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In cell biology, the meristem is a structure composed of specialized tissue found in plants, consisting of , known as meristematic cells, which are undifferentiated cells capable of continuous cell division. These meristematic cells play a fundamental role in plant growth, regeneration, and acclimatization, as they serve as the source of all differentiated plant tissues and Plant organ. They contribute to the formation of structures such as fruits, leaves, and seeds, as well as supportive tissues like stems and roots.
Meristematic cells are totipotent, meaning they have the ability to differentiate into any plant cell type. As they divide, they generate new cells, some of which remain meristematic cells while others differentiate into specialized cells that typically lose the ability to divide or produce new cell types. Due to their active division and undifferentiated nature, meristematic cells form the foundation for the formation of new plant organs and the continuous expansion of the plant body throughout the plant's life cycle.
Meristematic cells are small cells, with thin primary cell walls, and small or no vacuoles. Their protoplasm is dense, filling the entire cell, and they lack intercellular spaces. Instead of mature plastids such as chloroplasts or chromoplasts, they contain proplastids, which later develop into fully functional plastids.
Meristematic tissues are classified into three main types based on their location and function: apical meristems, found at the tips of roots and shoots; intercalary or basal meristems, located in the middle regions of stems or leaves, enabling Plant regrowth; and lateral meristems or cambium, responsible for secondary growth in woody plants. At the summit of the meristem, a small group of slowly dividing cells, known as the central zone, acts as a reservoir of stem cells, essential for maintaining meristem activity. The growth and proliferation rates of cells vary within the meristem, with higher activity at the periphery compared to the central region.
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").Galun, Esra (2007). Plant Patterning: Structural and Molecular Genetic Aspects. World Scientific Publishing Company. p. 333. It is derived , in recognition of its inherent function.
There are two main types of apical meristems: shoot apical meristem (SAM) and root apical meristem (RAM). The SAM is located at the tips of shoots and produces leaves, stems, and flowers, while the RAM is found at the tips of roots and generates new root tissues. Both types consist of rapidly-dividing cells that remain indeterminate, meaning they continuously produce new cells without a predefined final state, similar to stem cells in animals, which have an analogous behavior and function.
Structurally, apical meristems are organized into distinct zones. The central zone serves as a reservoir of undifferentiated cells, while the peripheral zone generates new organs and tissues. The medullary meristem contributes to vascular development, forming the medullary tissue, which makes up the plant's central structure. The meristem layers also vary depending on the plant type. The outermost layer, called the tunica, determines the leaf edge and margin in monocots, whereas in dicots, the second layer of the corpus influences leaf characteristics.
Apical meristems are generally found at the tips of roots and stems, but in some arctic plants, they are located in the lower or middle parts of the plant. This adaptation is believed to provide advantages in extreme environmental conditions.
The shoot apical meristem is the site of most of the embryogenesis in flowering plants. 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. CLV1 and CLV2 are predicted to form a receptor complex (of the LRR receptor-like kinase family) to which CLV3 is a ligand. CLV3 shares some homology with the ESR proteins of maize, with a short 14 amino acid region being conserved between the proteins. Proteins that contain these conserved regions have been grouped into the CLE family of proteins.
CLV1 has been shown to interact with several proteins that are most likely involved in downstream signalling. For example, the CLV complex has been found to be associated with GTPase. 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. KAPP is a kinase-associated protein phosphatase that has been shown to interact with CLV1. KAPP is thought to act as a negative regulator of CLV1 by dephosphorylating it.
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. WUS is expressed in the cells below the stem cells of the meristem and its presence prevents the differentiation of the stem cells. CLV1 acts to promote cellular differentiation by repressing WUS activity outside of the central zone containing the stem cells.
The function of WUS in the shoot apical meristem is linked to the Plant hormone cytokinin. Cytokinin activates which then Phosphorylation histidine phosphotransfer proteins. 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. Therefore, A-ARRs do not contribute to the activation of transcription, and by competing for phosphates from phosphotransfer proteins, inhibit B-ARRs function. In the SAM, B-ARRs induce the expression of WUS which induces stem cell identity. WUS then suppresses A-ARRs. 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.
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 and . 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.Lohmann, J. U. et al. (2001) A Molecular Link between Stem Cell Regulation and Floral Patterning in Arabidopsis Cell 105: 793-803 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. Once AG is activated it represses expression of WUS leading to the termination of the meristem.
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.
The mechanism of apical dominance is based on , types of plant growth regulators. These are produced in the apical meristem and transported towards the roots in the vascular 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.
Recent investigations into apical dominance and the control of branching have revealed a new plant hormone family termed . 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.
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. 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.
The growth of nitrogen-fixing on Legume 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 Vacuole. 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.
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.
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.
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.
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