A vacuole () is a

membrane-bound organelle In cell biology, 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 organs are to the body, hence ''organelle,'' the ...

which is present 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 exclu ...

and

fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...

cells and some

protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

, and

bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...

l cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in

solution Solution may refer to: * Solution (chemistry), a mixture where one substance is dissolved in another * Solution (equation), in mathematics ** Numerical solution, in numerical analysis, approximate solutions within specified error bounds * Soluti ...

, though in certain cases they may contain solids which have been engulfed. Vacuoles are formed by the fusion of multiple membrane

vesicles Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...

and are effectively just larger forms of these. The organelle has no basic shape or size; its structure varies according to the requirements of the cell.

Discovery

Contractile vacuole A contractile vacuole (CV) is a sub-cellular structure (organelle) involved in osmoregulation. It is found predominantly in protists and in unicellular algae. It was previously known as pulsatile or pulsating vacuole. Overview The contractile v ...

s ("stars") were first observed by

Spallanzani Lazzaro Spallanzani (; 12 January 1729 – 11 February 1799) was an Italian Catholic priest (for which he was nicknamed Abbé Spallanzani), biologist and physiologist who made important contributions to the experimental study of bodily function ...

(1776) in protozoa, although mistaken for respiratory organs.

Dujardin Dujardin is a French surname, meaning "from the garden", and may refer to: * Charlotte Dujardin, British dressage rider * Édouard Dujardin, French writer * Félix Dujardin (1801–1860), French biologist * Jean Dujardin, French actor and comedia ...

(1841) named these "stars" as ''vacuoles''. In 1842,

Schleiden Schleiden is a town in North Rhine-Westphalia, Germany. It lies in the Eifel hills, in the district of Euskirchen, and has 12,998 inhabitants as of 30 June 2017. Schleiden is connected by a tourist railway to Kall, on the Eifel Railway between ...

applied the term for plant cells, to distinguish the structure with cell sap from the rest of the protoplasm. In 1885,

de Vries De Vries is one of the most common Dutch surnames. It indicates a geographical origin: "Vriesland" is an old spelling of the Dutch province of Friesland (Frisia). Hence, "de Vries" means "the Frisian". The name has been modified to "DeVries", "d ...

named the vacuole membrane as tonoplast.

Function

The function and significance of vacuoles varies greatly according to the type of cell in which they are present, having much greater prominence in the cells of plants, fungi and certain protists than those of animals and bacteria. In general, the functions of the vacuole include: * Isolating materials that might be harmful or a threat to the cell * Containing waste products * Containing water in plant cells * Maintaining internal

hydrostatic pressure Fluid statics or hydrostatics is the branch of fluid mechanics that studies the condition of the equilibrium of a floating body and submerged body " fluids at hydrostatic equilibrium and the pressure in a fluid, or exerted by a fluid, on an imm ...

turgor Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall. It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilibri ...

within the cell * Maintaining an acidic internal pH * Containing small molecules * Exporting unwanted substances from the cell * Allows plants to support structures such as leaves and flowers due to the pressure of the central vacuole * By increasing in size, allows the germinating plant or its organs (such as leaves) to grow very quickly and using up mostly just water. * In seeds, stored proteins needed for germination are kept in 'protein bodies', which are modified vacuoles. Vacuoles also play a major role in autophagy, maintaining a balance between

biogenesis Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise ...

(production) and degradation (or turnover), of many substances and cell structures in certain organisms. They also aid in the lysis and recycling of misfolded proteins that have begun to build up within the cell. Thomas Boller and others proposed that the vacuole participates in the destruction of invading

and Robert B. Mellor proposed organ-specific forms have a role in 'housing' symbiotic bacteria. In protists, vacuoles have the additional function of storing food which has been absorbed by the organism and assisting in the digestive and waste management process for the cell. In animal cells, vacuoles perform mostly subordinate roles, assisting in larger processes of exocytosis and endocytosis. Animal vacuoles are smaller than their plant counterparts but also usually greater in number. There are also animal cells that do not have any vacuoles. Exocytosis is the extrusion process of proteins and lipids from the cell. These materials are absorbed into secretory granules within 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 ...

before being transported to the cell membrane and secreted into the extracellular environment. In this capacity, vacuoles are simply storage vesicles which allow for the containment, transport and disposal of selected proteins and lipids to the extracellular environment of the cell. Endocytosis is the reverse of exocytosis and can occur in a variety of forms.

Phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...

("cell eating") is the process by which bacteria, dead tissue, or other bits of material visible under the microscope are engulfed by cells. The material makes contact with the cell membrane, which then invaginates. The

invagination Invagination is the process of a surface folding in on itself to form a cavity, pouch or tube. In developmental biology, invagination is a mechanism that takes place during gastrulation. This mechanism or cell movement happens mostly in the vegeta ...

is pinched off, leaving the engulfed material in the membrane-enclosed vacuole and the cell membrane intact.

Pinocytosis In cellular biology, pinocytosis, otherwise known as fluid endocytosis and bulk-phase pinocytosis, is a mode of endocytosis in which small molecules dissolved in extracellular fluid are brought into the cell through an invagination of the cell ...

("cell drinking") is essentially the same process, the difference being that the substances ingested are in solution and not visible under the microscope. Phagocytosis and pinocytosis are both undertaken in association with

lysosomes 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 prote ...

which complete the breakdown of the material which has been engulfed. '' Salmonella'' is able to survive and reproduce in the vacuoles of several mammal species after being engulfed. The vacuole probably evolved several times independently, even within the

Viridiplantae Viridiplantae (literally "green plants") are a clade of eukaryotic organisms that comprise approximately 450,000–500,000 species and play important roles in both terrestrial and aquatic ecosystems. They are made up of the green algae, which ar ...

Vacuole types

Gas vacuoles

Gas vesicle Gas vesicles, also known as gas vacuoles, are nanocompartments in certain prokaryotic organisms, which help in buoyancy. Gas vesicles are composed entirely of protein; no lipids or carbohydrates have been detected. Function Gas vesicles occur ...

s, also known as gas vacuoles, are nanocompartments which are freely permeable to gas, and occur mainly in Cyanobacteria, but are also found in other bacteria species and some archaea. Gas vesicles allow the bacteria to control their buoyancy. They are formed when small biconical structures grow to form spindles. The vesicle walls are composed of a hydrophobic gas vesicle protein A (GvpA) which form a cylindrical hollow, proteinaceous structure that fills with gas. Small variances in the amino acid sequence produce changes in morphology of the gas vesicle, for example, GvpC, is a larger protein.

Central vacuoles

Most mature plant cells have one large vacuole that typically occupies more than 30% of the cell's volume, and that can occupy as much as 80% of the volume for certain cell types and conditions. Strands of

cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...

often run through the vacuole. A vacuole is surrounded by a membrane called the tonoplast (word origin: Gk tón(os) + -o-, meaning “stretching”, “tension”, “tone” + comb. form repr. Gk plastós formed, molded) and filled with cell sap. Also called the vacuolar membrane, the tonoplast is the cytoplasmic membrane surrounding a vacuole, separating the vacuolar contents from the cell's cytoplasm. As a membrane, it is mainly involved in regulating the movements of ions around the cell, and isolating materials that might be harmful or a threat to the cell. Transport of protons from the cytosol to the vacuole stabilizes cytoplasmic pH, while making the vacuolar interior more acidic creating a

proton motive force Chemiosmosis is the movement of ions across a semipermeable membrane bound structure, down their electrochemical gradient. An important example is the formation of adenosine triphosphate (ATP) by the movement of hydrogen ions (H+) across a membra ...

which the cell can use to transport nutrients into or out of the vacuole. The low pH of the vacuole also allows

degradative enzyme A degradative enzyme is an enzyme (in a broader sense a protein) which degrades biological molecules. Some examples of degradative enzymes: *Lipase, which digests lipids, *Carbohydrases, which digest carbohydrates (e.g., sugars), *Proteases, which d ...

s to act. Although single large vacuoles are most common, the size and number of vacuoles may vary in different tissues and stages of development. For example, developing cells in the

meristems The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...

contain small provacuoles and cells of the

vascular cambium The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xy ...

have many small vacuoles in the winter and one large one in the summer. Aside from storage, the main role of the central vacuole is to maintain

pressure against the cell wall. Proteins found in the tonoplast (

aquaporins Aquaporins, also called water channels, are channel proteins from a larger family of major intrinsic proteins that form pores in the membrane of biological cells, mainly facilitating transport of water between cells. The cell membranes of a va ...

) control the flow of water into and out of the vacuole through

active transport In cellular biology, ''active transport'' is the movement of molecules or ions across a cell membrane from a region of lower concentration to a region of higher concentration—against the concentration gradient. Active transport requires cellul ...

, pumping

potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosph ...

(K⁺)

ion An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conve ...

s into and out of the vacuolar interior. Due to

osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region o ...

, water will diffuse into the vacuole, placing pressure on the cell wall. If water loss leads to a significant decline in turgor pressure, the

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 ...

will plasmolyze. Turgor pressure exerted by vacuoles is also required for cellular elongation: as the cell wall is partially degraded by the action of

expansin Expansins are a family of closely related nonenzymatic proteins found in the plant cell wall, with important roles in plant cell growth, fruit softening, abscission, emergence of root hairs, pollen tube invasion of the stigma and style, meristem fun ...

s, the less rigid wall is expanded by the pressure coming from within the vacuole. Turgor pressure exerted by the vacuole is also essential in supporting plants in an upright position. Another function of a central vacuole is that it pushes all contents of the cell's cytoplasm against the cellular membrane, and thus keeps the

chloroplasts 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 ...

closer to light. Most plants store chemicals in the vacuole that react with chemicals in the cytosol. If the cell is broken, for example by a herbivore, then the two chemicals can react forming toxic chemicals. In garlic,

alliin Alliin is a sulfoxide that is a natural constituent of fresh garlic. It is a derivative of the amino acid cysteine. When fresh garlic is chopped or crushed, the enzyme alliinase converts alliin into allicin, which is responsible for the aroma ...

and the enzyme

alliinase In enzymology, an alliin lyase () is an enzyme that catalyzes the chemical reaction :an ''S''-alkyl-L-cysteine ''S''-oxide \rightleftharpoons an alkyl sulfenate + 2-aminoacrylate Hence, this enzyme has one substrate, ''S''-alkyl-L-cysteine ...

are normally separated but form

allicin Allicin is an organosulfur compound obtained from garlic, a species in the family Alliaceae. It was first isolated and studied in the laboratory by Chester J. Cavallito and John Hays Bailey in 1944. When fresh garlic is chopped or crushed, th ...

if the vacuole is broken. A similar reaction is responsible for the production of

syn-propanethial-S-oxide ''syn''-Propanethial ''S''-oxide (or (''Z'')-propanethial ''S''-oxide), a member of a class of organosulfur compounds known as thiocarbonyl ''S''-oxides (formerly "sulfines"), is a volatile liquid that acts as a lachrymatory agent (triggers teari ...

when

onion An onion (''Allium cepa'' L., from Latin ''cepa'' meaning "onion"), also known as the bulb onion or common onion, is a vegetable that is the most widely cultivated species of the genus ''Allium''. The shallot is a botanical variety of the onio ...

s are cut. Vacuoles in fungal cells perform similar functions to those in plants and there can be more than one vacuole per cell. In

yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...

cells the vacuole (

Vac7 Vacuolar segregation protein 7 is a protein that in yeast is encoded by the ''VAC7'' gene. VAC7 is a component of the PI(3,5)P2 regulatory complex, composed of ATG18, FIG4, FAB1, VAC14 and VAC7. Function The PI(3,5)P2 regulatory complex reg ...

) is a

dynamic Dynamics (from Greek δυναμικός ''dynamikos'' "powerful", from δύναμις ''dynamis'' "power") or dynamic may refer to: Physics and engineering * Dynamics (mechanics) ** Aerodynamics, the study of the motion of air ** Analytical dyna ...

structure that can rapidly modify its

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

. They are involved in many processes including the

homeostasis In biology, homeostasis (British also homoeostasis) (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physical, and chemical conditions maintained by living systems. This is the condition of optimal functioning for the organism and ...

of cell pH and the concentration of ions,

osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...

, storing amino acids and

polyphosphate Polyphosphates are salts or esters of polymeric oxyanions formed from tetrahedral PO4 (phosphate) structural units linked together by sharing oxygen atoms. Polyphosphates can adopt linear or a cyclic ring structures. In biology, the polyphosphate e ...

and degradative processes. Toxic ions, such as strontium (),

cobalt Cobalt is a chemical element with the symbol Co and atomic number 27. As with nickel, cobalt is found in the Earth's crust only in a chemically combined form, save for small deposits found in alloys of natural meteoric iron. The free element, p ...

(II) (), and

lead Lead is a chemical element with the symbol Pb (from the Latin ) and atomic number 82. It is a heavy metal that is denser than most common materials. Lead is soft and malleable, and also has a relatively low melting point. When freshly cu ...

(II) () are transported into the vacuole to isolate them from the rest of the cell.

Contractile vacuoles

Contractile Vacuoles A contractile vacuole (CV) is a sub-cellular structure (organelle) involved in osmoregulation. It is found predominantly in protists and in unicellular algae. It was previously known as pulsatile or pulsating vacuole. Overview The contractile ...

is a specialized osmoregulatory organelle that is present in many free-living protists. The contractile vacuole is part of the contractile vacuole complex which includes radial arms and a spongiome. The contractile vacuole complex works periodically contracts to remove excess water and ions from the cell to balance water flow into the cell. When the contractile vacuole is slowly taking water in, the contractile vacuole enlarges, this is called diastole and when it reaches its threshold, the central vacuole contracts then contracts (systole) periodically to release water.

Food vacuoles

Food vacuole The food vacuole, or digestive vacuole, is an organelle found in simple eukaryotes such as protists. This organelle is essentially a lysosome. During the stage of the symbiont parasites' lifecycle where it resides within a human (or other mammali ...

s (also called digestive vacuole) are organelles found in

Ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...

s, and Plasmodium falciparum, a protozoan parasite that causes

Malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...

Histopathology

In histopathology,

vacuolization Vacuolization is the formation of vacuoles or vacuole-like structures, within or adjacent to cells. Perinuclear vacuolization of epidermal keratinocytes is most likely inconsequential when not observed in combination with other pathologic find ...

is the formation of vacuoles or vacuole-like structures, within or adjacent to cells. It is an unspecific sign of disease.

References

External links

{{Authority control Vesicles Plant anatomy Cell anatomy