Vacuole
Vacuoles are
membrane-bounded compartments within some
eukaryotic cells that can serve a variety of secretory, excretory, and storage functions. Vacuoles and their contents are considered to be distinct from the
cytoplasm, and are classified as
ergastic according to some authors (Esau, 1965). Vacuoles are especially conspicuous in most
plant cells.
In general, vacuole functions include:
* capturing food materials
* removing unwanted structural debris surrounding the cell
* sequestering materials that might be toxic to the cell
* containment of waste products
* maintaining internal
hydrostatic pressure or
turgor within the cell
* maintaining an
acidic internal pH
* storing small molecules
* exporting unwanted substances from the cell.
* enabling the cell to elongate rapidly or otherwise alter relative cell size.
Vacuoles also play a major role in
autophagy, maintaining a balance between
biogenesis (production) and degradation (or turnover), of many substances and cell structures. They also aid in destruction of invading
bacteria or of misfolded proteins that have begun to aggregate within the cell. Autophagy is especially prominent in insects that undergo complete
metamorphosis; for example,
larval tissue is recycled to become appendages in an adult insect.
Some
protists and
macrophages use
food vacuoles as a stage in
phagocytosis—the intake of large molecules, particles, or even other cells, by the cell for
digestion.
A
contractile vacuole is used to pump excess water out of the cell to reduce
osmotic pressure and keep the cell from bursting, which is referred to as
cytolysis or
osmotic lysis. Contractile vacuoles are found in some freshwater
protozoa, such as
paramecium.
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The vacuole of Rhoeo discolor is more easily visible... |
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...when it's shrunk during plasmolysis. |
Most mature
plant cells have a single
central vacuole which often takes up more than 80% of the cell interior. It is surrounded by a membrane called the
tonoplast.
This vacuole houses large amounts of a liquid called
cell sap, composed of water,
enzymes, inorganic ions (like K
+ and Cl
-), salts (such as
calcium), and other substances, including toxic byproducts removed from the cytosol to avoid interference with
cell metabolism. Toxins located in the vacoule may also help to protect some plants from predators. Transport of
protons from cytosol to vacuole aids in keeping cytoplasmic
pH stable, while making the vacuolar interior more acidic, allowing degradative enzymes to act. Although having a large central vacuole is the most common case, the size and number of vacuoles may vary in different tissues and stages of development. Cells of the
vascular cambium, for example, have many small vacuoles in winter, and one large one in summer.
Aside from storage, the main role of the central vacuole is to maintain
turgor pressure against the
cell wall. Proteins found in the tonoplast control the flow of water into and out of the vacuole through
active transport, pumping
potassium (K
+)
ions into and out of the vacuolar interior. Due to
osmosis, 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 will
plasmolyse. Turgor pressure exerted by vacuoles is also helpful for cellular elongation: as the cell wall is partially degraded by the action of
auxins, the less rigid wall is expanded by the pressure coming from within the vacuole. Vacuoles can help some plant cells to reach considerable size. 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.
The vacuole also stores the
pigments in
flowers.
In budding
yeast cells, vacuoles are the storage compartments of
amino acids and the detoxification compartment, which are described in previous topics. Under conditions of starvation, proteins are degraded in vacuoles; this is called autophagy. First,
cytoplasms,
mitochondria, and small
organelles are covered with multiplex plasma membranes called autophagosomes. Next, the autophagosomes fuse the vacuoles. Finally, the cytoplasms and the organelles are degraded.
In a vacuole of budding yeast, a black particle sometimes appears. It is called a dancing body. The dancing body moves actively in the vacuole and appears and disappears within 10 minutes to several hours. In previous research, it was suggested but not proven that the main component of the dancing body is
polyphosphate acid. But the main component has been determined to be crystallized sodium polyphosphate and its function has been studied. It is thought that its function is to supply and store phosphates in budding yeast cells.
*Esau, K. 1965.
Plant Anatomy, 2nd Edition. John Wiley & Sons. 767 pp.