Comparative anatomy of embryogenesis in three species of Podostemaceae and evolution of the loss of embryonic shoot and root meristems

During embryogenesis in angiosperms, the embryonic shoot and root meristems are created at opposite poles of the embryo, establishing a vertical body plan. However, the aquatic eudicot family Podostemaceae exhibits an unusual horizontal body plan, which is attributed to the loss of embryonic shoot and root meristems. To infer the embryogenetic changes responsible for the loss of these meristems, we examined the embryogenesis of three podostemads with different meristem characters, that is, Terniopsis brevis with distinct shoot and root meristems, Zeylanidium lichenoides with reduced shoot and no root meristems, and Hydrobryum japonicum with no shoot and no root meristems. In T. brevis, as in other eudicots, the putative organizing center (OC) and L1 layer (=the epidermal cell layer) arose to generate a distinct shoot meristem initial, and the hypophysis formed the putative quiescent center (QC) of a root meristem. Z. lichenoides had a morphologically unrecognizable shoot meristem, because a distinct L1 layer did not develop, whereas the putative OC precursor arose normally. In H. japonicum, the vertical divisions of the apical cells of eight-cell embryo prevented putative OC initiation. In Z. lichenoides and H. japonicum, the putative QC failed to initiate because the hypophysis repeated longitudinal divisions during early embryogenesis. Based on their phylogenetic relationships, we infer that the conventional embryonic shoot meristem was lost in Podostemaceae via two steps, that is, the loss of a distinct L1 layer and the loss of the OC, whereas the loss of the embryonic root meristem occurred once by misspecification of the hypophysis.     

The Difference Between Open and Closed Meristems

An open and a closed root meristem have been compared by investigatingthe cell kinetics of small regions of the apices of Helianthusand Zea. The cells of the stelar pole are quiescent in both and thereis no exchange of cells between stele and cortex or stele andcap. The immediately distal cells in the closed meristem (Zea)are also quiescent and the few divisions that do occur can betransverse or longitudinal. In the open meristem (Helianthus)these cells are not quiescent, but they go out of cycle transiently,prolonging the potential cell-doubling time. Their divisionsare transverse. It is a consequence of these differences thatclosed meristems form root caps discrete from the cortex whereasopen meristems force instability in the boundary between theperipheral part of the cap and the cortex. Another consequencein roots with open meristems is a succession of columella complexestransversely displaced from each other by the state of fluxin the meristem during the non-cycling phase of the proximaltier of cells, those immediately distal to the stelar pole. The results are discussed in relation to the ontogenetic onsetof quiescence and the evidence for switches between open andclosed operation of meristems. meristem, root apex, Helianthus annuus, Zea mays L.     

Apical meristem exhaustion during determinate primary root growth in the moots koom 1 mutant of Arabidopsis thaliana

An indeterminate developmental program allows plant organs to grow continuously by maintaining functional meristems over time. The molecular mechanisms involved in the maintenance of the root apical meristem are not completely understood. We have identified a new Arabidopsis thaliana mutant named moots koom 1 (mko1) that showed complete root apical meristem exhaustion of the primary root by 9?days post-germination. MKO1 is essential for maintenance of root cell proliferation. In the mutant, cell division is uncoupled from cell growth in the region corresponding to the root apical meristem. We established the sequence of cellular events that lead to meristem exhaustion in this mutant. Interestingly, the SCR and WOX5 promoters were active in the mko1 quiescent center at all developmental stages. However, during meristem exhaustion, the mutant root tip showed defects in starch accumulation in the columella and changes in auxin response pattern. Therefore, contrary to many described mutants, the determinate growth in mko1 seedlings does not appear to be a consequence of incorrect establishment or affected maintenance of the quiescent center but rather of cell proliferation defects both in stem cell niche and in the rest of the apical meristem. Our results support a model whereby the MKO1 gene plays an important role in the maintenance of the root apical meristem proliferative capacity and indeterminate root growth, which apparently acts independently of the SCR/SHR and WOX5 regulatory pathways.     

The Arabidopsis OBERON1 and OBERON2 genes encode plant homeodomain finger proteins and are required for apical meristem maintenance

Maintenance of the stem cell population located at the apical meristems is essential for repetitive organ initiation during the development of higher plants. Here, we have characterized the roles of OBERON1 (OBE1) and its paralog OBERON2 (OBE2), which encode plant homeodomain finger proteins, in the maintenance and/or establishment of the meristems in Arabidopsis. Although the obe1 and obe2 single mutants were indistinguishable from wild-type plants, the obe1 obe2 double mutant displayed premature termination of the shoot meristem, suggesting that OBE1 and OBE2 function redundantly. Further analyses revealed that OBE1 and OBE2 allow the plant cells to acquire meristematic activity via the WUSCHEL-CLAVATA pathway, which is required for the maintenance of the stem cell population, and they function parallel to the SHOOT MERISTEMLESS gene, which is required for preventing cell differentiation in the shoot meristem. In addition, obe1 obe2 mutants failed to establish the root apical meristem, lacking both the initial cells and the quiescent center. In situ hybridization revealed that expression of PLETHORA and SCARECROW, which are required for stem cell specification and maintenance in the root meristem, was lost from obe1 obe2 mutant embryos. Taken together, these data suggest that the OBE1 and OBE2 genes are functionally redundant and crucial for the maintenance and/or establishment of both the shoot and root meristems.     

Activation of cell division in the quiescent center of excised maize root tip

The phenomenon of activation of cell proliferation in the quiescent center of excised maize roots is described. The root tips were grown on wet filter paper in Petri dishes. This phenomenon was observed in 8 to 14 maize cultivars and was absent in excised Arabidopsis root tips. The distribution of mitoses in meristems greatly varied in roots of individual seedlings from the same seed lot and seedlings of different cultivars. Meristem opening was observed after the removal of small root tips not longer than 3 mm and intact seminal roots. Sucrose (2%) and 10(-6)-10(-8) M indole-3-acetic acid did not prevent meristem opening. These findings indicate that the state of quiescent center is maintained by a system of intercellular and interorgan relations, which are to be clarified.     

ABA promotes quiescence of the quiescent centre and suppresses stem cell differentiation in the Arabidopsis primary root meristem

It is well known that abscisic acid (ABA) can halt meristems for long periods without loss of meristem function, and can also promote root growth at low concentrations, but the mechanisms underlying such regulation are largely unknown. Here we show that ABA promotes stem cell maintenance in Arabidopsis root meristems by both promoting the quiescence of the quiescent centre (QC) and suppressing the differentiation of stem cells and their daughters. We demonstrate that these two mechanisms of regulation by ABA involve distinct pathways, and identify components in each pathway. Our findings demonstrate a cellular mechanism for a positive role for ABA in promoting root meristem maintenance and root growth in Arabidopsis.     

Diversity of cell lengths in terminal portions of roots: location of the proliferative cell population

Terminal meristems are responsible for all primary growth of roots. It has been asserted that all cells of root meristems are actively dividing and that the stem cell (proliferative) population expands exponentially. Lengths of cells in roots just proximal to the root cap/root initial boundary were used to determine the numbers of cortex and stele cells in the meristem. Meristem cells were defined as cells that did not have significantly different cell lengths from initial cells at the boundary. Data show that, for five of the six species (Allium cepa, Pisum sativum, Pyrus communis, Triticum aestivum, Vicia faba, and Zea mays) tested, only the first 15 stele and the first 10-35 cortex cells in median longitudinal sections would be in the meristem. For T. aestivum, no discrete meristem was found because all cells proximal to initial cells were longer than initial cells. In addition to this subject area, distributions of lengths of cells in the root meristem using this definition, for the six species were compared with a theoretical cell-age distribution for exponentially dividing cells, to determine if distributions of cell lengths were similar to a theoretical distribution of exponentially dividing cells. For all species tested, distributions of cell lengths were not similar to a theoretical cell-age distribution. From the data of this study with six plant species, we conclude that either contiguous proliferative cell populations of root meristems are very small or the proliferative cell population is not continuous. In addition, such populations do not resemble a theoretical exponential cell-age distribution. Moreover, it seems that the proliferative capacities of cells within terminal root segments differ markedly among species and are not easily characterized.     

A developmental study of the epidermis of young roots ofZea mays L.

Summary In the apical meristems of main and young lateral roots of corn the uniseriate epidermis is clearly continuous with the most distal cell tier of the quiescent centre. These cells are characterized by the presence on their outer periclinal walls of material which forms the thin root cap junction layer over the apical pole and which thickens appreciably over the flanks of the meristem to form a distinctive extracellular deposit on the young epidermal cells. This material is polysaccharide in nature as indicated by strong periodic acid Schiff's positivity but its autofluorescence also suggests the presence of phenolic compounds.During their development the epidermal cells undergo marked shape change from periclinally flattened, polygonal at the root pole, through columnar on the meristem flank to tabular in the root hair zone. The mucigel thins markedly as cells become tabular but initiation of a root hair is characterized by deposition of polysaccharide on the inside of the periclinal wall where the hair will develop.     

Stem cells in the root and the problem of stem cells in plants

Plant cells are capable of reversible transition from the proliferating to the stem state. This transition is determined by a system of cell-cell interactions and interelationships between plant parts. Stem cells defined as the cells preserving the capacity to divisions and differentiation for a long time arise repeatedly during development of the root and shoot primordial, rather than are clones of a population of stem cells laid down at a certain stage of embryogenesis. The quiescent center cells, rather than the surrounding actively dividing cells, best correspond to the characteristics of stem cells according to Loeffler and Potten. The factors that determine the quiescent center formation and maintenance in the root have been analyzed. The available data suggest that among these factors, indoleacetic acid transport and cap influence are of paramount significance. The cap formation precedes the quiescent center formation both during the root development and in the course of meristem regeneration after the root decapitation. The capacity of stem cell formation by the meristem suggests that not only meristem arises from the stem cells, but also that stem cells are formed from actively dividing cells. Repeated formation of stem cells allows long-term preservation of the capacity of plants for open morphogenesis and vegetative propagation.     

Tolerance of dividing cells to replication stress in UVB-irradiated Arabidopsis roots: requirements for DNA translesion polymerases eta and zeta

In tissues of multicellular organisms, DNA lesions that block replication can disrupt division of the transiently amplifying (TA) cells and stem cells that drive growth. To study how tissue growth is maintained despite DNA damage, stem cells and other cell types must be clearly identifiable. In plants, root growth depends directly on cell divisions in the root meristem. In Arabidopsis thaliana, cell identities in root meristems are unambiguously defined by position relative to the quiescent center and are readily visualized by microscopy. We evaluated roles of two DNA translesion polymerases, AtPoleta (Eta) and AtPolzeta (Zeta), in resistance of dividing root cells to a model genotoxin, UVB-radiation. The major UV photoproducts in DNA, cyclobutane pyrimidine dimers (CPDs), were induced to roughly 0.03CPD/kb by a threshold dose (0.28 kJ m(-2)) that minimally affected wild-type roots. In roots lacking AtPoleta and/or AtPolzeta, this dose inhibited cell division and tissue growth and specifically killed stem cells; severities of all three phenotypes increased in the order eta-相似文献   

Stem cells in the root and the problem of stem cells in plants

Plant cells are capable of reversible transition from the proliferating to the stem state. This transition is determined by a system of cell-cell interactions and interrelationships between plant parts. Stem cells defined as the cells preserving the capacity to divisions and differentiation for a long time arise repeatedly during development of the root and shoot primordial, rather than are clones of a population of stem cells laid down at a certain stage of embryogenesis. The quiescent center cells, rather than the surrounding actively dividing cells, best correspond to the characteristics of stem cells according to Loeffler and Potten. The factors that determine the quiescent center formation and maintenance in the root have been analyzed. The available data suggest that among these factors, indoleacetic acid transport and cap influence are of paramount significance. The cap formation precedes the quiescent center formation both during the root development and in the course of meristem regeneration after the root decapitation. The capacity of tem cell formation by the meristem suggests that not only meristem arises from the stem cells, but also that stem cells are formed from actively dividing cells. Repeated formation of stem cells allows long-term preservation of the capacity of plants for open morphogenesis and vegetative propagation.     

Cell division activation in the quiescent center of excised maize root tip

The phenomenon of activating cell proliferation in the quiescent center of excised maize roots is described. The root tips were grown on wet filter paper in Petri dishes. This phenomenon was observed in 8 to 14 maize cultivars and was absent in excised Arabidopsis root tips. The distribution of mitoses in meristems greatly varied in individual seedlings roots from the same seed lot and seedlings of different cultivars. Meristem opening was observed after the removal of small root tips not longer than 3 mm and intact seminal roots. Sucrose (2%) and 10⁻⁶–10⁻⁸ M indole-3-acetic acid did not prevent meristem opening. These findings indicate that the state of quiescent center is maintained by a system of intercellular and interorgan relations, which are to be clarified.     

Nutrient Sensing in Plant Meristems

Plants need nutrient to grow and plant cells need nutrient to divide. The meristems are the factories and cells that are left behind will expand and differentiate. However, meristems are not simple homogenous entities; cells in different parts of the meristem do different things. Positional cues operate that can fate cells into different tissue domains. However, founder/stem cells persist in specific locations within the meristem e.g. the quiescent centre of root apical meristem (RAM) and the lower half of the central zone of the shoot apical meristem (SAM). Given the complexity of meristems, do their cells simply respond to a diffusing gradient of photosynthate? This in turn begs the question, why do stem cell populations tend to have longer cell cycles than their immediate descendants given that like all other cells they are directly in the path of diffusing nutrient? In this review, we have examined the extent to which nutrient sensing might be operating in meristems. The scene is set for sugar sensing, the plant cell cycle, SAMs and RAMs. Special emphasis is given to the metabolic regulator, SnRK1 (SNF1-related protein kinase 1), hexokinase and the trehalose pathway in relation to sugar sensing. The unique plant cell cycle gene, cylin-dependent kinase B1;1 may have evolved to be particularly responsive to sugar signalling pathways. Also, the homeobox gene, STIMPY, emerges strongly as a link between sugar sensing, plant cell proliferation and development. Flowering can be influenced by sucrose and glucose levels and both meristem identity and organ identity genes could well be differentially sensitive to sucrose and glucose signals. We also describe how meristems deal with extra photosynthate as a result of exposure to elevated CO₂. What we review are numerous instances of how developmental processes can be affected by sugars/nutrients. However, given the scarcity of knowledge we are unable to provide uncontested links between nutrient sensing and specific activities in meristems.     

Correlations Between the Dimensions of Different Zones of Grass Root Apices, and their Implications for Morphogenesis and Differentiation in Roots

The sizes of different zones within root apices of nine speciesof grass were estimated, and statistically significant correlationswere found between certain of them. The volume of the cap isrelated to the volume of the meristem of the root proper. Thecortical and stelar portions of the meristem are also related,and their lengths and volumes correlate with the volume of thequiescent centre. The volume of the quiescent centre also correlateswith the length of the zone in which periclinal divisions arefound in the inner cortex; these divisions generate the rowsof cells across the cortex. Diameter of the procambial cylinder,quiescent centre volume and vascular complexity are related,though from correlations alone it is not possible to say whetherone of these characters directly influences another as has beensuggested by other workers. All the zones within the root apexprobably form a tightly-integrated developmental unit. Root structure seems to be independent of cell size, thoughcell size correlates with nuclear DNA content which is a species-specificfeature. Gramineae, meristems, morphogenesis, root apices     

Fluridone affects quiescent centre division in the Arabidopsis thaliana root stem cell niche

Plants undergo cell division throughout their life in order to maintain their growth. It is well known that root and shoot tip of plants possess meristems, which contain quiescent cells. Fluridone (1-methyl-3-phenyl-5-(3-trifluoromethyl (phenyl))-4-(1H)-pyridinone) is an established inhibitor of both ABA and carotenoid biosynthesis. However, the other functions of fluridone remain undiscovered. In this report, we provide experimental evidence that fluridone plays a role in the division of the quiescent centre of the Arabidopsis root meristem. This study examined the effects of exogenous fluridone and ABA on the development of the stem cell niche in Arabidopsis root. We show that fluridone promoted the division of stem cells in the quiescent centre, whereas exogenous ABA suppressed quiescent centre division. Furthermore, we established a novel regulatory function for fluridone by demonstrating that it plays an important role in postembryonic development.     

Plant stem cell niches

Stem cells are required to support the indeterminate growth style of plants. Meristems are a plants stem cell niches that foster stem cell survival and the production of descendants destined for differentiation. In shoot meristems, stem cell fate is decided at the populational level. The size of the stem cell domain at the meristem tip depends on signals that are exchanged with cells of the organizing centre underneath. In root meristems, individual stem cells are controlled by direct interaction with cells of the quiescent centre that lie in the immediate neighbourhood. Analysis of the interactions and signaling processes in the stem cell niches has delivered some insights into the molecules that are involved and revealed that the two major niches for plant stem cells are more similar than anticipated.     

The response of apical meristems of primary roots of Vicia faba L. to colchicine treatments

The effects of 0.5% and 0.025% solutions of colchicine on the passage of cells through the mitotic cycle in apical meristems of primary roots of Vicia faba have been examined. Both treatments affected cell progression through the mitotic cycle in the same way: S and G1 were shorter, and G2 and mitosis longer, than the corresponding control values. The duration of the various phases of the mitotic cycle were similar to those reported previously for apical meristems of lateral roots though cycle time itself was longer. Recovery of root proliferating tissues from colchicine-induced inhibition of growth is correlated with the presence of quiescent cells. Meristems which have no quiescent cells do not recover from eolchicine treatment, while meristems which contain many quiescent cells recover faster than those which contain few. The growth fraction and the proportion of proliferating cells with a short cycle time are linearly related to the duration of the S period in root meristems.     

CELL DIVISION KINETICS IN THE SHOOT APICAL MERISTEM OF CERATOPTERIS THALICTROIDES BRONG. WITH SPECIAL REFERENCE TO THE APICAL CELL

The duration of mitosis and the cell cycle were determined for defined cell populations of the shoot apical meristem of Ceratopteris thalictroides Brong. by using the colchicine-induced metaphase accumulation technique. The results indicate that the apical cell is mitotically active and cycles at an apparently greater frequency than the cells of subjacent populations. Duration of mitosis was similar for all cells of the meristem. These results are correlated with mitotic indices of control apices, the geometry of the apex, and the mean number of cells in the meristem. Shoot apices from adult plants were examined to determine mitotic indices within the meristem; mitotic activity was again noted for the apical cell. These results contradict recent proposals that the pteridophyte apical cell serves as a unicellular quiescent center which lacks histogenic potential and offer experimental support for the classical concept of apical cell function in those fern shoot meristems which terminate in a single apical cell.