植物高效利用磷机制的研究进展

缺磷是限制目前农林业产量的一个重要因子,传统的农林业生产主要通过施肥和土壤改良来满足植物对磷的需求,近年来人们开始发掘磷高效利用植物来替代传统方法提高磷的利用效率。本文综述了国内外有关植物高效利用磷的形态学、生理学及遗传学作用机制,植物高效利用磷的机制主要包括：(1)磷高效利用植物能通过根系形态变化(包括根伸长、根轴变细、根毛数量和密度增大、侧根幼根数量增加及形成排根等)和根冠问的物质分配改变等形态学机制来适应磷胁迫;(2)在缺磷环境下磷高效利用植物能通过根系分泌物增加、菌根侵染、根系吸收动力学特征变化及植物磷素内循环加强等生理学机制来适应磷胁迫;(3)在磷亏缺的长期选择压力下,植物可通过某些“沉默”基因的诱导表达或DNA序列的特定形成相对稳定的磷营养遗传性状,由此通过遗传学机制来增加对土壤难溶态性磷的利用,使植物表现出较高的磷素利用效率。     

低分子量有机酸对土壤磷活化及其机制研究进展

近30年来,过量施用磷肥导致土壤磷素累积继而引起水体富营养化等问题备受关注。植物根系分泌的低分子量有机酸能活化土壤积累态磷,提高土壤磷素有效性,已成为研究热点问题之一。本文结合国内外已有研究,从低分子量有机酸的类型、添加浓度、土壤类型和土壤磷素水平等方面总结了低分子量有机酸活化土壤无机磷与有机磷的效果,并通过对比土壤磷活化试验前后各形态磷的变化探讨了低分子量有机酸对土壤磷的活化机制。低分子量有机酸对土壤无机磷的活化主要是促进了土壤中有效性低的无机磷形态向有效性较高的形态转化,而低分子量有机酸对土壤有机磷的活化结论尚不一致,活化机制也不明确,仍需进一步研究。未来研究应关注低分子量有机酸与磷肥之间的协同增效机制,并进一步探索低分子量有机酸对土壤磷素(特别是有机磷)的活化机制。     

在控制条件下云南松幼苗根系对低磷胁迫的响应

磷是控制生命过程的重要元素,植物在生长过程中需要大量的磷,低磷常导致一些植物发生适应性变化。云南松(PinusyunnanensisFranch.)以云南高原为起源和分布中心,其对低磷土壤环境表现出了很强的适应能力,广泛分布并正常生长于贫瘠的低磷红壤上,研究云南松对低磷环境的适应机制,对人类探索高效利用有限的磷素资源的方法具有现实意义。本实验通过对不同磷处理水平下培养的云南松幼苗根系生物量和根冠比等的研究,分析了云南松幼苗根系对低磷胁迫的响应。实验所用云南松种子采集自云南省通海县秀山森林公园内的健壮云南松林。结果表明:当磷浓度下降到0.5mmol/L时,云南松幼苗主根长度开始随磷浓度的降低而增加,根冠比随磷浓度的降低而增大,而侧根发生数没有随磷浓度的降低而显示出显著的增减规律,根系生物量也没有随磷浓度的降低而呈现出有规律的增减,根系生物量始终保持在一定的水平。进一步的分析表明:低磷胁迫下,云南松幼苗保证了物质分配对根的优先地位,以维持其根的生物量在一定水平,进而维持整个生命;云南松幼苗主要是靠主根长度的增加而不是靠侧根数量的增加来适应低磷环境。     

植物低磷胁迫适应机制的研究进展

磷对植物生长发育起到至关重要作用。但是土壤中存在大量植物不可利用的磷。根际周围非常低的有效磷含量是限制植物生长的因素。植物在遭受低磷胁迫时,其根形态、磷酸酶、有机酸、磷转运体等方面会产生一些适应机制。论述了这些机制最新研究进展．并对这一问题的研究方向进行展望。     

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。     

不同基因型春蚕豆对磷胁迫的适应性反应

利用不同作物或品种吸收利用土壤磷能力的差异提高磷素营养效率,是解决磷资源短缺的重要生物学途径.选择西北地区重要经济作物春蚕豆作为研究对象,选用3个不同春蚕豆品种(系),采用严重缺磷的碱性灌淤土,利用盆栽法研究了在不同供磷水平下不同基因型蚕豆的根系形态特征、酸性磷酸酶活性(APase)及产量的表现, 探讨不同基因型蚕豆对低磷胁迫的适应性反应.结果表明在整个生长过程中根长、根半径、根比表面积和根冠比变动最明显的是临蚕5号,分别为36.40%,65.10%、65.27%和13. 46%;缺磷条件下,蚕豆主要通过减小根半径,增加根长、根表面积,提高根冠比及体内酸性磷酸酶活性来实现对低磷胁迫的适应;不同基因型对低磷胁迫的适应能力不同;缺磷胁迫明显诱导各基因型蚕豆体内酸性磷酸酶活性的上升,临蚕5号增加最快为24.9%,8409为7. 79%,8354为7.29%;同一基因型的不同器官中酸性磷酸酶活性大小表现为根系>茎部>叶片 .根系酸性磷酸酶和根系形态参数可分别作为蚕豆耐低磷品种筛选的选择指标;缺磷导致作物减产,并且不同的基因型作物减产的幅度不同,临蚕5号缺磷比施磷减产30.98%,而8354 的产量在两个磷水平下变化不明显,说明临蚕5号对磷素的反应最强烈,为磷低效基因型,而 8354反应比较迟钝,为磷高效基因型.     

磷高效利用野生大麦基因型筛选及其根际土壤无机磷组分特征

通过土培盆栽试验,研究了16份野生大麦种质资源在相同供磷水平下磷素吸收利用的基因型差异,探讨磷高效野生大麦根际土壤无机磷组分特征.结果表明：拔节期和扬花期磷素干物质生产效率（CV=11.6%、12.4%）、成熟期磷素籽粒生产效率（CV=13.7%）基因型间差异较大.不同生育时期磷高效基因型IS-22-30和IS-22-25生物量、磷积累量和磷素干物质生产效率均显著高于低效基因型IS-07-07,且高效基因型的籽粒产量分别是低效基因型的3.10和3.20倍.不施磷、施磷30 mg·kg^-1条件下,不同磷素利用效率野生大麦根际土壤有效磷和水溶性磷含量均显著低于非根际土壤,且高效基因型较低效基因型根际土壤水溶性磷亏缺量更大.根际与非根际土壤无机磷组分含量为Ca₁₀-P＞O-P＞Fe-P＞Al-P＞Ca₂-P＞Ca₈-P.在拔节期和扬花期,施磷30 mg·kg^-1条件下,磷高效基因型根际土壤Ca₈-P含量显著高于低效基因型,而Ca₂-P含量显著低于低效基因型;不施磷条件下,高效基因型根际土壤Ca₂-P和Ca₈-P含量均显著高于低效基因型,且根际土壤Ca₁₀-P均减少.施磷30 mg·kg^-1条件下,根际土壤Fe-P和O-P含量均表现为高效基因型显著高于低效基因型,Al-P含量则呈现相反的趋势;不施磷条件下,高效基因型根际土壤Al-P、Fe-P和O-P含量均显著低于低效基因型.低磷胁迫下,高效基因型活化吸收Ca₂-P、Al-P的能力强于低效基因型.     

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。     

高效利用土壤磷素的植物营养学研究

论述磷在农业及生态系统中的重要地位,磷资源的危机以及磷土壤中的理化特性,探讨土壤的供磷潜力,不同植物品种以及相同品种不同基因型对土壤潜在磷资源的吸收利用差异,营养机制,遗传特性及其利用不同作物的基因型来活化,吸收利用土壤难溶态磷的可行性对策,从而为充分发挥植物自身潜力,提高土壤难溶态磷的生物有效性,高效利用土壤潜在磷资源,加快土壤磷素有效循环,节省资源,保护环境,真正达到生态系统的稳定与现代化农业     

三峡库区消落带4种适生植物根际与非根际土壤养分与酶活性特征研究

三峡库区消落带植被修复过程中,物种的更替对库区土壤的地球化学循环产生潜在影响。以三峡库区忠县石宝寨汝溪河消落带植被修复示范基地165-170 m海拔段人工种植狗牙根、牛鞭草、落羽杉以及立柳根际与非根际土为试验对象,探究其根际与非根际土壤的养分含量及酶活性差异,以阐明不同物种的生长适应性及其根际养分利用策略,比较不同物种对库区土壤的营养改良作用。结果表明：（1）三峡库区消落带4种适生植物根系活动导致根际与非根际土壤养分因子以及土壤酶活性产生差异,不同物种的栽植均在一定程度上使库区土壤营养条件得以改善;（2）碳、氮两种元素在4种适生植物根际土壤中发生不同程度的富集,但磷素与钾素在不同物种根际与非根际土壤之间的变化不一致;（3）蔗糖酶、脲酶以及酸性磷酸酶在4种适生植物根际土中均表现出一定程度的根际正效应（R/S>1）,且狗牙根对3种土壤酶的根际活化效果最为明显,其根际效应分别高达2.39、1.89和2.7;（4）在植物根系的调控下,根际土中有机质与氮素、磷素以及钾素的相关性更为显著,而非根际土壤中,仅钾素与有效氮、有效磷呈显著负相关,其余各土壤养分因子之间均无显著相关性;（5）与落羽杉和立柳两木本植物相比,狗牙根与牛鞭草两草本植物根际具有更为合理的养分调节模式,对库区土壤的改良效果更好。     

Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability

Low phosphorus availability stimulates root hair elongation in many plants, which may have adaptive significance in soil phosphorus acquisition. We investigated the effect of low phosphorus on the elongation of Arabidopsis thaliana root hairs. Arabidopsis thaliana plants were grown in plant culture containing high (1000 mmol m^?3) or low (1 mmol m^?3) phosphorus concentrations, and root hair elongation was analysed by image analysis. After 15d of growth, low-phosphorus plants developed root hairs averaging 0.9 mm in length while high-phosphorus plants of the same age developed root hairs averaging 0.3 mm in length. Increased root hair length in low-phosphorus plants was a result of both increased growth duration and increased growth rate. Root hair length decreased logarithmically in response to increasing phosphorus concentration. Local changes in phosphorus availability influenced root hair growth regardless of the phosphorus status of the plant. Low phosphorus stimulated root hair elongation in the hairless axr2 mutant, exogenously applied IAA stimulated root hair elongation in wild-type high-phosphorus plants and the auxin antagonist CM PA inhibited root hair elongation in low-phosphorus plants. These results indicate that auxin may be involved in the low-phosphorus response in root hairs.     

The efficiency of Arabidopsis thaliana (Brassicaceae) root hairs in phosphorus acquisition

Arabidopsis thaliana root hairs grow longer and denser in response to low-phosphorus availability. In addition, plants with the root hair response acquire more phosphorus than mutants that have root hairs that do not respond to phosphorus limiting conditions. The purpose of this experiment was to determine the efficiency of root hairs in phosphorus acquisition at high- and low-phosphorus availability. Root hair growth, root growth, root respiration, plant phosphorus uptake, and plant phosphorus content of 3-wk-old wild-type Arabidopsis (WS) were compared to two root hair mutants (rhd6 and rhd2) under high (54 mmol/m) and low (0.4 mmol/m) phosphorus availability. A cost-benefit analysis was constructed from the measurements to determine root hair efficiency. Under high-phosphorus availability, root hairs did not have an effect on any of the parameters measured. Under low-phosphorus availability, wild-type Arabidopsis had greater total root surface area, shoot biomass, phosphorus per root length, and specific phosphorus uptake. The cost-benefit analysis shows that under low phosphorus, wild-type roots acquire more phosphorus for every unit of carbon respired or unit of phosphorus invested into the roots than the mutants. We conclude that the response of root hairs to low-phosphorus availability is an efficient strategy for phosphorus acquisition.     

翅荚木人工林不同径阶间细根主要功能性状与根际土壤养分的关系

细根能敏感地感知土壤环境变化,对植物生长发育具有重要影响.以6年生翅荚木人工林为对象,对其不同径阶的细根主要功能性状与根际土壤养分特征及两者间关系进行分析.结果表明:细根生物量、根长密度与根体积密度均随径阶增加而增加,比根长与比根面积则随径阶增加呈先升高再下降后升高的趋势,根组织密度则与径阶大小不相关.不同径阶翅荚木根际土壤的pH值及含水率、全碳、全磷、铵态氮、硝态氮和总有效氮含量均存在显著差异,大径阶林木的根际土壤全碳、全氮、硝态氮、总有效氮含量相对较高,小径阶林木的根际土壤含水率、土壤全磷、铵态氮含量相对较高.土壤全氮、全碳、硝态氮和总有效氮含量与林木细根的生物量、根长密度、根体积密度呈显著正相关;土壤全磷与林木细根的根组织密度呈显著正相关,与比根长、比根面积呈显著负相关;土壤含水率与林木细根的生物量和根体积密度均呈显著正相关;根际土壤pH和林木细根的比根长、比根面积呈显著正相关,与根组织密度则呈显著负相关.研究结果可为翅荚木优良种质资源选育提供科学依据.     

Localized supply of phosphorus induces root morphological and architectural changes of rice in split and stratified soil cultures

A localized supply of phosphorus may affect root morphology and architecture, and thereby affect phosphorus uptake by rice plants. In the present study, we attempted to test this hypothesis using two rice cultivars representing upland and lowland ecotypes grown in specially designed split and stratified soil cultures with a low-phosphorus red soil. Our data indicate that a localized supply of phosphorus increased both total root length and root fineness, particularly in the high-phosphorus zone. In split culture, plants roots tended to preferentially grow on the high-phosphorus zone, with about 70–75% of the total root length allocated to the high-phosphorus compartment. The total root length on the high-phosphorus side in the split-phosphorus treatment was significantly longer than that in the homogenously high-phosphorus treatment, implying that a phosphorus-deficiency signal from the low-phosphorus side may stimulate the growth of the roots located in the high-phosphorus zone. In stratified soil culture, changes in root morphology and architecture were also observed as indicated by increased total root length, root fineness and relative root allocation in the high-phosphorus layers, again suggesting altered root morphology and preferential root proliferation in the high-phosphorus regions. The induced changes in root morphology and architecture by localized phosphorus supply may have both physiological significance and practical implications in that plants can meet the demand for phosphorus with parts of the roots reaching the high-phosphorus zone, hence localized fertilization methods such as side dressing or banded application of phosphorus fertilizers may both minimize phosphorus fixation by the soil and increase phosphorus uptake efficiency from the fertilizers.     

稻、麦根系H~ 的分泌与介质磷水平的关系

水稻、小麦根系H~ 的分泌量随供磷水平的降低而增加,并存在明显的昼夜变化。在自然光照下H~ 分泌量随光强度增加而增多,同时强光比黑暗时H~ 分泌对磷供应水平更为敏感。磷供应不足还诱导水稻根系柠檬酸分泌量增加,而苹果酸则差异不明显。难溶性磷的溶解率与根系H~ 和柠檬酸分泌所导致的根际pH下降有密切联系。因此,在有效磷不足的条件下可明显提高稻、麦根际土壤中难溶性磷的利用率,其中丰产型小麦和粳稻品种对土壤中磷利用的根际效应更为显著。     

Acquisition of phosphorus and nitrogen in the rhizosphere and plant growth promotion by microorganisms

The rhizosphere is a complex environment where roots interact with physical, chemical and biological properties of soil. Structural and functional characteristics of roots contribute to rhizosphere processes and both have significant influence on the capacity of roots to acquire nutrients. Roots also interact extensively with soil microorganisms which further impact on plant nutrition either directly, by influencing nutrient availability and uptake, or indirectly through plant (root) growth promotion. In this paper, features of the rhizosphere that are important for nutrient acquisition from soil are reviewed, with specific emphasis on the characteristics of roots that influence the availability and uptake of phosphorus and nitrogen. The interaction of roots with soil microorganisms, in particular with mycorrhizal fungi and non-symbiotic plant growth promoting rhizobacteria, is also considered in relation to nutrient availability and through the mechanisms that are associated with plant growth promotion.     

一株柠条内生解磷菌的分离鉴定及实时荧光定量PCR检测

结合形态观察与16S rDNA序列测定对柠条根系内分离筛选得到的解磷细菌C9进行鉴定,Blast比对结果表明C9为泛菌(Pantoea vagans).在无机磷液体培养基培养条件下,用钼锑抗比色法研究它的解磷能力.结果表明,随着时间的延长,培养液中速效磷含量逐渐增加到4.45mg/L,溶液pH可降至4.2.进一步利用实时荧光定量PCR检测柠条根系中、柠条根际土壤与柠条根围土壤中该细菌存在的相对基因拷贝数,结果发现该基因在3种样品中的数量为:柠条根系＞柠条根际土壤＞柠条根围土壤,表明泛菌解磷细菌能聚集生长在柠条根系内,随着与根系接触距离的增加而呈逐级递减趋势.     

Optimizing reproductive phenology in a two-resource world: a dynamic allocation model of plant growth predicts later reproduction in phosphorus-limited plants

Background and Aims

Timing of reproduction is a key life-history trait that is regulated by resource availability. Delayed reproduction in soils with low phosphorus availability is common among annuals, in contrast to the accelerated reproduction typical of other low-nutrient environments. It is hypothesized that this anomalous response arises from the high marginal value of additional allocation to root growth caused by the low mobility of phosphorus in soils.

Methods

To better understand the benefits and costs of such delayed reproduction, a two-resource dynamic allocation model of plant growth and reproduction is presented. The model incorporates growth, respiration, and carbon and phosphorus acquisition of both root and shoot tissue, and considers the reallocation of resources from senescent leaves. The model is parameterized with data from Arabidopsis and the optimal reproductive phenology is explored in a range of environments.

Key Results

The model predicts delayed reproduction in low-phosphorus environments. Reproductive timing in low-phosphorus environments is quite sensitive to phosphorus mobility, but is less sensitive to the temporal distribution of mortality risks. In low-phosphorus environments, the relative metabolic cost of roots was greater, and reproductive allocation reduced, compared with high-phosphorus conditions. The model suggests that delayed reproduction in response to low phosphorus availability may be reduced in plants adapted to environments where phosphorus mobility is greater.

Conclusions

Delayed reproduction in low-phosphorus soils can be a beneficial response allowing for increased acquisition and utilization of phosphorus. This finding has implications both for efforts to breed crops for low-phosphorus soils, and for efforts to understand how climate change may impact plant growth and productivity in low-phosphorus environments.     

毛竹根际与非根际土壤重金属、理化性质及酶活性特征

于2014年2月至2015年1月在浙江省衢州市铅锌矿附近毛竹林地采集土样,对土壤重金属含量、pH以及土壤酶活性等的动态变化特征进行了研究。结果表明:根际土与非根际土的p H在1a中变化均不明显,且变化范围都处在4.50—4.85之间;非根际土的土壤含水率大于根际土,两者的变化幅度较大,最低值均出现在6月和10月;根际土中Zn和Cd的有效态含量远高于非根际土,而Pb的有效态含量则表现出非根际土大于根际土的特点;过氧化氢酶、磷酸酶、脲酶和脱氢酶在根际土中的活性明显高于非根际土;从土壤重金属有效态与土壤酶活性相关性来看,根际土中,除有效态Pb与过氧化氢酶呈显著负相关外,其余重金属元素有效态与土壤酶活性相关性均没有显著相关性,而在非根际土壤中,有效态Cd与过氧化氢酶呈极显著负相关,与脲酶呈显著负相关,Zn和Cd则与土壤酶活性之间没有显著相关性。综合研究结果表明,毛竹根际土壤质量总体上明显优于非根际土,这为毛竹今后作为植物修复的材料提供了一定的理论依据。