Background and Aims
The root apical meristem (RAM) is the plant stem cell niche which provides for the formation and continuous development of the root. Auxin is the main regulator of RAM functioning, and auxin maxima coincide with the sites of RAM initiation and maintenance. Auxin gradients are formed due to local auxin biosynthesis and polar auxin transport. The PIN family of auxin transporters plays a critical role in polar auxin transport, and two mechanisms of auxin maximum formation in the RAM based on PIN-mediated auxin transport have been proposed to date: the reverse fountain and the reflected flow mechanisms.
The two mechanisms are combined here in in silico studies of auxin distribution in intact roots and roots cut into two pieces in the proximal meristem region. In parallel, corresponding experiments were performed in vivo using DR5::GFP Arabidopsis plants.
The reverse fountain and the reflected flow mechanism naturally cooperate for RAM patterning and maintenance in intact root. Regeneration of the RAM in decapitated roots is provided by the reflected flow mechanism. In the excised root tips local auxin biosynthesis either alone or in cooperation with the reverse fountain enables RAM maintenance.
The efficiency of a dual-mechanism model in guiding biological experiments on RAM regeneration and maintenance is demonstrated. The model also allows estimation of the concentrations of auxin and PINs in root cells during development and under various treatments. The dual-mechanism model proposed here can be a powerful tool for the study of several different aspects of auxin function in root.
Auxin response; root apical meristem; patterning; reverse fountain; reflected flow; mathematical model; Arabidopsis thaliana
In plant roots, auxin is critical for patterning and morphogenesis. It regulates cell elongation and division, the development and maintenance of root apical meristems, and other processes. In Arabidopsis, auxin distribution along the central root axis has several maxima: in the root tip, in the basal meristem and at the shoot/root junction. The distal maximum in the root tip maintains the stem cell niche. Proximal maxima may trigger lateral or adventitious root initiation.
We propose a reflected flow mechanism for the formation of the auxin maximum in the root apical meristem. The mechanism is based on auxin's known activation and inhibition of expressed PIN family auxin carriers at low and high auxin levels, respectively. Simulations showed that these regulatory interactions are sufficient for self-organization of the auxin distribution pattern along the central root axis under varying conditions. The mathematical model was extended with rules for discontinuous cell dynamics so that cell divisions were also governed by auxin, and by another morphogen Division Factor which combines the actions of cytokinin and ethylene on cell division in the root. The positional information specified by the gradients of these two morphogens is able to explain root patterning along the central root axis.
We present here a plausible mechanism for auxin patterning along the developing root, that may provide for self-organization of the distal auxin maximum when the reverse fountain has not yet been formed or has been disrupted. In addition, the proximal maxima are formed under the reflected flow mechanism in response to periods of increasing auxin flow from the growing shoot. These events may predetermine lateral root initiation in a rhyzotactic pattern. Another outcome of the reflected flow mechanism - the predominance of lateral or adventitious roots in different plant species - may be based on the different efficiencies with which auxin inhibits its own transport in different species, thereby distinguishing two main types of plant root architecture: taproot vs. fibrous.
The coordination of plant cell division and expansion controls plant morphogenesis, development, and growth. Cyclin-dependent kinases (CDKs) are not only key regulators of cell division but also play an important role in cell differentiation. In plants, CDK activity is modulated by the binding of INHIBITOR OF CDK/KIP-RELATED PROTEIN (ICK/KRP). Previously, ICK2/KRP2 has been shown to mediate auxin responses in lateral root initiation. Here are analysed the roles of all ICK/KRP genes in root growth. Analysis of ick/krp null-mutants revealed that only ick3/krp5 was affected in primary root growth. ICK3/KRP5 is strongly expressed in the root apical meristem (RAM), with lower expression in the expansion zone. ick3/krp5 roots grow more slowly than wildtype controls, and this results not from reduction of division in the proliferative region of the RAM but rather reduced expansion as cells exit the meristem. This leads to shorter final cell lengths in different tissues of the ick3/krp5 mutant root, particularly the epidermal non-hair cells, and this reduction in cell size correlates with reduced endoreduplication. Loss of ICK3/KRP5 also leads to delayed germination and in the mature embryo ICK3/KRP5 is specifically expressed in the transition zone between root and hypocotyl. Cells in the transition zone were smaller in the ick3/krp5 mutant, despite the absence of endoreduplication in the embryo suggesting a direct effect of ICK3/KRP5 on cell growth. It is concluded that ICK3/KRP5 is a positive regulator of both cell growth and endoreduplication.
Arabidopsis; CDK; cell cycle; cell division; cell expansion; development; root growth
Entry into mitosis is regulated by cyclin dependent kinases that in turn are phosphoregulated. In most eukaryotes, phosphoregulation is through WEE1 kinase and CDC25 phosphatase. In higher plants a homologous CDC25 gene is unconfirmed and hence the mitotic inducer Schizosaccharomyces pombe (Sp) cdc25 has been used as a tool in transgenic plants to probe cell cycle function. Expression of Spcdc25 in tobacco BY-2 cells accelerates entry into mitosis and depletes cytokinins; in whole plants it stimulates lateral root production. Here we show, for the first time, that alterations to cytokinin and ethylene signaling explain the rooting phenotype elicited by Spcdc25 expression in Arabidopsis.
Expressing Spcdc25 in Arabidopsis results in increased formation of lateral and adventitious roots, a reduction of primary root width and more isodiametric cells in the root apical meristem (RAM) compared with wild type. Furthermore it stimulates root morphogenesis from hypocotyls when cultured on two way grids of increasing auxin and cytokinin concentrations. Microarray analysis of seedling roots expressing Spcdc25 reveals that expression of 167 genes is changed by > 2-fold. As well as genes related to stress responses and defence, these include 19 genes related to transcriptional regulation and signaling. Amongst these was the up-regulation of genes associated with ethylene synthesis and signaling. Seedlings expressing Spcdc25 produced 2-fold more ethylene than WT and exhibited a significant reduction in hypocotyl length both in darkness or when exposed to 10 ppm ethylene. Furthermore in Spcdc25 expressing plants, the cytokinin receptor AHK3 was down-regulated, and endogenous levels of iPA were reduced whereas endogeous IAA concentrations in the roots increased.
We suggest that the reduction in root width and change to a more isodiametric cell phenotype in the RAM in Spcdc25 expressing plants is a response to ethylene over-production. The increased rooting phenotype in Spcdc25 expressing plants is due to an increase in the ratio of endogenous auxin to cytokinin that is known to stimulate an increased rate of lateral root production. Overall, our data reveal important cross talk between cell division and plant growth regulators leading to developmental changes.
Plants continuously generate new tissues and organs through the activity of populations of undifferentiated stem cells, called meristems. Here, we discuss the so-called shoot apical meristem (SAM), which generates all the aerial parts of the plant. It has been known for many years that auxin plays a central role in the functioning of this meristem. Auxin is not homogeneously distributed at the SAM and it is thought that this distribution is interpreted in terms of differential gene expression and patterned growth. In this context, auxin transporters of the PIN and AUX families, creating auxin maxima and minima, are crucial regulators. However, auxin transport is not the only factor involved. Auxin biosynthesis genes also show specific, patterned activities, and local auxin synthesis appears to be essential for meristem function as well. In addition, auxin perception and signal transduction defining the competence of cells to react to auxin, add further complexity to the issue. To unravel this intricate signaling network at the SAM, systems biology approaches, involving not only molecular genetics but also live imaging and computational modeling, have become increasingly important.
Auxin dynamically regulates patterning at the shoot apical meristem. Transporters and local biosynthesis are involved in the control of its distribution at the shoot apex, where it is required for formation of new buds.
The difference between indeterminate and determinate growth in plants consists of the presence or absence of an active meristem in the fully developed organ. Determinate root growth implies that the root apical meristem (RAM) becomes exhausted. As a consequence, all cells in the root tip differentiate. This type of growth is widely found in roots of many angiosperm taxa and might have evolved as a developmental adaptation to water deficit (in desert Cactaceae), or low mineral content in the soil (proteoid roots in various taxa).
Scope and Conclusions
This review considers the mechanisms of determinate root growth to better understand how the RAM is maintained, how it functions, and the cellular and genetic bases of these processes. The role of the quiescent centre in RAM maintenance and exhaustion will be analysed. During root ageing, the RAM becomes smaller and its organization changes; however, it remains unknown whether every root is truly determinate in the sense that its RAM becomes exhausted before senescence. We define two types of determinate growth: constitutive where determinacy is a natural part of root development; and non-constitutive where determinacy is induced usually by an environmental factor. Determinate root growth is proposed to include two phases: the indeterminate growth phase, when the RAM continuously produces new cells; and the termination growth phase, when cell production gradually decreases and eventually ceases. Finally, new concepts regarding stem cells and a stem cell niche are discussed to help comprehend how the meristem is maintained in a broad taxonomic context.
Angiosperms; determinate root growth; indeterminate growth; meristem maintenance; quiescent centre; root apical meristem; root development; stem cells; stem cell niche
The structure of roots has been studied for many years, but despite their importance to the growth and well-being of plants, most researchers tend to ignore them. This is unfortunate, because their simple body plan makes it possible to study complex developmental pathways without the complications sometimes found in the shoot. In this illustrated essay, my objective is to describe the body plan of the root and the root apical meristem (RAM) and point out the control points where differentiation and cell cycle decisions are made. Hopefully this outline will assist plant biologists in identifying the structural context for their observations.
Scope and Conclusions
This short paper outlines the types of RAM, i.e. basic-open, intermediate-open and closed, shows how they are similar and different, and makes the point that the structure and shape of the RAM are not static, but changes in shape, size and organization occur depending on root growth rate and development stage. RAMs with a closed organization lose their outer root cap layers in sheets of dead cells, while those with an open organization release living border cells from the outer surfaces of the root cap. This observation suggests a possible difference in the mechanisms whereby roots with different RAM types communicate with soil-borne micro-organisms. The root body is organized in cylinders, sectors (xylem and phloem in the vascular cylinder), cell files, packets and modules, and individual cells. The differentiation in these root development units is regulated at control points where genetic regulation is needed, and the location of these tissue-specific control points can be modulated as a function of root growth rate. In Arabidopsis thaliana the epidermis and peripheral root cap develop through a highly regulated series of steps starting with a periclinal division of an initial cell, the root cap/protoderm (RCP) initial. The derivative cells from the RCP initial divide into two cells, the inner cell divides again to renew the RCP and the other cell divides through four cycles to form 16 epidermal cells in a packet; the outer cell divides through four cycles to form the 16 cells making up the peripheral root cap packet. Together, the epidermal packet and the peripheral root cap packet make up a module of cells which are clonally related.
Root apical meristem; RAM; cell cycle; differentiation; peripheral root cap; closed RAM organization; open RAM organization; epidermis; module; determination; levels of organization; plasmodesmata; T-division; root cap/protoderm initial; columella initial
Local, efflux-dependent auxin gradients and maxima mediate organ and tissue development in plants. The auxin-efflux pattern is regulated by dynamic expression and asymmetric subcellular localization of PIN auxin-efflux proteins during plant organogenesis. Thus, the question of how the expression and subcellular localization of PIN proteins are controlled goes to the heart of plant development. It has been shown that PIN expression and polarity are established not only through a self-organizing auxin-mediated polarization mechanism, but also through other means such as cell-fate determination. We found that the Arabidopsis NO VEIN (NOV) gene, encoding a novel, plant-specific nuclear factor, is required for leaf vascular development, cellular patterning and stem-cell maintenance in the root meristem and cotyledon outgrowth and separation. NOV function underlies cell-fate decisions associated with auxin gradients and maxima, thereby establishing cell type-specific PIN expression and polarity. We propose that NOV mediates cell acquisition of the competence to undergo auxin-dependent coordinated cell specification and patterning, thereby educing context-dependent auxin-mediated developmental responses.
Arabidopsis; auxin; PIN; organ development; vascular development; stem-cell maintenance; NO VEIN
Plant hormones regulate many aspects of plant growth and development. Both auxin and cytokinin have been known for a long time to act either synergistically or antagonistically to control several significant developmental processes, such as the formation and maintenance of meristem. Over the past few years, exciting progress has been made to reveal the molecular mechanisms underlying the auxin–cytokinin action and interaction. In this review, we shall briefly discuss the major progress made in auxin and cytokinin biosynthesis, auxin transport, and auxin and cytokinin signaling. The frameworks for the complicated interaction of these two hormones in the control of shoot apical meristem and root apical meristem formation as well as their roles in in vitro organ regeneration are the major focus of this review.
Auxin; cytokinin; interaction; shoot meristem; root meristem; development
Several different D-type cyclins can be found in plants and in maize, four of these have been characterized: CycD2;1, CycD4;1, CycD5;1 and CycD5;2. These cyclins appear to form complexes with Cdks, with PCNA and also with KRP proteins and in these kinase activity can be measured. The expression of the corresponding genes during maize germination is highly stimulated by phytohormones like auxin and cytokinin, however this is not followed by an equivalent increase in the amount of the corresponding proteins; nonetheless, auxins do stimulate the associated kinase activity, particularly at early germination times. Thus, auxins appear to stimulate the cell cycle during germination at two levels, transcription and kinase activation. Both auxins and cytokinins appear to shorten the G1 phase during germination and stimulate DNA synthesis, but apparently they do it in different ways as the simultaneous addition of both to germinating maize axes eliminates DNA synthesis stimulation. Therefore, similar actions may be achieved by different paths.
maize; cell cycle; germination; D cyclins; phytohormones
Plants continuously generate new organs through the activity of populations of stem cells called meristems. The shoot apical meristem initiates leaves, flowers, and lateral meristems in highly ordered, spiralled, or whorled patterns via a process called phyllotaxis. It is commonly accepted that the active transport of the plant hormone auxin plays a major role in this process. Current hypotheses propose that cellular hormone transporters of the PIN family would create local auxin maxima at precise positions, which in turn would lead to organ initiation. To explain how auxin transporters could create hormone fluxes to distinct regions within the plant, different concepts have been proposed. A major hypothesis, canalization, proposes that the auxin transporters act by amplifying and stabilizing existing fluxes, which could be initiated, for example, by local diffusion. This convincingly explains the organised auxin fluxes during vein formation, but for the shoot apical meristem a second hypothesis was proposed, where the hormone would be systematically transported towards the areas with the highest concentrations. This implies the coexistence of two radically different mechanisms for PIN allocation in the membrane, one based on flux sensing and the other on local concentration sensing. Because these patterning processes require the interaction of hundreds of cells, it is impossible to estimate on a purely intuitive basis if a particular scenario is plausible or not. Therefore, computational modelling provides a powerful means to test this type of complex hypothesis. Here, using a dedicated computer simulation tool, we show that a flux-based polarization hypothesis is able to explain auxin transport at the shoot meristem as well, thus providing a unifying concept for the control of auxin distribution in the plant. Further experiments are now required to distinguish between flux-based polarization and other hypotheses.
Plants continuously generate new organs through the activity of populations of stem cells called meristems. The shoot apical meristem (SAM) initiates leaves, flowers, and lateral organs in highly ordered, spiraled, or whorled arrangements via a process called phyllotaxis. Auxin, a plant hormone, plays an essential role in this process. It is actively transported from cell to cell by specific membrane-associated transporters. In the SAM, this coordinated transport creates organized auxin fluxes resulting in hormone accumulation at precise positions, where organ formation is triggered. One key question in this process is to understand how auxin transport is coordinated. To address this issue, we have investigated a specific hypothesis, the canalization hypothesis, whereby every cell senses and attempts to stabilize existing hormone fluxes. Because such a patterning process would require the interaction of hundreds of cells, it is impossible to estimate on a purely intuitive basis whether it would be able to generate the observed organ positions. We, therefore, developed a computational approach to test this hypothesis and showed that a flux-based mechanism is indeed able to generate phyllotactic patterns and is consistent with biological data describing meristem development.
The hormones auxin and cytokinin are key regulators of plant growth and development. As they are active at minute concentrations and regulate dynamic processes, cell and tissue levels of the hormones are finely controlled developmentally, diurnally, and in response to environmental variables. This fine control, along with a regulation of the capacity to respond ensures that the appropriate type, duration and intensity of responses are elicited. We have recently discovered that cytokinin and auxin regulate the synthesis of each other, demonstrating a mechanism for mutual feed back and feed forward control of auxin and cytokinin levels. This regulatory loop could be important for many developmental processes in plants, i.e., in fine-tuning plant hormone levels in the developing meristems of the root and shoot apex. These findings could also give a molecular explanation for earlier observations of auxin and cytokinin effects on cell cultures,1 where specific auxin and cytokinin ratios have been used to trigger different morphological events.
auxin; cytokinin; biosynthesis; metabolism; signaling; root development; interactions
In plants, the phytohormone auxin is a crucial regulator sustaining growth and development. At the cellular level, auxin is interpreted differentially in a tissue- and dose-dependent manner. Mechanisms of auxin signalling are partially unknown and the contribution of the AUXIN BINDING PROTEIN 1 (ABP1) as an auxin receptor is still a matter of debate.
Here we took advantage of the present knowledge of the root biological system to demonstrate that ABP1 is required for auxin response. The use of conditional ABP1 defective plants reveals that the protein is essential for maintenance of the root meristem and acts at least on the D-type CYCLIN/RETINOBLASTOMA pathway to control entry into the cell cycle. ABP1 affects PLETHORA gradients and confers auxin sensitivity to root cells thus defining the competence of the cells to be maintained within the meristem or to elongate. ABP1 is also implicated in the regulation of gene expression in response to auxin.
Our data support that ABP1 is a key regulator for root growth and is required for auxin-mediated responses. Differential effects of ABP1 on various auxin responses support a model in which ABP1 is the major regulator for auxin action on the cell cycle and regulates auxin-mediated gene expression and cell elongation in addition to the already well known TIR1-mediated ubiquitination pathway.
Shoot branching is regulated by competition between branches to export the phytohormone auxin into the main stem. The phytohormone strigolactone balances shoot system growth by making auxin export harder to establish, thus modulating the auxin transport network.
Plants continuously extend their root and shoot systems through the action of meristems at their growing tips. By regulating which meristems are active, plants adjust their body plans to suit local environmental conditions. The transport network of the phytohormone auxin has been proposed to mediate this systemic growth coordination, due to its self-organising, environmentally sensitive properties. In particular, a positive feedback mechanism termed auxin transport canalization, which establishes auxin flow from active shoot meristems (auxin sources) to the roots (auxin sinks), has been proposed to mediate competition between shoot meristems and to balance shoot and root growth. Here we provide strong support for this hypothesis by demonstrating that a second hormone, strigolactone, regulates growth redistribution in the shoot by rapidly modulating auxin transport. A computational model in which strigolactone action is represented as an increase in the rate of removal of the auxin export protein, PIN1, from the plasma membrane can reproduce both the auxin transport and shoot branching phenotypes observed in various mutant combinations and strigolactone treatments, including the counterintuitive ability of strigolactones either to promote or inhibit shoot branching, depending on the auxin transport status of the plant. Consistent with this predicted mode of action, strigolactone signalling was found to trigger PIN1 depletion from the plasma membrane of xylem parenchyma cells in the stem. This effect could be detected within 10 minutes of strigolactone treatment and was independent of protein synthesis but dependent on clathrin-mediated membrane trafficking. Together these results support the hypothesis that growth across the plant shoot system is balanced by competition between shoot apices for a common auxin transport path to the root and that strigolactones regulate shoot branching by modulating this competition.
Plants can adapt their form to suit the environment in which they are growing. For example, genetically identical plants can develop as a single unbranched stem or as a highly ramified bush. This broad developmental potential is possible because the shoot system is produced continuously by growing tips, known as shoot meristems. Meristems produce the stem and leaves of a shoot, and at the base of each leaf, a new meristem is formed. This meristem can remain dormant as a small bud or activate to produce a branch. Thus, the shoot system is a community of shoot meristems, the combined activity and inactivity of which shape shoot form. Here we provide evidence that growth is balanced across the Arabidopsis shoot system by competition between the shoot meristems. This competition is likely mediated by the requirement of meristems to export the plant hormone auxin in order to activate bud outgrowth. In our model, auxin in the main stem, exported from active branches, can prevent auxin export by dormant buds, thus preventing their activation. Our findings show that a second hormone, strigolactone, increases the level of competition between branches by making auxin export harder to establish. Together, these hormones balance growth across the shoot system, adjusting it according to the environmental conditions in which a plant is growing.
• Background and Aims The root apical meristems (RAM) of flowering plant roots are organized into recognizable pattern types. At present, there are no known ecological or physiological benefits to having one RAM organization type over another. Although there are phylogenetic distribution patterns in plant groups, the possible evolutionary advantages of different RAM organization patterns are not understood. Root caps of many flowering plant roots are known to release living border cells into the rhizosphere, where the cells are believed to have the capacity to alter conditions in the soil and to interact with soil micro-organisms. Consequently, high rates of border cell production may have the potential to benefit plant growth and development greatly, and to provide a selective advantage in certain soil environments. This study reports the use of several approaches to elucidate the anatomical and developmental relationships between RAM organization and border cell production.
• Methods RAM types from many species were compared with numbers of border cells released in those species. In addition, other species were grown, fixed and sectioned to verify their organization type and capacity to produce border cells. Root tips were examined microscopically to characterize their pattern and some were stained to determine the viability of root cap cells.
• Key Results The first report of a correlation between RAM organization type and the production and release of border cells is provided: species exhibiting open RAM organization produce significantly more border cells than species exhibiting closed apical organization. Roots with closed apical organization release peripheral root cap cells in sheets or large groups of dead cells, whereas root caps with open organization release individual living border cells.
• Conclusions This study, the first to document a relationship between RAM organization, root cap behaviour and a possible ecological benefit to the plant, may yield a framework to examine the evolutionary causes for the diversification of RAM organization types across taxa.
Border cells; root caps; root apical organization; root meristem
It has been known for many decades that auxin inhibits the activation of axillary buds, and hence shoot branching, while cytokinin has the opposite effect. However, the modes of action of these two hormones in branching control is still a matter of debate, and their mechanisms of interaction are equally unresolved.
Here we review the evidence for various hypotheses that have been put forward to explain how auxin and cytokinin influence axillary bud activity. In particular we discuss the roles of auxin and cytokinin in regulating each other's synthesis, the cell cycle, meristem function and auxin transport, each of which could affect branching. These different mechanisms have implications for the main site of hormone action, ranging from systemic action throughout the plant, to local action at the node or in the bud meristem or leaves. The alternative models have specific predictions, and our increasing understanding of the molecular basis for hormone transport and signalling, cell cycle control and meristem biology is providing new tools to enable these predictions to be tested.
Shoot branching; axillary bud; dormancy; auxin; cytokinin; canalization; polar auxin transport stream; cell cycle
Using transcript profile analysis, we explored the nature of the stem cell niche in roots of maize (Zea mays). Toward assessing a role for specific genes in the establishment and maintenance of the niche, we perturbed the niche and simultaneously monitored the spatial expression patterns of genes hypothesized as essential. Our results allow us to quantify and localize gene activities to specific portions of the niche: to the quiescent center (QC) or the proximal meristem (PM), or to both. The data point to molecular, biochemical and physiological processes associated with the specification and maintenance of the niche, and include reduced expression of metabolism-, redox- and certain cell cycle-associated transcripts in the QC, enrichment of auxin-associated transcripts within the entire niche, controls for the state of differentiation of QC cells, a role for cytokinins specifically in the PM portion of the niche, processes (repair machinery) for maintaining DNA integrity and a role for gene silencing in niche stabilization. To provide additional support for the hypothesized roles of the above-mentioned and other transcripts in niche specification, we overexpressed, in Arabidopsis, homologs of representative genes (eight) identified as highly enriched or reduced in the maize root QC. We conclude that the coordinated changes in expression of auxin-, redox-, cell cycle- and metabolism-associated genes suggest the linkage of gene networks at the level of transcription, thereby providing additional insights into events likely associated with root stem cell niche establishment and maintenance.
Electronic supplementary material
The online version of this article (doi:10.1007/s00425-009-1059-3) contains supplementary material, which is available to authorized users.
Quiescent center; Root; Stem cell; Stem cell niche; Zea mays
The small ubiquitin modifier (SUMO) conjugation/deconjugation is an important regulatory progress in plant development and responses to abiotic stresses. However, much less is known about the roles of sumoylation in plant root development. Cytokinin and auxin play crucial roles in determining the balance between cell proliferation and cell differentiation in Arabidopsis roots. The SUMO E3 ligase AtMMS21 is a homologue of human NSE2/MMS21, which modulates DNA damage and DNA repair in human cells. This addendum summarizes our recent paper on the AtMMS21 mediating cytokinin signaling to regulate the root meristem cell proliferation. The mms21-1 roots had reduced responses to exogenous cytokinins and decreased expression of the cytokinin-induced genes ARR3, ARR4, ARR5 and ARR7, compared with the wild type. Furthermore, the expression of CRE1 and ARR1, which are both the receptor and positive regulator of cytokinin signaling, was also reduced in the mms21-1 mutant plants.
Arabidopsis thaliana; AtMMS21; cytokinin; root meristem; SUMO E3 ligase
The shoot apical meristem (SAM) is a group of proliferating, embryonic-type cells that generates the aerial parts of the plant. SAMs are highly organized and stable structures that can function for years or even centuries. This is in apparent contradiction to the behaviour of their constituent cells, which continuously proliferate and differentiate. To reconcile the dynamic nature of the cells with the stability of the overall system the existence of elaborate signalling networks has been proposed. This is supported by recent work suggesting that the exchange of signals between cells, rather than a rigidly predetermined genetic program, is required for the establishment and functioning of an organized meristem. Together these interactions form a stable network, set up during embryogenesis, that assures the coordination of cell behaviour throughout development. Besides meristem-specific signalling cascades such as the CLAVATA receptor kinase pathway, which controls meristem size, these interactions involve plant hormones. In particular, cytokinins and auxins are implicated in the maintenance of meristem identity and phyllotaxis, respectively.
The effect of mechanical stress on the root apical meristem (RAM) organization of Zea mays was investigated. In the experiment performed, root apices were grown through a narrowing of either circular (variant I) or elliptical (variant II) shape. This caused a mechanical impedance distributed circumferentially or from the opposite sides in variant I and II, respectively. The maximal force exerted by the growing root in response to the impedance reached the value of 0.15 N for variant I and 0.08 N for variant II. Significant morphological and anatomical changes were observed. The changes in morphology depended on the variant and concerned diminishing and/or deformation of the cross-section of the root apex, and buckling and swelling of the root. Anatomical changes, similar in both variants, concerned transformation of the meristem from closed to open, an increase in the number of the cell layers at the pole of the root proper, and atypical oblique divisions of the root cap cells. After leaving the narrowing, a return to both typical cellular organization and morphology of the apex was observed. The results are discussed in terms of three aspects: the morphological response, the RAM reorganization, and mechanical factors. Assuming that the orientation of division walls is affected by directional cues of a tensor nature, the changes mentioned may indicate that a pattern of such cues is modified when the root apex passes through the narrowing, but its primary mode is finally restored.
Mechanical stress; root apical meristem organization; tiers of initials; Zea mays
CLE, which is the term for the CLV3/ESR-related gene family, is thought to participate in CLAVATA3-WUSCHEL (CLV3-WUS) and CLV3-WUS-like signaling pathways to regulate meristem activity in plant. Although some CLE genes are expressed in meristems, many CLE genes appear to express in a variety of tissues/cells. Here we report that CLE14 and CLE20 express in various specific tissues/cells outside the shoot/root apical meristem (SAM/RAM), including in highly differentiated cells, and at different developmental stages. Overexpressing CLE14 or CLE20 also causes multiple phenotypes, which is consistent with its expression pattern in Arabidopsis. These results suggest that CLE genes may play multiple roles and involve other signaling cascades in addition to the CLV3-WUS and CLV3-WUS-like pathways.
CLE; CLAVATA3-WUSCHEL; cell signaling and development; root apical meristem; arabidopsis
Cytokinin promotes cell differentiation in plant roots by repressing auxin transport and responses at the meristem boundary.
The hormone cytokinin promotes cell differentiation in plant roots by repressing both auxin transport and responses to auxin at the boundary between the meristem and the root elongation zone.
On the search for sparingly available nutrients, plants may alter their root architecture to improve soil exploration. So far, the examples for root system modifications induced by a heterogeneous availability of nutrients have been reported for the macronutrients nitrogen (N) and phosphorous (P). In an attempt to extend this type of knowledge to other nutrients, we recently provided evidence that Arabidopsis roots are able to sense a local availability of the micronutrient iron (Fe) and to respond with lateral root elongation into the Fe-containing patch. This specific root response was caused by enhanced elongation of cells leaving the root meristem and was dependent on an AUX1-mediated auxin accumulation in the lateral root apices. In this report, we compare mechanisms underlying this response with those known for other nutrients and show that a substantial genotypic variation exists among accessions of A. thaliana in the responsiveness of lateral roots toward localized Fe supplies.
AUX1; auxin; iron sensing; lateral root development; natural variation; root architecture
We investigated expression patterns of DNA repair genes such as the CPD photolyase, UV-DDB1, CSB, PCNA, RPA32 and FEN-1 genes by northern hybridization analysis and in situ hybridization using a higher plant, rice (Oryza sativa L. cv. Nipponbare). We found that all the genes tested were expressed in tissues rich in proliferating cells, but only CPD photolyase was expressed in non-proliferating tissue such as the mature leaves and elongation zone of root. The removal of DNA damage, cyclobutane pyrimidine dimers and (6–4) photoproducts, in both mature leaves and the root apical meristem (RAM) was observed after UV irradiation under light. In the dark, DNA damage in mature leaves was not repaired efficiently, but that in the RAM was removed rapidly. Using a rice 22K custom oligo DNA microarray, we compared global gene expression patterns in the shoot apical meristem (SAM) and mature leaves. Most of the excision repair genes were more strongly expressed in SAM. These results suggested that photoreactivation is the major DNA repair pathway for the major UV-induced damage in non-proliferating cells, while both photoreactivation and excision repair are active in proliferating cells.
Background and Aims
Most studies of exodermal structure and function have involved species with a uniseriate exodermis. To extend this work, the development and apoplastic permeability of Iris germanica roots with a multiseriate exodermis (MEX) were investigated. The effects of different growth conditions on MEX maturation were also tested. In addition, the exodermises of eight Iris species were observed to determine if their mature anatomy correlated with habitat.
Plants were grown in soil, hydroponics (with and without a humid air gap) or aeroponics. Roots were sectioned and stained with various dyes to detect MEX development from the root apical meristem, Casparian bands, suberin lamellae and tertiary wall thickenings. Apoplastic permeability was tested using dye (berberine) and ionic (ferric) tracers.
The root apical meristem was open and MEX development non-uniform. In soil-grown roots, the exodermis started maturing (i.e. Casparian bands and suberin lamellae were deposited) 10 mm from the tip, and two layers had matured by 70 mm. In both hydro- and aeroponically grown roots, exodermal maturation was delayed. However, in areas of roots exposed to an air gap in the hydroponic system, MEX maturation was accelerated. In contrast, maturation of the endodermis was not influenced by the growth conditions. The mature MEX had an atypical Casparian band that was continuous around the root circumference. The MEX prevented the influx and efflux of berberine, but had variable resistance to ferric ions due to their toxic effects. Iris species living in well-drained soils developed a MEX, but species in water-saturated substrates had a uniseriate exodermis and aerenchyma.
MEX maturation was influenced by the roots' growth medium. The MEX matures very close to the root tip in soil, but much further from the tip in hydro- and aeroponic culture. The air gap accelerated maturation of the second exodermal layer. In Iris, the type of exodermis was correlated with natural habitat suggesting that a MEX may be advantageous for drought tolerance.
Iris germanica; roots; culture conditions; development; anatomy; apoplastic tracers; multiseriate exodermis; endodermis; root apical meristem