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Meristems and Differentiation
• Embryonic tissue regions, primarily concerned
with formation of new cells, are the meristems. • Greek merismos, division cell division activity as a characteristic of a meristematic tissue • Those cells that maintain the meristem as a continuing source of new cells are referred to as the initiating cells, or meristematic initials, or simply initials. • Their products, which after a variable number of cell divisions give rise to the body cells, are the derivatives of the initials. • So, a dividing cell is a precursor of derivatives. • A given initial in a meristem is a precursor of two derivatives, one of which is a new initial and the other a precursor of body cells. • Initials are not inherently different from their derivatives. The apical meristems of shoot and root certain cells in the meristems act as initials mainly because they occupy the proper position for such activity. • Initials are totipotent (from the Latin totus, entire); they have the capacity to produce the entire spectrum of cell types, even to develop into complete plants. the development and organization of a plant may be characterized as having plasticity Classification of Meristems • A Common Classification of Meristems Is Based on Their Position in the Plant Body • There are apical meristems, that is, meristems located at the apices of main and lateral shoots and roots, and lateral meristems, that is, meristems arranged parallel with the sides of the axis, usually that of stem and root. The vascular and cork cambia are lateral meristems. • The third term based on the position of a meristem is intercalary meristem. This term refers to meristematic tissue derived from the apical meristem and continuing meristematic activity some distance from that meristem. • The word intercalary implies that the meristem is inserted (intercalated) between tissues that are no longer meristematic. The best known examples of intercalary meristems are those in internodes and leaf sheaths of monocots, particularly grasses. • The initiating cells and their most recent derivatives are frequently distinguished from the partly differentiated but still meristematic subjacent tissues under the name of promeristem (or protomeristem The subjacent meristematic tissues are classified according to the tissue systems that are derived from them, namely into protoderm, which differentiates into the epidermis; procambium (also called provascular tissue), which gives rise to the primary vascular tissues; and the ground meristem, precursor of the fundamental or ground tissue system primary meristems • The terms protoderm, procambium, and ground meristem correlated with the equally simple and convenient classification of mature tissues into the three tissue systems, epidermal, vascular, and fundamental Meristems Are Also Classified According to the Nature of Cells That Give Origin to Their Initial Cells • If the initials are direct descendants of embryonic cells that never ceased to be concerned with meristematic activity, the resulting meristem is called primary. If, however, the initials originate from cells that had differentiated, then resumed meristematic Characteristics of Meristematic Cells • Meristematic cells are fundamentally similar to young parenchyma cells. During cell division the cells at the shoot apices are relatively thin- walled, rather poor in storage materials, and their plastids are in proplastid stages. The endoplasmic reticulum is small in amount and mitochondria have few cristae. Golgi bodies and microtubules are present as is characteristic of cells with growing cell walls. The vacuoles are small and dispersed. Deeper layers of apical meristems may be more highly vacuolated and contain starch • During periods of cell division, the vascular cambium cells are highly vacuolated, with one or two large vacuoles limiting the dense cytoplasm to a thin parietal layer , which contains rough endoplasmic reticulum and other cell components. ratio of cell size to nuclear size—the cytonuclear ratio—varies considerably The nuclei show characteristic structural variations during changes in mitotic activitymeristematic cells vary in size, shape, and cytoplasmic features. In recognition of this variability, the term eumeristem (true meristem) has been suggestsed for designating a meristem composed of small, approximately isodiametric cells with thin walls and rich in cytoplasm Growth Patterns in Meristems • Meristems and meristematic tissues show varied arrangements of cells resulting from differences in patterns of cell division and enlargement. Apical meristems having only one initiating cell occupying the very apex (Equisetum and many ferns) have an orderly distribution of the recently formed, still meristematic cells In seed plants the pattern of cell division appears less precise. It is not random, however, for an apical meristem grows as an organized whole and the divisions and enlargement of individual cells are related to the internal distribution of growth and to the external form of the apex. These correlative influences bring about a differentiation of distinctive zones in the meristems. In some parts of the meristem, the cells may divide sluggishly and attain considerable dimensions; in others they may divide frequently and remain small. • Some cell complexes divide in various planes (volume growth), and others only by walls at right angles to the surface of the meristem (anticlinal divisions, surface growth). The lateral meristems are particularly distinguished by divisions parallel with the nearest surface of the organ (periclinal divisions), which result in establishment of rows of cells parallel with the radii of the axes (radial seriation or alignment) and an increase in thickness of the organ. In cylindrical bodies, such as stems and roots, the term tangential division (or tangential longitudinal) is commonly used instead of periclinal division. If the anticlinal division occurs parallel with the radius of the cylinder, it is referred to as a radial anticlinal (or radial longitudinal) division . If the new wall is laid down at right angles to the longitudinal axis of the cylinder, the anticlinal division is transverse. • Organs arising at the same apical meristem may assume varied forms because the still- meristematic derivatives of the apical meristem (primary meristems) often exhibit distinct patterns of growth. Some of these patterns are so characteristic that the meristematic tissues showing them have received special names. These are mass meristem (or block meristem), rib meristem (or file meristem), and plate meristem. • The mass meristem grows by divisions in all planes and produces bodies that are isodiametric or spheroidal or have no definite shape. Such growth occurs, for example, during the formation of spores, sperms (in seedless vascular plants), and endosperm. Divisions in various planes are associated with the spheroidal shape of many angiosperm embryos at a certain stage of development. • The rib meristem gives rise to a complex of parallel longitudinal fi les (“ribs”) of cells by divisions at right angles to the longitudinal axis of the cell row. This pattern of growth occurs in elongating cylindrical plant parts as illustrated by the cortex of roots and pith and cortex of stems. • The plate meristem shows chiefly anticlinal divisions so that the number of layers originally established in the young organ does not increase any further, and a plate-like structure is produced. The fl at blades of angiosperm leaves exemplify the result of growth by a plate meristem (Fig. 5.5). The plate meristem and the rib meristem are growth forms that occur mainly in the ground meristem. They are associated with the two basic forms of the plant body, the thin spreading lamina (blade) of the leaf-like organs and the elongated cylindrical plant parts such as root, stem, and petiole. Meristematic Activity and Plant Growth • Meristems and their meristematic derivatives are concerned with growth in the broad sense of the term, meaning irreversible increase in size, including volume and surface. • In multicellular plants growth is based on two processes, cell division and cell enlargement. The recent derivatives of meristematic initials produce other derivatives by cell division, and the successive generations of derivatives increase in size. • Cell enlargement becomes dominant over cell division and, in time, replaces it entirely. • When the derivatives farthest from the meristem initials cease to divide and to expand, they acquire the characteristic specific for the tissues in which they are located; that is, the cells differentiate and eventually mature. Although cell division as such does not contribute to the volume of the growing entity • Cell division and cell expansion are different stages of the growth process, in which cell enlargement determines the final size of the plant and its parts. • In ontogeny (development of an individual entity), the enlargement of a cell serves as the transition between the stages of division and maturation. Cell division rarely occurs without being accompanied by cell enlargement. • The primary growth of the plant, initiated in the apical meristems, expands the plant body, determines its height, and through increase in its surface, the area of contact with air and soil. Eventually primary growth also gives rise to the reproductive organs. • The secondary growth resulting from the activity of the cambia increases the volume of conducting tissues and forms supportive and protective tissues. Usually not all of the apical meristems present on a plant are active simultaneously. Suppression of lateral bud growth while the terminal shoot is actively growing (apical dominance). • Then the activity of cambia varies in intensity, may show seasonal fluctuations in meristematic activity. DIFFERENTIATION
• Differentiation refers to the succession of
changes in form, structure, and function of progenies of meristematic derivatives, and their organization into tissues and organs. One may speak of differentiation of a single cell, a tissue (histogenesis), an organ (organogenesis), and the plant as a whole. • Differentiation also refers to processes by which a fertilized egg cell produces a progeny of cells showing increasing heterogeneity, specialization, and patterned organization. • The degree of differentiation and specialization (structural adaptation to a particular function) vary greatly. • Some cells diverge relatively little from their meristematic precursors and retain the power to divide (various parenchyma cells). • Others are more profoundly modified and lose most, or all, of their meristematic potentialities (sieve elements, laticifers, tracheary elements, various sclereids). Thus cells differentiating in a multicellular body become different from their meristematic precursors, as well as from cells in other tissues of the same plant. Mature cells with complete protoplasts may resume meristematic activity when properly stimulated. The stimulation can be induced by accidental wounding, invasion by parasites, and infection with agents of diseases on resumption of meristematic activity by nonmeristematic cells the terms dedifferentiation— loss of previously acquired characteristics—and redifferentiation— acquisition of new characteristics—are often used. The entire process is referred to as transdifferentiation. • During its development, a plant assumes a specific form. Thus the plant undergoes morphogenesis (from Greek words for shape and origin). The term commonly is used with reference to both external form and internal organization, and just like differentiation, morphogenesis is revealed at all levels of organization, from cell components to whole plants. Senescence (Programmed Cell Death) • The natural termination of life of a plant as a result of senescence may be regarded as a normal stage in plant development, a sequel to the events of differentiation and maturation. • The term senescence specifically refers to the series of changes in a living organism that lead to its death. • Senescence may affect the whole organism or some of its organs, tissues, or cells. Annual plants that bloom only once in their lifetime (monocarpy: fruiting once only) senesce within one season. • In deciduous trees the leaves commonly senesce at the end of seasonal growth. Since senescence occurs in orderly sequences in the life of the plant and is an active degenerative process, it is considered to be genetically controlled, or programmed—a process of programmed cell death. Cellular Changes in Differentiation • During differentiation, histologic diversity results from changes in the characteristics of individual cells and from alterations in intercellular relationships. The common features of more or less differentiated cells, including the structure and function of individual cell components, • The changes in cell wall structure during cell differentiation • Differential increase in thickness of primary and secondary cell walls, changes in cell wall texture and chemistry, and development of special sculptural patterns introduce differences among cells. • Endopolyploidy refers to a condition that arises from a DNA replication cycle within the nuclear envelope and without spindle formation. Hence the newly formed strands of DNA remain in the same nucleus, and the nucleus becomes polyploid. This type of DNA replication cycle is called an endocycle One of the Early Visible Changes in Differentiating Tissues Is the Unequal Increase in Cell Size • The increase in cell size may be relatively uniform along all diameters, but frequently the cell enlarges more in one direction than in others. Such cells may become strikingly different in shape from their meristematic precursors. Intercellular Adjustment in Differentiating Tissue Involves Coordinated and Intrusive Growth • Enlargement and change in shape of cells in a differentiating tissue are accompanied by more or less profound changes in the spatial relations between cells. • With regard to the growth of cell walls during tissue differentiation, two kinds of intercellular adjustments are visualized: (1) contiguous growing wall layers belonging to two neighboring cells do not separate but expand jointly; (2) contiguous wall layers do separate and the growing cells intrude into the resulting spaces. The fi rst method of growth, originally named symplastic Growth The intercellular adjustment that involves an intrusion of cells among others is called intrusive growth or interpositional growthThe elongating cells grow at their apices (apical intrusive growth), usually at both ends. CAUSAL FACTORS IN DIFFERENTIATION • Use of growth-regulating chemicals, surgical procedures, exposure to radiation, confinement to controlled temperatures and light, interference with normal gravity effects, and growth under selected day length conditions. • Polarity refers to the orientation of activities in space. It is an essential component of biological pattern formation. • Polarity manifests itself early in the life of the plant. It is displayed in the egg cell, in which the nucleus is located in the chalazal end and a large vacuole in the micropylar end, and in the bipolar development of the embryo from the zygote. It is later expressed in the organization of the plant into root and shoot and is also evident in various phenomena at the cellular level. • Polarity is related to the phenomenon of gradients, since the differences between the two poles of the plant axis appear in graded series. There are physiological gradients, for example, those expressed in the rates of metabolic processes, in concentration of auxins, and in concentration of sugar in the conducting system; there are also gradients in anatomical differentiation and in the development of the external features Plant Cells Differentiate According to Position • PLANT HORMONES from the Greek horman, meaning to set in motion Tissues may require different amounts of hormones. Such differences are referred to as differences in sensitivity
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