Secondary Growth
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In
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
, secondary growth is the growth that results from
cell division Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...
in the
cambia Cambia is an Australian-based global non-profit social enterprise focusing on open science, biology, innovation system reform and intellectual property. Its projects include The Lens, formerly known as Patent Lens, and the Biological Innovat ...
or lateral meristems and that causes the stems and
root In vascular plants, the roots are the plant organ, 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 bel ...
s 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 Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are flowering plants whose seeds contain only one embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but with various ranks a ...
usually lack secondary growth. If they do have secondary growth, it differs from the typical pattern of other seed plants. The formation of secondary vascular tissues from the cambium is a characteristic feature of dicotyledons and
gymnosperms The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...
. In certain monocots, the vascular tissues are also increased after the primary growth is completed but the cambium of these plants is of a different nature. In the living
pteridophyte A pteridophyte is a vascular plant (with xylem and phloem) that reproduces by means of spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as " cryptogams", meaning that their means of reproduction is ...
s this feature is extremely rare, only occurring in '' Isoetes''.


Lateral meristems

In many vascular plants, secondary growth is the result of the activity of the two lateral meristems, the
cork cambium Cork cambium (: 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 s ...
and
vascular cambium The vascular cambium is the main growth tissue in the stems and roots of many plants exhibiting secondary growth, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular ...
. Arising from ''lateral'' meristems, secondary growth increases the width of the plant root or stem, rather than its length. As long as the lateral meristems continue to produce new cells, the stem or root will continue to grow in diameter. In
woody plant A woody plant is a plant that produces wood as its structural tissue and thus has a hard stem. In cold climates, woody plants further survive winter or dry season above ground, as opposed to Herbaceous plant, herbaceous plants that die back to t ...
s, this process produces
wood Wood is a structural tissue/material found as xylem in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulosic fibers that are strong in tension and embedded in a matrix of lignin t ...
, and shapes the plant into a
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, e.g., including only woody plants with secondary growth, only ...
with a thickened trunk. Because this growth usually ruptures the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
of the stem or roots, plants with secondary growth usually also develop a
cork cambium Cork cambium (: 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 s ...
. The cork cambium gives rise to thickened cork cells to protect the surface of the plant and reduce water loss. If this is kept up over many years, this process may produce a layer of cork. In the case of the cork oak it will yield harvestable cork.


In nonwoody plants

Secondary growth also occurs in many nonwoody plants, e.g.
tomato The tomato (, ), ''Solanum lycopersicum'', is a plant whose fruit is an edible Berry (botany), berry that is eaten as a vegetable. The tomato is a member of the nightshade family that includes tobacco, potato, and chili peppers. It originate ...
,
potato The potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...
,
carrot The carrot ('' Daucus carota'' subsp. ''sativus'') is a root vegetable, typically orange in colour, though heirloom variants including purple, black, red, white, and yellow cultivars exist, all of which are domesticated forms of the wild ...
taproot A taproot is a large, central, and dominant root from which other roots sprout laterally. Typically a taproot is somewhat straight and very thick, is tapering in shape, and grows directly downward. In some plants, such as the carrot, the taproot ...
and
sweet potato The sweet potato or sweetpotato (''Ipomoea batatas'') is a dicotyledonous plant in the morning glory family, Convolvulaceae. Its sizeable, starchy, sweet-tasting tuberous roots are used as a root vegetable, which is a staple food in parts of ...
tuberous root. A few long-lived leaves also have secondary growth.


Abnormal secondary growth

Abnormal secondary growth does not follow the pattern of a single vascular cambium producing
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
to the inside and
phloem 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 ...
to the outside as in ancestral lignophytes. Some
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, ...
s have anomalous secondary growth, e.g. in
Bougainvillea ''Bougainvillea'' ( , ) is a genus of thorny ornamental vines, bushes, and trees belonging to the family, Nyctaginaceae. They are native to Brazil, Bolivia, Paraguay, Peru, and Argentina. There are between 4 and 22 species in the genus. The i ...
a series of cambia arise outside the oldest phloem. Ancestral
monocots Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are flowering plants whose seeds contain only one embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but with various ranks a ...
lost their secondary growth and their stele has changed in a way it could not be recovered without major changes that are very unlikely to occur. Monocots either have no secondary growth, as is the ancestral case, or they have an "anomalous secondary growth" of some type, or, in the case of palms, they enlarge their diameter in what is called a sort of secondary growth or not depending on the definition given to the term.
Palm Palm most commonly refers to: * Palm of the hand, the central region of the front of the hand * Palm plants, of family Arecaceae ** List of Arecaceae genera **Palm oil * Several other plants known as "palm" Palm or Palms may also refer to: Music ...
trees increase their trunk diameter due to division and enlargement of
parenchyma upright=1.6, Lung parenchyma showing damage due to large subpleural bullae. Parenchyma () is the bulk of functional substance in an animal organ such as the brain or lungs, or a structure such as a tumour. In zoology, it is the tissue that ...
cells, which is termed "primary gigantism" because there is no production of secondary xylem and phloem tissues, or sometimes "diffuse secondary growth". In some other monocot stems as in ''
Yucca ''Yucca'' ( , YUCK-uh) is both the scientific name and common name for a genus native to North America from Panama to southern Canada. It contains 50 accepted species. In addition to yucca, they are also known as Adam's needle or Spanish-bayon ...
'' and '' Dracaena'' with anomalous secondary growth, a cambium forms, but it produces vascular bundles and parenchyma internally and just parenchyma externally. Some monocot stems increase in diameter due to the activity of a primary thickening meristem, which is derived from the apical meristem.


See also

* Bark * Cambium (botany) *
Root In vascular plants, the roots are the plant organ, 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 bel ...
* Stem * Tylosis *
Dendrochronology Dendrochronology (or tree-ring dating) is the scientific method of chronological dating, dating tree rings (also called growth rings) to the exact year they were formed in a tree. As well as dating them, this can give data for dendroclimatology, ...
* Herbchronology *
Anatomy Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...


References

{{botany Plant anatomy Plant physiology Tissues (biology)