土壤解磷微生物促进植物磷素吸收策略研究进展

随着全球磷肥需求增加和磷矿资源储量短缺矛盾逐渐加重,土壤解磷微生物在促进磷循环方面的重要性日益突显,因此有必要对其种类和促进植物磷素吸收策略进行全面梳理总结。荟萃分析了土壤解磷微生物种类并构建了系统进化树,重点论述了土壤解磷微生物促进植物磷素吸收主要策略。土壤解磷微生物主要通过矿化和溶解作用直接活化难溶性磷,但其也能与植物根系互作间接活化磷素。其间接途径主要包括解磷微生物与根系分泌物和丛枝菌根真菌互作,它们通过碳磷交换间接活化磷素;其次包括一些解磷微生物可以通过固氮作用使植物生长受到磷的限制,从而调节植物磷酸盐转运系统间接活化土壤磷;解磷微生物也能通过分泌植物激素和生物防治剂促进植物根系生长间接促进植物磷素吸收。解磷微生物还能通过磷素固定减少磷流失,也可以通过加速自身磷素周转促进植物磷素吸收。对完善和发展解磷微生物主导的土壤磷循环和植物磷素吸收利用理论体系具有重要意义。     

菌根真菌促进植物磷吸收研究进展

土壤中低浓度的有效磷水平成为限制植物生长发育的主要因素。植物—真菌菌根共生可以显著提高植物吸收土壤中磷的能力,促进植物生长发育。该文对土壤中磷酸盐的形式、丛枝状菌根和外生菌根两种菌根类型的形态学特征和促磷吸收的发生机制、植物中已克隆的菌根特异性或诱导性磷转运蛋白,以及丛枝状真菌共生信号转导途径等进行了综述。     

植物菌根共生磷酸盐转运蛋白

大多数植物能和丛枝菌根（arbuscular mycorrhiza, AM）真菌形成菌根共生体。AM能够促进植物对土壤中矿质营养的吸收,尤其是磷的吸收。磷的吸收和转运由磷酸盐转运蛋白介导。总结了植物AM磷酸盐转运蛋白及其结构特征,分析其分类及系统进化,并综述了AM磷酸盐转运蛋白介导的磷的吸收和转运过程及其基因的表达调控。植物AM磷酸盐转运蛋白属于Pht1家族成员,它不仅对磷的吸收和转运是必需的,而且对AM共生也至关重要,为进一步了解菌根形成的分子机理及信号转导途径提供了理论基础。     

植物磷稳态的调控机制

磷是植物必需的重要营养元素之一,是生物大分子的重要组成部分,在植物生命过程中发挥着不可或缺的作用。维持体内磷稳态对于植物的生长发育和环境应答至关重要。多种信号分子参与调控植物对磷的吸收和转运。植物维持磷稳态主要包括土壤磷的活化、磷的吸收、转运、存储和再利用等过程,涉及磷胁迫响应、转录因子调节、miRNA调节、菌根共生、细胞器间转移等磷调节机制。未来的磷营养机制研究需要跨学科知识的融合,由模式植物研究转向栽培作物。全面总结了植物细胞磷吸收和转运的核心分子及其作用机制的研究进展,旨在为作物品种改良和遗传育种提供重要借鉴。     

VA菌根降低植物对重金属镉的吸收

高等植物在漫长的进化过程中对环境产生种种适应机制。菌根的形成即是对自然土壤中有效磷不足的一种适应。菌根真菌与寄主根系共生形成菌根后,真菌的菌丝可以远远伸出根际范围从而扩大了植物对土壤中难以移动的磷元素的吸收范围而改善植物的磷素营养。因此,地球上90％的陆生植物都可形成菌根。菌根的形成,不仅促进了植物对磷的吸收,而且也影响到植物对其它元素包括重金属的吸收。在重金属污染的土壤中,菌根对植物重金属的吸收将影响到植物对重金属的抗性和农产品品质。本文拟研究在添加镉的土壤上菌根对植物吸收Cd的影响。     

植物磷营养状况对丛枝菌根真菌生长及代谢活性的调控*

采用四室隔网培养装置,以玉米为宿主植物,通过在植物生长室设0、50、250和500 mgPkg-1 4个施磷水平,研究了植株体内的磷营养状况对AM真菌Glomus sinuosum和Glomus intraradices生长及活性的影响。研究发现在不施磷条件下,接种AM真菌G. intraradices显著促进了植物生长和磷的吸收;低磷条件(50 mgPkg-1)下,接种菌根真菌显著促进了植物对磷的吸收,但对植物生长没有明显的影响;而在高磷条件(250 mgPkg-1 和500 mgPkg-1)下,接种菌根真菌不但没有促进植物的生长和磷的吸收,反而对其有抑制作用。随着施磷水平的提高, AM真菌根内菌丝的碱性磷酸酶活性显著下降;与不施磷相比,低量(50 mgPkg-1)供磷增加了AM真菌土壤中根外菌丝的密度,高磷(250 mgPkg-1 和500 mgPkg-1)降低了土壤中根外菌丝的密度。上述结果说明:⑴ 给宿主植物施用磷肥引起的植物磷营养状况的改变,对AM真菌生长和代谢活性具有一定的调控作用;⑵ G. sinuosum和G. intraradices两种AM真菌的生长和代谢活性对施磷水平的响应程度无显著性差异;⑶ 高磷抑制AM真菌生长和代谢活性,使真菌吸磷量减少,可能是造成菌根效应降低的原因之一     

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。     

菌根菌的作用

相当多的植物的根系可与某些真菌共生,形成菌根。菌根植物比非菌根植物有几个显著优势:①菌根菌可代替根毛的作用,但菌丝体的扩展范围比根毛大得多,用放射性磷进行试验,证明菌根菌吸收磷的范围在根周围直径可达8-20厘米。因而扩大了吸收各种元素的范围;②促进了各种矿物质的吸收,特别是土壤中难     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响, 尤其是加剧了植物生长的磷限制, 从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响, 认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量, 有利于植物的磷吸收并增加植物磷含量。同时, 森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性, 该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制: 1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力; 3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述, 外源氮输入使植物磷含量发生改变, 首要原因是土壤有效磷含量的改变, 其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响,尤其是加剧了植物生长的磷限制,从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响,认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量,有利于植物的磷吸收并增加植物磷含量。同时,森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性,该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制:1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力;3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述,外源氮输入使植物磷含量发生改变,首要原因是土壤有效磷含量的改变,其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

Influence of aquatic macrophytes on phosphorus cycling in lakes

Emergent macrophytes take up their phosphorus exclusively from the sediment. Submerged species obtain phosphorus both from the surrounding water and from the substrate, but under normal pore and lake water phosphorus concentrations, substrate uptake dominates. Release of phosphorus from actively growing macrophytes (both submerged and emergent) is minimal and epiphytes obtain phosphorus mainly from the water. Decaying macrophytes may act as an internal phosphorus source for the lake and add considerable quantities of phosphorus to the water. A large part of the released phosphorus is often retained by the sediments. In perennial macrophytes the amount of phosphorus released from decaying shoots is dependent on the degree of phosphorus conservation within the plant. Macrophyte stands may also be a permanent phosphorus sink due to burial of plant litter. Macrophytes affect the chemical environment (oxygen, pH), which in turn has effects on the phosphorus cycling in lakes. However, the impact of aquatic macrophytes on whole-lake phosphorus cycling is largely unknown. Controlled full-scale harvesting, herbicide or herbivory experiments are almost totally lacking. Emergent macrophytes respond positively to eutrophication, but fertilization experiments have shown that nitrogen rather than phosphorus may be the key element. Submerged macrophytes are adversely affected by a large increase in the external phosphorus input to a lake. This effect may be caused by epiphyte shading, phytoplankton shading or deposition of unfavourable sediments.     

Benthic phosphorus regeneration in the Potomac River Estuary

The flux of dissolved reactive phosphate from Potomac riverine and estuarine sediments is controlled by processes occurring at the water-sediment interface and within surficial sediment.In situ benthic fluxes (0.1 to 2.0 mmoles m⁻² day⁻¹) are generally five to ten times higher than calculated diffusive fluxes (0.020 to 0.30 mmoles m⁻² day⁻¹). The discrepancy between the two flux estimates is greatest in the transition zone (river mile 50 to 70) and is attributd to macrofaunal irrigation. Bothin situ and diffusive fluxes of dissolved reactive phosphate from Potomac tidal river sediments are low while those from anoxic lower estuarine sediments are high. The net accumulation rate of phosphorus in benthic sediment exhibits an inverse pattern. Thus a large fraction of phosphorus is retained by Potomac tidal river sediments, which contain a surficial oxidized layer and oligochaete worms tolerant of low oxygen conditions, and a large fraction of phosphorus is released from anoxic lower estuary sediments. Tidal river sediment pore waters are in equilibrium with amorphous Fe (OH)₃ while lower estuary pore waters are significantly undersaturated with respect to this phase. Benthic regeneration of dissolved reactive phosphorus is sufficient to supply all the phosphorus requirements for net primary production in the lower tidal river and transition-zone waters of the Potomac River Estuary. Benthic regeneration supplies approximately 25% as much phosphorus as inputs from sewage treatment plants and 10% of all phosphorus inputs to the tidal Potomac River. When all available point source phosphorus data are put into a steady-state conservation of mass model and reasonable coefficients for uptake of dissolved phosphorus, remineralization of particulate phosphorus, and sedimentation of particulate phosphorus are used in the model, a reasonably accurate simulation of dissolved and particulate phosphorus in the water column is obtained for the summer of 1980.     

Aspects of the phosphorus cycle in Hartbeespoort Dam (South Africa) Phosphorus loading and seasonal distribution of phosphorus in the reservoir

Hartbeespoort Dam, a hypertrophic, warm monomictic impoundment in South Africa, receives extremely high phosphorus loads (14.6–25.9 g m^–2 a^–1) that are dominated by point source discharges from municipal wastewater treatment works. The reduced state of the phosphorus discharged from the works has led to the dominance of the dissolved phosphorus pool by low molecular weight orthophosphates which are analytically detectable as soluble reactive phosphorus (SRP; 60% of total phosphorus pool). Seasonality in the in-lake total phosphorus pool is regulated by a combination of abiotic and hydrological processes; biotic processes appear to play a minor role. Mass balance calculations indicate that between 62 and 77% of the annual total phosphorus inflow load is retained within the impoundment each year.     

岩溶石漠化区不同植被恢复模式土壤无机磷形态特征及影响因素

为分析岩溶石漠化区不同植被恢复模式土壤无机磷的形态特征,评价植被恢复的土壤供磷潜力,阐明有机碳及钙素在无机磷形态转化中的作用,选取研究区内8种有代表性的样地,采用蒋柏藩无机磷分级方法对土壤无机磷形态特征及影响因素进行研究。结果表明:研究区土壤全磷与速效磷含量分别在0.25—1.35 g/kg、1.05—53.01 mg/kg范围,无机磷总量在123.94—934.61 mg/kg,耕地与退耕地以及各退耕地之间全磷、速效磷、各形态无机磷含量水平差异明显,各退耕地磷素含量水平介于耕地与次生马尾松林地之间,退耕地中桃林地、花椒林地磷素含量水平较高、樟树林地、柳杉林地、撂荒草地次之、撂荒灌丛地较低。各样地土壤无机磷占全磷比例在51.2%—72.4%,不同形态的无机磷含量表现为O-PFe-PCa-PAl-P,其中Ca2-P、Al-P对速效磷的贡献率大,Fe-P、Ca8-P贡献较小,O-P、Ca10-P献率最小。不同活性土壤有机碳与不同形态钙素对各形态无机磷在总无机磷中比例的影响较大,p H、容重、粘粒含量、含水量等其它理化性质影响较小。     

社会经济系统磷物质流分析--以安徽省含山县为例

氮、磷等营养物质过量输入是造成我国湖泊富营养化问题日益严重的根源,磷作为水体富营养化过程关键限制元素,主要来自流域社会经济系统中的人类活动排放,因此,定量刻画社会经济系统内的磷流动路径是追踪水体外源磷来源和进行有效控制磷排放量的前提。以巢湖流域的安徽省含山县为例,构建社会经济系统磷流分析框架,建立磷流核算模型,并在实地调查和数据统计分析的基础上定量刻画了含山县2008年度社会经济系统磷流路径。结果表明,2008年含山县社会经济系统向水体排放的磷总量为1592t,其中农业种植子系统的排放所占比例最大(77%),该子系统的磷利用效率也较低(45%)。因此,含山县富营养化治理的重点是优化农业种植系统的磷流路径,主要措施包括合理施肥、科学排灌等。     

II. Quantitative aspects of phosphorus absorption in ruminants

Phosphorus absorption in ruminants was analysed from a database described in a previous article. For common values of ingested phosphorus (2.5-5.0 g x kg(-1) of DM), 0.73 of dietary phosphorus is absorbed. The remaining variability is probably due to phosphorus quality. Phosphorus absorbed from silage, cereal, cereal by-products and hay differs greatly. The current true absorption coefficient used to calculate daily phosphorus supply is a constant value in the current systems and often it underestimates the true absorption resulting in an excess of phosphorus being supplied in the diets. Adjusting the true absorption coefficient values requires better characterisation of the phosphorus supplied by each feedstuff. Dietary influences (phytate phosphorus, crude fibre, etc.) were investigated but trials assessing the ration effect on phosphorus absorption are lacking. Since rumen microbes have specific phosphorus requirements, particularly for cell-wall degradation, the feedstuffphosphorus availability for the rumen ecosystem is discussed.     

Sources and bioavailability of phosphorus fractions in freshwaters: a British perspective

This paper seeks a perspective on the forms of phosphorus which promote aquatic eutrophication, with the particular quest of establishing their sources. A short background traces the development of understanding of nutrient enrichment and the suppositions about the relative contributions of agriculture, sewage and detergent residues. Most aquatic systems, and their primary producers, are naturally deficient in biologically-available phosphorus. Aquatic plants have evolved very efficient phosphorus uptake mechanisms. The biomass responses to an increase in the supply of phosphorus are stoichiometrically predictable. The most bioavailable forms of phosphorus are in solution, as orthophosphate ions, or are readily soluble or elutable from loose combinations. Ready bioavailability coincides well with what is measurable as molybdate-reactive (MRP) or soluble-reactive phosphorus (SRP). Most other forms, including phosphates of the alkaline earth metals, aluminium and iron are scarcely available at all. Orthophosphate ions sorbed to metal oxides and hydroxides are normally not biologically available either, except through weak dissociation ('desorption'). The production of alkaline phosphatase provides organisms with an additional mechanism for accelerating the sequestration of phosphate from organic compounds. Bioavailable phosphate is liberated when redox- or alkali-sensitive metal hydroxides dissolve but these processes are minor contributors to the biological responses to nutrient enrichment. Most of the familiar eutrophication is attributable to the widespread application of secondary sewage treatment methods to the wastes emanating from a burgeoning and increasingly urbanised human population. The use of polyphosphate-based detergents, now in decline, has contributed to the problem. In aquatic systems, the additional phosphorus raises the biological supportive capacity, sometimes to the capacity of the next limiting factor (carbon, light, hydraulic retention or of another nutrient). At high orthophosphate loadings, the straight stoichiometric yield relationship between biomass yield and phosphorus a vailability is lost. Movements of phosphorus and its recycling within aquatic systems do not prevent the slow gravitation of phosphorus to the bottom substrata. The phosphorus retentivity of sediments depends upon their chemical composition. While oxide-hydroxide binding capacity in the surface sediments persists, they act as a sink for phosphorus and a control on further cycling. Iron-rich and clay-rich sediments perform best in these conditions; calcareous sediments least so. Eutrophication may lead to the exhaustion of sediment P-binding capacity. Non-sorbed phosphate is readily recyclable if primary producers have access to it. Recycling is most rapid in shallow waters (where sediment disturbance, by flow, by wind action and through bioturbation, is frequent and least in deep ventilated sediments. The contributions of phosphorus from catchments are assessed. The slow rate of weathering of (mostly apatitic) minerals, the role of chemical binding in soils and the incorporation and retentivity bv forested terrestrial ecosystems each contribute to the minimisation of phosphorus leakage to drainage waters. Palaeolimnological and experimental evidence confirms that clearance of land and ploughing its surface weakens the phosphorus retentivity of catchments. The phosphorus transferred from arable land to drainage remains dominated by sorbed fractions which are scarcely bioavailable. Some forms of intensive market gardening or concentrated stock rearing may mobilise phosphates to drainage but it is deduced that drainage from agricultural land is not commonly a major source of readily bioavailable phosphorus in water. Careful budgeting of the phosphates in run-off from over-fertilised soils may nevertheless show that a proportionately small loss of bioavailable phosphorus can still be highly significant in promoting aquatic plant production. The bioavailable-phosphorus (BAP) load achieving the OECD threshold of lake eutrophy (35 mg P m(-3)) is calculated to be equivalent to a terrestrial loss rate of approximately 17.5 kg BAP km(-2) year(-1)), or only 1-2% of a typical fertiliser application. The output is shown to be comparable with the P yield from secondary treatment of the sewage produced by a resident population of 30-44 persons km(-2). With tertiary treatment, the equivalence is with approximately 200 persons km(-2).     

Tissue culture studies; the relationship of incorporation of radioactive phosphorus in nucleic acid to the growth of cells in vitro

Growing cultures are shown to incorporate more radioactive phosphorus into ribo- and desoxyribonucleic acid than non-growing cultures. Strong indications are found that the incorporation of inorganic phosphorus into desoxyribonucleic acid parallels the mitotic coefficient and serves as a sensitive index of synthesis of new cells. There are indications that the phosphorus compounds from which the DNA molecule was synthesized were in equilibrium with the inorganic phosphorus of the medium. The effect of the implant upon growth of new cells and the assay of growth is discussed. Metabolic turnover was increased in adequate medium.     

Nuisance cyanobacteria in an urbanized impoundment: interacting internal phosphorus loading, nitrogen metabolism, and polymixis

Severe nuisance blooms of cyanobacteria, mainly Aphanizomenon and Microcystis, historically have plagued polymictic Ford Lake, one of the most productive warm-water sport fishing lakes in Michigan, U.S.A. Biomass development in the lake is known to be limited by phosphorus. Alternative theories ascribed nuisance conditions either to allochthonous inputs or to internal loading of phosphorus from anoxic sediments. From 2003 to 2009, there was a strong linear relationship between allochthonous total phosphorus income and phosphorus retained within the basin. The relationship had a negative intercept, implying negative retention, or positive export, of phosphorus from the lake from May to September. Mass balance calculations at lake inlet and outlet were consistent with rates of sedimentary phosphorus accumulation measured from sediment cores. Release rates of soluble reactive phosphorus from anoxic sediments were half that of allochthonous inputs. However, severe declines in nitrogen to phosphorus ratio developed in the anoxic, nitrate-poor hypolimnion because accumulation of ammonium was only tenfold that of phosphate. The result was a steep decline in ratio of total nitrogen to total phosphorus during July and August throughout the lake after episodic mixing events, followed by and coinciding with development of heterocystous Aphanizomenon populations. Lake sediment composition determined by X-ray fluorescence in addition to results of sediment core experiments indicates that phosphorus release is governed by an iron trap mechanism such that phosphate and iron are released only when both oxygen and nitrate are depleted.     

Phosphorus in soil,water and sediment: an overview

The geochemistry, availability and abundance of different forms of phosphorus in soil, water and sediments are reviewed. The present knowledge of phosphorus pathways in ecosystems and their regulation is discussed.In a drainage basin, anthropogenic phosphorus is brought into the system mainly as fertilizers and detergents. Sewer systems and outwash processes transfer the phosphorus from the terrestrial environment to the aquatic part of the ecosystem where an accumulation occurs in the sediments of the watercourse.A great part of the phosphates in soil is sorbed to soil particles or incorporated into soil organic matter. The release and export of phosphorus from uncultivated soil is a function of the geology and soil composition, but also of the air temperature, precipitation and the hydrological condition, pH etc.The solubility of phosphates is controlled by either sorption-desorption or precipitation-dissolution reactions depending on the environment in the soil or sediments. In soil and sediments with large amounts of iron and aluminium hydrous oxides, sorption-desorption reactions are largely responsible for determining the level of orthophosphate in the solution at equilibrium.Algal availability of phosphorus associated with soil-derived materials present in aquatic systems deserves more research. In addition, processes responsible for transport of phosphorus from cropland to aquatic systems and chemical and microbial transformations of phosphorus in lakes and streams deserve more attention.