues (such as cell–cell interaction) and intrinsic differences (such as those caused by the uneven distribution ofmolecules duringdivision).[49]

Eukaryotic cell types include those that make upanimals,plants,fungi,algae, andprotists. All of which have many different species and cell differences.

Animal cells

Further information:Animal embryonic development andCell types
Structure of an animal cell

All the cells in an animal body develop from onetotipotentdiploid cell called azygote. During theembryonic development of ananimal, the cellsdifferentiate into the specialisedtissues andorgans of the organism. Different groups of cells differentiate from thegerm layers. Thesponge has only one layer. Some other animals known asdiploblasts have two germ layers theectoderm, and theendoderm. More advanced animals have an extra layer, the middlemesodermal layer, and are known astriploblastic. Triploblastic animals make up the largeclade ofBilateria. Differentiation results in structural or functional changes tostem cells, andprogenitor cells. There are an estimated 200 differentcell types in the human body. The estimatedcell count in a typical adult human body is around 30 trillion cells, 36 trillion in an adult male, and 28 trillion in a female.[50]

Structure

An animal cell has acell membrane that surrounds a gel-likecytoplasm. The cytoplasm contains thecytoskeleton, the cell nucleus, the endoplasmic reticulum, ribosomes, the Golgi apparatus, mitochondria,lysosomes,peroxisomes,endosomes, vacuoles and vesicles, andvaults. An animal cell structure, as other eukaryotes, includes anendomembrane system encompassing all the membranes of the organelles and the cell membrane, excluding the mitochondria.The whole system cooperates in the modification, packaging, and transport of proteins and lipids.[47]

Cell membrane

Main article:Cell membrane
Colored illustration showing the membrane of an animal cell surrounded by tissue. The nucleus, mitochondria, gogli, ER, and lysosomes are labelled.
Diagram of cell membrane detailing thelipid bilayer
Colored illustration showing the cell membrane and membrane proteins.
Diagram of cell membrane detailing membrane proteins

Thecell membrane, or plasma membrane, is aselectively permeablemembrane as an outer boundary of the cell that encloses the cytoplasm.[51] The membrane serves to separate and protect a cell from its surrounding environment and is made mostly from alipid bilayer ofphospholipids, which areamphiphilic (partlyhydrophobic and partlyhydrophilic). It has been best described in thefluid mosaic model.[52] Embedded within the cell membrane is amacromolecular structure called theporosome the universal secretory portal in cells and a variety ofprotein molecules that act as channels and pumps that move different molecules into and out of the cell.[23] The membrane is semi-permeable, and selectively permeable, in that it can either let a substance (molecule orion) pass through freely, to a limited extent or not at all.[53]Cell surface receptors embedded in the membrane allow cells to detect external signaling molecules such ashormones.[54]

Underlying, and attached to the cell membrane is thecell cortex, the outermost part of the actin cytoskeleton.[55]

Cytoplasm

The cell membrane encloses thecytoplasm of the cell that surrounds all of the cell's organelles.[56][57] It is made up of two main components, theprotein filaments that make up thecytoskeleton, and thecytosol.[56][57] The network of filaments and microtubules of the cytoskeleton gives shape and support to the cell, and has a part in organising the cell components. The cytosol is the main site ofprotein synthesis, anddegradation.[57]

The cytosol is a gel-like substance made up of water, ions, and non-essential biomolecules. The acidity (pH) of the cytosol is near neutral, and transporters in the cell membrane regulate this. Different proteins in the cytoplasm operate optimally at different pHs.[58] The cytosol forms30%–50% of the cell's volume.[59]

Cytoskeleton

Thecytoskeleton acts to organize and maintain the cell's shape; anchors organelles in place; helps duringendocytosis, and in the uptake of external materials by a cell.The cytoskeleton is composed ofmicrotubules,intermediate filaments andmicrofilaments. There are a great number of proteins associated with them, each controlling a cell's structure by directing, bundling, and aligning filaments. The outermost part of the cytoskeleton is thecell cortex, oractin cortex, a thin layer of cross-linkedactomyosins.[55] Its thickness varies with cell type and physiology.[55] It directs the transport through theER and theGolgi apparatus.[60] The cytoskeleton in the animal cell also plays a part incytokinesis, in the formation of thespindle apparatus duringcell division, the separation of daughter cells.

Organelles

Main article:Organelle

Organelles arecompartments of the cell that are specialized for carrying out one or more functions, analogous to theorgans, such as the heart, and lungs.[23] There are several types of organelles held in the cytoplasm. Most organelles are membrane-bounded, and vary in size and number based on the growth of the host cell.[61] Organelles include the nucleus, mitochondria, endoplasmic reticulum, Golgi apparatus, vesicles, and vacuoles.[62] Non membrane-bounded organelles include the centrosome, and typically the ribosome.[62]

Nucleus

Diagram of the nucleus showing theribosome-studdedouter nuclear membrane,nuclear pores,DNA (complexed aschromatin), and thenucleolus.
Deoxyribonucleic acid (DNA)

Thecell nucleus is the largest organelle in the animal cell.[50] It houses the cell'schromosomes, and is the place where almost allDNA replication andRNA synthesis (transcription) occur. The nucleus is spherical and separated from the cytoplasm by a double-membranednuclear envelope. A space between the membranes is called the perinuclear space. The nuclear envelope isolates and protects a cell's DNA from various molecules that could accidentally damage its structure or interfere with its processing. During processing,DNA istranscribed, or copied into a specialRNA, calledmessenger RNA (mRNA). This mRNA is then transported out of the nucleus, where it is translated into a specific protein molecule. Thenucleolus is a specialized region within the nucleus where ribosome subunits are assembled.[23] Cells use DNA for their long-term information storage that isencoded in itsDNA sequence.[23] RNA is used for information transport (e.g.,mRNA) andenzymatic functions (e.g.,ribosomal RNA).Transfer RNA (tRNA) molecules are used to add amino acids during proteintranslation.[63]

TheDNA of each cell is its genetic material, and is organized in multiple linear molecules, calledchromosomes, that are coiled aroundhistoneproteins and housed in thecell nucleus.[44][64] In humans, the nuclear genome is divided into 46 linear chromosomes, including 22homologous chromosome pairs and a pair ofsex chromosomes. The nucleus is amembrane-bound organelle. Other organelles in the cell have specific functions such asmitochondria which provide the cell's energy.[65]

Endoplasmic reticulum

Theendoplasmic reticulum (ER) is a transport network for molecules targeted for certain modifications and specific destinations, as compared to molecules that float freely in the cytoplasm. The ER has two forms: therough endoplasmic reticulum (RER), which hasribosomes on its surface that secrete proteins into the ER, and thesmooth endoplasmic reticulum (SER), which lacks ribosomes.[23] The smooth ER plays a role in calcium sequestration and release, and helps in synthesis oflipid.[66]

Golgi apparatus

TheGolgi apparatus processes and packagesproteins, andlipids, that are synthesized by the cell. It is organized as a stack of plate-like structures known ascisternae.[67]

Mitochondria

Mitochondria generate energy for the cell. Mitochondria are self-replicating double membrane-bound organelles that occur in various numbers, shapes, and sizes in the cytoplasm of the cell.[23]Respiration occurs in the cell mitochondria, which generate the cell's energy byoxidative phosphorylation, usingoxygen to release energy stored in cellular nutrients (typically pertaining toglucose) to generateATP (aerobic respiration).[68] Mitochondria multiply bybinary fission.[69] Mitochondria have their own DNA (mitochondrial DNA).[70] The mitochondrial genome is a circular DNA molecule distinct from nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes,[23] it codes for 13 proteins involved in mitochondrial energy production and specific tRNAs.[71]

Lysosomes

Lysosomes contain over 60 differenthydrolytic enzymes.[72] They digest excess or worn-out organelles, food particles, and engulfedviruses orbacteria. Lysosomes are optimally active in an acidic environment. The cell could not house these destructive enzymes if they were not contained in a membrane-bound system.[23][73]

Peroxisomes

Peroxisomes, aremicrobodies bounded by a single membrane. A peroxisome has no DNA or ribosomes and the proteins that it needs are encoded in the nucleus, and selectively imported from the cytosol. Some proteins enter via the endomembrane reticulum.[74] They have enzymes that rid the cell of toxicperoxides. The enzymatic content of the peroxisomes varies widely across the species, as it can in an individual organism.[75][74] The peroxisomes in animal cells are concentrated in theliver cells andadipocytes.[75]

Vacuoles

Vacuoles sequester waste products. Some cells, most notablyAmoeba, have contractile vacuoles, which can pump water out of the cell if there is too much water.[76]

Centrosome

Thecentrosome is a non membrane-bounded organelle composed ofpericentriolar material and the twocentrioles.[77][62] The centrosome is the mainmicrotubule organizing center in the animal cell that produces themicrotubules key components of the cytoskeleton. Centrosomes are composed of twocentrioles which lie perpendicular to each other in which each has an organization like acartwheel, which separate duringcell division and help in the formation of themitotic spindle.[78]

Ribosomes

Aribosome is a large complex ofRNA andprotein molecules often considered as anon-membrane-bound organelle.[23] They each consist of two subunits, one larger than the other, and act as an assembly line where RNA from the nucleus is used to synthesise proteins from amino acids. Ribosomes can be found either floating freely or bound to a membrane of the rough endoplasmatic reticulum.[79]

Vaults

Avault is a large ribonuclear protein particle, anon-membrane-bound organelle, three times the size of a ribosome but with only three proteins in contrast to the near hundred in the ribosome.[80] Most human cells have around 10,000 vaults, and in some types ofimmune cell there may be up to 100,000.Macrophages have the greatest number of vaults of any human cell.[81] Vaults are largely overlooked because their functions are purely speculative. They may play a role in transport from the nucleus to the cytoplasm, and may serve as scaffolds for signal transduction proteins. They are present in normal tissues, and more so in secretory and excretory epithelial cells.[80][81]

Animal cell types

See also:Cell type

Some types of specialised cell are localised to a particular animal group.Vertebrates for example have specialised, structurally changed cells includingmuscle cells. The cell membrane of askeletal muscle cell or of acardiac muscle cell is termed thesarcolemma.[82] And the cytoplasm is termed thesarcoplasm. Skeletal muscle cells also becomemultinucleated. Populations of animal groupsevolve to become distinct species, where sexual reproduction is isolated. The manyspecies of vertebrates for example have other unique characteristics by way of additional specialised cells. In some species ofelectric fish for example modified muscle cells or nerve cells have specialised to becomeelecterocytes capable of creating and storing electrical energy for future release, as in stunning prey, or use inelectrolocation.[83] These are large flat cells in theelectric eel, andelectric ray in which thousands are stacked into anelectric organ comparable to avoltaic pile.[84]

Many animal cells areciliated and most cells except red blood cells haveprimary cilia. Primary cilia play important roles inchemosensation andmechanosensation.[85][86] Each cilium may be "viewed as a sensory cellularantennae that coordinates a large number of cellular signaling pathways, sometimes coupling the signaling to ciliary motility or alternatively to cell division and differentiation."[87] The cilia in other cells are motile organelles, and in therespiratory epithelium play an important role in themovement of mucus. In the reproductive system ciliated epithelium in thefallopian tubes move the egg from the uterus to the ovary. Motile cilia also known as flagella, drive the sperm cells.[88]Invertebrateplanarians have ciliated excretoryflame cells.[89] Other excretory cells also found in planarians aresolenocytes that are long and flagellated.

Plant cells

Main article:Plant cell
Structure of a typicalplant cell
Onion (Allium cepa) root cells in different phases of thecell cycle (drawn byE.B. Wilson, 1900)

Other types of organelle specific toplant cells, arepigment-containingplastids, especiallychloroplasts that containchlorophyll. Chloroplasts capture the sun's energy to make carbohydrates throughphotosynthesis.[90]Chromoplasts contain fat-solublecarotenoid pigments such as orangecarotene and yellowxanthophylls which helps in synthesis and storage.Leucoplasts are non-pigmented plastids and helps in storage of nutrients.[91]

Plastids divide by binary fission.Vacuoles in plant cells store water, and are surrounded by a membrane.[92] The vacuoles of plant cells are usually larger than those of animal cells. The vacuole membrane transports ions against concentration gradients.[93]

The plant cytoskeleton is a dynamic structure that has a scaffold of microtubules and microfilaments, but not the intermediate filaments.[94] Themicrotubule organizing center in plant cells is often sited underneath the cell membrane where nucleated microtubules often form sheet-like semi-parallel arrays.[95]

There are two types of peroxisomes in plants. One type is in the leaves where it takes part inphotorespiration. The other type is in germinating seeds where they take part in the conversion of fatty acids into sugars for the plant's growth.[74] In this peroxisome type the enzymatic content is so different than in other groups that it has an alternative name ofglyoxysome, their enzymes are of theglyoxylate cycle.[75]

Algal cells

Further information:Eukaryotic algae

Algae members arephotoautotrophs able to usephotosynthesis to produce energy. Photosynthesis is made possible by the use ofplastids, organelles in the cytoplasm known aschloroplasts. Algal photoautotrophs includered algae.[96]

Alginate is apolysaccharide found in the matrix of the cell walls ofbrown algae, and has many important uses in the food industry, and inpharmacology.[97]

Fungal cells

Main article:Fungus

The cells offungi have in addition to the shared eukaryotic organelles aspitzenkörper in their endomembrane system, associated withhyphal tip growth. It is aphase-dark body that is composed of an aggregation of membrane-bound vesicles containing cell wall components, serving as a point of assemblage and release of such components intermediate between the Golgi and the cell membrane. The spitzenkörper is motile and generates new hyphal tip growth as it moves forward.[98]

The cell walls of fungi are uniquely made of achitin-glucan complex.[99]

Protist cells

Further information:Protist § Common types

The cells ofprotists may be bounded only by acell membrane, or may in addition have acell wall, or may be covered by apellicle (inciliates), atest (intestate amoebae), or afrustule (indiatoms).

Some protists such asamoebae mayfeed on other organisms and ingest food byphagocytosis. Vacuoles known asphagosomes in the cytoplasm may be used to draw in and incorporate the captured particles. Other types of protists arephotoautotrophs, providing themselves with energy by photosynthesis.[100] Most single-celled protists are motile, andgenerate movement with cilia, flagella, orpseudopodia.[101]

Ciliates have two different sorts ofnuclei: a tiny,diploidmicronucleus (the "generative nucleus", which carries thegermline of the cell), and a large,ampliploidmacronucleus (the "vegetative nucleus", which takes care of general cell regulation.[102][103]

Physiology

See also:Cell cycle andCell physiology
Prokaryotes divide bybinary fission, whileeukaryotes divide bymitosis ormeiosis.

Replication

Main article:Cell division
Human cancer cells, specificallyHeLa cells, with DNA stained blue. The central and rightmost cell are ininterphase, so their DNA is diffuse and the entire nuclei are labelled. The cell on the left is going throughmitosis and its chromosomes have condensed.

Duringcell division, a single cell, themother cell divides into two daughter cells. This leads to the growth oftissue inmulticellular organisms.Prokaryotic cells divide bybinary fission, whileeukaryotic cells usually undergo a process of nuclear division, calledmitosis, followed by division of the cell, calledcytokinesis. Adiploid cell may undergomeiosis to produce haploid cells, usually four.Haploid cells serve asgametes in multicellular organisms, fusing to form new diploid cells.[citation needed]

DNA replication, or the process of duplicating a cell's genome,[23] always happens when a cell divides through mitosis or binary fission.[citation needed] This occurs during theS (synthesis) phase of thecell cycle.[citation needed]

In meiosis, the DNA is replicated only once, while the cell divides twice. DNA replication only occurs beforemeiosis I. DNA replication does not occur when the cells divide the second time, inmeiosis II.[104] Replication, like all cellular activities, requires specialized proteins.[23]

Signaling

Main article:Cell signaling

Cell signaling is the process by which a cell interacts with itself, other cells, and the environment. Typically, the signaling process involves three components: the first messenger (the ligand), thereceptor, and the signal itself.[105] Most cell signaling is chemical in nature, and can occur with neighboring cells or more distant targets. Signal receptors are complex proteins or tightly bound multimer of proteins,located in the plasma membrane orwithin the interior.[106]

Each cell is programmed to respond to specific extracellular signal molecules, and this process is the basis ofdevelopment,tissue repair,immunity, andhomeostasis. Individual cells are able to manage receptor sensitivity including turning them off, and receptors can become less sensitive when they are occupied for long durations.[106] Errors in signaling interactions may cause diseases such ascancer,autoimmunity, anddiabetes.[107]

Protein targeting

Protein targeting or protein sorting is thebiological mechanism by whichproteins are transported to their appropriate destinations within or outside the cell.[108][109]

Proteins can be targeted to the inner space of anorganelle, different intracellularmembranes, theplasma membrane, or to the exterior of the cell viasecretion.[108][109] Information contained in the protein itself directs this delivery process.[109][110] Correct sorting is crucial for the cell; errors or dysfunction in sorting have been linked to multiple diseases.[109][111][112]

DNA repair

Main article:DNA repair

All cells contain enzyme systems that scan forDNA damage and carry outrepair. Diverse repair processes have evolved in all organisms. Repair is vital to maintain DNA integrity, avoid cell death and errors of replication that could lead tomutation. Repair processes includenucleotide excision repair,DNA mismatch repair,non-homologous end joining of double-strand breaks,recombinational repair and light-dependent repair (photoreactivation).[113]

Growth and metabolism

Main articles:Cell growth,Metabolism, andPhotosynthesis

Between successive cell divisions, cells grow through the functioning of cellular metabolism. Cell metabolism is the process by which individual cells process nutrient molecules. Metabolism has two distinct divisions:catabolism, in which the cell breaks down complex molecules to produce energy andreducing power, andanabolism, in which the cell uses energy and reducing power to construct complex molecules and perform other biological functions.[114]

Complex sugars can be broken down into simpler sugar molecules calledmonosaccharides such asglucose. Once inside the cell, glucose is broken down to make adenosine triphosphate (ATP),[23] a molecule that possesses readily available energy, through two different pathways. In plant cells,chloroplasts create sugars byphotosynthesis, using the energy of light to join molecules of water andcarbon dioxide.[115]

Protein synthesis

Main article:Protein biosynthesis

Cells are capable of synthesizing new proteins, which are essential for the modulation and maintenance of cellular activities. This process involves the formation of new protein molecules fromamino acid building blocks based on information encoded in DNA/RNA. Protein synthesis generally consists of two major steps:transcription andtranslation.[63]

Transcription is the process where genetic information in DNA is used to produce a complementary RNA strand. This RNA strand is then processed to givemessenger RNA (mRNA), which is free to migrate into the cytoplasm. mRNA molecules bind to protein-RNA complexes calledribosomes located in thecytosol, where they are translated into polypeptide sequences. The ribosome mediates the formation of a polypeptide sequence based on the mRNA sequence. The mRNA sequence directly relates to the polypeptide sequence by binding totransfer RNA (tRNA) adapter molecules in binding pockets within the ribosome.[63] The new polypeptide then folds into a functional three-dimensional protein molecule.

Motility

Main article:Motility

Unicellular organisms can move in order to find food or escape predators. Common mechanisms of motion includeflagella andcilia.[36]

In multicellular organisms, cells can move during processes such as wound healing, the immune response andcancer metastasis. For example, in wound healing in animals, white blood cells move to the wound site to kill the microorganisms that cause infection. Cell motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.[116] The process is divided into three steps: protrusion of the leading edge of the cell, adhesion of the leading edge and de-adhesion at the cell body and rear, and cytoskeletal contraction to pull the cell forward. Each step is driven by physical forces generated by unique segments of the cytoskeleton.[117][116]

Navigation, control and communication

See also:Cybernetics § In biology

In August 2020, scientists described one way cells—in particular cells of a slime mold and mouse pancreatic cancer-derived cells—are able tonavigate efficiently through a body and identify the best routes through complex mazes: generating gradients after breaking down diffusedchemoattractants which enable them to sense upcoming maze junctions before reaching them, including around corners.[118][119][120]

Cell death

Main article:Cell death

Cell death occurs when a cell ceases to carry out its functions, as a result of ageing, or types ofcell injury (necrosis).Programmed cell death, includingapoptosis, andautophagy is a natural process of replacing dead cells with new ones.[121][122]

A separate mode of cellular death is known as amitotic catastrophe, which occurs duringmitosis, following the improper progression of, or entrance to thecell cycle. This mechanism operates to prevent genomic instability.[123][124]

Staining of anematodeCaenorhabditis elegans highlights thenuclei of its cells.

Origins

Main article:History of life
Further information:Abiogenesis andEvolution of cells
Stromatolites are left behind bycyanobacteria, known as blue-green algae. They are among the oldest fossils of life on Earth. This one-billion-year-old fossil is fromGlacier National Park in the United States.

The origin of cells has to do with theorigin of life, which began thehistory of life on Earth. Small molecules needed for life may have been carried to Earth on meteorites, created atdeep-sea vents, orsynthesized by lightning in a reducing atmosphere. There is little experimental data defining what the first self-replicating forms were.RNA may have beenthe earliest self-replicating molecule, as it can both store genetic information and catalyze chemical reactions.[125] This process required an enzyme tocatalyze the RNA reactions, which may have been the earlypeptides that formed inhydrothermal vents.[126]

Cells emerged around 4 billion years ago.[127][128] The first cells were most likelyheterotrophs. The early cell membranes were probably simpler and more permeable than later ones, with only a single fatty acid chain per lipid. Lipids spontaneously form bilayeredvesicles in water, and could have preceded RNA.[129][130]

In the theory ofsymbiogenesis, a merger of anarchaean and an aerobic bacterium created the eukaryotes, with aerobicmitochondria, some 2.2 billion years ago. A second merger, 1.6 billion years ago, addedchloroplasts, creating thegreen plants.[131]

Eukaryotic cells were created some 2.2 billion years ago in a process calledeukaryogenesis. This is widely agreed to have involvedsymbiogenesis, in which anarchaean and abacterium came together to create thefirst eukaryotic common ancestor.[131] It evolved into a population of single-celled organisms that included thelast eukaryotic common ancestor, gaining capabilities along the way.[132][133]

This cell had a new level of complexity, with a nucleus[134][132] and facultatively aerobicmitochondria.[131] It featured at least onecentriole andcilium, sex (meiosis andsyngamy),peroxisomes, and a dormantcyst with a cell wall ofchitin and/orcellulose.[135][133] The last eukaryotic common ancestor gave rise to the eukaryotes'crown group, containing the ancestors ofanimals,fungi,plants, and a diverse range of single-celled organisms.[136][137] Thegreen plants were created around 1.6 billion years ago with a second episode of symbiogenesis that addedchloroplasts, derived fromcyanobacteria.[131]

Multicellularity

Multicellular behavior is demonstrated bymicroorganisms that arecloned from a single cell and form visiblemicrobial colonies. Amicrobial consortium of two or more species can form abiofilm by the secretion ofextracellular polymeric substances (EPSs).[138]Slime molds consist of different groups of microorganisms grouped together in a multicellular-like fashion.

The first evidence of multicellularity in an organism comes fromcyanobacteria-like organisms that lived between 3 and 3.5 billion years ago.[139] Cyanobacteria are variable in morphology,filamentous forms exhibit functional cell differentiation such asheterocysts (for nitrogen fixation),akinetes (resting stage cells), andhormogonia (reproductive, motile filaments). These, together with the intercellular connections they possess, are considered the first signs of multicellularity.[140]

Staining of anematodeCaenorhabditis elegans highlights thenuclei of its cells.

Multicellularity was made possible by the development of theextracellular matrix (ECM) similar in function to the bacterial ECM that consists of extracellular polymeric substances.[141] EPS enables microbial cell adhesion, and is believed to be the first evolutionary step toward multicellular organisms.[142]Basement membranes are a type of specialized extracellular matrix that surrounds mostanimal tissues, and are essential in their formation. Their emergence coincided with the origin of multicellularity.[143]

The evolution of multicellularity from unicellular ancestors has been replicated in the laboratory, inevolution experiments using predation as theselective pressure.[139]

History of research

Main article:Cell theory § Discovery of cells
Robert Hooke's drawing of cells incork, 1665

In 1665,Robert Hooke examined a thin slice of cork under hismicroscope, and saw a structure of small enclosures. He wrote "I could exceeding plainly perceive it to be all perforated and porous, much like ahoneycomb, but that the pores of it were not regular".[144] To further support his theory,Matthias Schleiden andTheodor Schwann studied cells of both animal and plants. What they discovered were significant differences between the two types of cells. This put forth the idea that cells were fundamental to both plants and animals.[145]

See also

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