Understanding plant responses to phosphorus starvation for improvement of plant tolerance to phosphorus deficiency by biotechnological approaches

Abstract

In both prokaryotes and eukaryotes, including plants, phosphorus (P) is an essential nutrient that is involved in various biochemical processes, such as lipid metabolism and the biosynthesis of nucleic acids and cell membranes. P also contributes to cellular signaling cascades by function as mediators of signal transduction and it also serves as a vital energy source for a wide range of biological functions. Due to its intensive use in agriculture, P resources have become limited. Therefore, it is critically important in the future to develop scientific strategies that aim to increase P use efficiency and P recycling. In addition, the biologically available soluble form of P for uptake (phosphate; Pi) is readily washed out of topsoil layers, resulting in serious environmental pollution. In addition to this environmental concern, the wash out of Pi from topsoil necessitates a continuous Pi supply to maintain adequate levels of fertilization, making the situation worse. As a coping mechanism to P stress, plants are known to undergo drastic cellular changes in metabolism, physiology, hormonal balance and gene expression. Understanding these molecular, physiological and biochemical responses developed by plants will play a vital role in improving agronomic practices, resource conservation and environmental protection as well as serving as a foundation for the development of biotechnological strategies, which aim to improve P use efficiency in crops. In this review, we will discuss a variety of plant responses to low P conditions and various molecular mechanisms that regulate these responses. In addition, we also discuss the implication of this knowledge for the development of plant biotechnological applications.     

32P uptake and transport to shoots in Pinuus serotina seedlings under aerobic and hypoxic growth conditions

We examined the effects o long-term hypoxic growth conditions on net uptake and transport of P to shoots of pond Pine (Pinus serotina Michx.), a moderately flood-tolerant southern pine. Seedlings were grown under aerobic orhypoxic solution conditions for 4–5 weeks in continuously flowing solution culture containing 100 μM P. Short – and long-term ³²P. experiments were then concluded with intact seedlings to determine rates of ³²P influx, efflux and net transport to the shoot. Shoot fresh weight/root fresh weight ratios were significantly higher under hypoxic gorwth conditions, reflecting the larger reduction in root growth than shoot growth, despite extensive aerechyma formation in roots. Estimates for the unidirectional influx of ³²P in aerobic and hypoxic seedlings were 1.43 and 3.20 μmol P (g_FW root)_?1 h_?1, respectively. However, ³²P accumulation between the two treatments became similar within 8 h, suggesting that efflux was also higer in seedlings from the hypoxic treatment. Indeed in a separate experiment, hypoxic growth conditions increased efflux by over 60%. Transport of ³²P to shoots was significantly reduced under hypoxic growth conditions, despite higher root P concentrations and lower shoot P concentrations. After 48 h, ³²P accumulation in roots was similar between the two treatments. Yet total accumulation of seedling ³²P decrcased by 31% under the hypoxic treatment, largely because of reduced transport of ³²p to the shoot. The lower accumulation of ³² by shoots of seedlings in the hypoxic treatment may be the result of a direct inhibition on the transport process in O²-defident tissues, but could also reflect a slower turnover or labeling of the ool available for transport. Indeed, the percentage of total ³²P in. roots present in the soluble P. (or transportable form of P) was about 33% lower in seedlings from the hypoxic treatment, probably reflecting increased assimilation into organic compounds as well as chelation with iron. Our results suggest that P transport to the shoots of acclimated seedlings may be more sensitive to hypoxic solution conditions than influx at the root Plasmalemma.     

哈泥泥炭地落叶和常绿两类灌木对氮磷输入增加响应的对比

大气氮沉降及磷输入正影响泥炭地的生态过程,而其重要植物成分灌木对氮磷输入如何响应尚鲜有研究。在哈泥泥炭地,通过连续6年的人工氮、磷添加后对3种落叶灌木和3种常绿灌木生长状况进行测定,研究泥炭地两种类型灌木对氮磷输入的响应。落叶灌木(多度和盖度下降,高度上升)与常绿灌木(多度、盖度和重要值下降)对氮输入增加的响应存在差异,对磷输入增加的响应一致。从物种的角度看,存在种间差异,施氮抑制落叶灌木金露梅和笃斯越桔的生长,其多度和盖度均下降,对落叶灌木油桦有促进作用,使其高度、盖度和重要值增加。施氮仅对常绿灌木甸杜产生了影响,降低了其多度和重要值,对常绿灌木小叶杜鹃和狭叶杜香无影响。施磷缓解了部分灌木的养分限制,提高了落叶灌木油桦的盖度、常绿灌木小叶杜鹃的高度和重要值以及常绿灌木甸杜的多度。同时,氮磷对落叶灌木存在交互作用,未施氮肥和低浓度施氮时,施磷对落叶灌木的高度和重要值无影响,在高浓度施氮情况下,施磷使落叶灌木的高度和重要值增加,即氮和磷只有在一定比例下才会促进灌木植物的生长。研究表明,氮磷输入可影响泥炭地灌木间的竞争平衡,使落叶灌木油桦获得生长优势,抑制其他灌木的生长,进而对泥炭地的植被格局产生深远影响。     

Functions of Calcium-bound Phosphorus in Relation to Characteristics of Phosphorus Releasing Bacteria in Sediment of a Chinese Shallow Lake (Lake Wabu)

Sediment phosphorus (P) release accelerates lake eutrophication, while retention capacity and release potential of different P fractions, calcium-bound P (CaCO₃～P) in particular, still remains unclear. Fractionation and sorption behaviors of phosphorus were studied in sediment of a Chinese shallow lake (Lake Wabu) and two inflowing rivers from December 2011 to December 2012. Abundance of P releasing bacteria was analyzed, and their main species were isolated using a culture-dependent method and identified by their 16S rDNA sequences. CaCO₃～P release abilities of these bacteria were also tested. In sediments of both the lake and rivers studied, the rank order of the different P extracts was CaCO₃～P > iron-bound P > acid-soluble organic P > hot NaOH-extractable organic P. At the same time, CaCO₃～P content and equilibrium P concentration (EPC₀) values in river sediments were significantly higher than those in the lake. Additionally, EPC₀ changes non-monotonically with increasing CaCO₃～P content, forming a V-shaped curve, with the lowest EPC₀ at an intermediate CaCO₃～P content (around 180 mg kg^?1). Below this threshold, CaCO₃～P was a component strengthening P retention; moreover, CaCO₃～P became an active species responsible for P release. Noticeably, between the two parts divided by this threshold, the differences in abundance of inorganic phosphorus solubilizing bacteria (IPB) and organic phosphorus mineralizing bacteria (OPB) were insignificant and the dominant IPB species clustered together. By contrast, OPB was distinguished from each other, whose dominant species isolated from the part with higher CaCO₃～P content, namely Novosphingobium sp., exhibited a stronger ability to solubilize CaCO₃～P. Shortly, with lower content, CaCO₃～P tends to stabilize P in sediment; while with higher content or under eutrophic condition, it shifted into P source, with some OPB species becoming the main factors to drive its release.     

Effects of phosphate stress on the rate of phosphate uptake during resupply to deficient tomato plants

Plants of Lycopersicon esculentum Mill. P. I. 120262 show an increased phosphate uptake rate per unit dry weight of root after as little as one day of growth in solutions lacking phosphate. The reversibility of this response in plants experiencing various degrees of phosphate stress was investigated by measuring the depletion of phosphate from solutions over 3-h intervals. Measurements were made at three times in the first 30 h after phosphate was resupplied. Reversibility decreased as the level of phosphate stress increased. The phosphate uptake rate was returned to the level of controls after 30 h of phosphate resupply in plants grown without phosphate for one or three days. Plants grown without phosphate for five or seven days had uptake rates 26 and 40% higher than controls, respectively, after the same period of phosphate resupply. Internal phosphate concentrations after 30 h of phosphate resupply were equal to or greater than the controls in all plants. These results are consistent with a simple reversible feedback of phosphate status on phosphate transport in slightly stressed plants, but such a mechanism seems inadequate to explain the responses observed in more severely stressed plants.     

A simple method to release polyphosphate from activated sludge for phosphorus reuse and recycling

In enhanced biological phosphorus removal processes, activated sludge microorganisms accumulate large quantities of polyphosphate (polyP). It was discovered that nearly all of the polyP could be released from activated sludge simply by heating it at 70 degrees C for about 1 h. The chain length of released polyP ranged from 100 to 200 phosphate (P(i)) residues. The addition of CaCl(2) precipitated approximately 75% of the total phosphorus without pH adjustment. The formed precipitate contained more P and less Ca than typical natural phosphorite deposits. Hence, in combination with enhanced biological phosphorus removal, the present method has potential for the development of a simple process for recovering phosphorus in a reusable form from wastewater.     

Arbuscular mycorrhiza improves plant adaptation to phosphorus deficiency through regulating the expression of genes relevant to carbon and phosphorus metabolism

Aims Arbuscular mycorrhizal (AM) symbiosis plays an important role in plant adaptation to phosphorus (P) deficiency. The mycorrhizal fungi can directly regulate P stress response of the host plants, and can also indirectly influence neighbor plants via AM exudates. This study aimed to reveal the regulation mechanisms of plant response to P deficiency by AM associations. Methods In a compartmentation cultivation experiment with Zea mays ‘B73’ and AM fungus Rhizophagus irregularis ‘DAOM197198’, we investigated mycorrhizal effects on plant P nutrition and the expression of plant and fungal genes related to P and carbon (C) metabolisms under both low P (10 mg?kg^-1) and high P (100 mg?kg^-1) conditions. The cultivation system consisted of three compartments, namely donor compartment, buffer compartment and receiver compartment divided by two pieces of microporous filters with pore size of 0.45 μm. Maize plant in donor compartment inoculated with AM fungus served as a source of AM exudates. The microporous filters could restrict the development of extraradical mycelium of AM fungi, but allow diffusion of AM exudates. Real-time PCR was performed to quantify the gene expression levels both in maize plants and AM fungi. Important findings The experimental results indicated that under low P conditions mycorrhizal colonization increased plant dry weight and P concentration in donor plants, and up-regulated plant genes encoding P transporters Pht1;2, Pht1;6, phosphoenolpiruvate carboxylase (PEPC), inorganic pyrophosphatase (TC289), glycerol-3-phosphate transporter (G3PT) and malate synthase (MAS1). The expression of AM fungal genes encoding P transporter (GiPT), GlcNAc transporter (NGT1), GlcNAc kinase (HXK1b), GlcNAc phosphomutase (AGM1), UDP GlcNAc pyrophosphorylase (UAP1), chitin synthase (CHS1), GlcNAc-6-phosphate deacetylase (DAC1) and glucosamine-6-phosphate isomerase (NAG1) was significantly higher under low P conditions compared with high P conditions. However, for the receiver plants, plant dry mass and P concentration were only significantly increased by higher P addition, while inoculation treatment significantly up-regulated the expression of P transporter genes Pht1;2 and Pht1;6, C metabolism related genes G3PT, PEPC, TC289 and MAS1. The study proved that AM exudates could potentially stimulate plant response to P deficiency by regulating functional genes relevant to P and C metabolisms in the mycorrhizal associations.     

大豆种子储藏性磷对幼苗适应低磷胁迫的作用

选用种子大小不同、磷效率不同的两个大豆品种‘巴西10号’(B10)和‘本地2号’(L2),在不同供磷条件下进行营养液浇灌沙培,从大豆萌发至2片三出复叶完全展开期测定植株主要器官总磷、可溶性磷浓度、子叶可溶性蛋白、酸性磷酸酶比活性、植酸酶比活性的变化动态,探讨储藏性磷在大豆幼苗期适应磷胁迫中的作用。结果发现:(1)磷效率不同的两个大豆品种的种子中磷含量差异显著。(2)大豆萌发和幼苗生长过程中子叶的磷逐渐转入根、茎、叶中,并以转入叶中的磷最多,其中磷高效品种B10在发芽过程中子叶磷向各个器官转移的总磷量要高于磷低效品种L2,且持续时间长。(3)大豆萌发和幼苗生长过程中外源供磷水平显著影响子叶磷的转移,在外源供磷充足条件下各器官中总磷均高于低供磷条件,子叶中磷和外源磷存在补偿关系。(4)磷高效品种B10子叶中酸性磷酸酶活性在低磷条件下显著高于高磷条件,但磷低效品种L2在高、低磷间无显著差异。研究表明,大豆种子储藏性磷在幼苗期耐低磷能力建立方面具有重要作用。     

Net and gross sedimentation in relation to the phosphorus budget of Eau Galle Reservoir,Wisconsin

We compared P retention with direct measures of Psedimentation, estimated fromsediment cores (annual P sedimentation) and sedimenttraps (daily P sedimentation),to quantify P sedimentation in Eau Galle Reservoir,Wisconsin. Mean annual Pretention was similar to mean annual P sedimentation,as estimated from sediment corerates integrated over the entire lake basin,indicating that annual P mass balanceapproximated annual net P sedimentation in thisreservoir. However, sediment trap Prates, measured over the summer stratified period,overestimated P retention ratesdetermined over the same period, suggestingsubstantial deposition of internally-derived P.Inclusion of measured internal P loadings from avariety of sources in EauGalle Reservoir in a P mass balance only accounted foran additional 24% of thesummer sediment trap P rate, indicating substantialuncertainty in the overall P budget.Imbalances in the P budget may also suggest depositionof sediment from other as yetunquantified internal sources. Potential internalsources of P include sedimentresuspension and chemical release and direct uptake ofP from the sediment byphytoplankton.     

秸秆还田及磷细菌对土壤微生态及豆角产量的影响

为研究秸秆还田及外源添加磷细菌对土壤微生态及农作物产量的影响,以玉米秸秆和具有解磷能力的恶臭假单胞菌(Pseudomonas putida)为研究对象,通过在设施大棚中栽培豆角,研究了不同处理对土壤和豆角根际解磷类细菌数量、土壤有效磷含量、解磷能力以及豆角产量的影响。研究结果显示,玉米秸秆(处理1)、玉米秸秆+磷细菌(处理2)、磷细菌(处理3)三个处理的土壤中解磷类细菌数量有较大差异,其中处理2数量最高,比对照区高31.89%,差异显著(P<0.05);三个处理均能明显增加豆角根际解磷类细菌数量,差异达到显著水平(P<0.05),其中处理2最高,比对照高86.30%,差异达到极显著水平(P<0.01);三个处理土壤有效磷含量均明显高于对照,差异显著(P<0.05),其中处理2最高,比对照高9.8%;三个处理土壤解磷能力差异较大,处理1和处理2可明显提高土壤解磷能力,分别比对照高50.2%和65.2%,差异极显著(P<0.01);三个处理对豆角均有增产效果,但差异较大,处理2比处理1、处理3产量增加明显,分别增产6.8%、10.3%,差异达到显著水平(P&l...     

异质低磷胁迫下马尾松家系根构型和磷效率的遗传变异

以7个马尾松(Pinus massoniana)一代种子园自由授粉家系为材料, 设置同质低磷(P)胁迫和异质低P胁迫模拟的盆栽试验, 系统研究马尾松家系对不同类型低P胁迫的适应机制和P效率变异规律。结果表明, 参试马尾松家系的苗高、地径和生物量等P效率指标均表现出显著的家系变异, 主要P效率指标的家系遗传力均较高, 干物质积累量的广义遗传力大于0.80, 揭示了马尾松P营养效率的较大遗传改良潜力。马尾松对不同类型低P胁迫的适应机制有所差异。在同质低P胁迫下, ‘3201’、‘1217’等高P效率家系的根系主要参数均高于低P效率家系, 表明整体根系参数的适应性变化是P效率和生物量形成的决定因素。在异质低P胁迫下, 高P效率马尾松家系在表层富P介质的根系分布量、分布比例均显著增加, 表层根系参数与马尾松家系P效率呈显著正相关, 揭示根系空间构型的适应性变化是决定马尾松高P效率的重要生物学基础。表层根系生物量、表层根相对比例的家系遗传力达0.88和0.72, 证实了以马尾松根构型的适应变化为突破口, 选育具有理想根构型和较高P效率的马尾松家系。     

磷高效基因型小麦对缺磷胁迫的根际适应性反应

采用奶箱分隔栽培试验法,进行了磷高效与磷低效小麦基因型根际土壤ＰＨ与有效磷变化的研究,结果表明：小麦根际土壤ＰＨ和有效磷含量皆明显低于外围土壤,表现出明显的根际效应特征;磷高效基因型小麦的根际ＰＨ和有效磷含量明显低于磷低效基因型,ＰＨ变异范围和磷素亏缺区也表现明显较大。为了进一步验证磷高效小麦基因型这的这一根际特征,同时进行了施用水溶性,枸溶性磷肥的试验研究,结果表明,以水溶性磷肥对根际ＰＨ和有效     

Response of early and late semiarid seral species to nitrogen and phosphorus gradients

Above- and below-ground biomass production, nitrogen (N) and phosphorus (P) tissue concentrations, and root: shoot ratios were examined for five species that are characteristic of a semiarid successional sequence under controlled greenhouse conditions. In two simultaneous experiments, seedlings of one forb, two grass, and two shrub species important in a sagebrush successional sere, were subjected to seven levels of N and P. Results of the experiments suggest distinct differences in nutrient response patterns between early and late seral species. Early seral species produced more biomass but had lower tissue nutrient concentrations than late seral species. As N and P availabilities decreased, late seral species displayed characteristics indicative of increasing competitive advantage over those of early seral species. Root: shoot ratios of the five species primarily reflected patterns related to lifeform, but with some early and late seral characteristics. Results from this study 1) confirm that nutrient use pattern, nutrient availability, and seral position relationships characteristic of mesic ecosystems hold equally true for semiarid systems, and 2) suggest that nutrients are important organizing factors in semiarid ecosystems.     

甘蔗幼苗对难溶性磷的吸收及其根系对低磷胁迫的响应

为明确甘蔗适应低磷胁迫的生理生化机制,挖掘甘蔗对磷素的利用潜力,揭示甘蔗对低磷胁迫适应的可能机制,该研究以ROC22和ROC10两个甘蔗品种为材料,采用水培和土培的方法研究了甘蔗幼苗对难溶性磷的吸收及其在低磷胁迫下根构型和根系的生理反应。结果表明:(1)培养在以难溶性磷(Ca-P和Al-P)为磷源的培养液中的甘蔗的叶片数、地上部干重、生物量较缺磷(-P)处理显著增加,与对照(+P)的相当,甘蔗总磷积累量也显著提高,达到对照(+P)处理磷积累量的30%～77%。(2)在低磷条件下,甘蔗幼苗的根系有向土壤深层分布的趋势,根的总体积增大、最长根长变长、浅根系分布增多。(3)甘蔗幼苗在低磷环境下,根际环境明显酸化,且根系分泌物能溶解难溶性的铝磷,植株体内酸性磷酸酶的活性也明显增强。以上表明甘蔗幼苗有较强的吸收利用难溶性磷的能力,而低磷条件下根系数量增加、主根的向地性、浅根系分布增多、根际酸化以及植株体内酸性磷酸酶活性的增强可能是甘蔗幼苗适应缺磷环境的重要机制。     

Response and correlation of above- and below-ground functional traits of Leymus chinensis to nitrogen and phosphorus additions

AimsLeymus chinensis is a constructive and dominant species in typical steppe of northern China. The structure and functions of L. chinensis grassland ecosystem has been degenerated seriously due to long-term overgrazing in recent decades. As an effective measure to restore the degraded grasslands, the effects of nutrient addition on plant growth and ecosystem structure and functioning have been paid more attention in manipulation experimental research. The effects of nutrient addition, especially P addition on the above- and below-ground functional traits of L. chinensis have rarely been studied; particularly the underpinning mechanisms remain unclear. Our objective is to examine the responses and adaptive mechanisms of L. chinensis to different levels of N and P additions. MethodsWe conducted a culture experiment in the greenhouse, with three levels of N (50, 100 and 250 mg N·kg^-1) and P (5, 10 and 25 mg P·kg^-1) addition treatments. The above- and below-ground biomass, leaf traits (e.g., specific leaf area, leaf N and P contents) and root traits (e.g., specific root length, root N and P contents) of L. chinensis were determined in this study.Important findings Our results showed that: 1) the aboveground biomass and total biomass of L. chinensis were mostly affected by N addition, while the belowground biomass was mainly affected by P addition. N addition greatly enhanced the aboveground biomass of L. chinensis, while P addition reduced the belowground biomass at the moderate and high N levels. The root-shoot ratio of L. chinensis was influenced by both N and P additions, and root-shoot ratio decreased with increasing N and P levels. N and P additions promoted more biomass and N and P allocations to aboveground and leaf biomass. 2) Leymus chinensis showed different responses and adaptive mechanisms to P addition at low and high N levels. At low N level, L. chinensis exhibited high photosynthetic rate and specific root length (SRL) to improve photosynthetic capacity and root N acquisition, which promoted aboveground biomass. High root P content was favorable for belowground biomass. At high N level, P addition did not significantly affect plant growth of L. chinensis, even reduced its belowground biomass. Leymus chinensis showed high specific leaf area (SLA) and SRL to improve light interception and N acquisition in order to maintain stable aboveground biomass. 3) P addition greatly impacted below-ground than above-ground functional traits. SLA exhibited a weakly positive correlation with SRL, indicating L. chinensis exhibited relatively independence of resource acquirement and utilization between leaf and root functional traits.     

Response of field-grown chickpea (Cicer arietinum L.) to phosphorus fertilization for yield and nitrogen fixation

Application of phosphorus at 40, 60, 80 and 100 kg P₂O₅ ha^–1 in the presence of a uniform dressing of nitrogen (N) and potash (K₂O) each applied at 20 and 24 kg ha^–1 to chickpea (CM-88) grown in sandy loam soil in a replicated field experiment improved the nodulation response of the crop, increased its grain yield (ka ha^–1) by 18, 59, 40 and 14 percent, biomass yield (ka ha^–1) by 32, 32, 54 and 14 percent, biomass N (kg ha^–1) by 31, 48, 49, 19 percent, and biomass P (kg ha^–1) by 26, 40, 41 and 11 percent, respectively. The effect of phosphorus on the nitrogenase activity of the excised roots of chickpea was, however, inconsistent.     

桉树幼苗对难溶性磷的吸收及其根系对低磷胁迫的响应

以‘广林9号’桉树幼苗为试验材料,采用水培和土培试验方法研究了桉树幼苗对难溶性磷酸盐的吸收及其在低磷胁迫下的根构型和根系的生理反应,以揭示桉树高效吸收磷素的机制。结果显示:(1)桉树幼苗在含磷酸铝的缺磷培养液中吸收的磷达4.24mg/株,与供应水溶性磷和磷酸钙处理的相当。(2)土壤缺磷或仅在上土层(0~20cm)施磷肥处理均有利于桉树幼苗浅层根的分布,使根表面积及根数在上土层与下层(20~40cm)比值明显增高。(3)桉树幼苗根尖的H+-ATPase活性在缺磷处理15d后显著提高,其根尖周围的溴甲酚紫指示剂变黄,根基环境明显酸化;根尖分泌的酸性磷酸酶活性在低磷胁迫也显著提升,且随着处理时间(10、15、20d)的延长而进一步提高;铝和低磷胁迫能明显诱导桉树根系分泌草酸,其分泌量显著高于对照和缺磷处理。研究结果表明,桉树幼苗具有较强的难溶性磷吸收能力,而在缺磷及磷铝胁迫下根系的浅层化、根尖酸化及根分泌的酸性磷酸酶及草酸量增加可能是桉树幼苗适应酸性土壤铝毒和缺磷环境的重要机制。     

增温对南亚热带森林土壤磷形态的影响及其对有效磷的贡献

磷(P)是森林生态系统生产力的重要限制性元素。土壤磷的有效性取决于磷的存在形态及其转化过程。目前有关增温如何调控磷形态转化过程, 从而促进土壤有效磷含量增加的机制尚未明确。该研究以南亚热带森林为研究对象, 采用沿海拔高度从300 m下降至30 m以模拟温度自然上升的方法, 采集该林型0-10、10-20、20-40 cm的土壤, 并用适用于酸性土壤的连续浸提方法分离不同形态磷, 研究增温对土壤不同形态磷含量的影响, 探讨土壤不同形态磷与有效磷的关系, 识别对土壤有效磷在增温背景下增加有重要贡献的磷组分。结果表明增温使0-10 cm的无机钙磷(Ca-Pi)及20-40 cm的无机铁磷(Fe-Pi)和总无机磷含量分别显著增加了65.5%、17.9%和18.5%, 但对总有机磷及各有机磷组分含量均无显著影响。土壤不同形态磷与有效磷含量的相关分析表明, 有效磷与无机态的不同形态磷及有机铝磷、有机铁磷含量均显著正相关, 其中与Fe-Pi含量的相关性最强。通过土壤不同形态磷与有效磷含量的通径分析进一步发现, 无机铝磷、Fe-Pi是土壤磷转化过程中的重要中间过渡性磷组分, 且Fe-Pi是促进有效磷含量增加最重要的直接贡献磷组分。结合前期研究结果, 增温可能增大了凋落物磷对土壤磷的输入, 还可能强化了土壤的吸附和沉淀过程, 使得更多进入到土壤的溶解态磷转化为Ca-Pi、Fe-Pi等缓效磷源, 其中Fe-Pi可能成为南亚热带森林在气候变暖背景下最重要的有效磷来源。     

羊草的地上-地下功能性状对氮磷施肥梯度的响应及关联

羊草(Leymus chinensis)是我国北方典型草原群落的主要建群种和优势种, 由于长期的过度放牧, 羊草草原生态系统的结构和功能严重退化。养分添加作为恢复草地生态系统的一种管理措施, 其应用目前还处于实验性研究阶段。关于羊草的地上-地下功能性状对养分添加, 尤其是P添加的响应研究较少, 相关机制尚不十分清楚。为此, 该文以羊草为研究对象, 通过温室栽培进行N (50, 100, 250 mg N·kg^-1)和P (5, 10, 25 mg P·kg^-1)各3个水平的养分添加实验, 研究羊草的地上-地下功能性状对N、P添加的响应及适应机制。主要研究结果表明: 1)羊草的地上生物量和总生物量主要受N添加的影响, N添加显著提高了羊草的地上生物量, 而地下生物量主要受P添加的影响, 尤其在中N和高N水平, P添加显著降低了羊草的地下生物量。羊草的根冠比受N、P添加的共同影响, 随着N、P添加梯度加大, 根冠比显著降低, N、P添加促进了羊草生物量向地上部分的分配和N、P向叶片的分配。2)在低N和高N水平, 羊草对P添加的响应与适应机制不同。低N水平, 羊草主要通过增加光合速率和比根长(SRL), 提高光合能力和根系对N的获取能力促进地上部分的生长, 而根系对P的吸收有利于地下部分的生长; 在高N水平, P添加对羊草的个体生长无明显促进作用, 甚至地下生物量明显受到P素抑制, 羊草主要通过保持较高的比叶面积(SLA)和SRL, 提高对光资源的截获能力和根系对N的获取和吸收能力, 维持地上部分的生长。3)相对于地上性状, P添加对羊草的地下性状影响更大, 羊草的SLA与SRL呈较弱的正相关关系, 表明叶片与根系在资源获取和利用方面具有相对独立性。