植物根系向地性感应的分子机理与养分吸收

植物根系向地性是决定根系空间生长趋势的主要因素之一, 对于养分吸收具有重要影响。认识根系向地性感应和根系生长变化的分子机理及其与养分吸收的关系, 可为遗传改良根系性状、提高植物养分吸收效率提供理论依据。本文从重力感应、信号转导和生长素非对称分布等方面总结了植物根系向地性感应的分子机理, 探讨了根系在养分胁迫下(特别是磷胁迫下)向地性变化的生理基础及其与养分吸收(特别是磷吸收)的关系, 最后对根系向地性研究的若干问题进行了展望。     

植物根构型特性与磷吸收效率

植物根构型,即根系在生长介质中的空间造型和分布,与磷吸收效率密切相关;认识植物根构型,可为植物磷效率的遗传改良提供依据。长期以来,人们试图定量描述植物根构型,确立一个能客观全面地描述根系三维立体构型的综合指标。试验指出,植物主要通过向地性变化和根冠之间的碳源分配来改变根构型,从而影响磷吸收效率;根系向地性变化可由缺磷等因素所诱导,且存在着一定的遗传变异性。有证据表明,根构型对低磷胁迫的适应性变化是     

植物根构型特性与磷吸收效率

植物根构型，即根系在生长介质中的空间造型和分布，与磷吸收效率密切相关；认识植物根构型，可为植物磷效率的遗传改良提供依据。长期以来，人们试图定量描述植物根构型，确立一个能客观全面地描述根系三维立体构型的综合指标。试验指出，植物主要通过向地性变化和根冠之间的碳源分配来改变根构型, 从而影响磷吸收效率；根系向地性变化可由缺磷等因素所诱导，且存在着一定的遗传变异性。有证据表明，根构型对低磷胁迫的适应性变化是受基因调控的一个生理过程，其中乙烯可能是一种重要的生理调节物质。迄今已在一些植物上定位到了部分控制根构型的数量性状座位，为该性状的分子生物学改良提供了基础。随着现代技术的进展，植物根构型研究将取得更大的突破。     

植物miRNA在调节低磷胁迫响应中的作用

植物MicroRNA(miRNA)是一类内源性非编码小分子RNA,它们参与调节植物的生长、发育和代谢过程中多种基因的表达。近期的研究发现miRNA参与调节磷的吸收和利用,对植物适应低磷胁迫具有重要作用。本文概述了植物磷吸收和转运的机制,介绍了低磷胁迫下miRNA的表达水平变化,重点对miRNA在植物响应低磷胁迫中的作用,如改变根系结构、提高磷的转运和再利用效率、参与花青素和抗氧化物生物合成等进行了综述,以期为揭示植物低磷胁迫响应分子机制,提高植物对磷的吸收效率提供借鉴。     

植物磷获取机制及其对全球变化的响应

磷是植物生长的必需元素,而陆地生态系统普遍存在磷限制,全球变化可能会影响土壤磷循环过程,进一步加剧磷限制,探讨植物磷获取策略对科学预测生态系统生产力如何适应全球变化具有重要意义。该文通过收集和梳理相关文献,从4个方面综述植物的磷获取机制及其对全球变化的响应:1)植物的磷饥饿响应机制;2)植物的磷获取途径和策略;3)土壤微生物对植物磷吸收的影响; 4)植物磷吸收对全球变化(温度升高、氮沉降和降水变化)的响应及其机制。该综述有助于深入理解全球变化背景下植物适应低磷胁迫的机理,也可为养分资源管理实践提供理论依据。     

利用抑制性扣除杂交技术克隆水稻磷饥饿诱导基因

磷素是植物生长所必需的重要元素.在缺磷环境中,植物能够调节自身的形态、生理生化和基因表达水平来适应环境的变化.为研究水稻(Oryza sativa L.)耐低磷胁迫的分子机理,采用抑制性扣除杂交技术(SSH)构建磷饥饿诱导的水稻根系扣除cDNA文库.通过文库筛选和测序获得18个已知基因和47个功能未知基因.这些基因参与了不同的代谢过程,包括磷吸收和转运、信号传导、蛋白质合成和降解、碳水化合物代谢和胁迫反应.Northern杂交结果表明,在磷饥饿胁迫下这些基因呈现不同的表达模式,并且不同代谢过程中的基因对磷饥饿有着不同的反应.     

部分根系供磷对黄瓜根系和幼苗生长及根系酸性磷酸酶活性影响

通过分根处理研究了部分根系供磷对黄瓜幼苗生长、植株体内的含磷量及根系酸性磷酸酶活性的影响。结果表明 ,2 0 %根系缺磷 (1条根缺磷 ,4条根供磷 )可以促进根系及植株地上部的生长 ,其根系及地上部的生物量分别是正常生长植株的 1.39倍和 1.2 1倍。2 0 %根系缺磷 ,还可以促进其它供磷根系对磷的吸收。分根处理后 ,2 0 %根系缺磷不影响植物对磷营养的需要 ,但却表现出了R/S比增大的典型缺磷反应 ,说明植物感应缺磷根系起着比地上部更为重要的作用。分根处理后不供磷根系的酸性磷酸酶活性显著高于供磷根系的酸性磷酸酶活性 ,并且根系的酸性磷酸酶活性只与根系的含磷量显著相关 ,与地上部的磷营养状况关系不明显。这说明 ,缺磷条件下 ,黄瓜植株根系分泌酸性磷酸酶活性的增高 ,是黄瓜根系对低磷胁迫的适应性机理 ,而不是地上部改善体内磷营养的调控机理。     

菜豆根构型对低磷胁迫的适应性变化及基因型差异

利用特殊设计的营养袋纸培养和分层式磷控释砂培等根系生长系统结合计算机图像分析技术,以基根根长在生长介质各层的相对分布和基根平均生长角度为指标,定量测定菜豆（Ｐｈａｓｅｏｌｕｓ ｖｕｌｇａｒｉｓ Ｌ．）根构型在低磷胁迫下的适应性变化及其与磷效率的关系。结果表明,菜豆根构型对低磷胁迫具有适应性反应,在缺磷条件下基根向地性减弱,基根在生长介质表层相对分布增多、基根平均生长角度（与水平线夹角）变小,从而导     

植物根系响应低磷胁迫的机理研究

磷是植物生长的必需营养元素之一。但大部分土壤中有效磷含量较低,难以满足植物生长的需求。作物磷效率遗传改良是解决土壤磷供应不足的有效途径。根系是植物吸收矿质营养元素的主要器官,其性状决定了植物对土壤磷的吸收利用效率。解析根系对低磷胁迫的响应机制是进行作物磷效率遗传改良的基础。主要介绍了近年来关于植物根系响应低磷胁迫机理的重要研究成果。     

利用抑制性扣除杂交技术克隆水稻磷饥饿诱导基因

磷素是植物生长所必需的重要元素。在缺磷环境中,植物能够调节自身的形态、生理生化和基因表达水平来适应环境的变化。为研究水稻(Oryzn sativa L.)耐低磷胁迫的分子机理,采用抑制性扣除杂交技术(SSH)构建磷饥饿诱导的水稻根系扣除cDNA文库。通过文库筛选和测序获得18个已知基因和47个功能未知基因。这些基因参与了不同的代谢过程,包括磷吸收和转运、信号传导、蛋白质合成和降解、碳水化合物代谢和胁迫反应。Northern杂交结果表明,在磷饥饿胁迫下这些基因呈现不同的表达模式,并且不同代谢过程中的基因对磷饥饿有着不同的反应。     

The importance of root gravitropism for inter-root competition and phosphorus acquisition efficiency: results from a geometric simulation model

We have observed that low soil phosphorus availability alters the gravitropic response of basal roots in common bean (Phaseolus vulgaris L.), resulting in a shallower root system. In this study we use a geometric model to test the hypotheses that a shallower root system is a positive adaptive response to low soil P availability by (1) concentrating root foraging in surface soil horizons, which generally have the highest P availability, and (2) reducing spatial competition for P among roots of the same plant. The growth of nine root systems contrasting in gravitropic response over 320 h was simulated in SimRoot, a dynamic three-dimensional geometric model of root growth and architecture. Phosphorus acquisition and inter-root competition were estimated with Depzone, a program that dynamically models nutrient diffusion to roots. Shallower root systems had greater P acquisition per unit carbon cost than deeper root systems, especially in older root systems. This was due to greater inter-root competition in deeper root systems, as measured by the volume of overlapping P depletion zones. Inter-root competition for P was a significant fraction of total soil P depletion, and increased with increasing values of the P diffusion coefficient (De), with root age, and with increasing root gravitropism. In heterogenous soil having greater P availability in surface horizons, shallower root systems had greater P acquisition than deeper root systems, because of less inter-root competition as well as increased root foraging in the topsoil. Root P acquisition predicted by SimRoot was validated against values for bean P uptake in the field, with an r ² between observed and predicted values of 0.75. Our results support the hypothesis that altered gravitropic sensitivity in P-stressed roots, resulting in a shallower root system, is a positive adaptive response to low P availability by reducing inter-root competition within the same plant and by concentrating root activity in soil domains with the greatest P availability. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effect of phosphorus availability on basal root shallowness in common bean

Root gravitropism may be an important element of plant response to phosphorus availability because it determines root foraging in fertile topsoil horizons, and thereby phosphorus acquisition. In this study we seek to test this hypothesis in both two dimensional paper growth pouch and three-dimensional solid media of sand and soil cultures. Five common bean (Phaseolus vulgaris L.) genotypes with contrasting adaptation to low phosphorus availability were evaluated in growth pouches over 6 days of growth, and in sand culture and soil culture over 4 weeks of growth. In all three media, phosphorus availability regulated the gravitropic response of basal roots in a genotype-dependent manner. In pouches, sand, and soil, the phosphorus-inefficient genotype DOR 364 had deeper roots with phosphorus stress, whereas the phosphorus-efficient genotype G19833 responded to phosphorus stress by producing shallower roots. Genotypes were most responsive to phosphorus stress in sand culture, where relative root allocation to the 0–3- and 3–6-cm horizons increased 50% with phosphorus stress, and varied 300% (3–6 cm) to 500% (0–3 cm) among genotypes. Our results indicate that (1) phosphorus availability regulates root gravitropic growth in both paper and solid media, (2) responses observed in young seedlings continue throughout vegetative growth, (3) the response of root gravitropism to phosphorus availability varies among genotypes, and (4) genotypic adaptation to low phosphorus availability is correlated with the ability to allocate roots to shallow soil horizons under phosphorus stress.     

Root system adjustments: regulation of plant nutrient uptake and growth responses to elevated CO2

Nutrients such as nitrogen (N) and phosphorus (P) often limit plant growth rate and production in natural and agricultural ecosystems. Limited availability of these nutrients is also a major factor influencing long-term plant and ecosystem responses to rising atmospheric CO₂ levels, i.e., the commonly observed short-term increase in plant biomass may not be sustained over the long-term. Therefore, it is critical to obtain a mechanistic understanding of whether elevated CO₂ can elicit compensatory adjustments such that acquisition capacity for minerals increases in concert with carbon (C) uptake. Compensatory adjustments such as increases in (a) root mycorrhizal infection, (b) root-to-shoot ratio and changes in root morphology and architecture, (c) root nutrient absorption capacity, and (d) nutrient-use efficiency can enable plants to meet an increased nutrient demand under high CO₂. Here we examine the literature to assess the extent to which these mechanisms have been shown to respond to high CO₂. The literature survey reveals no consistent pattern either in direction or magnitude of responses of these mechanisms to high CO₂. This apparent lack of a pattern may represent variations in experimental protocol and/or interspecific differences. We found that in addressing nutrient uptake responses to high CO₂ most investigators have examined these mechanisms in isolation. Because such mechanisms can potentially counterbalance one another, a more reliable prediction of elevated CO₂ responses requires experimental designs that integrate all mechanisms simultaneously. Finally, we present a functional balance (FB) model as an example of how root system adjustments and nitrogen-use efficiency can be integrated to assess growth responses to high CO₂. The FB model suggests that the mechanisms of increased N uptake highlighted here have different weights in determining overall plant responses to high CO₂. For example, while changes in root-to-shoot biomass allocation, r, have a small effect on growth, adjustments in uptake rate per unit root mass, [`(n)]\bar \nu , and photosynthetic N use efficiency, p*, have a significantly greater leverage on growth responses to elevated CO₂ except when relative growth rate (RGR) reaches its developmental limit, maximum RGR (RGR_max).     

Molecular mechanisms mediating root hydrotropism: what we have observed since the rediscovery of hydrotropism

Roots display directional growth toward moisture in response to a water potential gradient. Root hydrotropism is thought to facilitate plant adaptation to continuously changing water availability. Hydrotropism has not been as extensively studied as gravitropism. However, comparisons of hydrotropic and gravitropic responses identified mechanisms that are unique to hydrotropism. Regulatory mechanisms underlying the hydrotropic response appear to differ among different species. We recently performed molecular and genetic analyses of root hydrotropism in Arabidopsis thaliana. In this review, we summarize the current knowledge of specific mechanisms mediating root hydrotropism in several plant species.

     

The role of nodules in the tolerance of common bean to iron deficiency

Iron is vital for the establishment and function of symbiotic root nodules of legumes. Although abundant in the environment, Fe is often a limiting nutrient for plant growth due to its low solubility and availability in some soils. We have studied the mechanism of iron uptake in the root nodules of common bean to evaluate the role of nodules in physiological responses to iron deficiency. Based on experiments using full or partial submergence of nodulated roots in the nutrient solution, our results show that the nodules were affected only slightly under iron deficiency, especially when the nodules were submerged in nutrient solution in the tolerant cultivar. In addition, fully submerged root nodules showed enhanced acidification of the nutrient solution and showed higher ferric chelate reductase activity than that of partially submerged roots in plants cultivated under Fe deficiency. The main results obtained in this work suggest that in addition to preferential Fe allocation from the root system to the nodules, this symbiotic organ probably develops some mechanisms to respond to iron deficiency. These mechanisms were implied especially in nodule Fe absorption efficiency and in the ability of this organ to take up Fe directly from the medium.     

Nutrient uptake and allocation at steady-state nutrition

Ingestad, T. and Ågren, G. I. 1988. Nutrient uptake and allocation at steady-state nutrition. - Physiol. Plant. 72: 450–459. Net nutrient uptake and translocation rates are discussed for conditions of steady-state nutrition and growth. Under these conditions, the relative uptake rate is equal to the relative growth rate, for whole plants as well as for plant parts, since the root/shoot ratio and internal concentrations remain stable. The nutrient productivity and the minimum internal concentration are parameters characteristic for the plant and the nutrient. A conceptual, mathematical model, based on these two fundamental parameters is used for calculation and prediction of the net nutrient uptake rate, which is required to maintain steady-state nutrition at a specified internal nutrient concentration or relative growth rate. When uptake rate is expressed on the basis of the root growth rate, there is, up to optimum, a strong linear relationship between uptake rate and the internal concentration of the limiting nutrient. More complicated and less consistent relationships are obtained when uptake rate is related to root biomass. The limiting factor for suboptimum uptake is the amount of nutrients becoming available at the root surface. When replenishment is efficient, e.g. with vigorous stirring, the concentration requirement at the root surface appears to be extremely low, even at optimum. In the suboptimum range of nutrition, the effect of nutrient status on root growth rate is a critical factor with a strong feed-back on nutrition, growth and allocation. At supraoptimum conditions, the uptake mechanism is interpreted as a protection against too high uptake rates and internal concentrations at high external concentration. In birch (Betula pendula Roth.), the allocation of nitrogen to the shoots is high compared to that of potassium and also to that of phosphorus at low nitrogen or phosphorus status. With decreasing stress, phosphorus allocation becomes more and more similar to nitrogen allocation. The formulation of a mathematical model for calculation of allocation of biomass and nutrients requires more exact information on the quantitative dependence of the growth-regulating processes on nutrition.     

植物根毛的发生、发育及养分吸收

根毛是植物吸收养分的重要器官,认识根毛的发生、发育规律及其与养分吸收的关系,可为植物养分吸收效率的遗传改良提供依据。介绍了植物根毛的形态特性、发生和发育过程及其调控机制,并结合本实验室的工作,讨论了根毛对养分吸收的贡献、根毛受养分有效性的调节及其与其他根系形态构型性状间的关系,阐述了根毛中养分转运等植物营养过程及其生理和分子生物学基础。最后提出了关于根毛研究中的一些问题和研究前景。