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 an ...

, 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

organ Organ and organs may refer to: Biology * Organ (biology), a group of tissues organized to serve a common function * Organ system, a collection of organs that function together to carry out specific functions within the body. Musical instruments ...

s are to the

body Body may refer to: In science * Physical body, an object in physics that represents a large amount, has mass or takes up space * Body (biology), the physical material of an organism * Body plan, the physical features shared by a group of anim ...

, hence ''organelle,'' the suffix ''-elle'' being a

diminutive A diminutive is a word obtained by modifying a root word to convey a slighter degree of its root meaning, either to convey the smallness of the object or quality named, or to convey a sense of intimacy or endearment, and sometimes to belittle s ...

. Organelles are either separately enclosed within their own lipid bilayers (also called membrane-bounded organelles) or are spatially distinct functional units without a surrounding lipid bilayer (non-membrane bounded organelles). Although most organelles are functional units within cells, some functional units that extend outside of cells are often termed organelles, such as

cilia The cilium (: cilia; ; in Medieval Latin and in anatomy, ''cilium'') is a short hair-like membrane protrusion from many types of eukaryotic cell. (Cilia are absent in bacteria and archaea.) The cilium has the shape of a slender threadlike proj ...

, the

flagellum A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...

and

archaellum The archaellum (: archaella; formerly archaeal flagellum) is a unique structure on the cell surface of many archaea that allows for swimming motility. The archaellum consists of a rigid helix, helical filament that is attached to the cell membr ...

, and the

trichocyst A trichocyst is an organelle found in certain ciliates and dinoflagellate The Dinoflagellates (), also called Dinophytes, are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered pr ...

(these could be referred to as membrane bound in the sense that they are attached to (or bound to) the membrane). Organelles are identified by

microscopy Microscopy is the technical field of using microscopes to view subjects too small to be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of microscopy: optical mic ...

, and can also be purified by

cell fractionation In cell biology, cell fractionation is the process used to separate cell organelle, cellular components while preserving individual functions of each component. This is a method that was originally used to demonstrate the cellular location of vari ...

. There are many types of organelles, particularly in eukaryotic cells. They include structures that make up the

endomembrane system The endomembrane system is composed of the different membranes (endomembranes) that are suspended in the cytoplasm within a eukaryotic cell. These membranes divide the cell into functional and structural compartments, or organelles. In eukaryote ...

(such as the

nuclear envelope 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 membran ...

endoplasmic reticulum The endoplasmic reticulum (ER) is a part of a transportation system of the eukaryote, eukaryotic cell, and has many other important functions such as protein folding. The word endoplasmic means "within the cytoplasm", and reticulum is Latin for ...

, and

Golgi apparatus The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic Cell (biology), cells. Part of the endomembrane system in the cytoplasm, it protein targeting, packages proteins ...

), and other structures such as

mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...

and

plastid A plastid is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. Plastids are considered to be intracellular endosymbiotic cyanobacteria. Examples of plastids include chloroplasts ...

s. While

prokaryote A prokaryote (; less commonly spelled procaryote) is a unicellular organism, single-celled organism whose cell (biology), cell lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Gree ...

s do not possess eukaryotic organelles, some do contain

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...

-shelled

bacterial microcompartment Bacterial microcompartments (BMCs) are organelle-like structures found in bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large do ...

s, which are thought to act as primitive

prokaryotic organelles A prokaryote (; less commonly spelled procaryote) is a single-celled organism whose cell lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Greek (), meaning 'before', and (), meaning 'nut' o ...

; and there is also evidence of other membrane-bounded structures. Also, the prokaryotic

which protrudes outside the cell, and its motor, as well as the largely extracellular

pilus A pilus (Latin for 'hair'; : pili) is a hair-like cell-surface appendage found on many bacteria and archaea. The terms ''pilus'' and '' fimbria'' (Latin for 'fringe'; plural: ''fimbriae'') can be used interchangeably, although some researchers ...

, are often spoken of as organelles.

History and terminology

In biology, ''

s'' are defined as confined functional units within an

organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...

. The

analogy Analogy is a comparison or correspondence between two things (or two groups of things) because of a third element that they are considered to share. In logic, it is an inference or an argument from one particular to another particular, as oppose ...

of bodily organs to microscopic cellular substructures is obvious, as from even early works, authors of respective textbooks rarely elaborate on the distinction between the two. In the 1830s, Félix Dujardin refuted Ehrenberg's theory that microorganisms have the same organs as multicellular animals, only smaller. Credited as the first to use a

of ''organ'' (i.e., little organ) for cellular structures was German zoologist Karl August Möbius (1884), who used the term ''organula'' (plural of ''organulum'', the diminutive of

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

''organum''). In a footnote, which was published as a correction in the next issue of the journal, he justified his suggestion to call organs of unicellular organisms "organella" since they are only differently formed parts of one cell, in contrast to multicellular organs of multicellular organisms.

Types

In the broadest definition, an organelle is any part of the cell that acts as a distinct functional unit. This includes ''membrane-bounded'' as well as ''non-membrane-bounded'' organelles. In a more restrictive definition, only membrane-bounded ones are included. In the most restrictive definition, only the endosymbiotic membrane-bounded ones are included. The membrane-bounded organelles include the endosymbiotic organelles (

and

s) and components formed by the

such as the

lysosome A lysosome () is a membrane-bound organelle that is found in all mammalian cells, with the exception of red blood cells (erythrocytes). There are normally hundreds of lysosomes in the cytosol, where they function as the cell’s degradation cent ...

. An endomembrane system and mitochondria are found in almost all

eukaryotes The eukaryotes ( ) constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, seaweeds, and many unicellular organisms are eukaryotes. They constitute a major group of ...

. Plants, algae, and some protists additionally have chloroplasts. A very small minority of bacteria also have a sort-of endomembrane system. The non-membrane bounded organelles, also called large biomolecular complexes, are large assemblies of

macromolecule A macromolecule is a "molecule of high relative molecular mass, the structure of which essentially comprises the multiple repetition of units derived, actually or conceptually, from molecules of low relative molecular mass." Polymers are physi ...

s that carry out particular and specialized functions, but they lack membrane boundaries. Many of these are referred to as "proteinaceous organelles" as their main structure is made of proteins. Such cell structures include: * large RNA and protein complexes:

ribosome Ribosomes () are molecular machine, macromolecular machines, found within all cell (biology), cells, that perform Translation (biology), biological protein synthesis (messenger RNA translation). Ribosomes link amino acids together in the order s ...

spliceosome A spliceosome is a large ribonucleoprotein (RNP) complex found primarily within the nucleus of eukaryotic cells. The spliceosome is assembled from small nuclear RNAs ( snRNA) and numerous proteins. Small nuclear RNA (snRNA) molecules bind to sp ...

, vault * large

protein complex A protein complex or multiprotein complex is a group of two or more associated polypeptide chains. Protein complexes are distinct from multidomain enzymes, in which multiple active site, catalytic domains are found in a single polypeptide chain. ...

es:

proteasome Proteasomes are essential protein complexes responsible for the degradation of proteins by proteolysis, a chemical reaction that breaks peptide bonds. Enzymes that help such reactions are called proteases. Proteasomes are found inside all e ...

DNA polymerase III holoenzyme DNA polymerase III holoenzyme is the primary enzyme complex involved in prokaryotic DNA replication. It was discovered by Thomas Kornberg (son of Arthur Kornberg) and Malcolm Gefter in 1970. The complex has high processivity (i.e. the number ...

RNA polymerase II holoenzyme RNA polymerase II holoenzyme is a form of eukaryotic RNA polymerase II that is recruited to the promoters of protein-coding genes in living cells. It consists of RNA polymerase II, a subset of general transcription factors, and regulatory proteins ...

, symmetric viral

capsid A capsid is the protein shell of a virus, enclosing its genetic material. It consists of several oligomeric (repeating) structural subunits made of protein called protomers. The observable 3-dimensional morphological subunits, which may or m ...

s, complex of

GroEL GroEL is a protein which belongs to the chaperonin family of Chaperone (protein), molecular chaperones, and is found in many bacteria. It is required for the proper protein folding, folding of many proteins. To function properly, GroEL requires ...

and

GroES Heat shock 10 kDa protein 1 (Hsp10), also known as chaperonin 10 (cpn10) or early-pregnancy factor (EPF), is a protein that in humans is encoded by the ''HSPE1'' gene. The homolog in ''Escherichia coli, E. coli'' is GroES that is a chaperonin ...

; membrane protein complexes: porosome,

photosystem I Photosystem I (PSI, or plastocyanin–ferredoxin oxidoreductase) is one of two photosystems in the Light-dependent reactions, photosynthetic light reactions of algae, plants, and cyanobacteria. Photosystem I is an integral membrane ...

ATP synthase ATP synthase is an enzyme that catalyzes the formation of the energy storage molecule adenosine triphosphate (ATP) using adenosine diphosphate (ADP) and inorganic phosphate (Pi). ATP synthase is a molecular machine. The overall reaction catalyzed ...

* large DNA and protein complexes:

nucleosome A nucleosome is the basic structural unit of DNA packaging in eukaryotes. The structure of a nucleosome consists of a segment of DNA wound around eight histone, histone proteins and resembles thread wrapped around a bobbin, spool. The nucleosome ...

centriole In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers ( Pinophyta), flowering plants ( angiosperms) and most fungi, an ...

and microtubule-organizing center (MTOC) *

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 ...

nucleolus The nucleolus (; : nucleoli ) is the largest structure in the cell nucleus, nucleus of eukaryote, eukaryotic cell (biology), cells. It is best known as the site of ribosome biogenesis. The nucleolus also participates in the formation of signa ...

* stress granule *

germ cell A germ cell is any cell that gives rise to the gametes of an organism that reproduces sexually. In many animals, the germ cells originate in the primitive streak and migrate via the gut of an embryo to the developing gonads. There, they unde ...

granule * neuronal transport granule The mechanisms by which such non-membrane bounded organelles form and retain their spatial integrity have been likened to liquid-liquid

phase separation Phase separation is the creation of two distinct Phase (matter), phases from a single homogeneous mixture. The most common type of phase separation is between two immiscible liquids, such as oil and water. This type of phase separation is kn ...

Eukaryotic organelles

Eukaryotic The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

cells are structurally complex, and by definition are organized, in part, by interior compartments that are themselves enclosed by lipid membranes that resemble the outermost

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

. The larger organelles, such as the

nucleus Nucleus (: nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucleu ...

and vacuoles, are easily visible with the

light microscope The optical microscope, also referred to as a light microscope, is a type of microscope that commonly uses visible spectrum, visible light and a system of lens (optics), lenses to generate magnified images of small objects. Optical microscopes ...

. They were among the first biological discoveries made after the invention of the

microscope A microscope () is a laboratory equipment, laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic ...

. Not all eukaryotic cells have each of the organelles listed below. Exceptional organisms have cells that do not include some organelles (such as mitochondria) that might otherwise be considered universal to eukaryotes. The several

s including

chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...

s are distributed among some but not all eukaryotes. There are also occasional exceptions to the number of membranes surrounding organelles, listed in the tables below (e.g., some that are listed as double-membrane are sometimes found with single or triple membranes). In addition, the number of individual organelles of each type found in a given cell varies depending upon the function of that cell. The cell membrane and cell wall are not organelles. Other related structures: *

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 ...

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 nanometer, nm and have an inner diameter bet ...

Prokaryotic organelles

Prokaryote A prokaryote (; less commonly spelled procaryote) is a unicellular organism, single-celled organism whose cell (biology), cell lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Gree ...

s are not as structurally complex as eukaryotes, and were once thought to have little internal organization, and lack

cellular compartment Cellular compartments in cell biology comprise all of the closed parts within the cytosol of a eukaryotic cell, usually surrounded by a single or double lipid layer membrane. These compartments are often, but not always, defined as membrane ...

s and internal membranes; but slowly, details are emerging about prokaryotic internal structures that overturn these assumptions. An early false turn was the idea developed in the 1970s that bacteria might contain

folds termed mesosomes, but these were later shown to be artifacts produced by the chemicals used to prepare the cells for

electron microscopy An electron microscope is a microscope that uses a beam of electrons as a source of illumination. It uses electron optics that are analogous to the glass lenses of an optical light microscope to control the electron beam, for instance focusing i ...

. However, there is increasing evidence of compartmentalization in at least some prokaryotes. Research has revealed that at least some prokaryotes have microcompartments, such as

carboxysome Carboxysomes are bacterial microcompartments (BMCs) consisting of polyhedral protein shells filled with the enzymes ribulose-1,5-bisphosphate carboxylase/oxygenase ( RuBisCO)—the predominant enzyme in carbon fixation and the rate limiting ...

s. These subcellular compartments are 100–200 nm in diameter and are enclosed by a shell of proteins. Even more striking is the description of membrane-bounded magnetosomes in bacteria, reported in 2006. The bacterial phylum Planctomycetota has revealed a number of compartmentalization features. The Planctomycetota cell plan includes intracytoplasmic membranes that separates the cytoplasm into paryphoplasm (an outer ribosome-free space) and pirellulosome (or riboplasm, an inner ribosome-containing space). Membrane-bounded anammoxosomes have been discovered in five Planctomycetota "anammox" genera, which perform anaerobic ammonium oxidation. In the Planctomycetota species '' Gemmata obscuriglobus'', a nucleus-like structure surrounded by lipid membranes has been reported. Compartmentalization is a feature of prokaryotic

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

structures.

Purple bacteria Purple bacteria or purple photosynthetic bacteria are Gram-negative proteobacteria that are phototrophic, capable of producing their own food via photosynthesis. They are pigmented with bacteriochlorophyll ''a'' or ''b'', together with various ...

have "chromatophores", which are reaction centers found in

invagination Invagination is the process of a surface folding in on itself to form a cavity, pouch or tube. In developmental biology, invagination of Epithelium, epithelial sheets occurs in many contexts during Animal embryonic development, embryonic developme ...

s of the cell membrane.

Green sulfur bacteria The green sulfur bacteria are a phylum, Chlorobiota, of obligately anaerobic photoautotrophic bacteria that metabolize sulfur. Green sulfur bacteria are nonmotile (except ''Chloroherpeton thalassium'', which may glide) and capable of anoxyg ...

have

chlorosome A chlorosome is a photosynthetic antenna complex found in green sulfur bacteria (GSB) and many green non-sulfur bacteria (GNsB), together known as green bacteria. They differ from other antenna complexes by their large size and lack of protei ...

s, which are photosynthetic antenna complexes found bonded to cell membranes.

Cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...

have internal

thylakoid Thylakoids are membrane-bound compartments inside chloroplasts and cyanobacterium, cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a #Membrane, thylakoid membrane surrounding a #Lumen, ...

membranes for light-dependent photosynthesis; studies have revealed that the cell membrane and the thylakoid membranes are not continuous with each other.

References

External links