In vascular plant
Vascular plant
Vascular plants are those plants that have lignified tissues for conducting water, minerals, and photosynthetic products through the plant. Vascular plants include the clubmosses, Equisetum, ferns, gymnosperms and angiosperms...

s, phloem is the living tissue
Biological tissue
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...

 that carries organic nutrient
A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...

s (known as photosynthate), in particular, glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

, a sugar, to all parts of the plant where needed. In tree
A tree is a perennial woody plant. It is most often defined as a woody plant that has many secondary branches supported clear of the ground on a single main stem or trunk with clear apical dominance. A minimum height specification at maturity is cited by some authors, varying from 3 m to...

s, the phloem is the innermost layer of the bark
Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines and shrubs. Bark refers to all the tissues outside of the vascular cambium and is a nontechnical term. It overlays the wood and consists of the inner bark and the outer bark. The inner...

, hence the name, derived from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 word (phloos) "bark". The phloem is concerned mainly with the transport of soluble organic material made during photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

. This is called translocation.


Phloem tissue consists of less specialized and nucleate parenchyma
Parenchyma is a term used to describe a bulk of a substance. It is used in different ways in animals and in plants.The term is New Latin, f. Greek παρέγχυμα - parenkhuma, "visceral flesh", f. παρεγχεῖν - parenkhein, "to pour in" f. para-, "beside" + en-, "in" + khein, "to pour"...

cells, sieve-tube cells, and companion cells (in addition albuminous cells, fibres and sclereids).

Sieve tubes

The sieve-tube cells lack a nucleus
Cell nucleus
In cell biology, the nucleus is a membrane-enclosed organelle found in eukaryotic cells. It contains most of the cell's genetic material, organized as multiple long linear DNA molecules in complex with a large variety of proteins, such as histones, to form chromosomes. The genes within these...

, have very few vacuole
A vacuole is a membrane-bound organelle which is present in all plant and fungal cells and some protist, animal and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain...

s, but contain other organelles such as ribosome
A ribosome is a component of cells that assembles the twenty specific amino acid molecules to form the particular protein molecule determined by the nucleotide sequence of an RNA molecule....

s. The sieve tube is an elongated rank of individual cells, called sieve-tube members, arranged end to end. The endoplasmic reticulum
Endoplasmic reticulum
The endoplasmic reticulum is an organelle of cells in eukaryotic organisms that forms an interconnected network of tubules, vesicles, and cisternae...

 is concentrated at the lateral walls. Sieve-tube members are joined end to end to form a tube that conducts food materials throughout the plant. The end walls of these cells have many small pores and are called sieve plates and have enlarged plasmodesmata
Plasmodesmata are microscopic channels which traverse the cell walls of plant cells and some algal cells, enabling transport and communication between them. Species that have plasmodesmata include members of the Charophyceae, Charales and Coleochaetales , as well as all embryophytes, better known...


Companion cells

The survival of sieve-tube members depends on a close association with the companion cells. All of the cellular functions of a sieve-tube element are carried out by the (much smaller) companion cell, a typical plant cell
Plant cell
Plant cells are eukaryotic cells that differ in several key respects from the cells of other eukaryotic organisms. Their distinctive features include:...

, except the companion cell usually has a larger number of ribosomes and mitochondria. This is because the companion cell is more metabolically active than a 'typical' plant cell. The 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...

 of a companion cell is connected to the sieve-tube element by plasmodesmata.

There are three types of companion cell.
  1. Ordinary companions cells, which have smooth walls and few or no plasmodesmata connections to cells other than the sieve tube.
  2. Transfer cells
    Transfer cells
    Transfer cells are specialized parenchyma cells that have an increased surface area, due to infoldings of the plasma membrane. They facilitate the transport of sugars from a sugar source, mainly leaves, to a sugar sink, often developing fruits....

    , which have much folded walls that are adjacent to non-sieve cells, allowing for larger areas of transfer. They are specialised in scavenging solutes from those in the cell walls that are actively pumped requiring energy.
  3. Intermediary cells, which have smooth walls and numerous plasmodesmata connecting them to other cells.

The first two types of cell collect solutes through apoplast
Within a plant, the apoplast is the free diffusional space outside the plasma membrane. It is interrupted by the Casparian strip in roots, air spaces between plant cells and the cuticula of the plant....

ic (cell wall) transfers, whilst the third type can collect solutes via the symplast
The symplast of a plant is the inner side of the plasma membrane in which water can freely diffuse.The plasmodesmata allow the direct flow of small molecules such as sugars, amino acids, and ions between cells...

 through the plasmodesmata connections.


Unlike xylem
Xylem is one of the two types of transport tissue in vascular plants. . The word xylem is derived from the Classical Greek word ξυλον , meaning "wood"; the best-known xylem tissue is wood, though it is found throughout the plant...

 (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 sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

s made by the photosynthetic areas. These sugars are transported to non-photosynthetic parts of the plant, such as the roots, or into storage structures, such as tuber
Tubers are various types of modified plant structures that are enlarged to store nutrients. They are used by plants to survive the winter or dry months and provide energy and nutrients for regrowth during the next growing season and they are a means of asexual reproduction...

s or bulbs.

The Pressure flow hypothesis was a hypothesis proposed by Ernst Münch
Ernst Munch
Ernst Münch was a German plant physiologist who proposed the Pressure Flow Hypothesis in 1926.He worked in Aschaffenburg and Munich with Robert Hartig. He worked in a number of fields including forest pathology, resin production, and fungi...

 in 1930 that explained the mechanism of phloem translocation. A high concentration of organic substance inside 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....

s of the phloem at a source, such as a leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

, creates a diffusion gradient
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...

 that draws water into the cells. Movement occurs by bulk flow; phloem sap moves from sugar sources to sugar sinks by means of turgor pressure gradient. A sugar source is any part of the plant that is producing or releasing sugar.

During the plant's growth period, usually during the spring, storage organs such as the root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...

s 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 meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....

s are dormant, the leaves
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 are sources, and storage organs are sinks. Developing seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

-bearing organs (such as fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

) 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. Cells in a sugar source "load" a sieve-tube element by actively transporting
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...

 solute molecules into it. This causes water to move into the sieve-tube element by 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...

, creating pressure that pushes the sap down the tube. In sugar sinks, cells actively transport solutes out of the sieve-tube elements, producing the exactly opposite effect.

Some plants however appear not to load phloem by active transport. In these cases a mechanism known as the polymer trap mechanism was proposed by Robert Turgeon. In this case small sugars such as sucrose move into intermediary cells through narrow plasmodesmata, where they are polymerised to raffinose
Raffinose is a trisaccharide composed of galactose, fructose, and glucose. It can be found in beans, cabbage, brussels sprouts, broccoli, asparagus, other vegetables, and whole grains. Raffinose can be hydrolyzed to D-galactose and sucrose by the enzyme α-galactosidase , an enzyme not found in the...

 and other larger oligosaccharides. Now they are unable to move back, but can proceed through wider plasmodesmata into the sieve tube element.

The symplastic phloem loading (polymer trap mechanism above) is confined mostly to plants in tropical rain forests and is seen as more primitive. The actively-transported apoplastic phloem loading is viewed as more advanced, as it is found in the later-evolved plants, and particularly in those in temperate and arid conditions. This mechanism may, therefore, have allowed plants to colonise the cooler locations.

Organic molecule
A molecule is an electrically neutral group of at least two atoms held together by covalent chemical bonds. Molecules are distinguished from ions by their electrical charge...

s such as sugars, amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...

s, certain hormone
A hormone is a chemical released by a cell or a gland in one part of the body that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. In essence, it is a chemical messenger that transports a signal from one...

s, and even messenger RNA
Messenger RNA
Messenger RNA is a molecule of RNA encoding a chemical "blueprint" for a protein product. mRNA is transcribed from a DNA template, and carries coding information to the sites of protein synthesis: the ribosomes. Here, the nucleic acid polymer is translated into a polymer of amino acids: a protein...

s are transported in the phloem through sieve tube element
Sieve tube element
Sieve tubes are mainly to transport sugars and nutrients up and down the plant. In plant anatomy, sieve vascular tissue tube elements, also called sieve tube members, are a type of elongated sclerenchyma cells in phloem tissue. The ends of these cells are connected with other sieve elements, and...



Because phloem tubes sit on the outside of the xylem
Xylem is one of the two types of transport tissue in vascular plants. . The word xylem is derived from the Classical Greek word ξυλον , meaning "wood"; the best-known xylem tissue is wood, though it is found throughout the plant...

 in most plants, a tree or other plant can be effectively killed by stripping away the bark in a ring on the trunk or stem. With the phloem destroyed, nutrients cannot reach the roots, and the tree/plant will die. Trees located in areas with animals such as beavers are vulnerable since beavers chew off the bark at a fairly precise height. This process is known as girdling, and can be used for agricultural purposes. For example, enormous fruits and vegetables seen at fairs and carnivals are produced via girdling. A farmer would place a girdle at base of a large branch, and remove all but one fruit/vegetable from that branch. Thus, all the sugars manufactured by leaves on that branch have no sinks
Carbon dioxide sink
A carbon sink is a natural or artificial reservoir that accumulates and stores some carbon-containing chemical compound for an indefinite period. The process by which carbon sinks remove carbon dioxide from the atmosphere is known as carbon sequestration...

 to go to but the one fruit/vegetable, which thus expands to many times normal size.


The phloem originates, and grows outwards from, meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....

atic cells in 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...

. Phloem is produced in phases. Primary phloem is laid down by the apical meristem and develops from the procambium. Secondary phloem is laid down by the vascular cambium to the inside of the established layer(s) of phloem.

In some eudicot families (Apocynaceae
The Apocynaceae or dogbane family is a family of flowering plants that includes trees, shrubs, herbs, and lianas.Many species are tall trees found in tropical rainforests, and most are from the tropics and subtropics, but some grow in tropical dry, xeric environments. There are also perennial herbs...

, Convolvulaceae
Convolvulaceae, known commonly as the bindweed or morning glory family, are a group of about 60 genera and more than 1,650 species of mostly herbaceous vines, but also trees, shrubs and herbs.- Description :...

, Cucurbitaceae
The plant family Cucurbitaceae consists of various squashes, melons, and gourds, including crops such as cucumber, pumpkins, luffas, and watermelons...

, Solanaceae
Solanaceae are a family of flowering plants that include a number of important agricultural crops as well as many toxic plants. The name of the family comes from the Latin Solanum "the nightshade plant", but the further etymology of that word is unclear...

, Myrtaceae
The Myrtaceae or Myrtle family are a family of dicotyledon plants, placed within the order Myrtales. Myrtle, clove, guava, feijoa, allspice, and eucalyptus belong here. All species are woody, with essential oils, and flower parts in multiples of four or five...

, Asteraceae
The Asteraceae or Compositae , is an exceedingly large and widespread family of vascular plants. The group has more than 22,750 currently accepted species, spread across 1620 genera and 12 subfamilies...

), phloem also develops on the inner side of the vascular cambium; in this case, a distinction between external phloem and internal phloem or intraxylary phloem is made. Internal phloem is mostly primary, and begins differentiation later than the external phloem and protoxylem, though it is not without exceptions. In some other families (Amaranthaceae
The flowering plant family Amaranthaceae, the Amaranth family, contains about 176 genera and 2,400 species.- Description :Most of these species are herbs or subshrubs; very few are trees or climbers. Some species are succulent....

, Nyctaginaceae
Nyctaginaceae, the Four O'Clock Family, is a family of around 33 genera and 290 species of flowering plants, widely distributed in tropical and subtropical regions, with a few representatives in temperate regions...

, Salvadoraceae
Salvadoraceae is a family in the plant order Brassicales, comprising 3 genera totalling around 12 species. They occur in Africa, including Madagascar; South East Asia; and have also been found on Java, suggesting they are probably found in much of Malesia...

), the cambium also periodically forms inward strands or layers of phloem, embedded in the xylem: Such phloem strands are called included phloem or interxylary phloem.

Nutritional use

Phloem of pine
Pines are trees in the genus Pinus ,in the family Pinaceae. They make up the monotypic subfamily Pinoideae. There are about 115 species of pine, although different authorities accept between 105 and 125 species.-Etymology:...

 trees has been used in Finland
Finland , officially the Republic of Finland, is a Nordic country situated in the Fennoscandian region of Northern Europe. It is bordered by Sweden in the west, Norway in the north and Russia in the east, while Estonia lies to its south across the Gulf of Finland.Around 5.4 million people reside...

 as a substitute food in times of famine
A famine is a widespread scarcity of food, caused by several factors including crop failure, overpopulation, or government policies. This phenomenon is usually accompanied or followed by regional malnutrition, starvation, epidemic, and increased mortality. Every continent in the world has...

, and even in good years in the northeast, where supplies of phloem from earlier years helped stave off starvation somewhat in the great famine of the 1860s
Finnish famine of 1866-1868
The Famine of 1866–1868 was the last famine in Finland and northern Sweden, and the last major naturally caused famine in Europe. In Finland the famine is known as "the great hunger years", or . About 15% of the entire population died; in the hardest-hit areas up to 20%. The total death toll was...

. Phloem is dried and milled to flour (pettu in Finnish
Finnish language
Finnish is the language spoken by the majority of the population in Finland Primarily for use by restaurant menus and by ethnic Finns outside Finland. It is one of the two official languages of Finland and an official minority language in Sweden. In Sweden, both standard Finnish and Meänkieli, a...

) and mixed with rye
Rye is a grass grown extensively as a grain and as a forage crop. It is a member of the wheat tribe and is closely related to barley and wheat. Rye grain is used for flour, rye bread, rye beer, some whiskeys, some vodkas, and animal fodder...

to form a hard dark bread. Recently, pettu has again become available as a curiosity, and some have made claims of health benefits.
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