Vascular cambium

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Helianthus stem in section. The cells of the vascular cambium (F) divide to form phloem on the outside, located beneath the bundle cap (E), and xylem (D) on the inside. Most of the vascular cambium is here in vascular bundles (ovals of phloem and xylem together) but it is starting to join these up as at point F between the bundles. Helianthus stem 2 L.jpg
Helianthus stem in section. The cells of the vascular cambium (F) divide to form phloem on the outside, located beneath the bundle cap (E), and xylem (D) on the inside. Most of the vascular cambium is here in vascular bundles (ovals of phloem and xylem together) but it is starting to join these up as at point F between the bundles.

The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xylem inwards, towards the pith, and secondary phloem outwards, towards the bark.

Contents

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

Occurrence

Vascular cambia are found in all seed plants except for five angiosperm lineages which have independently lost it; Nymphaeales, Ceratophyllum , Nelumbo , Podostemaceae, and monocots. [1] In dicot and gymnosperm trees, the vascular cambium is the obvious line separating the bark and wood; they also have a cork cambium. For successful grafting, 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 meristematic 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:

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 meristems, 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. [2]

Hormonal regulation

The phytohormones that are involved in the vascular cambial activity are auxins, ethylene, gibberellins, cytokinins, 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, 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 bundles. 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.[ citation needed ]

Cambium as food

The cambium of most trees are edible. In Scandinavia, it was historically used as a flour to make bark bread. [3]

See also

Related Research Articles

<span class="mw-page-title-main">Root</span> Basal organ of a vascular plant

In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the surface of the soil, but roots can also be aerial or aerating, that is, growing up above the ground or especially above water.

<span class="mw-page-title-main">Phloem</span> Sugar transport tissue in vascular plants

Phloem 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. Different types of phloem can be distinguished. The early phloem formed in the growth apices is called protophloem. Protophloem eventually becomes obliterated once it connects to the durable phloem in mature organs, the metaphloem. Further, secondary phloem is formed during the thickening of stem structures.

<span class="mw-page-title-main">Tissue (biology)</span> Group of similar cells performing a specific function

In biology, tissue is an assembly of similar cells and their extracellular matrix from the same embryonic origin that together carry out a specific function. Tissues occupy a biological organizational level between cells and a complete organ. Accordingly, organs are formed by the functional grouping together of multiple tissues.

<span class="mw-page-title-main">Apical dominance</span> Phenomenon where the central stem grows stronger than other stems

In botany, apical dominance is the phenomenon whereby the main, central stem of the plant is dominant over other side stems; on a branch the main stem of the branch is further dominant over its own side twigs.

<span class="mw-page-title-main">Cork cambium</span> Party of a plant

Cork cambium 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 secondary growth that replaces the epidermis in roots and stems. It is found in woody and many herbaceous dicots, gymnosperms and some monocots. It is one of the plant's meristems – the series of tissues consisting of embryonic disk cells from which the plant grows. The function of cork cambium is to produce the cork, a tough protective material.

<span class="mw-page-title-main">Meristem</span> Type of plant tissue involved in cell proliferation

In cell biology, the meristem is a type of tissue found in plants. It consists of undifferentiated cells capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells continue to divide until they become differentiated and lose the ability to divide.

<span class="mw-page-title-main">Plant hormone</span> Chemical compounds that regulate plant growth and development

Plant hormones are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of organ size, pathogen defense, stress tolerance and reproductive development. Unlike in animals each plant cell is capable of producing hormones. Went and Thimann coined the term "phytohormone" and used it in the title of their 1937 book.

<span class="mw-page-title-main">Auxin</span> Plant hormone

Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essential for plant body development. The Dutch biologist Frits Warmolt Went first described auxins and their role in plant growth in the 1920s. Kenneth V. Thimann became the first to isolate one of these phytohormones and to determine its chemical structure as indole-3-acetic acid (IAA). Went and Thimann co-authored a book on plant hormones, Phytohormones, in 1937.

<span class="mw-page-title-main">Cytokinin</span> Class of plant hormones promoting cell division

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 leaf senescence.

The pericycle is a cylinder of parenchyma or sclerenchyma cells that lies just inside the endodermis and is the outer most part of the stele of plants.

<span class="mw-page-title-main">Epidermis (botany)</span> Layer of cells that covers leaves, flowers, roots of plants

The epidermis is a single layer of cells that covers the leaves, flowers, roots and stems of plants. It forms a boundary between the plant and the external environment. The epidermis serves several functions: it protects against water loss, regulates gas exchange, secretes metabolic compounds, and absorbs water and mineral nutrients. The epidermis of most leaves shows dorsoventral anatomy: the upper (adaxial) and lower (abaxial) surfaces have somewhat different construction and may serve different functions. Woody stems and some other stem structures such as potato tubers produce a secondary covering called the periderm that replaces the epidermis as the protective covering.

<span class="mw-page-title-main">Vascular tissue</span> Conducting tissue in vascular plants

Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There are also two meristems associated with vascular tissue: the vascular cambium and the cork cambium. All the vascular tissues within a particular plant together constitute the vascular tissue system of that plant.

<span class="mw-page-title-main">Secondary growth</span> Type of growth in plants

In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of stems and roots, causing them to elongate, and gives rise to primary tissue. Secondary growth occurs in most seed plants, but monocots usually lack secondary growth. If they do have secondary growth, it differs from the typical pattern of other seed plants.

<span class="mw-page-title-main">Primordium</span> Organ in the earliest recognizable stage of embryonic development

A primordium in embryology, is an organ or tissue in its earliest recognizable stage of development. Cells of the primordium are called primordial cells. A primordium is the simplest set of cells capable of triggering growth of the would-be organ and the initial foundation from which an organ is able to grow. In flowering plants, a floral primordium gives rise to a flower.

<span class="mw-page-title-main">Lateral root</span> Plant root

Lateral roots, emerging from the pericycle, extend horizontally from the primary root (radicle) and over time makeup the iconic branching pattern of root systems. They contribute to anchoring the plant securely into the soil, increasing water uptake, and facilitate the extraction of nutrients required for the growth and development of the plant. Lateral roots increase the surface area of a plant's root system and can be found in great abundance in several plant species. In some cases, lateral roots have been found to form symbiotic relationships with rhizobia (bacteria) and mycorrhizae (fungi) found in the soil, to further increase surface area and increase nutrient uptake.

The unifacial cambium produces cells to the interior of its cylinder. These cells differentiate into xylem tissue. Unlike the more common bifacial cambium found in later woody plants, the unifacial cambium does not produce phloem to its exterior. Also in contrast to the bifacial cambium, the unifacial cambium is unable to expand its circumference with anticlinal cell division. Cell elongation provides a limited amount of expansion.

Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues. By contrast, an animal embryo will very early produce all of the body parts that it will ever have in its life. When the animal is born, it has all its body parts and from that point will only grow larger and more mature. However, both plants and animals pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.

<span class="mw-page-title-main">Plant stem</span> Structural axis of a vascular plant

A stem is on of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, photosynthesis takes place here, stores nutrients, and produces new living tissue. The stem can also be called halm or haulm or culms.

Primary growth in plants is growth that takes place from the tips of roots or shoots. It leads to lengthening of roots and stems and sets the stage for organ formation. It is distinguished from secondary growth that leads to widening. Plant growth takes place in well defined plant locations. Specifically, the cell division and differentiation needed for growth occurs in specialized structures called meristems. These consist of undifferentiated cells capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells continue to divide until they differentiate and then lose the ability to divide. Thus, the meristems produce all the cells used for plant growth and function.

<span class="mw-page-title-main">Cambium</span> Layer of plant tissue with cells for growth

A cambium, 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 which phloem, xylem, or cork grows by division, resulting in secondary thickening. It forms parallel rows of cells, which result in secondary tissues.

References

  1. Povilus, Rebecca A.; DaCosta, Jeffrey M.; Grassa, Christopher; Satyaki, Prasad R. V.; Moeglein, Morgan; Jaenisch, Johan; Xi, Zhenxiang; Mathews, Sarah; Gehring, Mary; Davis, Charles C.; Friedman, William E. (2020-04-14). "Water lily ( Nymphaea thermarum ) genome reveals variable genomic signatures of ancient vascular cambium losses". Proceedings of the National Academy of Sciences. 117 (15): 8649–8656. Bibcode:2020PNAS..117.8649P. doi: 10.1073/pnas.1922873117 . ISSN   0027-8424. PMC   7165481 . PMID   32234787.
  2. Etchells, J. Peter; Mishra, Laxmi S.; Kumar, Manoj; Campbell, Liam; Turner, Simon R. (April 2015). "Wood Formation in Trees Is Increased by Manipulating PXY-Regulated Cell Division". Current Biology. 25 (8): 1050–1055. doi:10.1016/j.cub.2015.02.023. PMC   4406943 . PMID   25866390.
  3. "So You Want to Eat a Tree". 20 May 2016.