Organelles

In cell biology, an **organelle** is a specialized subunit within a cell that has a specific function. Organelles are either separately enclosed within their own lipid bilayers (also called membrane-bound organelles) or are spatially distinct functional units without a surrounding lipid bilayer (non-membrane bound organelles).

The name *organelle* comes from the idea that these structures are parts of cells, as organs are to the body, hence *organelle,* the suffix *-elle* being a diminutive. Organelles are identified by microscopy, and can also be purified by cell fractionation. There are many types of organelles, particularly in eukaryotic cells. While prokaryotes do not possess organelles *per se*, some do contain protein-based bacterial microcompartments, which are thought to act as primitive organelles.

## History and terminology

1. Nucleolus
2. Nucleus
3. Ribosome (little dots)
4. Vesicle
5. Rough endoplasmic reticulum
6. Golgi apparatus (or, Golgi body)
7. Cytoskeleton
8. Smooth endoplasmic reticulum
9. Mitochondrion
10. Vacuole
11. Cytosol (fluid that contains organelles, comprising the cytoplasm)
12. Lysosome
13. Centrosome
14. Cell membrane

In biology *organs* are defined as confined functional units within an organism. The analogy 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 theory which said that microorganisms have the same organs of multicellular animals, only minor.

Credited as the first to use a diminutive 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 *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

While most cell biologists consider the term *organelle* to be synonymous with cell compartment, a space often bound by one or two lipid bilayers, some cell biologists choose to limit the term to include only those cell compartments that contain deoxyribonucleic acid (DNA), having originated from formerly autonomous microscopic organisms acquired via endosymbiosis.

Under this definition, there would only be two broad classes of organelles (i.e. those that contain their own DNA, and have originated from endosymbiotic bacteria):

* mitochondria (in almost all eukaryotes)
* plastids (e.g. in plants, algae, and some protists).

Other organelles are also suggested to have endosymbiotic origins, but do not contain their own DNA (notably the flagellum – see evolution of flagella).

A second, less restrictive definition of organelles is that they are membrane-bound structures. However, even by using this definition, some parts of the cell that have been shown to be distinct functional units do not qualify as organelles. Therefore, the use of organelle to also refer to non-membrane bound structures such as ribosomes is common and accepted. This has led many texts to delineate between **membrane-bound** and **non-membrane** bound organelles. The non-membrane bound organelles, also called large biomolecular complexes, are large assemblies of macromolecules that carry out particular and specialized functions, but they lack membrane boundaries. Many of these are referred to as "proteinaceous organelles" as there many structure is made of proteins. Such cell structures include:

* large RNA and protein complexes: ribosome, spliceosome, vault
* large protein complexes: proteasome, DNA polymerase III holoenzyme, RNA polymerase II holoenzyme, symmetric viral capsids, complex of GroEL and GroES; membrane protein complexes: photosystem I, ATP synthase
* large DNA and protein complexes: nucleosome
* centriole and microtubule-organizing center (MTOC)
* cytoskeleton
* flagellum
* nucleolus
* stress granule
* germ cell granule
* neuronal transport granule

The mechanisms by which such non-membrane bound organelles form and retain their spatial integrity have been likened to liquid-liquid phase separation.

### Eukaryotic organelles

Eukaryotic 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 larger organelles, such as the nucleus and vacuoles, are easily visible with the light microscope. They were among the first biological discoveries made after the invention of the microscope.

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

Mitochondria and plastids, including chloroplasts, have double membranes and their own DNA. According to the endosymbiotic theory, they are believed to have originated from incompletely consumed or invading prokaryotic organisms.

Other related structures:

* cytosol
* endomembrane system
* nucleosome
* microtubule
* cell membrane

### Prokaryotic organelles

Prokaryotes are not as structurally complex as eukaryotes, and were once thought not to have any internal structures enclosed by lipid membranes. In the past, they were often viewed as having little internal organization, but slowly, details are emerging about prokaryotic internal structures. An early false turn was the idea developed in the 1970s that bacteria might contain membrane folds termed mesosomes, but these were later shown to be artifacts produced by the chemicals used to prepare the cells for electron microscopy.

However, more recent research has revealed that at least some prokaryotes have microcompartments such as carboxysomes. 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-bound magnetosomes in bacteria, reported in 2006, as well as the nucleus-like structures of the *Planctomycetes* that are surrounded by lipid membranes, reported in 2005.