Vacuole
Encyclopedia
Biological membrane
A biological membrane or biomembrane is an enclosing or separatingmembrane that acts as a selective barrier, within or around a cell. It consists of a lipid bilayer with embedded proteins that may constitute close to 50% of membrane content...
-bound organelle
Organelle
In cell biology, an organelle is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid bilayer....
which is present in all plant
Plant cell
Plant cells are eukaryotic cells that differ in several key respects from the cells of other eukaryotic organisms. Their distinctive features include:...
and fungal cells
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....
and some protist
Protist
Protists are a diverse group of eukaryotic microorganisms. Historically, protists were treated as the kingdom Protista, which includes mostly unicellular organisms that do not fit into the other kingdoms, but this group is contested in modern taxonomy...
, animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
and bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
l cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain cases they may contain solids which have been engulfed. Vacuoles are formed by the fusion of multiple membrane vesicles
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...
and are effectively just larger forms of these. The organelle has no basic shape or size; its structure varies according to the needs of the cell.
The function and importance 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 or turgor within the cell
- Maintaining an acidAcidAn acid is a substance which reacts with a base. Commonly, acids can be identified as tasting sour, reacting with metals such as calcium, and bases like sodium carbonate. Aqueous acids have a pH of less than 7, where an acid of lower pH is typically stronger, and turn blue litmus paper red...
ic internal pHPHIn chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline... - 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
- In seeds, stored proteins needed for germination are kept in 'protein bodies', which are modified vacuoles.
Vacuoles also play a major role in autophagy
Autophagy
In cell biology, autophagy, or autophagocytosis, is a catabolic process involving the degradation of a cell's own components through the lysosomal machinery. It is a tightly regulated process that plays a normal part in cell growth, development, and homeostasis, helping to maintain a balance...
, maintaining a balance between biogenesis
Biogenesis
Biogenesis is the law that living things come only from other living things, e.g. a spider lays eggs, which develop into spiders. It may also refer to biochemical processes of production in living organisms.-Spontaneous generation:...
(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 bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
and Robert B Mellor
Robert B Mellor
Robert Brooke Mellor is a British biologist, known for his "unified vacuole theory" which states that in endosymbioses generally, the micro-symbiont and the macro-symbiont share their lytic vacuolar compartments...
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.
Bacteria
Large vacuoles are found in three generaGenera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
of filamentous sulfur bacteria, the Thioploca
Thioploca
Thioploca is a genus of filamentous sulfur bacteria which occurs along of coast off the west of South America. A large vacuole occupies more than 80% of their cell volume and contains sulfide and nitrate which they use to make energy for their metabolism by oxidising sulfate with nitrate...
, Beggiatoa
Beggiatoa
Beggiatoa is a genus of bacteria in the order Thiotrichales. They are named after the Italian medic and botanist F.S. Beggiato. The organisms live in sulfur-rich environments...
and Thiomargarita. The cytosol is extremely reduced in these genera and the vacuole can occupy between 40–98% of the cell. The vacuole contains high concentrations of nitrate
Nitrate
The nitrate ion is a polyatomic ion with the molecular formula NO and a molecular mass of 62.0049 g/mol. It is the conjugate base of nitric acid, consisting of one central nitrogen atom surrounded by three identically-bonded oxygen atoms in a trigonal planar arrangement. The nitrate ion carries a...
ions and is therefore thought to be a storage organelle
Organelle
In cell biology, an organelle is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid bilayer....
.
Gas vacuoles, which are freely permeable to gas, are present in some species of Cyanobacteria. They allow the bacteria to control their buoyancy.
Plants
Plant cell
Plant cells are eukaryotic cells that differ in several key respects from the cells of other eukaryotic organisms. Their distinctive features include:...
s have one large central 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
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...
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). 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 proton
Proton
The proton is a subatomic particle with the symbol or and a positive electric charge of 1 elementary charge. One or more protons are present in the nucleus of each atom, along with neutrons. The number of protons in each atom is its atomic number....
s from the cytosol to the vacuole stabilizes cytoplasmic pH
PH
In chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline...
, while making the vacuolar interior more acidic creating a proton motive force which the cell can use to transport nutrients into or out of the vacuole. The low pH of the vacuole also allows degradative enzymes to act. Although single large central 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 contain small provacuoles and cells of the vascular cambium
Vascular cambium
The vascular cambium is a part of the morphology of plants. It consists of cells that are partly specialized, for the tissues that transport water solutions, but have not reached any of the final forms that occur in their branch of the specialization graph...
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 turgor pressure against the cell wall
Cell wall
The cell wall is the tough, usually flexible but sometimes fairly rigid layer that surrounds some types of cells. It is located outside the cell membrane and provides these cells with structural support and protection, and also acts as a filtering mechanism. A major function of the cell wall is to...
. Proteins found in the tonoplast (aquaporins) control the flow of water into and out of the vacuole through active transport
Active transport
Active transport is the movement of a substance against its concentration gradient . In all cells, this is usually concerned with accumulating high concentrations of molecules that the cell needs, such as ions, glucose, and amino acids. If the process uses chemical energy, such as from adenosine...
, pumping potassium
Potassium
Potassium is the chemical element with the symbol K and atomic number 19. Elemental potassium is a soft silvery-white alkali metal that oxidizes rapidly in air and is very reactive with water, generating sufficient heat to ignite the hydrogen emitted in the reaction.Potassium and sodium are...
(K+) ion
Ion
An ion is an atom or molecule in which the total number of electrons is not equal to the total number of protons, giving it a net positive or negative electrical charge. The name was given by physicist Michael Faraday for the substances that allow a current to pass between electrodes in a...
s into and out of the vacuolar interior. Due to osmosis
Osmosis
Osmosis is the movement of solvent molecules through a selectively permeable membrane into a region of higher solute concentration, aiming to equalize the solute concentrations on the two sides...
, 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 (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....
will plasmolyze
Plasmolysis
Plasmolysis is the process in plant cells where the cytoplasm pulls away from the cell wall due to the loss of water through osmosis. The reverse process, cytolysis, can occur if the cell is in a hypotonic solution resulting in a higher external osmotic pressure and a net flow of water into the cell...
. Turgor pressure exerted by vacuoles is also required for cellular elongation: as the cell wall is partially degraded by the action of expansin
Expansin
Expansin refers to 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 function, and other developmental processes...
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 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
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
, then the two chemicals can react forming toxic chemicals. In garlic, alliin
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 of fresh garlic...
and the enzyme alliinase
Alliinase
In enzymology, an alliin lyase is an enzyme that catalyzes the chemical reactionHence, this enzyme has one substrate, S-alkyl-L-cysteine S-oxide, and two products, alkyl sulfenate and 2-aminoacrylate....
are normally separated but form allicin if the vacuole is broken. A similar reaction is responsible for the production of syn-propanethial-S-oxide
Syn-propanethial-S-oxide
syn-Propanethial S-oxide is a gas that acts as a lachrymatory agent . The chemical is released from onions, Allium cepa, as they are sliced...
when onion
Onion
The onion , also known as the bulb onion, common onion and garden onion, is the most widely cultivated species of the genus Allium. The genus Allium also contains a number of other species variously referred to as onions and cultivated for food, such as the Japanese bunching onion The onion...
s are cut.
Fungi
Vacuoles in fungal cells perform similar functions to those in plants and there can be more than one vacuole per cell. In yeastSaccharomyces cerevisiae
Saccharomyces cerevisiae is a species of yeast. It is perhaps the most useful yeast, having been instrumental to baking and brewing since ancient times. It is believed that it was originally isolated from the skin of grapes...
cells the vacuole is a dynamic structure that can rapidly modify its morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
. They are involved in many processes including the homeostasis
Homeostasis
Homeostasis is the property of a system that regulates its internal environment and tends to maintain a stable, constant condition of properties like temperature or pH...
of cell pH and the concentration of ions, osmoregulation
Osmoregulation
Osmoregulation is the active regulation of the osmotic pressure of an organism's fluids to maintain the homeostasis of the organism's water content; that is it keeps the organism's fluids from becoming too diluted or too concentrated. Osmotic pressure is a measure of the tendency of water to move...
, storing amino acids and polyphosphate
Polyphosphate
Triphosphates are salts or esters of polymeric oxyanions formed from tetrahedral PO4 structural units linked together by sharing oxygen atoms. When two corners are shared the polyphosphate may have a linear chain structure or a cyclic ring structure. In biology the polyphosphate esters AMP, ADP...
and degradative processes. Toxic ions, such as strontium
Strontium
Strontium is a chemical element with the symbol Sr and the atomic number 38. An alkaline earth metal, strontium is a soft silver-white or yellowish metallic element that is highly reactive chemically. The metal turns yellow when exposed to air. It occurs naturally in the minerals celestine and...
, cobalt
Cobalt
Cobalt is a chemical element with symbol Co and atomic number 27. It is found naturally only in chemically combined form. The free element, produced by reductive smelting, is a hard, lustrous, silver-gray metal....
(II) , and lead
Lead
Lead is a main-group element in the carbon group with the symbol Pb and atomic number 82. Lead is a soft, malleable poor metal. It is also counted as one of the heavy metals. Metallic lead has a bluish-white color after being freshly cut, but it soon tarnishes to a dull grayish color when exposed...
(II) are transported into the vacuole to isolate them from the rest of the cell.
Animals
In animal cells, vacuoles perform mostly subordinate roles, assisting in larger processes of exocytosisExocytosis
Exocytosis , also known as 'The peni-cytosis', is the durable process by which a cell directs the contents of secretory vesicles out of the cell membrane...
and endocytosis
Endocytosis
Endocytosis is a process by which cells absorb molecules by engulfing them. It is used by all cells of the body because most substances important to them are large polar molecules that cannot pass through the hydrophobic plasma or cell membrane...
.
Animal vacuoles are smaller that their vegetable 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
Golgi apparatus
The Golgi apparatus is an organelle found in most eukaryotic cells. It was identified in 1898 by the Italian physician Camillo Golgi, after whom the Golgi apparatus is named....
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 ("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
Invagination means to fold inward or to sheath. In biology, this can refer to a number of processes.* Invagination is the morphogenetic processes by which an embryo takes form, and is the initial step of gastrulation, the massive reorganization of the embryo from a simple spherical ball of cells,...
is pinched off, leaving the engulfed material in the membrane-enclosed vacuole and the cell membrane intact. Pinocytosis
Pinocytosis
In cellular biology, pinocytosis is a form of endocytosis in which small particles are brought into the cell—forming an invagination, and then suspended within small vesicles that subsequently fuse with lysosomes to hydrolyze, or to break down, the particles...
("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 which complete the breakdown of the material which has been engulfed.
Salmonella
Salmonella
Salmonella is a genus of rod-shaped, Gram-negative, non-spore-forming, predominantly motile enterobacteria with diameters around 0.7 to 1.5 µm, lengths from 2 to 5 µm, and flagella which grade in all directions . They are chemoorganotrophs, obtaining their energy from oxidation and reduction...
is able to survive and reproduce in the vacuoles of several mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
species after being engulfed.