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Phloem (, FLOH-əm) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as photosynthates, in particular the sugar sucrose, to the rest of the plant. This transport process is called translocation. In trees, the phloem is the innermost layer of the bark, hence the name, derived from the Ancient Greek word φλοιός (phloiós), meaning "bark". The term was introduced by Carl Nägeli in 1858.Structure
Phloem tissue consists of conducting cells, generally called sieve elements, parenchyma cells, including both specialized companion cells or albuminous cells and unspecialized cells and supportive cells, such as fibres and sclereids.Conducting cells (sieve elements)
Sieve elements are the type of cell that are responsible for transporting sugars throughout the plant. At maturity they lack a nucleus and have very few organelles, so they rely on companion cells or albuminous cells for most of their metabolic needs. Sieve tube cells do contain vacuoles and other organelles, such as ribosomes, before they mature, but these generally migrate to the cell wall and dissolve at maturity; this ensures there is little to impede the movement of fluids. One of the few organelles they do contain at maturity is the rough endoplasmic reticulum, which can be found at the plasma membrane, often nearby the plasmodesmata that connect them to their companion or albuminous cells. All sieve cells have groups of pores at their ends that grow from modified and enlarged plasmodesmata, called sieve areas. The pores are reinforced by platelets of a polysaccharide called callose.Supportive cells
Although its primary function is transport of sugars, phloem may also contain cells that have a mechanical support function. These are sclerenchyma cells which generally fall into two categories: fibres and sclereids. Both cell types have a secondary cell wall and are dead at maturity. The secondary cell wall increases their rigidity and tensile strength, especially because they contain lignin.Function
Unlike xylem (which is composed primarily of dead cells), the phloem is composed of still-living cells that transport sap. The sap is a water-based solution, but rich in sugars made by photosynthesis. These sugars are transported to non-photosynthetic parts of the plant, such as the roots, or into storage structures, such as tubers or bulbs.
During the plant's growth period, usually during the spring, storage organs such as the roots are sugar sources, and the plant's many growing areas are sugar sinks. The movement in phloem is multidirectional, whereas, in xylem cells, it is unidirectional (upward).After the growth period, when the meristems are dormant, the leaves are sources, and storage organs are sinks. Developing seed-bearing organs (such as fruit) are always sinks. Because of this multi-directional flow, coupled with the fact that sap cannot move with ease between adjacent sieve-tubes, it is not unusual for sap in adjacent sieve-tubes to be flowing in opposite directions.While movement of water and minerals through the xylem is driven by negative pressures (tension) most of the time, movement through the phloem is driven by positive hydrostatic pressures. This process is termed translocation, and is accomplished by a process called phloem loading and unloading.
Phloem sap is also thought to play a role in sending informational signals throughout vascular plants. "Loading and unloading patterns are largely determined by the conductivity and number of plasmodesmata and the position-dependent function of solute-specific, plasma membrane transport proteins. Recent evidence indicates that mobile proteins and RNA are part of the plant's long-distance communication signaling system. Evidence also exists for the directed transport and sorting of macromolecules as they pass through plasmodesmata."Organic molecules such as sugars, amino acids, certain phytohormones, and even messenger RNAs are transported in the phloem through sieve tube elements.Phloem is also used as a popular site for oviposition and breeding of insects belonging to the order Diptera, including the fruit fly Drosophila montana.
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