Contents 1 Apical meristems 1.1 Shoot apical meristems 1.2 Root apical meristem 1.3 Intercalary meristem 1.4 Floral meristem 1.5 Apical dominance 1.6 Diversity in meristem architectures 1.7 Role of the KNOX-family genes 2 Primary meristems 3 Secondary meristems 4 Indeterminate growth of meristems 5 Cloning 6 See also 7 References 8 Footnotes

Apical meristems[edit] Organisation of an apical meristem (growing tip) 1 - Central zone 2 - Peripheral zone 3 - Medullary (i.e. central) meristem 4 - Medullary tissue 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 determine the physical characteristics of the leaf edge and margin. In dicots, layer two of the corpus determine 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[edit] Shoot apical meristems of Crassula ovata (left). Fourteen days later, leaves have developed (right). The source of all above-ground organs. 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 4 distinct cell groups: Stem cells The immediate daughter cells of the stem cells A subjacent organising centre Founder cells for organ initiation in surrounding regions The four distinct zones mentioned above 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.[2] CLV1 and CLV2 are predicted to form a receptor complex (of the LRR receptor-like kinase family) to which CLV3 is a ligand.[3][4][5] CLV3 shares some homology with the ESR proteins of maize, with a short 14 amino acid region being conserved between the proteins.[6][7] Proteins that contain these conserved regions have been grouped into the CLE family of proteins.[6][7] 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.[2] 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.[8] KAPP is a kinase-associated protein phosphatase that has been shown to interact with CLV1.[9] KAPP is thought to act as a negative regulator of CLV1 by dephosphorylating it.[9] Another important gene in plant meristem maintenance is WUSCHEL (shortened to WUS), which is a target of CLV signalling.[10] WUS is expressed in the cells below the stem cells of the meristem and its presence prevents the differentiation of the stem cells.[10] CLV1 acts to promote cellular differentiation by repressing WUS activity outside of the central zone containing the stem cells.[10] SHOOT MERISTEMLESS (STM) also acts to prevent the differentiation of stem cells by repressing the expression of MYB genes that are involved in cellular differentiation.[2] Root apical meristem[edit] 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 produce most of the cells in an adult root.[11][12] 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.[13] 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[edit] In angiosperms, intercalary meristems occur only in monocot (in particular, grass) stems at the base of nodes and leaf blades. Horsetails 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. Floral meristem[edit] Further information: ABC model of flower development When plants begin the developmental process known as 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 exflorescence meristems, floral meristems cannot continue to grow indefinitely. Their future 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.[2] AG is necessary to prevent the conversion of floral meristems to inflorescence shoot meristems, but is not involved in the transition from shoot to floral meristem.[14] AG is turned on by the floral meristem identity gene LEAFY (LFY) and WUS and is restricted to the centre of the floral meristem or the inner two whorls.[15] 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.[15] Once AG is activated it represses expression of WUS leading to the termination of the meristem.[15] 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[edit] Apical dominance is phenomenon 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 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 behaviour after the removal of apical meristem, leading to a bushy growth. The mechanism of apical dominance is based on the plant hormone auxin. It is produced in the apical meristem and transported towards the roots in the cambium. If apical dominance is complete, it prevents 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 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.[16] Diversity in meristem architectures[edit] Is the mechanism of being indeterminate conserved in the SAMs of the plant world? 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 indeed 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.[17] Similarly, in Rice, the FON1-FON2 system seems to bear a close relationship with the CLV signaling system in Arabidopsis thaliana.[18] 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.[18] This example underlines the innovation that goes about in the living world all the time. Role of the KNOX-family genes[edit] Note the long spur of the above flower. Spurs attract pollinators and confer pollinator specificity. (Flower:Linaria dalmatica) Complex leaves of C. hirsuta are a result of 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 formation of interesting morphological features. For example, among members of Antirrhinae, only the species of 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 Antirrhinae,[19] 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.[20] 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.[21]

Primary meristems[edit] 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, secondary meristems. The cork cambium further differentiates into the phelloderm (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.[22] Ground meristem: develops into the cortex and the pith. Composed of parenchyma, collenchyma and sclerenchyma cells.[22] These meristems are responsible for primary growth, or an increase in length or height, which were discovered by scientist Joseph D. Carr of North Carolina in 1943.[citation needed]

Secondary meristems[edit] There are two types of secondary meristems, these are also called the lateral meristems because they surround the established stem of a plant and cause it to grow laterally (i.e., larger in diameter). 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 arborescent. 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.

Indeterminate growth of meristems[edit] 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 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[edit] 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 is also known as mericloning. 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.[23]

See also[edit] Secondary growth Stem cell Thallus Tissues

References[edit] ^ Galun, Esra (2007). Plant Patterning: Structural and Molecular Genetic Aspects. World Scientific Publishing Company. p. 333. ISBN 9789812704085 ^ a b c d Fletcher, J. C. (2002). "Shoot and Floral Meristem Maintenance in Arabidopsis". Annu. Rev. Plant Biol. 53: 45–66.  ^ Clark SE, Williams RW, Meyerowitz E (1997). "The CLAVATA1 gene encodes a putative receptor kinase that controls shoot and floral meristem size in Arabidopsis". Cell. 89 (4): 575–85. doi:10.1016/S0092-8674(00)80239-1. PMID 9160749.  ^ Jeong S, Trotochaud AE, Clark S (1999). "The Arabidopsis CLAVATA2 gene encodes a receptor-like protein required for the stability of the CLAVATA1 receptor-like kinase". Plant Cell. 11 (10): 1925–33. doi:10.1105/tpc.11.10.1925.  ^ Fletcher JC, Brand U, Running MP, Simon R, Meyerowitz EM (1999). "Signaling of cell fate decisions by CLAVATA3 in Arabidopsis shoot meristems". Science. 283 (5409): 1911–14. doi:10.1126/science.283.5409.1911.  ^ a b J. Mark Cock; Sheila McCormick (July 2001). "A Large Family of Genes That Share Homology with CLAVATA3". Plant Physiology. 126: 939–942. doi:10.1104/pp.126.3.939. PMC 1540125 . PMID 11457943.  ^ a b Karsten Oelkers, Nicolas Goffard, Georg F Weiller, Peter M Gresshoff, Ulrike Mathesius and Tancred Frickey (3 January 2008). "Bioinformatic Analysis of the CLE signalling peptide family". BMC Plant Biology. 8: 1. doi:10.1186/1471-2229-8-1. PMC 2254619 . PMID 18171480. CS1 maint: Multiple names: authors list (link) ^ Valster, A. H.; et al. (2000). "Plant GTPases: the Rhos in bloom". Trends in Cell Biology. 10 (4): 141–146. doi:10.1016/s0962-8924(00)01728-1.  ^ a b Stone, J. M.; et al. (1998). "Control of meristem development by CLAVATA1 receptor kinase and kinase-associated protein phosphatase interactions". Plant Physiology. 117: 1217–1225. doi:10.1104/pp.117.4.1217. PMC 34886 . PMID 9701578.  ^ a b c Mayer, K. F. X; et al. (1998). "Role of WUSCHEL in Regulating Stem Cell Fate in the Arabidopsis Shoot Meristem". Cell. 95: 805–815. doi:10.1016/S0092-8674(00)81703-1. PMID 9865698.  ^ Jose Sebastian and Ji-Young Lee (2013). Root Apical Meristems. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1002/9780470015902.a0020121.pub2] ^ Tom Bennett and Ben Scheres (2010). Root development-two meristems for the price of one? Current Topics in Developmental Biology. doi:10.1016/S0070-2153(10)91003-X ^ Renze Heidstra and Sabrina Sabatini (2014). Plant and animal stem cells: similar yet different. Nature Reviews Molecular Cell Biology 15(5):301-12 doi:10.1038/nrm3790 ^ Mizukami, Y and Ma, H (1997) Determination of Arabidopsis Floral Meristem identity by Agamous The Plant Cell, Vol. 9, 393- 408 ^ a b c Lohmann, J. U. et al. (2001) A Molecular Link between Stem Cell Regulation and Floral Patterning in Arabidopsis Cell 105: 793-803 ^ "Branching out: new class of plant hormones inhibits branch formation". Nature. 455 (7210). 2008-09-11. Retrieved 2009-04-30.  ^ Taguchi-Shiobara; Yuan, Z; Hake, S; Jackson, D; et al. (2001). "The fasciated ear2 gene encodes a leucine-rich repeat receptor-like protein that regulates shoot meristem proliferation in maize". Genes & Development. 15 (20): 2755–2766. doi:10.1101/gad.208501. PMC 312812 . PMID 11641280.  ^ a b Suzaki T.; Toriba, T; Fujimoto, M; Tsutsumi, N; Kitano, H; Hirano, HY (2006). "Conservation and Diversification of Meristem Maintenance Mechanism in Oryza sativa: Function of the FLORAL ORGAN NUMBER2 Gene". Plant and Cell Physiol. 47 (12): 1591–1602. doi:10.1093/pcp/pcl025. PMID 17056620.  ^ Golz J.F.; Keck, Emma J.; Hudson, Andrew (2002). "Spontaneous Mutations in KNOX Genes Give Rise to a Novel Floral Structure in Antirrhinum". Curr. Biol. 12 (7): 515–522. doi:10.1016/S0960-9822(02)00721-2.  ^ Hay and Tsiantis; Tsiantis, M (2006). "The genetic basis for differences in leaf form between Arabidopsis thaliana and its wild relative Cardamine hirsuta". Nat. Genet. 38 (8): 942–947. doi:10.1038/ng1835. PMID 16823378.  ^ Bharathan G, et al. (2002). "Homologies in Leaf Form Inferred from KNOXI Gene Expression During Development". Science. 296 (5574): 1858–1860. doi:10.1126/science.1070343. PMID 12052958.  ^ a b Evert, Ray, and Susan Eichhorn. Raven Biology of Plants. New York: W. H. Freeman and Company, 2013. Print. ^ Mackenzie, K.A.D; Howard, B.H (1986). "The Anatomical Relationship Between Cambial Regeneration and Root Initiation in Wounded Winter Cuttings of the Apple Rootstock M.26". Annals of Botany. 58: 649–661. 

Footnotes[edit] Wikimedia Commons has media related to Méristème. Wikisource has the text of The New Student's Reference Work article Meristem. Plant Anatomy Laboratory from University of Texas; the lab of JD Mauseth. Micrographs of plant cells and tissues, with explanatory text. Schoof, Heiko; Lenhard, M; Haecker, A; Mayer, KF; Jürgens, G; Laux, T (2000). "Arabidopsis shoot meristems is maintained by a regulatory loop between Clavata and Wuschel genes". Cell. 100 (6): 635–644. doi:10.1016/S0092-8674(00)80700-X. PMID 10761929.  Scofield and Murray (2006). The evolving concept of the meristem. Plant Molecular Biology 60:v–vii Research on meristems v t e Botany History of botany Subdisciplines Plant systematics Ethnobotany Paleobotany Plant anatomy Plant ecology Phytogeography Geobotany Flora Phytochemistry Plant pathology Bryology Phycology Floristics Dendrology Plant groups Algae Archaeplastida Bryophyte Non-vascular plants Vascular plants Spermatophytes Pteridophyte Gymnosperm Angiosperm Plant morphology (glossary) Plant cells Cell wall Phragmoplast Plastid Plasmodesma Vacuole Tissues Meristem Vascular tissue Vascular bundle Ground tissue Mesophyll Cork Wood Storage organs Vegetative Root Rhizoid Bulb Rhizome Shoot Stem Leaf Petiole Cataphyll Bud Sessility Reproductive (Flower) Flower development Inflorescence Umbel Raceme Bract Pedicellate Flower Whorl Floral symmetry Floral diagram Floral formula Receptacle Hypanthium (Floral cup) Perianth Tepal Petal Sepal Sporophyll Gynoecium Ovary Ovule Stigma Archegonium Androecium Stamen Staminode Pollen Tapetum Gynandrium Gametophyte Sporophyte Plant embryo Fruit Fruit anatomy Berry Capsule Seed Seed dispersal Endosperm Surface structures Epicuticular wax Plant cuticle Epidermis Stoma Nectary Trichome Prickle Plant physiology Materials Nutrition Photosynthesis Chlorophyll Plant hormone Transpiration Turgor pressure Bulk flow Aleurone Phytomelanin Sugar Sap Starch Cellulose Plant growth and habit Secondary growth Woody plants Herbaceous plants Habit Vines Lianas Shrubs Subshrubs Trees Succulent plants Reproduction Evolution Ecology Alternation of generations Sporangium Spore Microsporangia Microspore Megasporangium Megaspore Pollination Pollinators Pollen tube Double fertilization Germination Evolutionary development Evolutionary history timeline Hardiness zone Plant taxonomy History of plant systematics Herbarium Biological classification Botanical nomenclature Botanical name Correct name Author citation International Code of Nomenclature for algae, fungi, and plants (ICN) - for Cultivated Plants (ICNCP) Taxonomic rank International Association for Plant Taxonomy (IAPT) Plant taxonomy systems Cultivated plant taxonomy Citrus taxonomy cultigen cultivar Group grex Practice Agronomy Floriculture Forestry Horticulture Lists Related topics Botanical terms Botanists by author abbreviation Botanical expedition Category Portal WikiProject Retrieved from "" Categories: Plant anatomyPlant physiologyHidden categories: CS1 maint: Multiple names: authors listAll articles with unsourced statementsArticles with unsourced statements from February 2008

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