Notes from the Underground： Receptor-Like Kinases in Arabidopsis Root Development

During plant development, the frequency and context of cell division must be controlled, and cells must differentiate properly to perform their mature functions. In addition, stem cell niches need to be maintained as a reservoir for new cells. All of these processes require intercellular signaling, whether it is a cell relaying its position to other cells, or more mature cells signaling to the stem cell niche to regulate the rate of growth. Receptor-like kinases have emerged as a major component in these diverse roles, especially within the Arabidopsis root. In this review, the functions of receptor-like kinase signaling in regulating Arabidopsis root development will be examined in theareas of root apical meristem maintenance, regulation of epidermal cell fate, lateral root development and vascular differentiation.     

Determinate root growth and meristem maintenance in angiosperms

BACKGROUND: 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.     

Plant stem cells and their regulations in shoot apical meristems

Stem cells in plants, established during embryogenesis, are located in the centers of the shoot apical meristem (SAM) and the root apical meristem (RAM). Stem cells in SAM have a capacity to renew themselves and to produce new organs and tissues indefinitely. Although fully differentiated organs such as leaves do not contain stem cells, cells in such organs do have the capacity to re-establish new stem cells, especially under the induction of phytohormones in vitro. Cytokinin and auxin are critical in creating position signals in the SAM to maintain the stem cell organizing center and to position the new organ primordia, respectively. This review addresses the distinct features of plant stem cells and focuses on how stem cell renewal and differentiation are regulated in SAMs.     

Phosphorus acquisition from soil by white lupin (Lupinus albus L.) and soybean (Glycine max L.), species with contrasting root development

White lupin and soybean have contrasting root morphologies: white lupin develops proteoid or cluster roots, roots with discreet clusters of short, determinate branch roots (rootlets) while soybean develops a more fibrous root system with evenly distributed, longer branch roots. Growth and P acquisition by white lupin and soybean were compared in a soil high in bound, total P, with or without additional inorganic P applied in solution. Additional P increased biomass by 25% and doubled total P in soybean. In contrast, white lupin did not respond to additional P in biomass or total P. However added P decreased cluster development on proteoid roots indicating that white lupin sensed the added P. The reduction in cluster weight per plant was exactly countered by an increase in dry weight of other roots. Soybean root development responded to P application, proliferating branch roots with active meristems in the upper portion of the soil profile where P was applied, and reducing root weight to plant weight by 13%. White lupin did not proliferate roots in response to P application. When P was not added to soil, soybean and lupin acquired similar P per unit root dry weight. However, white lupin accumulated 4.8 times more P per unit root length, suggesting that P acquisition in these plants involved other mechanisms such as the exudation of P solubilizing compounds. Soybean accessed P by developing more root length thus colonising more soil volume than white lupin and, therefore, was better able to take advantage of the added P. Pericycle and root tip meristem activities were critical to the differences in root development between white lupin and soybean, and therefore their responses to plant and soil P.     

Why are laterals less affected than main axes by homogeneous unfavourable physical conditions? A model-based hypothesis

When plants develop in strong soils, growth of the root system is generally depressed. However, branching and elongation of branches are often less affected than growth of the main axes, whenever the whole root system encounters even-impeded conditions. On the basis of a model simulating root growth and architecture as related to assimilate availability, we propose a simple hypothesis to explain such behaviour. In the model, growth of each root depends on its own elongation potential, which is estimated by its apical diameter. The potential elongation rate–apical diameter relationship is the same for all the roots of the system and is described by a monomolecular function. Our hypothesis is that the effect of soil strength can be simulated by introducing an impedance factor in the definition of root maximum potential elongation rate, common to the whole root system. When such impedance factor is applied, it affects more the potential of larger roots (main axes) than that of thinner roots (secondary and tertiary branches). Simulations provided in high impedance conditions led to root systems characterised by short taproots, whereas growth of secondary roots was unaffected and growth of tertiary roots was enhanced. Actual branching density was also higher, although branching rules have been unchanged. Such simulated systems where similar to that observed in strong soils. Friction laws or pore size can be involved in the larger reduction of the potential growth of main axes. Moreover, when growth of main axes is restricted, assimilate availability becomes higher for branches and that could explain that their growth could be increased in a homogeneous strong soil. This revised version was published online in June 2006 with corrections to the Cover Date.     

The production and release of living root cap border cells is a function of root apical meristem type in dicotyledonous angiosperm plants

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.     

生长素在微生物调控根发育过程中的作用

很多微生物通过分泌生长素和生长素前体与植物建立了有益的关系并改变植物根系的形态结构,此外,微生物分泌的其他代谢产物也能改变植物生长素信号通路。因此,生长素和生长素信号通路在微生物调控植物根系发育的过程中起着至关重要的作用。该文从生长素合成、生长素信号和生长素极性运输3个方面总结了生长素在微生物调控植物根系发育过程中的作用,主要包括微生物增加了植物内源生长素的含量、增强了生长素的信号和调控PIN蛋白的表达水平,进而如何调控植物生理和分子水平来适应微生物对其根系的改变,为进一步开展该方面的研究奠定了基础。     

土壤湿度对香樟幼苗根系生长和构型的影响

以1年生香樟(Cinnamomum camphora)幼苗为试材,设置对照组(CK)、中度干旱处理(M)、重度干旱处理(S)三个处理,比较不同土壤湿度下香樟幼苗不同时期地上部分生长和根系构型,探究香樟幼苗根系对不同土壤湿度的适应性及其耐旱机制。结果表明,中度和重度干旱处理组的香樟根系及地上部分干物质积累、根系长度、根系表面积、根系直径和根尖数均显著小于对照组(P<0.05)。同时干旱显著增加香樟幼苗的根系拓扑指数,降低香樟根的分形维数和平均分枝角度(P<0.05)。可见土壤湿度程度及处理时间显著影响香樟根系的生长及在土壤中的布局。较低土壤湿度可显著抑制根长的延长、根表面积扩大和根的增殖,且随着土壤湿度的继续降低以及处理时间延长,香樟幼苗根系的生长受到水分亏缺的抑制作用加重,根系建成成本增高的同时,根系分枝的复杂性降低,根系必须通过朝着更陡、更深的方向生长伸长来提高水分吸收效率。建议在园林绿化工程养护过程中制定科学的水分管理策略,以满足香樟生长过程中对土壤水分的需要。