Meristem

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Tunica-corpus model of the apical meristem (growing tip). The epidermal (L1) and subepidermal (L2) layers form the outer layers called the tunica. The inner L3 layer is called the corpus. Cells in the L1 and L2 layers divide in a sideways fashion, which keeps these layers distinct, whereas the L3 layer divides in a more random fashion. Meristeme couches.png
Tunica-corpus model of the apical meristem (growing tip). The epidermal (L1) and subepidermal (L2) layers form the outer layers called the tunica. The inner L3 layer is called the corpus. Cells in the L1 and L2 layers divide in a sideways fashion, which keeps these layers distinct, whereas the L3 layer divides in a more random fashion.

In cell biology, 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 continue to divide until they become differentiated and lose the ability to divide.

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Differentiated plant cells generally cannot divide or produce cells of a different type. Meristematic cells are undifferentiated or incompletely differentiated. They are totipotent and capable of continued cell division. Division of meristematic cells provides new cells for expansion and differentiation of tissues and the initiation of new organs, providing the basic structure of the plant body. The cells are small, with small vacuoles or none, and protoplasm filling the cell completely. The plastids (chloroplasts or chromoplasts) are undifferentiated, but are present in rudimentary form (proplastids). Meristematic cells are packed closely together without intercellular spaces. The cell wall is a very thin primary cell wall.

The term meristem was first used in 1858 by Swiss botanist Carl Wilhelm von Nägeli (1817–1891) in his book Beiträge zur Wissenschaftlichen Botanik ("Contributions to Scientific Botany"). [1] It is derived from Greek μερίζειν (merizein) 'to divide', in recognition of its inherent function.[ citation needed ]

There are three types of meristematic tissues: apical (at the tips), intercalary or basal (in the middle), and lateral (at the sides also known as cambium). At the meristem summit, there is a small group of slowly dividing cells, which is commonly called the central zone. Cells of this zone have a stem cell function and are essential for meristem maintenance. The proliferation and growth rates at the meristem summit usually differ considerably from those at the periphery.

Primary meristems

Apical meristems give rise to the primary plant body and are responsible for primary growth, or an increase in length or height. [2] [3] Apical meristems may differentiate into three kinds of primary meristem:

Secondary meristems

After the primary growth, lateral meristems develop as secondary plant growth. This growth adds to the plant in diameter from the established stem but not all plants exhibit secondary growth. There are two types of secondary meristems: the vascular cambium and the cork cambium.

Apical meristems

Apical Meristems are the completely undifferentiated (indeterminate) meristems in a plant. These differentiate into three kinds of primary meristems. The primary meristems in turn produce the two secondary meristem types. These secondary meristems are also known as lateral meristems as they are involved in lateral growth.

Organisation of an apical meristem (growing tip) Central zone Peripheral zone Medullary (i.e. central) meristem Medullary tissue Meristeme coupe zones chiffres.png
Organisation of an apical meristem (growing tip)
  1. Central zone
  2. Peripheral zone
  3. Medullary (i.e. central) meristem
  4. Medullary tissue

There are two types of apical meristem tissue: shoot apical meristem (SAM), which gives rise to organs like the leaves and flowers, and root apical meristem (RAM), which provides the meristematic cells for future root growth. SAM and RAM cells divide rapidly and are considered indeterminate, in that they do not possess any defined end status. In that sense, the meristematic cells are frequently compared to the stem cells in animals, which have an analogous behavior and function.

The apical meristems are layered where the number of layers varies according to plant type. In general the outermost layer is called the tunica while the innermost layers are the corpus. In monocots, the tunica determines the physical characteristics of the leaf edge and margin. In dicots, layer two of the corpus determines the characteristics of the edge of the leaf. The corpus and tunica play a critical part of the plant physical appearance as all plant cells are formed from the meristems. Apical meristems are found in two locations: the root and the stem. Some arctic plants have an apical meristem in the lower/middle parts of the plant. It is thought that this kind of meristem evolved because it is advantageous in arctic conditions.[ citation needed ]

Shoot Apical Meristems

Shoot apical meristems of Crassula ovata (left). Fourteen days later, leaves have developed (right). Apical Meristems in Crassula ovata.png
Shoot apical meristems of Crassula ovata (left). Fourteen days later, leaves have developed (right).

Shoot apical meristems are the source of all above-ground organs, such as leaves and flowers. Cells at the shoot apical meristem summit serve as stem cells to the surrounding peripheral region, where they proliferate rapidly and are incorporated into differentiating leaf or flower primordia.

The shoot apical meristem is the site of most of the embryogenesis in flowering plants.[ citation needed ] Primordia of leaves, sepals, petals, stamens, and ovaries are initiated here at the rate of one every time interval, called a plastochron. It is where the first indications that flower development has been evoked are manifested. One of these indications might be the loss of apical dominance and the release of otherwise dormant cells to develop as auxiliary shoot meristems, in some species in axils of primordia as close as two or three away from the apical dome.

The shoot apical meristem consists of four distinct cell groups:

These four distinct zones are maintained by a complex signalling pathway. In Arabidopsis thaliana , 3 interacting CLAVATA genes are required to regulate the size of the stem cell reservoir in the shoot apical meristem by controlling the rate of cell division. [5] CLV1 and CLV2 are predicted to form a receptor complex (of the LRR receptor-like kinase family) to which CLV3 is a ligand. [6] [7] [8] CLV3 shares some homology with the ESR proteins of maize, with a short 14 amino acid region being conserved between the proteins. [9] [10] Proteins that contain these conserved regions have been grouped into the CLE family of proteins. [9] [10]

CLV1 has been shown to interact with several cytoplasmic proteins that are most likely involved in downstream signalling. For example, the CLV complex has been found to be associated with Rho/Rac small GTPase-related proteins. [5] These proteins may act as an intermediate between the CLV complex and a mitogen-activated protein kinase (MAPK), which is often involved in signalling cascades. [11] KAPP is a kinase-associated protein phosphatase that has been shown to interact with CLV1. [12] KAPP is thought to act as a negative regulator of CLV1 by dephosphorylating it. [12]

Another important gene in plant meristem maintenance is WUSCHEL (shortened to WUS), which is a target of CLV signaling in addition to positively regulating CLV, thus forming a feedback loop. [13] WUS is expressed in the cells below the stem cells of the meristem and its presence prevents the differentiation of the stem cells. [13] CLV1 acts to promote cellular differentiation by repressing WUS activity outside of the central zone containing the stem cells. [5]

The function of WUS in the shoot apical meristem is linked to the phytohormone cytokinin. Cytokinin activates histidine kinases which then phosphorylate histidine phosphotransfer proteins. [14] Subsequently, the phosphate groups are transferred onto two types of Arabidopsis response regulators (ARRs): Type-B ARRS and Type-A ARRs. Type-B ARRs work as transcription factors to activate genes downstream of cytokinin, including A-ARRs. A-ARRs are similar to B-ARRs in structure; however, A-ARRs do not contain the DNA binding domains that B-ARRs have, and which are required to function as transcription factors. [15] Therefore, A-ARRs do not contribute to the activation of transcription, and by competing for phosphates from phosphotransfer proteins, inhibit B-ARRs function. [16] In the SAM, B-ARRs induce the expression of WUS which induces stem cell identity. [17] WUS then suppresses A-ARRs. [18] As a result, B-ARRs are no longer inhibited, causing sustained cytokinin signaling in the center of the shoot apical meristem. Altogether with CLAVATA signaling, this system works as a negative feedback loop. Cytokinin signaling is positively reinforced by WUS to prevent the inhibition of cytokinin signaling, while WUS promotes its own inhibitor in the form of CLV3, which ultimately keeps WUS and cytokinin signaling in check. [19]

Root apical meristem

10x microscope image of root tip with meristem quiescent center calyptrogen (live rootcap cells) rootcap sloughed off dead rootcap cells procambium Root-tip-tag.png
10x microscope image of root tip with meristem
  1. quiescent center
  2. calyptrogen (live rootcap cells)
  3. rootcap
  4. sloughed off dead rootcap cells
  5. procambium

Unlike the shoot apical meristem, the root apical meristem produces cells in two dimensions. It harbors two pools of stem cells around an organizing center called the quiescent center (QC) cells and together produces most of the cells in an adult root. [20] [21] At its apex, the root meristem is covered by the root cap, which protects and guides its growth trajectory. Cells are continuously sloughed off the outer surface of the root cap. The QC cells are characterized by their low mitotic activity. Evidence suggests that the QC maintains the surrounding stem cells by preventing their differentiation, via signal(s) that are yet to be discovered. This allows a constant supply of new cells in the meristem required for continuous root growth. Recent findings indicate that QC can also act as a reservoir of stem cells to replenish whatever is lost or damaged. [22] Root apical meristem and tissue patterns become established in the embryo in the case of the primary root, and in the new lateral root primordium in the case of secondary roots.

Intercalary meristem

In angiosperms, intercalary (sometimes called basal) meristems occur in monocot (in particular, grass) stems at the base of nodes and leaf blades. Horsetails and Welwitschia also exhibit intercalary growth. Intercalary meristems are capable of cell division, and they allow for rapid growth and regrowth of many monocots. Intercalary meristems at the nodes of bamboo allow for rapid stem elongation, while those at the base of most grass leaf blades allow damaged leaves to rapidly regrow. This leaf regrowth in grasses evolved in response to damage by grazing herbivores and/or wildfires.

Floral meristem

When plants begin flowering, the shoot apical meristem is transformed into an inflorescence meristem, which goes on to produce the floral meristem, which produces the sepals, petals, stamens, and carpels of the flower.

In contrast to vegetative apical meristems and some efflorescence meristems, floral meristems cannot continue to grow indefinitely. Their growth is limited to the flower with a particular size and form. The transition from shoot meristem to floral meristem requires floral meristem identity genes, that both specify the floral organs and cause the termination of the production of stem cells. AGAMOUS (AG) is a floral homeotic gene required for floral meristem termination and necessary for proper development of the stamens and carpels. [5] AG is necessary to prevent the conversion of floral meristems to inflorescence shoot meristems, but is identity gene LEAFY (LFY) and WUS and is restricted to the centre of the floral meristem or the inner two whorls. [23] This way floral identity and region specificity is achieved. WUS activates AG by binding to a consensus sequence in the AG's second intron and LFY binds to adjacent recognition sites. [23] Once AG is activated it represses expression of WUS leading to the termination of the meristem. [23]

Through the years, scientists have manipulated floral meristems for economic reasons. An example is the mutant tobacco plant "Maryland Mammoth". In 1936, the department of agriculture of Switzerland performed several scientific tests with this plant. "Maryland Mammoth" is peculiar in that it grows much faster than other tobacco plants.

Apical dominance

Apical dominance is where one meristem prevents or inhibits the growth of other meristems. As a result, the plant will have one clearly defined main trunk. For example, in trees, the tip of the main trunk bears the dominant shoot meristem. Therefore, the tip of the trunk grows rapidly and is not shadowed by branches. If the dominant meristem is cut off, one or more branch tips will assume dominance. The branch will start growing faster and the new growth will be vertical. Over the years, the branch may begin to look more and more like an extension of the main trunk. Often several branches will exhibit this behavior after the removal of apical meristem, leading to a bushy growth.

The mechanism of apical dominance is based on auxins, types of plant growth regulators. These are produced in the apical meristem and transported towards the roots in the cambium. If apical dominance is complete, they prevent any branches from forming as long as the apical meristem is active. If the dominance is incomplete, side branches will develop.[ citation needed ]

Recent investigations into apical dominance and the control of branching have revealed a new plant hormone family termed strigolactones. These compounds were previously known to be involved in seed germination and communication with mycorrhizal fungi and are now shown to be involved in inhibition of branching. [24]

Diversity in meristem architectures

The SAM contains a population of stem cells that also produce the lateral meristems while the stem elongates. It turns out that the mechanism of regulation of the stem cell number might be evolutionarily conserved. The CLAVATA gene CLV2 responsible for maintaining the stem cell population in Arabidopsis thaliana is very closely related to the maize gene FASCIATED EAR 2(FEA2) also involved in the same function. [25] Similarly, in rice, the FON1-FON2 system seems to bear a close relationship with the CLV signaling system in Arabidopsis thaliana . [26] These studies suggest that the regulation of stem cell number, identity and differentiation might be an evolutionarily conserved mechanism in monocots, if not in angiosperms. Rice also contains another genetic system distinct from FON1-FON2, that is involved in regulating stem cell number. [26] This example underlines the innovation that goes about in the living world all the time.

Role of the KNOX-family genes

Note the long spur of the above flower. Spurs attract pollinators and confer pollinator specificity. (Flower: Linaria dalmatica) Linaria spur.jpg
Note the long spur of the above flower. Spurs attract pollinators and confer pollinator specificity. (Flower: Linaria dalmatica)
Complex leaves of Cardamine hirsuta result from KNOX gene expression Cardamine hirsuta.jpg
Complex leaves of Cardamine hirsuta result from KNOX gene expression

Genetic screens have identified genes belonging to the KNOX family in this function. These genes essentially maintain the stem cells in an undifferentiated state. The KNOX family has undergone quite a bit of evolutionary diversification while keeping the overall mechanism more or less similar. Members of the KNOX family have been found in plants as diverse as Arabidopsis thaliana, rice, barley and tomato. KNOX-like genes are also present in some algae, mosses, ferns and gymnosperms. Misexpression of these genes leads to the formation of interesting morphological features. For example, among members of Antirrhineae , only the species of the genus Antirrhinum lack a structure called spur in the floral region. A spur is considered an evolutionary innovation because it defines pollinator specificity and attraction. Researchers carried out transposon mutagenesis in Antirrhinum majus, and saw that some insertions led to formation of spurs that were very similar to the other members of Antirrhineae , [27] indicating that the loss of spur in wild Antirrhinum majus populations could probably be an evolutionary innovation.

The KNOX family has also been implicated in leaf shape evolution (See below for a more detailed discussion). One study looked at the pattern of KNOX gene expression in A. thaliana , that has simple leaves and Cardamine hirsuta , a plant having complex leaves. In A. thaliana, the KNOX genes are completely turned off in leaves, but in C.hirsuta, the expression continued, generating complex leaves. [28] Also, it has been proposed that the mechanism of KNOX gene action is conserved across all vascular plants, because there is a tight correlation between KNOX expression and a complex leaf morphology. [29]

Indeterminate growth of meristems

Though each plant grows according to a certain set of rules, each new root and shoot meristem can go on growing for as long as it is alive. In many plants, meristematic growth is potentially indeterminate, making the overall shape of the plant not determinate in advance. This is the primary growth. Primary growth leads to lengthening of the plant body and organ formation. All plant organs arise ultimately from cell divisions in the apical meristems, followed by cell expansion and differentiation. Primary growth gives rise to the apical part of many plants.

The growth of nitrogen-fixing root nodules on legume plants such as soybean and pea is either determinate or indeterminate. Thus, soybean (or bean and Lotus japonicus) produce determinate nodules (spherical), with a branched vascular system surrounding the central infected zone. Often, Rhizobium-infected cells have only small vacuoles. In contrast, nodules on pea, clovers, and Medicago truncatula are indeterminate, to maintain (at least for some time) an active meristem that yields new cells for Rhizobium infection. Thus zones of maturity exist in the nodule. Infected cells usually possess a large vacuole. The plant vascular system is branched and peripheral.

Cloning

Under appropriate conditions, each shoot meristem can develop into a complete, new plant or clone. Such new plants can be grown from shoot cuttings that contain an apical meristem. Root apical meristems are not readily cloned, however. This cloning is called asexual reproduction or vegetative reproduction and is widely practiced in horticulture to mass-produce plants of a desirable genotype. This process known as mericloning, has been shown to reduce or eliminate viruses present in the parent plant in multiple species of plants. [30] [31]

Propagating through cuttings is another form of vegetative propagation that initiates root or shoot production from secondary meristematic cambial cells. This explains why basal 'wounding' of shoot-borne cuttings often aids root formation. [32]

Induced meristems

Meristems may also be induced in the roots of legumes such as soybean, Lotus japonicus , pea, and Medicago truncatula after infection with soil bacteria commonly called Rhizobia.[ citation needed ] Cells of the inner or outer cortex in the so-called "window of nodulation" just behind the developing root tip are induced to divide. The critical signal substance is the lipo-oligosaccharide Nod factor, decorated with side groups to allow specificity of interaction. The Nod factor receptor proteins NFR1 and NFR5 were cloned from several legumes including Lotus japonicus, Medicago truncatula and soybean (Glycine max). Regulation of nodule meristems utilizes long-distance regulation known as the autoregulation of nodulation (AON). This process involves a leaf-vascular tissue located LRR receptor kinases (LjHAR1, GmNARK and MtSUNN), CLE peptide signalling, and KAPP interaction, similar to that seen in the CLV1,2,3 system. LjKLAVIER also exhibits a nodule regulation phenotype though it is not yet known how this relates to the other AON receptor kinases.

Lateral Meristems

Lateral meristems, the form of secondary plant growth, add growth to the plants in their diameter. This is primarily observed in perennial dicots that survive from year to year. There are two types of lateral meristems: vascular cambium and cork cambium.

In vascular cambium, the primary phloem and xylem are produced by the apical meristem. After this initial development, secondary phloem and xylem are produced by the lateral meristem. The two are connected through a thin layer of parenchymal cells which are differentiated into the fascicular cambium. The fascicular cambium divides to create the new secondary phloem and xylem. Following this the cortical parenchyma between vascular cylinders differentiates interfascicular cambium. This process repeats for indeterminate growth. [33]

Cork cambium creates a protective covering around the outside of a plant. This occurs after the secondary xylem and phloem has expanded already. Cortical parenchymal cells differentiate into cork cambium near the epidermis which lays down new cells called phelloderm and cork cells. These cork cells are impermeable to water and gases because of a substance called suberin that coats them. [34]

See also

Related Research Articles

Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem cells in the adult organism.

<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">Vascular cambium</span> Main growth tissue in the stems, roots of plants

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.

<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">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.

Florigens are proteins capable of inducing flowering time in angiosperms. The prototypical florigen is encoded by the FT gene and its orthologs in Arabidopsis and other plants. Florigens are produced in the leaves, and act in the shoot apical meristem of buds and growing tips.

<span class="mw-page-title-main">ABC model of flower development</span> Model for genetics of flower development

The ABC model of flower development is a scientific model of the process by which flowering plants produce a pattern of gene expression in meristems that leads to the appearance of an organ oriented towards sexual reproduction, a flower. There are three physiological developments that must occur in order for this to take place: firstly, the plant must pass from sexual immaturity into a sexually mature state ; secondly, the transformation of the apical meristem's function from a vegetative meristem into a floral meristem or inflorescence; and finally the growth of the flower's individual organs. The latter phase has been modelled using the ABC model, which aims to describe the biological basis of the process from the perspective of molecular and developmental genetics.

<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.

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.

Peptide signaling plays a significant role in various aspects of plant growth and development and specific receptors for various peptides have been identified as being membrane-localized receptor kinases, the largest family of receptor-like molecules in plants. Signaling peptides include members of the following protein families.

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.

The epigenetics of plant growth and development refers to the heritable changes in gene expression that occur without alterations to the DNA sequence, influencing processes in plants such as seed germination, flowering, and stress responses through mechanisms like DNA methylation, histone modification, and chromatin remodeling.

CLE peptides are a group of peptides found in plants that are involved with cell signaling. Production is controlled by the CLE genes. Upon binding to a CLE peptide receptor in another cell, a chain reaction of events occurs, which can lead to various physiological and developmental processes. This signaling pathway is conserved in diverse land plants.

A cytokinin signaling and response regulator protein is a plant protein that is involved in a two step cytokinin signaling and response regulation pathway.

References

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