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In
botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, secondary growth is the growth that results from
cell division Cell division is the process by which a parent 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 eukaryotes, there ...
in the cambia or lateral
meristems 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 ...
and that causes the
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
s and
root 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 su ...
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 plant 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 ...
s, but
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 ...
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 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, ...
and
gymnosperms 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, γυμν� ...
. 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 disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as " cryptogams", meaning that their means of reproduction is hidden. Fer ...
s this feature is extremely rare, only occurring in ''
Isoetes ''Isoetes'', commonly known as the quillworts, is the only extant genus of plants in the family Isoetaceae, which is in the class of lycopods. There are currently 192 recognized species, with a cosmopolitan distribution but with the individual ...
''.


Lateral meristems

In many
vascular plants 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 ...
, secondary growth is the result of the activity of the two lateral meristems, the
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 ...
and
vascular cambium 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 xy ...
. 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 opposite to herbaceous plants that die back to the ground until sp ...
s, this process produces
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...
, 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, including only woody plants with secondary growth, plants that are ...
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 hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rel ...
of the stem or roots, plants with secondary growth usually also develop 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 ...
. 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 ''Quercus suber'', commonly called the cork oak, is a medium-sized, evergreen oak tree in the section ''Quercus'' sect. ''Cerris''. It is the primary source of cork for wine bottle stoppers and other uses, such as cork flooring and as the core ...
it will yield harvestable
cork Cork or CORK may refer to: Materials * Cork (material), an impermeable buoyant plant product ** Cork (plug), a cylindrical or conical object used to seal a container ***Wine cork Places Ireland * Cork (city) ** Metropolitan Cork, also known as G ...
.


In nonwoody plants

Secondary growth also occurs in many nonwoody plants, e.g.
tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
,
potato The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae. Wild potato species can be found from the southern Uni ...
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
,
carrot The carrot ('' Daucus carota'' subsp. ''sativus'') is a root vegetable, typically orange in color, though purple, black, red, white, and yellow cultivars exist, all of which are domesticated forms of the wild carrot, ''Daucus carota'', na ...
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 taproo ...
and
sweet potato The sweet potato or sweetpotato ('' Ipomoea batatas'') is a dicotyledonous plant that belongs to the bindweed or morning glory family, Convolvulaceae. Its large, starchy, sweet-tasting tuberous roots are used as a root vegetable. The young ...
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 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 ...
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 c ...
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 four o' clock family, Nyctaginaceae. It is native to eastern South America, found from Brazil, west to Peru, and south to southern Argentina. ...
a series of cambia arise outside the oldest phloem. Ancestral
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 ...
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 * Several other plants known as "palm" Palm or Palms may also refer to: Music * Palm (ba ...
trees increase their trunk diameter due to division and enlargement of
parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word � ...
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'' is a genus of perennial shrubs and trees in the family Asparagaceae, subfamily Agavoideae. Its 40–50 species are notable for their rosettes of evergreen, tough, sword-shaped leaves and large terminal panicles of white or whitis ...
'' 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 Apical means "pertaining to an apex". It may refer to: *Apical ancestor, refers to the last common ancestor of an entire group, such as a species (biology) or a clan (anthropology) *Apical (anatomy), an anatomical term of location for features loc ...
meristem.


See also

*
Bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, e ...
*
Cambium (botany) 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 ...
*
Root 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 su ...
*
Stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
* Tylosis *
Dendrochronology Dendrochronology (or tree-ring dating) is the scientific method of dating tree rings (also called growth rings) to the exact year they were formed. As well as dating them, this can give data for dendroclimatology, the study of climate and atm ...
*
Herbchronology Herbchronology is the analysis of annual growth rings (or simply annual rings) in the secondary root xylem of perennial herbaceous plants. While leaves and stems of perennial herbs die down at the end of the growing season the root often persist ...
*
Anatomy Anatomy () is the branch of biology concerned with the study of the 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 science, having i ...


References

{{botany Plant anatomy Plant physiology Tissues (biology)