The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in

dicot The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...

s such as buttercups and oak trees,

gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό� ...

s such as pine trees, as well as in certain other

vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...

s. It produces secondary

xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...

inwards, towards the

pith Pith, or medulla, is a tissue in the stems of vascular plants. Pith is composed of soft, spongy parenchyma cells, which in some cases can store starch. In eudicotyledons, pith is located in the center of the stem. In monocotyledons, it ext ...

, and secondary phloem outwards, towards the bark. In herbaceous plants, it occurs in the vascular bundles which are often arranged like beads on a necklace forming an interrupted ring inside the stem. In woody plants, it forms a cylinder of unspecialized

meristem The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...

cells, as a continuous ring from which the new tissues are grown. Unlike the xylem and phloem, it does not transport water, minerals or food through the plant. Other names for the vascular

cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from w ...

are the main cambium, wood cambium, or bifacial cambium.

Occurrence

Vascular cambia are found in all

seed plants A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...

except for five

angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

lineages which have independently lost it;

Nymphaeales The Nymphaeales are an order (biology), order of flowering plants, consisting of three Family (biology), families of aquatic plants, the Hydatellaceae, the Cabombaceae, and the Nymphaeaceae (water lilies). It is one of the three orders of basal a ...

Ceratophyllum ''Ceratophyllum'' is a cosmopolitan genus of flowering plants including four accepted living species in 2016, commonly found in ponds, marshes, and quiet streams in tropical and in temperate regions. It is the only extant genus in the family Ce ...

Nelumbo ''Nelumbo'' is a genus of aquatic plants with large, showy flowers. Members are commonly called lotus, though the name is also applied to various other plants and plant groups, including the unrelated genus '' Lotus''. Members outwardly resemb ...

Podostemaceae Podostemaceae (riverweed family), a family in the order Malpighiales, comprise about 50 genera and species of more or less thalloid aquatic herbs. Distribution and habitat They are found mostly in tropical and subtropical areas worldwide. Ma ...

, and

monocots Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...

. A few leaf types also have a vascular cambium. In dicot and gymnosperm

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

s, the vascular cambium is the obvious line separating the bark and wood; they also have a

cork cambium Cork cambium (pl. cambia or cambiums) is a tissue found in many vascular plants as a part of the epidermis. It is one of the many layers of bark, between the cork and primary phloem. The cork cambium is a lateral meristem and is responsible for ...

. For successful

grafting Grafting or graftage is a horticultural technique whereby tissues of plants are joined so as to continue their growth together. The upper part of the combined plant is called the scion () while the lower part is called the rootstock. The succ ...

, the vascular cambia of the rootstock and scion must be aligned so they can grow together.

Structure and function

The cambium present between primary xylem and primary phloem is called the ''intrafascicular'' cambium (within vascular bundles). During secondary growth, cells of medullary rays, in a line (as seen in section; in three dimensions, it is a sheet) between neighbouring vascular bundles, become

atic and form new ''interfascicular'' cambium (between vascular bundles). The fascicular and interfascicular cambia thus join up to form a ring (in three dimensions, a tube) which separates the primary xylem and primary phloem, the ''cambium ring''. The vascular cambium produces secondary xylem on the inside of the ring, and secondary phloem on the outside, pushing the primary xylem and phloem apart. The vascular cambium usually consists of two types of cells: * Fusiform initials (tall, axially oriented) * Ray initials (smaller and round to angular in shape)

Maintenance of cambial meristem

The vascular cambium is maintained by a network of interacting signal feedback loops. Currently, both hormones and short peptides have been identified as information carriers in these systems. While similar regulation occurs in other plant

s, the cambial meristem receives signals from both the xylem and phloem sides for the meristem. Signals received from outside the meristem act to down regulate internal factors, which promotes cell proliferation and differentiation.

Hormonal regulation

The

phytohormones Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...

that are involved in the vascular cambial activity are auxins,

ethylene Ethylene (IUPAC name: ethene) is a hydrocarbon which has the formula or . It is a colourless, flammable gas with a faint "sweet and musky" odour when pure. It is the simplest alkene (a hydrocarbon with carbon-carbon double bonds). Ethylene i ...

gibberellin Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. GAs are one of the longest-known classes of p ...

cytokinin Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and le ...

s, abscisic acid and probably more to be discovered. Each one of these plant hormones is vital for regulation of cambial activity. Combination of different concentrations of these hormones is very important in plant metabolism. Auxin hormones are proven to stimulate

mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...

, cell production and regulate interfascicular and fascicular cambium. Applying auxin to the surface of a tree stump allowed decapitated shoots to continue secondary growth. The absence of auxin hormones will have a detrimental effect on a plant. It has been shown that mutants without auxin will exhibit increased spacing between the interfascicular cambiums and reduced growth of the

vascular bundle A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will inc ...

s. The mutant plant will therefore experience a decrease in water, nutrients, and photosynthates being transported throughout the plant, eventually leading to death. Auxin also regulates the two types of cell in the vascular cambium, ray and fusiform initials. Regulation of these initials ensures the connection and communication between xylem and phloem is maintained for the translocation of nourishment and sugars are safely being stored as an energy resource. Ethylene levels are high in plants with an active cambial zone and are still currently being studied. Gibberellin stimulates the cambial cell division and also regulates differentiation of the xylem tissues, with no effect on the rate of phloem differentiation. Differentiation is an essential process that changes these tissues into a more specialized type, leading to an important role in maintaining the life form of a plant. In poplar trees, high concentrations of gibberellin is positively correlated to an increase of cambial cell division and an increase of auxin in the cambial stem cells. Gibberellin is also responsible for the expansion of xylem through a signal traveling from the shoot to the root. Cytokinin hormone is known to regulate the rate of the cell division instead of the direction of cell differentiation. A study demonstrated that the mutants are found to have a reduction in stem and root growth but the secondary vascular pattern of the vascular bundles were not affected with a treatment of cytokinin.

Cambium as food

The cambium of most trees are edible. In Scandinavia, it was historically used as a flour to make

bark bread Bark bread is a traditional food made with cambium (phloem) flour. It has a history of use as famine food. History Bark bread seems to be a primarily Scandinavian tradition. Mention of it is found in medieval literature, and it may have an even ...

References

External links

* Pictures o
Vascular cambium

* {{Authority control Non-timber forest products Plant physiology Plant anatomy Plant cells