植物对有机氮源的利用及其在自然生态系统中的意义

近来大量实验研究表明,许多植物能够在不经矿化的情况下直接吸收、利用环境介质中的生物有机氮,尤其氨基酸类。而且,有些植物利用氨基酸的效率可以与矿质氮源(NH4 、NO3)相当或更高。自然界植物赖以生存的土壤生境中同时存在多种有机氮和矿质氮养分,这是导致植物(至少部分植物)进化产生利用各种不同氮源能力的环境驱动力。土壤中的游离氨基酸尽管含量不高,但其周转快、通量大,理论上可远大于植物的氮需求。尽管植物在与土壤微生物的有机氮源竞争中处于根本性劣势,但土壤中氨基酸的巨大潜在通量和植物相对于微生物的生命周期仍可使植物在长期竞争中获取数量可观的氮。基于植物根对氨基酸的吸收能力、土壤中游离氨基酸库的大小和通量、植物与土壤微生物对氨基酸氮源的竞争以及有关的原位实验结果,近来许多研究者都认为植物有机氮营养在多种生态系统中是重要或潜在重要的。尤其是在一些极地、高山、亚高山、北方针叶林或泰加林生态系统中,由于低温等因素限制有机氮矿化,土壤氨基酸浓度常超过矿质氮(NH4 、NO3-)浓度,氨基酸可能代表着植物的一个主要氮源。认识到现实生态系统中植物对有机氮源利用的重要性意味着传统的矿质营养观念的更新,这将在很大程度上改变人们对某些重要生态过程的理解,并导致对若干生态学中心问题的再认识。研究以森林生态系统为例,阐述了我国开展该领域研究的科学意义和基本框架。     

高等植物对氨基酸态氮的吸收与利用研究进展

植物能够在不经矿化的情况下直接吸收利用环境中的分子态氨基酸.氨基酸作为植物和微生物的优良碳源和氮源,二者对其吸收存在着激烈竞争,氨基酸态氮来源广、半衰期短的特点使其具有巨大的流通量.运用氮同位素示踪方法研究氨基酸对植物的氮营养贡献一直是国内外学者研究的热点,对揭示土壤肥力本质具有重要意义.本文对不同生态系统中氨基酸形态特征、代谢机制及营养贡献进行了简要综述,分析了氨基酸态氮在植物-土壤-微生物系统中的循环机制及生物有效性等方面研究现状和发展趋势,并提出了土壤氨基酸生物有效性环境调控、氨基酸碳-氮代谢及提高农田生态系统有机氮管理等待解决的科学问题.     

草地生态系统中土壤氮素矿化影响因素的研究进展

氮素是各种植物生长和发育所需的大量营养元素之一,也是牧草从土壤吸收最多的矿质元素．土壤中的氮大部分以有机态形式存在,而植物可以直接吸收利用的是无机态氮．这些有机态氮在土壤动物和微生物的作用下。由难以被植物直接吸收利用的有机态转化为可被植物直接吸收利用的无机态的过程就是土壤氮的矿化．氮素矿化受多种因子的影响,这些因子可以归结为生物因子和非生物因子．生物因子包括：土壤动物、土壤微生物和植物种类．土壤动物可以促进土壤有机质的矿化;土壤微生物种类、结构及功能与氮的分解、矿化有密切的关系;不同的植物种类对土壤氮素的矿化作用是不相同的,一般来说。有豆科植物生长的土壤比其它种类土氮素矿化的作用大．非生物因素一般可以分为环境因子和人类活动干扰．环境因子中土壤温度和含水量对土壤氮素矿化的影响是国内外众多科学家研究的方向．尽管如此,在此方面的研究还没有取得一致意见,仍然需要进行这方面的研究,而在其他诸如：不同的土壤质地与土壤类型方面,研究报道的结论也很不一致,草地生态系统中人类活动对土壤氮素矿化的影响主要包括,不同强度的放牧,割草以及施肥、火烧强度等．非生物因子对氮素矿化的影响非常直接和明显,尤其是人类活动．本文综述了近年来影响草地生态系统土壤氮素矿化有关因素的一些进展．     

土壤可溶性有机氮研究进展

土壤可溶性有机氮是土壤氮素的重要组成部分, 也是氮循环过程中最活跃的因子之一。土壤可溶性有机氮不仅可以直接被植物吸收利用, 也可以被土壤微生物利用。土壤可溶性有机氮可以随土壤水分向下迁移造成农业面源污染等环境问题, 所以引起众多研究者关注。从土壤可溶性有机氮的含量、土壤可溶性有机氮的来源和成分、土壤可溶性有机氮的迁移特征以及土壤可溶性有机氮的影响因素几个方面的研究进展进行综述, 最后对未来土壤可溶性有机氮研究方向进行了展望。     

土壤-植物-大气连续体系中氮的研究进展

氮素是植物最需要的重要养分元素之一.近年来,土壤-植物-大气这一连续体系(SPAC)中的氮循环成为研究的热点之一.大气中的氮素可以通过生物固定和N沉降等作用进入土壤和植物内,同时土壤和植物内的氮素又会以氨挥发和氮氧化物等方式排放到大气中.氮素通过生物固持和植物吸收等方式进入植物体内,植物器官脱落使植物损失一部分的氮素,另外雨水的淋洗和植物溢出液也会造成植物的N损失.植物氮素在植物体内的积累和分布随着生长时期和各营养器官而有所不同.另外,植物吸收氮素的过程又受到大气状况和土壤状况的制约.土壤中氮素经过矿化作用、硝化作用和反硝化作用进行转化,一部分把氮素转化成植物能吸收的营养形态,另一部分则从土壤中损失.凋落物的分解和N沉降能补充土壤中的氮素,而植物吸收、微生物固持、水文流失和N溢出等方式使氮素从土壤中损失出去.另外,凋落物的分解和根际土壤、CO2浓度和臭氧对氮素循环有着重要的作用.N污染、N沉降、碳氮循环的耦合作用是今后研究的热点问题.     

植物对氨基酸的吸收研究进展

氨基酸在提高植物产量、改善产品品质、增强植株抗逆性、保护生态环境等方面发挥着越来越重要的作用,在农业生产中越来越受到重视.本文简述了氨基酸含量、氨基酸种类和植物种类对植物吸收氨基酸的影响,并对氨基酸营养研究进行展望,以期提高人们对植物氨基酸营养的认识,促进氨基酸在农业中的应用和发展.     

丛枝菌根利用氮素研究进展

氮素是植物需求量最大的元素,丛枝菌根真菌与植物形成共生体后能从土壤中获取无机氮、简单的氨基酸,还能利用一些复杂的有机态氮.考虑到NH+4在土壤中的移动性低及丛枝菌根真菌的专性共生菌的特点,丛枝菌根真菌吸收NH+4对植物的贡献较大.近年来的研究发现丛枝菌根真菌内存在与氮素代谢有关的鸟氨酸循环,而精氨酸则是菌丝内氮素转移的主要形式.综述最近的AMF对氮素的吸收、转运、同化、交换等方面的文献,旨在揭示丛枝菌根真菌氮素利用特点,阐明丛枝菌根真菌在氮循环系统中的重要作用.     

高等植物对有机氮吸收与利用研究进展

主要综述植物氨基酸营养生理生化和分子生物学研究的最新进展。长期以来,人们一直认为植物只能吸收无机态N,有机N必须矿化为无机N后才能被植物吸收利用,而近年来越来越多实验证明植物能吸收有机N,特别是氨基酸,其吸收能力因植物种类而异,生长在有机N丰富的北极,高山和亚高山生态环境中的植物甚至嗜好氨基酸,因此,不应过分夸大有机N矿化的重要性,迄今一些植物细胞质膜上的氨基酸转运子基因已被描述并加以克隆。     

古尔班通古特沙漠一年生植物的氮吸收策略

同一生活型的植物可能通过吸收不同形态的氮来利用陆地生态系统中有限的氮, 避免和减少对资源的竞争, 从而完成共生。研究荒漠生态系统同一生活型植物对氮的利用是否存在生态位分离, 有助于深入了解荒漠植物的生存策略, 更好掌握氮利用对荒漠植物生存的影响。该研究利用¹⁵N同位素示踪法, 研究古尔班通古特沙漠中广泛分布的2种一年生植物——角果藜(Ceratocarpus arenarius)和碱蓬(Suaeda glauca)在不同月份和不同土壤深度对不同形态氮的吸收策略。结果显示, 在浅层土壤中, 2种植物7月的氮吸收速率均高于6月; 对比不同形态氮的吸收速率, 植物对无机氮的吸收均高于有机氮, 角果藜更偏好吸收硝态氮, 每克干根系最高氮吸收速率可达3.81 μg·h^-1, 碱蓬更偏好吸收铵态氮, 每克干根系最高氮吸收速率可达4.74 μg·h^-1; 从不同形态氮对总氮的贡献率看, 硝态氮是角果藜吸收氮的有利形态, 占比在35.7%-43.9%之间, 铵态氮是碱蓬吸收氮的有利形态, 占比最高可达48.3%, 最低也有40.0%。2种一年生植物不仅可以利用土壤中的无机氮, 也可以直接吸收利用土壤有机氮。研究结果表明: 在古尔班通古特沙漠生态系统中, 一年生植物对氮的吸收能力有着差异和多元化的特点, 且均可吸收土壤中的可溶性有机态氮源。     

陆地生态系统植物的氮源及氮素吸收

氮是植物生长发育所必需的营养元素,也是其主要的限制因子之一.陆地生态系统植物所需氮的来源及植物对氮素的吸收利用均受控于其种类和生长环境.环境条件的改变,一方面可能改变植物生长区原有氮的形态、浓度、赋存方式等,从而改变氮对植物的供给状况;另一方面可能引起植物生长区土壤质量、水分利用状况、光照等的改变,从而产生耦合现象,直接影响植物的生理生态特性,使植物对氮素的吸收利用发生改变,导致植物生长区的种群类型及物种多样性发生改变,并直接影响到生态系统的功能及演替.本文主要对陆地生态系统中高等植物生长发育所需氮素的来源及植物对氮素吸收利用过程中的影响因素进行了综述和讨论,并结合国内外在该领域的研究现状对其研究前景进行了展望.     

Plant acquisition of organic nitrogen in boreal forests

Research on plant nitrogen (N) uptake and metabolism has more or less exclusively concerned inorganic N, particularly nitrate. Nevertheless, recent as well as older studies indicate that plants may have access to organic N sources. Laboratory studies have shown that ectomycorrhizal and ericoid mycorrhizal plants can degrade polymeric N and absorb the resulting products. Recent studies have also shown that some non‐mycorrhizal plants are able to absorb amino acids. Moreover, amino acid transporters have been shown to be present in both plant roots and in mycorrhizal hyphae. Although both mycorrhizal and non‐mycorrhizal plants appear to have a capacity for absorbing a range of organic N compounds, is this capacity realized in the field? Several lines of evidence show that plants are outcompeted by microorganisms for organic N sources. Such studies, however, have not addressed the issue of spatial and temporal separation between plants and microorganisms. Moreover, competition studies have not been able to separate uptake by symbiotic and non‐symbiotic microorganisms. Qualitative assessment of organic N uptake by plants has been performed with dual‐labelled glycine in several studies. These studies arrive at different conclusions: some indicate that plants do not absorb this organic N source when competing with other organisms in soil, while others conclude that significant fractions of amino acid N are absorbed as intact amino acid. These variable results may reflect species differences in the ability to absorb glycine as well as differences in experimental conditions and analytical techniques. Although theoretical calculations indicate that organic N might add significant amounts of N to plant N uptake, direct quantitative assessment of the fraction of plant N derived from uptake by organic N sources is a challenge for future research.     

The Pitcher Plant Sarracenia purpurea Can Directly Acquire Organic Nitrogen and Short-Circuit the Inorganic Nitrogen Cycle

Background

Despite the large stocks of organic nitrogen in soil, nitrogen availability limits plant growth in many terrestrial ecosystems because most plants take up only inorganic nitrogen, not organic nitrogen. Although some vascular plants can assimilate organic nitrogen directly, only recently has organic nitrogen been found to contribute significantly to the nutrient budget of any plant. Carnivorous plants grow in extremely nutrient-poor environments and carnivory has evolved in these plants as an alternative pathway for obtaining nutrients. We tested if the carnivorous pitcher plant Sarracenia purpurea could directly take up intact amino acids in the field and compared uptake of organic and inorganic forms of nitrogen across a gradient of nitrogen deposition. We hypothesized that the contribution of organic nitrogen to the nitrogen budget of the pitcher plant would decline with increasing nitrogen deposition.

Methodology and Principal Findings

At sites in Canada (low nitrogen deposition) and the United States (high nitrogen deposition), individual pitchers were fed two amino acids, glycine and phenylalanine, and inorganic nitrogen (as ammonium nitrate), individually and in mixture. Plants took up intact amino acids. Acquisition of each form of nitrogen provided in isolation exceeded uptake of the same form in mixture. At the high deposition site, uptake of organic nitrogen was higher than uptake of inorganic nitrogen. At the low deposition site, uptake of all three forms of nitrogen was similar. Completeness of the associated detritus-based food web that inhabits pitcher-plant leaves and breaks down captured prey had no effect on nitrogen uptake.

Conclusions and Significance

By taking up intact amino acids, Sarracenia purpurea can short-circuit the inorganic nitrogen cycle, thus minimizing potential bottlenecks in nitrogen availability that result from the plant''s reliance for nitrogen mineralization on a seasonally reconstructed food web operating on infrequent and irregular prey capture.     

Amino acid uptake: a widespread ability among boreal forest plants

Amino acids constitute a potentially important source of nitrogen for plants in boreal forest ecosystems. Accordingly, it may be suggested that distinct plant species differing abilities to take up amino acids constitutes an important factor in determining plant ecosystem composition. Using GC-MS and isotopically labelled amino acids, we measured the simultaneous uptake of 15 different amino acids by 31 common boreal forest plant species. The results from this study show that all plant species tested, representing a wide variety of plant types, have the ability to take up amino acids from an incubation solution. Furthermore, uptake rates were unrelated to mycorrhizal associations as well as habitat soil amino acid concentrations and plant nitrogen availability dependence as measured by Ellenberg nitrogen indicator values. These results suggest that mycorrhiza is of minor importance for discrete plant amino acid uptake rates and further points out the potential importance of amino acids to plant nitrogen nutrition in boreal forest ecosystems.     

植物对重金属的吸收和分布

植物修复是利用植物来清除污染土壤中重金属的一项技术。该技术成功与否取决于植 物从土壤中吸取金属以及向地上部运输金属的能力。植物对金属的吸收主要取决于自由态离子活度。许多螯合剂能诱导植物对重金属的吸收。金属离子在液泡中的区域化分布是植物耐 重金属的主要原因。同时,细胞内的金属硫蛋白、植物螯合肽等蛋白质以及有机酸、氨基酸等在金属贮存和解毒方面也起重要作用。本文还论述了重金属在植物体内运输的生理及分子 方面的研究进展。     

植物对重金属的吸收和分布

植物修复是利用植物来清除污染土壤中重金属的一项技术。该技术成功与否取决于植物从土壤中吸取金属以及向地上部运输金属的能力。植物对金属的吸收主要取决于自由态离子活度。许多螯合剂能诱导植物对重金属的吸收。金属离子在液泡中的区域化分布是植物耐重金属的主要原因。同时,细胞内的金属硫蛋白、植物螯合脓等蛋白质以及有机酸、氨基酸等在金属贮存和解毒方面也起重要作用。本文还论述了重金属在植物体内运输的生理及分子方面的研究进展。     

植物对氨基酸的吸收及其生理效应

<正> 在古代的农业系统中,所用的肥料都是有机肥,人们并不知道植物依赖于何种物质得以生长繁衍。自1840年,德国化学家李比希(J.U.Liebig)发表了《化学在农业和植物生理学上应用》一书后,人们对植物的营养有了新的认识,认识到矿物质对植物的重要性。李比希在书中指出:“土壤中矿物质是一切绿色植物唯一的养料。植物可以完全依赖于无机物质而生长发育。这一观点即为“植物的矿质营养学说”。该学说后来为生产实践所充分证明,并成为农业化学     

Abstracts of Platform Presentations from the Phytoremediation Session at the 14th Annual West Coast Conference on Contaminated Soils,Sediments, and Water (March 15–18, San Diego,CA)

Previous field studies indicate that zucchini (Cucurbita pepo) has a unique ability to phytoextract persistent organic pollutants from soil. It is unlikely that C. pepo evolved a unique mechanism favoring POP extraction and uptake, but all plants have evolved means to facilitate nutrient acquisition from soil. We have hypothesized that the exudation of organic acids as a means to acquire phosphorus could facilitate the uptake of persistent organic pollutants by increasing contaminant bioavailability to the plants. In one study, we assessed DDE uptake and organic acid exudation by zucchini (an uptaker of POPs) and cucumber (a non-uptaker of POPs) under various cultivation and nutrient conditions. Under dense planting (5 plants in a 5-kg pot of DDE-contaminated soil), zucchini accumulated significant and expected amounts of DDE but surprisingly, under these stressed conditions, cucumber phytoextracted greater amounts of DDE. The cucumber rhizosphere concentrations of organic acids were significantly higher than that of zucchini, suggesting that the increased organic acid exudation promoted DDE uptake by cucumber. Conversely, under non-stressed conditions zucchini phytoextracted significantly greater quantities of pollutant than cucumber but no differences in organic acid content of the rhizosphere of the two species were observed. Separately, zucchini and other species were grown under field conditions and weekly amendments of different nutrients were made (nitrogen, phosphorus, nitrogen/phosphorus, aluminum sulfate to bind phosphorus in the soil). The uptake and translocation of the weathered pollutant and inorganic elements was found to vary with nutrient amendments. Lastly, data will be presented from rhizotron units constructed to facilitate not only the direct in situ isolation of exuded organic acids but also the isolation of xylem sap and rhizosphere soil pore water from individual plants. The role of cultivation conditions and nutrient availability in controlling root morphology, organic acid exudation, and contaminant uptake will be discussed.     

Comprehensive screening of Arabidopsis mutants suggests the lysine histidine transporter 1 to be involved in plant uptake of amino acids

Plant nitrogen (N) uptake is a key process in the global N cycle and is usually considered a "bottleneck" for biomass production in land ecosystems. Earlier, mineral N was considered the only form available to plants. Recent studies have questioned this dogma and shown that plants may access organic N sources such as amino acids. The actual mechanism enabling plants to access amino acid N is still unknown. However, a recent study suggested the Lysine Histidine Transporter 1 (LHT1) to be involved in root amino acid uptake. In this study, we isolated mutants defective in root amino acid uptake by screening Arabidopsis (Arabidopsis thaliana) seeds from ethyl methanesulfonate-treated plants and seeds from amino acid transporter T-DNA knockout mutants for resistance against the toxic D-enantiomer of alanine (Ala). Both ethyl methanesulfonate and T-DNA knockout plants identified as D-Ala resistant were found to be mutated in the LHT1 gene. LHT1 mutants displayed impaired capacity for uptake of a range of amino acids from solutions, displayed impaired growth when N was supplied in organic forms, and acquired substantially lower amounts of amino acids than wild-type plants from solid growth media. LHT1 mutants grown on mineral N did not display a phenotype until at the stage of flowering, when premature senescence of old leaf pairs occurred, suggesting that LHT1 may fulfill an important function at this developmental stage. Based on the broad and unbiased screening of mutants resistant to D-Ala, we suggest that LHT1 is an important mediator of root uptake of amino acids. This provides a molecular background for plant acquisition of organic N from the soil.     

The influence of living plants on mineralization and immobilization of nitrogen

Summary Changes in the pattern of distribution of the nitrogen of the soil and seedling grass plants have been investigated when the grass plants were grown in pots of sandy soil, from a pasture, at pH 5.7. Net mineralization of soil nitrogen was not observed during an experimental period of one month in the absence of added nitrogenous fertilizer (Table 2). Addition of labeled nitrogen (as ammonium sulphate) to the soil at the beginning of the experimental period resulted in a negative net mineralization during this period (Table 4b). When none of the fertilizer nitrogen remained in its original form in the soil it was found that approximately 12 per cent of the labeled nitrogen had been immobilized in soil organic compounds. Clipping of the grass at this date was followed by a decrease in the amount of labeled soil organic nitrogen, indicating that mineralization was not depressed by living plants. The application of unlabeled ammonium sulphate subsequent to the utilization of the labeled nitrogen did not decrease the amount of immobilized labeled nitrogen in the soil organic matter, as would be expected if the organic nitrogen compounds of the soil had been decomposed to ammonia. This was thought to be due to the fact that decomposition of organic nitrogen compounds in permanent grassland results in the production of peptides, amino acids etc. which are utilized by microorganisms without deamination taking place. In pots with ageing grass plants, labeled organic nitrogen compounds were found to be translocated from the grass shoots to the soil (Table 7). Net mineralization of soil organic nitrogen was positive in the contents of pots containing killed root systems (Table 3b). About 8 per cent of the labeled nitrogen added to the contents of such pots, in the form of ammonium sulphate, was found to be present in soil organic nitrogen compounds approximately 4 weeks after application, while a total of about twice this amount of soil organic nitrogen was mineralized during that period. From the results obtained in this investigation, it is concluded that the constant presence of living plants is responsible for the accumulation of nitrogen in organic compounds in permanent grassland. No evidence was obtained that the decomposition of such compounds in the soil is inhibited by living plants.