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D. The Origin of Vascular Plants
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Meristem



A meristem is the tissue in most plants containing undifferentiated cells (meristematic cells), found in zones of the plant where growth can take place.Meristematic cells give rise to various organs of the plant and keep the plant growing. The shoot apical meristem (SAM) gives rise to organs like the leaves and flowers, while the root apical meristem (RAM) provides the meristematic cells for the 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 term meristem was first used in 1858 by Karl Wilhelm von Nägeli (1817–1891) in his book Beiträge zur Wissenschaftlichen Botanik. It is derived from the Greek word merizein (μερίζειν), meaning to divide, in recognition of its inherent function.In general, differentiated plant cells cannot divide or produce cells of a different type. Therefore, cell division in the meristem is required to provide new cells for expansion and differentiation of tissues and initiation of new organs, providing the basic structure of the plant body.Meristematic cells are incompletely or not at all differentiated, and are capable of continued cellular division (youthful). Furthermore, the cells are small and protoplasm fills the cell completely. The vacuoles are extremely small. The cytoplasm does not contain differentiated plastids (chloroplasts or chromoplasts), although they are present in rudimentary form (proplastids). Meristematic cells are packed closely together without intercellular cavities. The cell wall is a very thin primary cell wall.Maintenance of the cells requires a balance between two antagonistic processes: organ initiation and stem cell population renewal.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 because they are involved in lateral growth.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.Meristems also are induced in the roots of legumes such as soybean, Lotus japonicus, pea, and Medicago truncatula after infection with soil bacteria commonly called Rhizobium. 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 commonly called ""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.
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