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.     

植物根毛生长发育及分子调控机理

植物根毛是植物吸收营养的主要器官, 了解根毛的发生、发育及遗传规律, 能对植物的养分吸收研究提供有利依据。文章旨在介绍植物根毛形态发生特性、发育生长过程及分子调控机理的研究进展, 利用比较基因组学方法研究农作物根毛形态和功能, 及有目的性的对根生长发育进行调控提供参考。研究发现植物根毛发育有反馈侧向抑制(lateral inhibition with feedback)和位置决定模式(position-dependent pattern of cell differentiation)两种方式。拟南芥根表皮细胞是以位置方式决定毛或非毛细胞发育类型, 已成为研究植物细胞命运和分化的模型。目前, 已经鉴定出控制根毛发育的基因, 包括一些转录因子如MYB家族蛋白TRIPTYCHON(TRY)、CAPRICE(CPC)和basic Helix-Loop-Helix (bHLH)蛋白GLABRA3、ENHANCER OF GLABRA3(EGL3)及WD-repeat蛋白等基因。最后针对根毛研究前景提出展望。     

根毛的生长发育及其遗传基础

根毛是根系特异化表皮细胞外伸形成的管状凸起物,是植物吸收矿质养分和水分的重要器官。根毛的发育可分为根毛细胞命运决定、根毛起始、根毛顶端生长和根毛成熟等阶段。本文对根毛发育生长过程中的细胞形态及其生理生化变化进行了综述,并从根表皮细胞命运决定分子机理, EXPANSIN、bHLH和MYB等转录因子以及小G蛋白和生长素/乙烯等方面简要说明了根毛生长发育的遗传基础。     

植物根毛发育调控机制的研究进展

根毛是植物根表皮细胞的管状延伸结构,在植物固着土壤、吸收水分和无机盐,以及协助植物根部和外界进行信息交流等过程中起到十分重要的作用。植物根毛的发育过程具有很强的可塑性,多种植物激素和环境因素都可以影响植物根毛的发育过程。得益于根毛结构和功能的特点,其也常被作为研究植物细胞顶端生长和命运分化的模式对象。因而,根毛的发育调控机制一直是植物学研究领域的热点。该文梳理了近20年来植物根毛发育调控领域的研究进展。     

菌根真菌与吸收根功能性状的关系：研究进展与评述北大核心CSCD

吸收根(absorption root)一般是指根枝系统末端少数几级具有初生结构、负责物质吸收的根。吸收根功能性状被广泛用于评价和预测植物个体到生态系统水平上的一系列功能和过程。菌根真菌侵染是吸收根的一个关键性状,它可以深刻影响吸收根的形态、结构,以及功能性状之间的关系。该文针对与吸收功能密切相关的菌根真菌与根毛和根直径之间的关系进行了研究综述,提出了真菌侵染、根毛和化学防御之间关系的一个假说;探讨了温带和热带不同类型的吸收根如何通过菌根真菌影响根的功能性状,从而适应不同的水热条件、养分状况和能量消耗;提出一些需要关注的议题和研究方向,以期为菌根真菌与吸收根功能性状之间关系的研究提供借鉴。     

Relationship between mycorrhizal fungi and functional traits in absorption roots: research progress and synthesis

吸收根(absorption root)一般是指根枝系统末端少数几级具有初生结构、负责物质吸收的根。吸收根功能性状被广泛用于评价和预测植物个体到生态系统水平上的一系列功能和过程。菌根真菌侵染是吸收根的一个关键性状, 它可以深刻影响吸收根的形态、结构, 以及功能性状之间的关系。该文针对与吸收功能密切相关的菌根真菌与根毛和根直径之间的关系进行了研究综述, 提出了真菌侵染、根毛和化学防御之间关系的一个假说; 探讨了温带和热带不同类型的吸收根如何通过菌根真菌影响根的功能性状, 从而适应不同的水热条件、养分状况和能量消耗; 提出一些需要关注的议题和研究方向, 以期为菌根真菌与吸收根功能性状之间关系的研究提供借鉴。     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

增加植物根毛长度可提高作物产量

面对气候变化，让作物更有效率地吸收营养和水从而增加产量显得越发重要。英国科学家揭示了如何增加植物根毛长度，而具较长根毛的植物能更有效地吸收水和养分，从而可能提高作物产量。     

丛枝菌根真菌与氮添加对不同根形态基因型水稻氮吸收的影响

植物主要依赖自身根系从土壤中获取矿质养分; 具有不同根形态的植物对于养分的吸收能力存在差异。丛枝菌根真菌(AMF)能与陆地植物根系形成共生关系, 帮助植物吸收矿质养分。但是, AMF对于植物根系养分吸收的促进效应是否会受根形态的影响还鲜有研究。该研究选取4种不同根形态基因型水稻(根毛缺陷突变体rhl1、侧根缺陷突变体iaa11、不定根缺失突变体arl1和野生型Kas)为研究对象, 设置2种施氮水平处理(低氮: 20 mg·kg^-1氨氮; 高氮: 100 mg·kg^-1氨氮), 利用稳定同位素¹⁵N示踪标记技术, 探究AMF和氮添加对不同根形态植物氮吸收的影响。研究结果发现, 相比低氮处理, 高氮处理下, rhl1、Kas、iaa11与arl1的茎叶¹⁵N浓度分别提高了60%、72%、128%与118%, 说明氮添加显著促进了水稻氮吸收, 且iaa11与arl1对氮添加的响应更强烈。在低氮水平下, AMF对rhl1、Kas、iaa11与arl1氮吸收的平均效应值分别为17%、31%、42%、51%, 表明AMF对于植物氮吸收的促进效应受根形态影响, iaa11与arl1对AMF的响应明显高于Kas与rhl1; 相较于低氮水平, 高氮水平下AMF对于不同根形态水稻氮吸收的促进效应都会显著降低, 表明氮添加削弱了AMF对植物氮吸收的促进效应。该研究阐明了4种不同根形态基因型水稻氮养分吸收存在显著差异, 其中氮吸收能力较弱的基因型水稻对AMF的响应更强, 该结果补充了植物与AMF在养分吸收上存在功能互补的控制实验证据。     

葡萄试管苗根形态学和解剖学研究

葡萄试管苗不定根形态解剖观察表明,试管苗根的形态结构与正常根有明显差异:无根冠和根毛;侧根少,以不定根为主要吸收面积;外皮层细胞形大,壁薄,质浓,代替了根毛的吸收功能;内皮层具有初生特征;维管束发育延迟等。为此,本文讨论了试管苗根解剖学特点与移栽成活率的关系。     

Conserved and unique features of the maize (Zea mays L.) root hair proteome

Root hairs are unicellular extensions of specialized epidermis cells. Under limiting conditions, they significantly increase the water and nutrient uptake capacity of plants by enlarging their root surface. Thus far, little is known about the initiation and growth of root hairs in the monocot model species maize. To gain a first insight into the protein composition of these specialized cells, the 2573 most abundant proteins of maize root hairs attached to four-day-old primary roots of the inbred line B73 were identified by combining 1DE with nanoLC-MS/MS in a shotgun proteomic experiment. Among the identified proteins, homologues of 252 proteins have been previously associated with root hair formation and development in other species. Comparison of the root hair reference proteome of the monocot species maize with the previously published root hair proteome of the dicot species soybean revealed conserved, but also unique, protein functions in root hairs of these two major groups of flowering plants.     

Proteoid root morphology and function inLupinus albus

Summary Current theories of phosphorus uptake by plants imply that they can augment diffusion to their root axes by the development of abundant root hairs or mycorrhizas. Some phosphorus efficient plants have root morphology with multi-branched roots and localised regions of densely packed root hairs, which we suggest is better suited to the retention of substances exuded by the roots than uptake of substances moving to the root by diffusion. Evidence of substantial exudation by the proteoid roots ofLupinus albus is presented.     

Identification of legume RopGEF gene families and characterization of a Medicago truncatula RopGEF mediating polar growth of root hairs

Root hairs play important roles in the interaction of plants with their environment. Root hairs anchor the plant in the soil, facilitate nutrient uptake from the rhizosphere, and participate in symbiotic plant-microbe interactions. These specialized cells grow in a polar fashion which gives rise to their elongated shape, a process mediated in part by a family of small GTPases known as Rops. RopGEFs (GEF, guanine nucleotide exchange factor) activate Rops to effect tip growth in Arabidopsis pollen and root hairs, but the genes mediating tip growth in legumes have not yet been characterized. In this report we describe the Rop and RopGEF gene families from the model legume Medicago truncatula and from the crop legume soybean. We find that one member of the M. truncatula gene family, MtRopGEF2, is required for root hair development because silencing this gene by RNA interference affects the cytosolic Ca2+ gradient and subcellular structure of root hairs, and reduces root hair growth. Consistent with its role in polar growth, we find that a GFP::MtRopGEF2 fusion protein localizes in the apex of emerging and actively growing root hairs. The amino terminus of MtRopGEF2 regulates its ability to interact with MtRops in yeast, and regulates its biological activity in vivo.     

The effect of the nutrient intensity and buffering power of a soil,and the absorbing power,size and root hairs of a root,on nutrient absorption by diffusion

Summary A portion of a single plant root is treated as an absorbing cylindrical sink to which nutrients move by diffusion. Assuming that the rate of uptake of nutrient is proportional to its concentration at the root surface, and that the nutrient, though reacting with the solid, moves only through the soil solution, standard diffusion equations are used to calculate the effect of soil and plant characteristics on the rate of uptake. The treatment is applicable to phosphorus and potassium. Among soil properties uptake should increase directly with the soil solution concentration. It should also increase, but only slowly, with increasing buffering power. It increases with increasing soil moisture. Among plant characteristics, uptake should increase with the root absorbing power until diffusion through the soil becomes limiting. Absorption by unit surface area of root increases as the root radius decreases. A root hair is shown to interfere quickly with the uptake of adjacent hairs. The hairs increase absorption by the root because they can exploit rapidly the soil between the hairs, and they have the effect of extending the effective root surface to their tips.     

OsCSLD1, a cellulose synthase-like D1 gene, is required for root hair morphogenesis in rice

Root hairs are long tubular outgrowths that form on the surface of specialized epidermal cells. They are required for nutrient and water uptake and interact with the soil microflora. Here we show that the Oryza sativa cellulose synthase-like D1 (OsCSLD1) gene is required for root hair development, as rice (Oryza sativa) mutants that lack OsCSLD1 function develop abnormal root hairs. In these mutants, while hair development is initiated normally, the hairs elongate less than the wild-type hairs and they have kinks and swellings along their length. Because the csld1 mutants develop the same density and number of root hairs along their seminal root as the wild-type plants, we propose that OsCSLD1 function is required for hair elongation but not initiation. Both gene trap expression pattern and in situ hybridization analyses indicate that OsCSLD1 is expressed in only root hair cells. Furthermore, OsCSLD1 is the only member of the four rice CSLD genes that shows root-specific expression. Given that the Arabidopsis (Arabidopsis thaliana) gene KOJAK/AtCSLD3 is required for root hair elongation and is expressed in the root hair, it appears that OsCSLD1 may be the functional ortholog of KOJAK/AtCSLD3 and that these two genes represent the root hair-specific members of this family of proteins. Thus, at least part of the mechanism of root hair morphogenesis in Arabidopsis is conserved in rice.     

A dual porosity model of nutrient uptake by root hairs

? The importance of root hairs in the uptake of sparingly soluble nutrients is understood qualitatively, but not quantitatively, and this limits efforts to breed plants tolerant of nutrient-deficient soils. ? Here, we develop a mathematical model of nutrient uptake by root hairs allowing for hair geometry and the details of nutrient transport through soil, including diffusion within and between soil particles. We give illustrative results for phosphate uptake. ? Compared with conventional 'single porosity' models, this 'dual porosity' model predicts greater root uptake because more nutrient is available by slow release from within soil particles. Also the effect of soil moisture is less important with the dual porosity model because the effective volume available for diffusion in the soil is larger, and the predicted effects of hair length and density are different. ? Consistent with experimental observations, with the dual porosity model, increases in hair length give greater increases in uptake than increases in hair density per unit main root length. The effect of hair density is less in dry soil because the minimum concentration in solution for net influx is reached more rapidly. The effect of hair length is much less sensitive to soil moisture.     

Nitrate induction of root hair density is mediated by TGA1/TGA4 and CPC transcription factors in Arabidopsis thaliana

Root hairs are specialized cells that are important for nutrient uptake. It is well established that nutrients such as phosphate have a great influence on root hair development in many plant species. Here we investigated the role of nitrate on root hair development at a physiological and molecular level. We showed that nitrate increases root hair density in Arabidopsis thaliana. We found that two different root hair defective mutants have significantly less nitrate than wild‐type plants, suggesting that in A. thaliana root hairs have an important role in the capacity to acquire nitrate. Nitrate reductase‐null mutants exhibited nitrate‐dependent root hair phenotypes comparable with wild‐type plants, indicating that nitrate is the signal that leads to increased formation of root hairs. We examined the role of two key regulators of root hair cell fate, CPC and WER, in response to nitrate treatments. Phenotypic analyses of these mutants showed that CPC is essential for nitrate‐induced responses of root hair development. Moreover, we showed that NRT1.1 and TGA1/TGA4 are required for pathways that induce root hair development by suppression of longitudinal elongation of trichoblast cells in response to nitrate treatments. Our results prompted a model where nitrate signaling via TGA1/TGA4 directly regulates the CPC root hair cell fate specification gene to increase formation of root hairs in A. thaliana.     

Heterogeneous nutrient supply promotes maize growth and phosphorus acquisition: additive and compensatory effects of lateral roots and root hairs

Background and AimsRoot proliferation is a response to a heterogeneous nutrient distribution. However, the growth of root hairs in response to heterogeneous nutrients and the relationship between root hairs and lateral roots remain unclear. This study aims to understand the effects of heterogeneous nutrients on root hair growth and the trade-off between root hairs and lateral roots in phosphorus (P) acquisition.MethodsNear-isogenic maize lines, the B73 wild type (WT) and the rth3 root hairless mutant, were grown in rhizoboxes with uniform or localized supply of 40 (low) or 140 (high) mg P kg⁻¹ soil.ResultsBoth WT and rth3 had nearly two-fold greater shoot biomass and P content under local than uniform treatment at low P. Significant root proliferation was observed in both WT and rth3 in the nutrient patch, with the WT accompanied by an obvious increase (from 0.7 to 1.2 mm) in root hair length. The root response ratio of rth3 was greater than that of WT at low P, but could not completely compensate for the loss of root hairs. This suggests that plants enhanced P acquisition through complementarity between lateral roots and root hairs, and thus regulated nutrient foraging and shoot growth. The disappearance of WT and rth3 root response differences at high P indicated that the P application reduced the dependence of the plants on specific root traits to obtain nutrients.ConclusionsIn addition to root proliferation, the root response to a nutrient-rich patch was also accompanied by root hair elongation. The genotypes without root hairs increased their investment in lateral roots in a nutrient-rich patch to compensate for the absence of root hairs, suggesting that plants enhanced nutrient acquisition by regulating the trade-off of complementary root traits.