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。     

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。     

铵减弱拟南芥主根向地性及其相关作用途径

向地性是决定植物根系空间构型的主要因素之一,对植物锚定和水分养分吸收至关重要。除了重力,根系向地性还受土壤环境因子影响。本文采用琼脂培养方法,研究了铵对拟南芥主根向地性反应的影响及相关作用途径。结果表明：短期内,不同浓度（NH4）2SO4均显著抑制主根向地性弯曲,但随着时间的延长,根尖向地性角度逐渐变小。而等（NH4）2SO4浓度的NaCl对主根向地性抑制效应较小,不同浓度的甘露醇不阻碍主根向地性弯曲。纽织化学染色结果显示铵处理12h以内,Col-0根尖没有淀粉体的快速降解过程,并且铵对淀粉体缺失突变体pgm—1主根向地性的影响同Col-0相似。铵处理部分恢复生长素转运载体突变体auxl-22和eir1-1主根向地性缺失。这些结果表明,铵对拟南芥主根向地性的影响独立于根尖淀粉体参与的重力感应途径。     

模拟氮沉降对低磷胁迫下马尾松无性系细根形态和氮磷效率的影响

根系是植物吸收土壤营养的关键部位, 不同径级根系的形态和功能差异不仅与植株自身的遗传因素有关, 而且受到土壤中营养元素分布和水平的影响。在我国亚热带高氮沉降和酸性红壤磷匮乏及不均一的土壤环境下, 研究林木不同径级根系对外界营养环境变化的响应有利于深入了解林木根系的觅养机制及规律。该文以马尾松(Pinus massoniana)无性系19-5 (高磷效率)和21-3 (低磷效率)为材料, 在同质低磷和异质低磷两种盆栽处理下, 设置3个氮水平(对照、中氮和高氮)的模拟氮沉降实验。结果表明: 1)马尾松无性系苗木的生长受磷环境、氮水平和无性系三因素共同影响, 模拟氮沉降显著促进了异质低磷下马尾松苗高和整株干物质量的增加, 而在同质低磷下氮效应不显著; 在异质低磷、高氮下, 无性系19-5的苗高和整株干物质量分别是无性系21-3的1.1倍和1.6倍。2)马尾松各径级细根长度和表面积随径级增大而减小, 模拟氮沉降促进了直径≤1.5 mm的细根的增生发育, 直径1.5-2.0 mm的细根和>2.0 mm的较粗根无明显变化; 另外, 直径≤1.5 mm的细根长度占总根长的比例保持在90.4%-92.8%范围内, 受氮影响较小。3)模拟氮沉降显著提高了异质低磷下无性系19-5≤1.5 mm的细根长度和表面积, 同时, 其根系氮、磷吸收效率较对照分别高出93.3%和148.4%; 无性系21-3的根系氮、磷吸收效率受氮影响较小; 根系氮、磷利用效率均无显著变化。上述结果表明, ≤1.5 mm的细根的增生发育和氮、磷吸收效率的提高可能是磷高效马尾松无性系应对高氮低磷环境的重要响应机制。     

植物磷营养高效的分子生物学研究进展

挖掘利用植物自身的磷高效营养遗传资源是农业可持续发展的关键。磷高效营养性状涉及根形态、根分泌物、膜与体内磷转运以及菌根等许多方面,表现为数量遗传性状及受多基因控制。近年来,许多高亲和磷转运子基因已被克隆, 磷向地上部转运和磷吸收负反馈调节的控制基因也被发现, 对于根系分泌有机酸和酸性磷酸酶的基因的控制也有了一定的了解, 但目前对于根毛、排根、根构型以及菌根的营养学意义性状的分子生物学研究进展缓慢。     

植物根构型的定量分析

植物根系具有锚定植株、吸收和运输土壤中的水分及养分、合成和贮藏营养物质等重要功能。根构型是根系在土壤中的空间造型和分布。对植物根构型进行定量分析,有助于人们了解根系结构和根系功能在生态系统中的重要作用。本文对植物根构型的概念及其定量分析研究进展进行了概述,并介绍了植物根构型的主要研究方法和定量分析技术。     

植物根构型的定量分析

植物根系具有锚定植株、吸收和运输土壤中的水分及养分、合成和贮藏营养物质等重要功能。根构型是根系在土壤中的空间造型和分布。对植物根构型进行定量分析, 有助于人们了解根系结构和根系功能在生态系统中的重要作用。本文对植物根构型的概念及其定量分析研究进展进行了概述, 并介绍了植物根构型的主要研究方法和定量分析技术。     

Adaptive Changes and Genotypic Variation for Root Architecture of Common Bean in Response to Phosphorus Deficiency

Root architectural responses to phosphorus (p) availability may be an important trait for P acquisition efficiency. In the present study, The authors examined the effects of P availability on root architectural responses of different common bean genotypes. Five common bean (Phaseolus vulgaris L.) genotypes representing different origins and ecotypic races were compared both in a specially designed paper pouch system and a stratified P buffer sand culture system with computer image analysis. The results showed that root architecture was regulated by P availability. P deficiency led to form a shallower root system, as indicated by increased relative distribution of basal root length in the upper layers and decreased the growth angle of basal roots. There was significant genetic variation in root architecture in response to P deficiency both in the paper pouch system and the stratified sand culture system. Under low P conditions some genotypes were more gravitropically sensitive to low P availability, resulting in producing a shallower root system and enhanced root exploration into the surface soil, where soil available P is more concentrated. G19833 and DOR364, which were most contrasting in P efficiency, were also very different in root architectural response to P availability. The results from this study suggest that P availability regulates root architecture and P deficiency leads to shallower root architecture in beans. The genetic potential of root architecture provides the possibility of selecting this trait for improving P acquisition efficiency in common bean.     

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.     

Topsoil foraging – an architectural adaptation of plants to low phosphorus availability

Low phosphorus availability is a primary constraint to plant productivity in many natural and agricultural ecosystems. Plants display a wide array of adaptive responses to low phosphorus availability that generally serve to enhance phosphorus mobility in the soil and increase its uptake. One set of adaptive responses is the alteration of root architecture to increase phosphorus acquisition from the soil at minimum metabolic cost. In a series of studies with the common bean, work in our laboratory has shown that architectural traits that enhance topsoil foraging appear to be particularly important for genotypic adaptation to low phosphorus soils (phosphorus efficiency). In particular, the gravitropic trajectory of basal roots, adventitious rooting, the dispersion of lateral roots, and the plasticity of these processes in response to phosphorus availability contribute to phosphorus efficiency in this species. These traits enhance the exploration and exploitation of shallow soil horizons, where phosphorus availability is greatest in many soils. Studies with computer models of root architecture show that root systems with enhanced topsoil foraging acquire phosphorus more efficiently than others of equivalent size. Comparisons of contrasting genotypes in controlled environments and in the field show that plants with better topsoil foraging have superior phosphorus acquisition and growth in low phosphorus soils. It appears that many architectural responses to phosphorus stress may be mediated by the plant hormone ethylene. Genetic mapping of these traits shows that they are quantitatively inherited but can be tagged with QTLs that can be used in plant breeding programs. New crop genotypes incorporating these traits have substantially improved yield in low phosphorus soils, and are being deployed in Africa and Latin America.     

Genotypic Variation in Root Morphological Characteristics of Common Bean in Relation to Phosphorus Efficiency (in English)

Root morphology in plants may be related to phosphorus (P) efficiency by affecting the absorption characteristics of the root system. However, genotypic variation in root morphological characteristics of common bean (Phaseolus vulgaris L.) as affected by P availability has not been well clarified. In the present study, systematic studies were conducted in a P-buffered sand culture system using three pairs of common bean parental materials with contrasting root traits in response to P deficiency. The results indicate that P availability significantly affects bean root morphology. Common bean tends to have smaller root system, shorter and coarser roots at low P availability. Genotypic variation in root morphology was observed among different genotypes in response to P availability. The P efficient genotypes appear to have larger, finer and longer root systems than the P inefficient genotypes, and such a variation was particularly obvious in the basal roots. From allomeric analysis, we found that morphological characteristics of the basal roots contribute more to P efficiency than those of the tap roots. Further studies with the F9 recombinant inbred lines derived from one of the most contrasting parental pairs, DOR364 and G19833, confirmed the above findings, indicating that those morphological characteristics are inheritable hence provide potential for genetic improvement. Root morphology in plants may be related to phosphorus (P) efficiency by affecting the absorption characteristics of the root system. However, genotypic variation in root morphological characteristics of common bean (Phaseolus vulgaris L.) as affected by P availability has not been well clarified. In the present study, systematic studies were conducted in a P-buffered sand culture system using three pairs of common bean parental materials with contrasting root traits in response to P deficiency. The results indicate that P availability significantly affects bean root morphology. Common bean tends to have smaller root system, shorter and coarser roots at low P availability. Genotypic variation in root morphology was observed among different genotypes in response to P availability. The P efficient genotypes appear to have larger, finer and longer root systems than the P inefficient genotypes, and such a variation was particularly obvious in the basal roots. From allomeric analysis, we found that morphological characteristics of the basal roots contribute more to P efficiency than those of the tap roots. Further studies with the F9 recombinant inbred lines derived from one of the most contrasting parental pairs, DOR364 and G19833, confirmed the above findings, indicating that those morphological characteristics are inheritable hence provide potential for genetic improvement.     

Rhizoeconomics: Carbon costs of phosphorus acquisition

Plants display a wide array of physiological adaptations to low soil phosphorus availability. Here we discuss metabolic and ecological costs associated with these strategies, focusing on the carbon costs of root traits related to phosphorus acquisition in crop plants. We propose that such costs are an important component of adaptation to low phosphorus soils. In common bean, genotypes with superior low phosphorus adaptation express traits that reduce the respiratory burden of root growth, including greater allocation to metabolically inexpensive root classes, such as adventitious roots, and greater formation of cortical aerenchyma, which reduces specific root respiration. Root hair formation increases phosphorus acquisition at minimal carbon cost, but may have other unknown ecological costs. Mycorrhizas and root exudates enhance phosphorus acquisition in some taxa, but at significant carbon cost. Root architectural patterns that enhance topsoil foraging enhance phosphorus acquisition but appear to incur tradeoffs for water acquisition and spatial competition. A better understanding of the metabolic and ecological costs associated with phosphorus acquisition strategies is needed for an intelligent deployment of such traits in crop improvement programs.     

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.     

菜豆根形态特性的基因型差异与磷效率

应用磷控释砂培以及计算机图象分析技术,研究了磷效率差异显的菜豆（Phaseolus vulgaris L.)亲本及其重组自交系后代的根形态特性及其与磷效率的关系。试验结果表明,供磷状况显影响菜豆根系形态学特性。在低磷胁迫下,菜豆根系总根长变短、根部生物量减少,根直径增大。菜豆根形态特性对低磷有效性的适应性反应具有显的基因型差异。在低磷条件下磷高效率基因型的根系比磷低效率基因型相对根部生物量较大、总根长较长,根表面积较大。异计分析表明,菜豆基根根形态特性在低磷条件下的适应性变化对磷效率的贡献远远大于主根,并且这些适应性变化是可以遗传的,表明通过对菜豆根形态特性进行遗传改良来提高磷效率有一定的可行性。