A contractile vacuole (CV) is a sub-cellular structure (

organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as Organ (anatomy), organs are to th ...

) involved in osmoregulation. It is found predominantly in

protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...

, including unicellular

algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...

. It was previously known as pulsatile or pulsating vacuole.

Overview

The contractile vacuole is a specialized type of

vacuole A vacuole () is a membrane-bound organelle which is present in Plant cell, plant and Fungus, fungal Cell (biology), cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water ...

that regulates the quantity of

water Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...

inside a cell. In

freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...

environments, the concentration of solutes is hypotonic, lower outside than inside the cell. Under these conditions, osmosis causes water to accumulate in the cell from the external environment. The contractile vacuole acts as part of a protective mechanism that prevents the cell from absorbing too much water and possibly lysing (rupturing) through excessive internal pressure. The contractile vacuole, as its name suggests, expels water out of the cell by contracting. The growth (water gathering) and contraction (water expulsion) of the contractile vacuole are periodical. One cycle takes several seconds, depending on the species and the osmolarity of the environment. The stage in which water flows into the CV is called diastole. The contraction of the contractile vacuole and the expulsion of water out of the cell is called systole. Water always flows first from outside the cell into the

cytoplasm The cytoplasm describes all the material within a eukaryotic or prokaryotic cell, enclosed by the cell membrane, including the organelles and excluding the nucleus in eukaryotic cells. The material inside the nucleus of a eukaryotic cell a ...

, and is only then moved from the cytoplasm into the contractile vacuole for expulsion. Species that possess a contractile vacuole typically always use the organelle, even at very hypertonic (high concentration of solutes) environments, since the cell tends to adjust its cytoplasm to become even more hyperosmotic than the environment. The amount of water expelled from the cell and the rate of contraction are related to the osmolarity of the environment. In hyperosmotic environments, less water will be expelled and the contraction cycle will be longer. The best-understood contractile vacuoles belong to the protists '' Paramecium'', '' Amoeba'', '' Dictyostelium'' and '' Trypanosoma'', and to a lesser extent the green alga '' Chlamydomonas''. Not all species that possess a contractile vacuole are freshwater

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 ...

s; some marine,

soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...

microorganism A microorganism, or microbe, is an organism of microscopic scale, microscopic size, which may exist in its unicellular organism, single-celled form or as a Colony (biology)#Microbial colonies, colony of cells. The possible existence of unseen ...

s and

parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...

s also have a contractile vacuole. The contractile vacuole is predominant in species that do not have a

cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...

, but there are exceptions (notably ''Chlamydomonas'') which do possess a cell wall. Through

evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

, the contractile vacuole has typically been lost in

multicellular A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...

organisms, but it still exists in the unicellular stage of several multicellular

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

, as well as in several types of cells in sponges ( amoebocytes, pinacocytes, and choanocytes). The number of contractile vacuoles per cell varies, depending on the

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

. ''Amoeba'' have one, '' Dictyostelium discoideum'', '' Paramecium aurelia'' and '' Chlamydomonas reinhardtii'' have two, and giant amoeba, such as '' Chaos carolinensis'', have many. The number of contractile vacuoles in each species is mostly constant and is therefore used for species characterization in

systematics Systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: phylogenetic trees, phylogenies). Phy ...

. The contractile vacuole has several structures attached to it in most cells, such as membrane folds, tubules, water tracts and small vesicles. These structures have been termed the spongiome; the contractile vacuole together with the spongiome is sometimes called the "contractile vacuole complex" (CVC). The spongiome serves several functions in water transport into the contractile vacuole and in localization and docking of the contractile vacuole within the cell. ''Paramecium'' and ''Amoeba'' possess large contractile vacuoles (average diameter of 13 and 45 μm, respectively), which are relatively comfortable to isolate, manipulate and assay. The smallest known contractile vacuoles belong to ''Chlamydomonas'', with a diameter of 1.5 μm. In ''Paramecium'', which has one of the most complex contractile vacuoles, the vacuole is surrounded by several canals, which absorb water by osmosis from the cytoplasm. After the canals fill with water, the water is pumped into the vacuole. When the vacuole is full, it expels the water through a pore in the cytoplasm which can be opened and closed. Other protists, such as '' Amoeba'', have CVs that move to the surface of the cell when full and undergo exocytosis. In ''Amoeba'' contractile vacuoles collect excretory waste, such as

ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...

, from the intracellular fluid by both

diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...

and

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 cellula ...

Water flow into the CV

The way in which water enters the CV had been a mystery for many years, but several discoveries since the 1990s have improved understanding of this issue. Water could theoretically cross the CV membrane by osmosis, but only if the inside of the CV is hyperosmotic (higher solute concentration) to the cytoplasm. The discovery of proton pumps in the CV membrane and the direct measurement of ion concentrations inside the CV using microelectrodes led to the following model: the pumping of protons either into or out of the CV causes different ions to enter the CV. For example, some proton pumps work as cation exchangers, whereby a proton is pumped out of the CV and a cation is pumped at the same time into the CV. In other cases, protons pumped into the CV drag anions with them (

carbonate A carbonate is a salt of carbonic acid, (), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word "carbonate" may also refer to a carbonate ester, an organic compound containing the carbonate group ...

, for example), to balance the pH. This ion flux into the CV causes an increase in CV osmolarity and as a result water enters the CV by osmosis. Water has been shown in at least some species to enter the CV through aquaporins. Acidocalcisomes have been implied to work alongside the contractile vacuole in responding to osmotic stress. They were detected in the vicinity of the vacuole in '' Trypanosoma cruzi'' and were shown to fuse with the vacuole when the cells were exposed to osmotic stress. Presumably the acidocalcisomes empty their ion contents into the contractile vacuole, thereby increasing the vacuole's osmolarity.

Unresolved issues

The CV does not exist in higher organisms, but some of its unique characteristics are used by them in their osmoregulatory mechanisms. Research on the CV can therefore help us understand how osmoregulation works in all species. Many issues regarding the CV remain, as of 2010, unsolved: * Contraction. It is not completely known what causes the CV membrane to contract, and whether it is an active process which costs

energy Energy () is the physical quantity, quantitative physical property, property that is transferred to a physical body, body or to a physical system, recognizable in the performance of Work (thermodynamics), work and in the form of heat and l ...

or a passive collapse of the CV membrane. Evidence for involvement of

actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ...

and myosin, prominent contractile

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 ...

s which are found in many cells, are ambiguous. * Membrane composition. Although it is known that several proteins decorate the CV membrane (V−H+−ATPases, aquaporins), a complete list is missing. The composition of the membrane itself and its similarities to and differences from other cellular membranes are also not clear. * Contents of the CV. Several studies have shown the ion concentrations inside some of the largest CVs but not in the smallest ones (such as in the important model organism ''Chlamydomonas rheinhardii''). The reasons and mechanisms for ion exchange between the CV and cytoplasm are not entirely clear.

References

{{DEFAULTSORT:Contractile Vacuole Organelles