Meristem

Apical meristems, also known as the primary meristem, give rise to the primary plant body and are responsible for primary growth, or an increase in length or height. Apical meristems may differentiate into three kinds of primary meristem: • Protoderm: lies around the outside of the stem and develops into the epidermis. • Procambium: lies just inside of the protoderm and develops into primary xylem and primary phloem. It also produces the vascular cambium, and cork cambium (part of the secondary meristems but descendants of apical meristematic cells). The cork cambium further differentiates into the phelloderm, or bark, (to the inside) and the phellem, or cork (to the outside). All three of these layers (cork cambium, phellem, and phelloderm) constitute the periderm. In roots, the procambium can also give rise to the pericycle, which produces lateral roots in eudicots. • Ground meristem: Composed of ground tissue parenchyma, collenchyma and sclerenchyma cells that develop into the cortex and the pith. ==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. • Vascular cambium, which produces secondary xylem and secondary phloem. This is a process that may continue throughout the life of the plant. This is what gives rise to wood in plants. Such plants are called arboraceous. This does not occur in plants that do not go through secondary growth, known as herbaceous plants. • Cork cambium, which gives rise to the periderm, which replaces the epidermis with bark and cork for example. ==Apical meristems== Apical meristems are the completely undifferentiated (indeterminate) meristems of a plant. They give rise to primary growth, enabling the elongation of shoots and roots. Apical meristems give rise to three types of primary meristems, which later develop into secondary or lateral meristems, contributing to the plant's lateral expansion. 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. Shoot Apical Meristems '' (left). Fourteen days later, leaves have developed (right). of a shoot apical meristem surrounded by leaf primordia of Arabidopsis thaliana. in Lycopodium clavatum (bar = 100 μm). 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. 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: • Stem cells • The immediate daughter cells of the stem cells • A subjacent organizing center • Founder cells for organ initiation in surrounding regions 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. 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. WUS is expressed in the cells below the stem cells of the meristem and its presence prevents the differentiation of the stem cells. 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. Root apical meristem 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. 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. 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. 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. 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. Similarly, in rice, the FON1-FON2 system seems to bear a close relationship with the CLV signaling system in Arabidopsis thaliana. 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. 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. 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. ==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. 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. == 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. 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. 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. ==See also==