植物磷获取机制及其对全球变化的响应
Plant phosphorus acquisition mechanisms and their response to global climate changes
通讯作者: * 范跃新: ORCID:0000-0001-8881-1956 (yxfan@fjnu.edu.cn)
编委: 陈保冬
责任编辑: 赵航
收稿日期: 2021-12-6 接受日期: 2022-06-6
基金资助: |
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Corresponding authors: * (yxfan@fjnu.edu.cn)
Received: 2021-12-6 Accepted: 2022-06-6
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磷是植物生长的必需元素, 而陆地生态系统普遍存在磷限制, 全球变化可能会影响土壤磷循环过程, 进一步加剧磷限制, 探讨植物磷获取策略对科学预测生态系统生产力如何适应全球变化具有重要意义。该文通过收集和梳理相关文献, 从4个方面综述植物的磷获取机制及其对全球变化的响应: 1)植物的磷饥饿响应机制; 2)植物的磷获取途径和策略; 3)土壤微生物对植物磷吸收的影响; 4)植物磷吸收对全球变化(温度升高、氮沉降和降水变化)的响应及其机制。该综述有助于深入理解全球变化背景下植物适应低磷胁迫的机理, 也可为养分资源管理实践提供理论依据。
关键词:
Phosphorus (P) is an essential but limited nutrient for plant growth, and global climate changes may affect soil P cycling and further aggravate P limitations in the soil. In this review, we focused on the response of plant P acquisition strategies to climate changes and subsequent influences on ecosystem productivity. By searching and analyzing the existing literatures, we summarized the P acquisition mechanism of plants and their response to global climate changes from following aspects: 1) plant P starvation response mechanisms; 2) plant P acquisition pathways and strategies; 3) involvements of soil microorganisms in plant P utilization; and 4) responses of plant P acquisition strategies to global climate changes (e.g., warming, nitrogen deposition and precipitation changes) and the underlying mechanisms. The review is expected to deepen our understanding of plant adaptation to low-P stress under the future climate scenario, and can also provide a theoretical basis for nutrient management in agriculture.
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引用本文
何敏, 许秋月, 夏允, 杨柳明, 范跃新, 杨玉盛.
HE Min, XU Qiu-Yue, XIA Yun, YANG Liu-Ming, FAN Yue-Xin, YANG Yu-Sheng.
土壤磷是植物获取磷的主要来源, 分为无机磷和有机磷两类。其中无机磷又分为水溶态磷、吸附态磷和矿物态磷; 有机磷则主要是磷酸肌醇、磷酯和核酸以及微生物生物量磷(McDowell & Stewart, 2006; Huang et al., 2017)。在多种磷形态中, 能够被植物直接吸收利用的部分是水溶态磷, 但其在土壤中含量极低, 且易被铁铝氧化物等吸附固定和优先被植物利用, 容易消耗(秦武明等, 2008)。因此, 其他形态的磷转化为水溶态磷是土壤维持磷供应的重要途径, 如吸附态磷在解吸附作用下释放磷酸根, 有机磷在磷酸酶水解之后释放无机磷等(Antelo et al., 2007)。探寻植物如何参与土壤磷转化并影响土壤磷的有效性, 对理解植物如何获取土壤磷至关重要。
微生物在植物磷获取过程中发挥重要作用, 能够帮助植物吸收根际之外的磷, 亦可促进难利用磷向有效磷的转化(朱倩等, 2018)。其中能够活化难利用磷的这部分微生物被称为溶磷微生物(phosphate-solubilizing microorganisms, PSMs), 主要通过以下途径参与磷的转化。一是PSMs具备分泌胞外酶的能力, 如磷酸酶、植酸酶、核酸酶、脱氢酶等(Richardson et al., 2001; 吉蓉, 2013), 在酶的作用下促进有机磷水解, 进而释放可被植物利用的无机磷(包鑫等, 2014)。二是PSMs通过分泌有机酸等物质, 促进吸附态磷和蓄闭态磷的解吸附并释放磷酸根。三是微生物直接参与磷转化, 微生物在生长过程中固定了部分磷(Turner et al., 2003), 在死亡后这部分磷归还至土壤, 成为植物的磷源之一。四是菌根真菌定植在植物上形成菌根共生体, 例如丛枝菌根和外生菌根, 能够有效增加植物从贫瘠土壤中获取流动性差的磷(Lambers et al., 2018)。
近几十年, 全球气候变化(温度升高、降水变化及氮沉降等)对植物生长及其养分获取的影响受到越来越多的关注, 气候因子的变化将直接或间接改变植物的磷吸收策略。许多研究通过设置野外控制实验来研究全球变化对土壤磷循环的影响, 取得了较大进展, 如Zhang等(2020)研究表明了氮沉降和降水格局变化都显著改变了土壤磷组分和磷循环。梳理全球变化对植物磷吸收过程的影响, 对理解植物未来如何适应气候变化具有重要意义。
已有研究表明植物获取磷是一个复杂的过程。在低磷条件下, 植物如何感受养分胁迫? 通过何种途径增加磷获取? 与土壤微生物如何协作? 如何响应全球变化? 针对以上问题, 本文综述已有相关研究, 从1)植物的磷饥饿响应; 2)植物的磷获取途径; 3)土壤微生物对植物磷吸收的影响; 4)植物磷吸收对全球变化(温度升高、氮沉降和降水变化)的响应及其机制等4个方面进行探讨, 以期为未来磷循环研究提供理论支持。
1 植物磷饥饿响应
土壤中磷的缺乏会使植物发生磷饥饿响应(phosphorus starvation responses, PSR), 具体表现在磷转运子表达变化, 细胞代谢途径和根系分泌物改变, 以及根系的形态和构型的调整等方面(李锋等, 2004; 图1)。在磷饥饿响应过程中, 植物首先感知磷缺乏, 然后产生信号级联来启动和协调适应反应(Vance et al., 2003; Franco-Zorrilla et al., 2004; Ticconi & Abel, 2004)。研究表明, 这些适应性反应受基因表达变化调控(Amtmann et al., 2005), 当植物生长受到磷限制时, 成千上万个基因的表达会发生改变(Wang et al., 2002; Wasaki et al., 2003; Wu et al., 2003)。这些基因被分为“早期”基因和“晚期”基因, “早期”基因反应迅速且短暂, 通常在植物面临磷饥饿的几小时之内反应, “晚期”基因在植物长时间处于磷饥饿时会改变植物形态、生理和代谢(Hammond et al., 2003)。已有研究发现, 磷饥饿响应基因有很大一部分受磷酸盐饥饿反应调控因子(如拟南芥(Arabidopsis thaliana)中的PHR1 (phosphate starvation response regulator 1))及其同源物(如水稻(Oryza sativa)中的PHR2)的调控(Wang et al., 2014b)。当植物受到低磷胁迫时, PHR1会从调控基因和负调控因子SPX1、SPX2的复合物中迅速解离出来, 对磷饥饿响应基因进行调控(González et al., 2005)。“早期”转录反应包括一般应激相关蛋白(如几丁质酶和过氧化物酶)的编码基因表达的增加(Wang et al., 2002; Hammond et al., 2003), 也包括编码各种转录因子(如HD-ZIP、WRKY转录因子、MYB-CC转录因子、bHLH DN结合蛋白)的基因表达的改变(Rubio et al., 2001; Wang et al., 2002; Hammond et al., 2003; Todd et al., 2004)、核糖体调节因子(如At4和TPS家族)的改变(Burleigh & Harrison, 1999), 以及细胞内信号级联的其他组成部分(如蛋白激酶和蛋白磷酸酶)的变化(Wang et al., 2002; Hammond et al., 2003)。“晚期”转录反应主要是适应性反应, 它能促进植物对土壤磷的吸收, 提高植物体内磷的再利用(Hammond et al., 2004)。例如在根系中, 编码PHT1磷转运蛋白家族成员(Smith et al., 2003; Wasaki et al., 2003)、胞内和胞外RNA酶(Wasaki et al., 2003)、磷酸酶(Stenzel et al., 2003; Wasaki et al., 2003)等基因的表达在磷饥饿过程中都逐渐增加。通过对十字花科植物根部的相关研究发现, 参与植物PSR相关的调控因子可能会影响植物微生物之间的磷养分竞争, 而微生物定植在植物体上会帮助植物积累磷, 影响PSR转录因子的表达水平, 同时刺激植物发生PSR (Castrillo et al., 2017)。
图1
根系分泌物是植物受磷胁迫的一种信号传递物质, 能够有效地帮助植物与土壤之间进行信息传递和物质交换(Chiou & Lin, 2011)。研究表明, 低磷环境会刺激植物根系分泌一系列代谢物, 包括初级代谢物(如糖类、氨基酸和有机酸)和次级代谢物(如酚类、黄酮类化合物、萜类化合物), 这些分泌物一方面可为微生物生长提供能量, 一方面作为信号物质调控植物根系和土壤微生物的相互作用(尹华军等, 2018)。如低磷条件下植物分泌的有机酸和酚类物质会显著增加, 如柠檬酸和酸性磷酸酶分泌增多促进了植物对磷的吸收和利用(许仙菊和张永春, 2018; 李靖怡, 2020)。张治宏等(2020)发现低磷处理下的菖蒲(Acorus calamus)、菱(Trapa bispinosa)、苦草(Vallisneria natans)和荇菜(Nymphoides peltatum)根系分泌的有机酸量均明显高于对照组。除有机酸外, 一些植物面对低磷胁迫时还会分泌质子, 如番茄(Lycopersicon esculentum)、鹰嘴豆(Cicer arietinum)、蚕豆(Vicia faba)通过增加质子的分泌来活化土壤难溶态磷, 从而提高了磷利用效率(Gardner et al., 1982; 张恩和等, 2004; 张瑜等, 2015)。此外, 根系分泌物与微生物关系密切, 初级代谢产物还会影响微生物多样性, 改变微生物群落结构, 进而影响土壤磷转化和磷有效性。同时土壤微生物也会影响根系分泌物的产生, 如通过释放特定化合物刺激根系代谢物如氨基酸等的变化(Lambers et al., 2018), 进而影响植物磷吸收。
植物根系构型的变化是植物磷饥饿响应的另一个特征, 如根和根毛的伸长和增多、菌根真菌侵染率增加等(刘灵等, 2008; Lynch, 2011)。研究发现, 磷饥饿下根系生长和结构的适应性变化受根和地上部分之间碳水化合物分配的影响, 这可能与植物激素、糖类等信号物质的产生有关(Phillips et al., 2004)。与缺磷响应类似, 生长素外部供应的改变会抑制初生根生长、促进侧根生长和根毛形成, 引起根表型的变化(Raghothama, 1999)。如磷饥饿时侧根的形成与转录调节因子(ARF19)调控的生长素变化有关, 而生长素通过赤霉素(GA)介导的生长抑制因子(DELLA蛋白: RGA和GAI)来影响根的生长(Nacry et al., 2005)。植物磷饥饿会降低GA的生物活性, 促进DELLA蛋白的积累(Fu & Harberd, 2003), 抑制了初生根的生长, 并促进了根毛的生长(Jiang et al., 2007)。还有研究发现, 初生根分生组织的停滞可能会触发糖的积累(Canarini et al., 2019)。当根尖分生组织衰竭时, 分生组织细胞的共质体连接遭到破坏(可能是由于胼胝体沉积), 根分泌物减少(Shishkova et al., 2008), 糖向根尖的持续运输会导致根系糖浓度的短暂增加。因此, 根系糖浓度可作为指示根系结构变化的一种信号分子(Canarini et al., 2019)。
2 植物磷获取途径
植物主要通过两种途径吸收土壤磷, 一是根表皮和根毛直接吸收, 另一种是通过菌根途径吸收土壤磷(Smith et al., 2004; Shi et al., 2021; 图2)。多数研究表明, 在低磷胁迫条件下, 植物通过增加根毛数量和长度、根系变细变长等方式, 扩大根系吸收面积以获取更多的磷(Lynch, 2011)。如低磷胁迫下大豆(Glycine max)的总根长、根表面积显著增加(孔令剑等, 2018); 棉花(Gossypium sp.)幼苗通过增加侧根数、比根长、分枝密度、根长密度和根毛长度, 同时降低根的平均直径, 促进根的伸展, 扩大根覆盖面积, 增加了磷的获取范围(Zhang et al., 2021)。然而, 在对驴食豆(Onobrychis viciifolia)新品系幼苗生长及生理特性的研究中, 低磷胁迫显著降低了驴食豆的根体积、根表面积、根长和根尖数(江海慧等, 2020)。除形态外, 根系还会发生生理上的变化。如Tang等(2013)发现低磷胁迫下, 白羽扇豆(Lupinus albus)根系对质子、柠檬酸和酸性磷酸酶的分泌增多。但不同植物分泌物的种类和数量会有不同, 可见, 不同植物的磷获取机制可能存在差异, 仅关注根系的形态和生理变化无法全面揭示植物应对低磷胁迫的策略。
图2
陆地上超过95%的植物可以和菌根真菌形成菌根共生体, 而菌根是植物获取土壤磷的另一个重要途径。由于菌丝可以达到植物根系触及不到的土壤区域, 可帮助植物扩大磷吸收面积(曹庆芹等, 2011)。因此在缺磷条件下, 植物更易与菌根真菌建立菌根共生体来增强自身从土壤中获取磷的能力(Raven et al., 2018)。如接种丛枝菌根真菌摩西管柄囊霉(Funneliformis mosseae, FM)和地表球囊霉(Glomus versiforme, GV)后, 南美蟛蜞菊(Sphagneticola trilobata)的磷含量分别增加了36.6%和40.7%, 这表明菌根共生体能促进植物的生长, 提高土壤难溶性磷的吸收效率(李琴等, 2020)。刘云龙等(2021)在研究丛枝菌根真菌对豆科作物磷吸收影响的实验中, 也证实了接种丛枝菌根真菌显著提高了大豆地上部分的磷含量和磷吸收量。此外, 不同菌根类型(外生菌根和丛枝菌根)促进植物磷吸收的机制也不相同(Rosling et al., 2016)。如外生菌根主要通过分泌磷酸酶来水解土壤中的有机磷(Häussling & Marschner, 1989), 而丛枝菌根分泌磷酸酶的能力相对较弱(Joner et al., 2000)。外生菌根真菌还可以产生低分子量有机酸和螯合剂以促进矿物磷的风化和无机磷释放(Plassard & Dell, 2010), 因此外生菌根植物通常具有比丛枝菌根真菌植物更强的土壤磷获取能力。
3 土壤微生物对植物磷获取的影响
PSMs对无机磷的溶解与小分子有机酸的分泌有关, 主要是葡萄糖酸、草酸、酮基葡萄糖酸、苹果酸、柠檬酸等(Marra et al., 2015; Sane & Mehta, 2015), 不同微生物类群合成的有机酸种类和含量不同, 溶磷效果也存在差异。如唐岷宸等(2020)从农田土壤中筛选出解磷能力最强的X-P18菌株, 并使用16S rDNA技术对该菌株进行了分析, 鉴定其为贝莱斯芽孢杆菌(Bacillus velezensis), 并通过测定发酵液中小分子有机酸种类发现其溶磷机理是分泌乙酸及其他少量小分子有机酸。有研究报道, 欧文氏菌(Erwinia)对矿物态磷的高效溶解作用, 其原理是葡萄糖脱氢酶(GDH)催化葡萄糖氧化后产生葡萄糖酸(Liu et al., 1992), 而葡萄糖酸对磷矿石进行溶解后释放大量无机磷(Goldstein, 1994)。但是, 骆韵涵等(2020)从罗源湾红树林根际土壤中分离溶磷菌, 筛选出洋葱伯克霍尔德菌(Burkholderia cepacia, NR113645.1), 发现其溶磷效果不佳, 原因可能与菌液中主要以丙酮酸、乳酸、乙酸和丁二酸等为主, 而葡萄糖酸量极低有关。部分溶磷微生物还可将葡萄糖酸进一步氧化为2-酮基葡萄糖酸, 其具有更强的溶磷效果(Goldstein, 1986)。然而一些微生物的GDH相关基因突变失活后就不能再溶解难溶性无机磷, 但可在吡咯喹啉醌(yr-roloquinoline quinone, PQQ)为辅基的条件下参与无机磷的溶解(Babu-Khan et al., 1995)。例如有人发现土壤细菌恶臭假单胞菌(Pseudomonas putida) KT2440可以通过分泌葡萄糖酸来溶解难溶性土壤磷酸盐, 同时还需要PQQ作为辅酶(An & Moe, 2016)。张健(2014)在分子水平上阐明了草酸青霉菌(Penicillium oxalicum) BK可通过提高编码苹果酸合成反应的催化酶基因表达, 增加苹果酸的分泌和积累, 从而缓解磷胁迫。何迪等(2020)发现草酸青霉菌HB1主要是通过分泌氢质子酸解含磷矿物来进行溶磷的, 结果也表明其对磷酸钙和磷矿粉都有较好的溶解效果。
土壤中有机磷一般不能被植物直接吸收利用, 而是要通过酸性磷酸酶、碱性磷酸酶、肌醇六磷酸酶等水解酶对有机磷进行分解后才能供植物吸收(陈哲等, 2009)。Torriani和Ludtka (1985)首次证实了大肠杆菌(Escherichia coli)中有关有机磷的溶解基因的存在。Thaller等(1994)根据分子大小将酸性磷酸酶分为A、B、C三类, 从革兰氏阴性杆菌(Gram negative bacillus)中分离克隆了酸性磷酸酶基因, 由acpA、phoC和napA等基因编码的酸性磷酸酶在pH 6.0时表现出了较强的溶磷活性。在农田土壤中筛选出的有机磷细菌GN1, 在不同磷源培养条件下基因表达差异明显, 长期低磷胁迫激发了GN1大量磷代谢基因的表达, 导致碱性磷酸酶活性持续升高(廖梓鹏, 2017)。来自动物排泄物和有机肥的肌醇六磷酸盐, 在肌醇六磷酸酶作用下水解后才可被植物利用(陈哲等, 2009)。已有研究从部分细菌中分离克隆出肌醇六磷酸酶基因(phy), 并证实了接种含有phyA基因的微生物明显提高了植物对肌醇六磷酸盐的利用能力(Asea et al., 1988)。
长期以来, 根系形态构型的改变、菌根真菌与植物的共生关系被认为可以有效改善磷的转运, 然而随着对细菌与植物的关联研究越来越深入, 发现内生菌也能够促进非结瘤植物在低营养环境中的生长(Santi et al., 2013)。有研究表明, 磷酸盐一旦进入植物体内, 很容易与植物中普遍存在的钙和其他金属离子发生反应, 形成不溶性磷酸盐, 而内生菌能够根据植物需求重新释放磷酸盐(Varga et al., 2020), 这说明了内生菌在植物磷的利用中起着重要作用。在植物组织内, 内生菌还能够从宿主中获取养分, 因此面临的营养限制较少, 并且不易受到环境胁迫(Gaiero et al., 2013)。Emami等(2020)报道了溶磷细菌对提升植物磷利用效率的作用, 对小麦(Triticum aestivum)同时接种根际细菌和内生细菌后, 两个小麦品种的磷效率指数分别提高29.5%和18.7%, 可能是接种细菌后有机酸含量和酶活性的增加提高了小麦的磷利用效率。由此可见, 无论是内生菌还是内生菌与根际微生物对植物的共同作用, 都对植物磷吸收有着重要意义。
4 全球变化对植物获取磷的影响
4.1 温度升高对土壤磷转化和植物磷吸收的影响
一是增温改变植物的磷吸收能力。如Zhou等(2021)对青藏高原高寒草甸进行了为期8年的增温实验, 发现地上和地下植物生物量中磷浓度都显著增加, 表明了增温条件下植物对磷的动员和同化能力提升; Rui等(2012)也观察到增温使得青藏高原草甸生态系统两种主要植物鹅绒委陵菜(Potentilla anserine)和麻花艽(Gentiana straminea)叶片中磷浓度显著增加。但Wang等(2021)的研究具有相反的结果, 对青藏高原草场进行连续增温3年后发现, 植物磷浓度下降了14.3%-49.4%; Yang等(2019)发现增温条件下亚热带杉木(Cunninghamia lanceolata)幼苗中的磷含量无显著变化。据此, 推测增温对植物磷吸收能力的影响因植被类型、增温时长等存在差异。
二是增温改变了土壤养分状况、pH等理化性质, 影响土壤磷循环(Yang et al., 2019; 付倩等, 2020)。由于土壤活性有机碳是微生物最主要的碳源, 有机碳含量变化将直接影响微生物活性, 进而影响土壤磷的转化(张恩平等, 2014)。此外, 有机碳含量高的土壤有利于降低土壤矿物对磷的固持, 使得土壤磷更容易被植物和微生物利用(陈智等, 2010)。因此, 土壤有机碳含量通常与磷有效性呈正相关关系(杨昊天等, 2018)。增温可能会促进土壤有机碳分解, 降低有机碳含量并影响磷的有效性。如王子龙等(2021)设置了不同水平的增温处理, 发现温度升高导致冻结期黑土有机碳含量显著降低。增温还会改变土壤养分计量比来影响土壤磷周转。如增温提高了南亚热带季风常绿阔叶林土壤的硝化速率, 增加了土壤氮有效性(赵建琪等, 2019), 有效氮含量的增加可能会改变氮磷养分平衡, 加剧土壤磷限制(Zhang et al., 2013)。
三是增温改变土壤微生物群落组成和活性, 进而影响土壤磷周转。研究表明, 温度升高能够增加某些特异性微生物的丰度, 从而可能直接促进根际磷的溶解和植物对磷的吸收(Jin et al., 2015)。例如Drigo等(2010)发现根际土壤中具有溶磷功能的假单胞菌的丰度在CO2浓度升高的条件下增加; Smith等(2004)发现温度升高增加了宿主植物根部的碳分配, 这刺激了丛枝菌根真菌丰度的增加, 菌丝网络扩大, 有助于宿主植物对磷的获取。这表明, 温度的升高可能有利于这些溶磷微生物的生长, 改变微生物及其活性。但这种效应的大小取决于土壤中含磷化合物和植物的种类, 而植物种类又决定其根际溶磷微生物的种类和丰度(Wenzel et al., 1994)。如增温改变了北方森林交错带微生物群落结构, 具体表现在变暖显著改变了细菌群落组成(如酸杆菌增加了11%, 放线菌增加了29%, 拟杆菌减少了29%)和外生菌根真菌群落的结构(以担子菌的变化为主, 其丰度减少了47%) (van Nuland et al., 2020); 但Fang等(2020)在中国黄土高原半干旱草原上的田间实验发现, 升温导致微生物代谢速率下降, 但未改变微生物的生物量。同时, 不可忽略的是, 增温诱导的微生物生物量和活性的增加将意味着这些微生物可能会与植物争夺更多的磷, 从而导致磷固化, 进一步影响植物对磷的吸收。
四是增温通过改变土壤磷酸酶活性调节有机磷矿化速率, 影响土壤有效磷含量(贝昭贤等, 2018)。不少研究发现增温可以提高土壤磷酸酶活性, 如Rui等(2012)发现酸性和碱性磷酸单酯酶的活性都因变暖而升高, 与此同时, 土壤有机磷矿化因温度和磷酸酶活性的增加而增加; 高CO2浓度诱导的升温下, 白毛羊胡子草(Eriophorum vaginatum)根际土壤中磷酸酶活性增加了254%, 使得在草丛中每年释放的磷增加了40%以上(Moorhead & Linkins, 1997)。但也有相反的结果, 如刘朝阳(2020)发现在农田土壤中, 酸性磷酸酶活性随温度升高呈下降趋势; 还有研究发现增温未显著影响高山草甸土壤的磷酸酶活性(Wang et al., 2014b)。不同的研究结果表明增温对土壤磷转化酶的影响机制相对复杂, 仍需要更为深入的研究探索其潜在机制。
4.2 氮沉降对土壤磷转化和植物磷吸收的影响
化石燃料的大量使用导致全球范围内氮沉降持续增加, 高额的氮输入可能改变土壤氮有效性、引起土壤酸化, 进而影响土壤磷转化(陈立新等, 2012)。首先, 氮沉降可能会刺激植物生长, 增加磷需求, 诱导植物的磷饥饿响应并驱动土壤不同磷组分之间的转化(Crowley et al., 2012; Fan et al., 2018)。如庞丽等(2014)发现氮沉降显著增加了马尾松(Pinus massoniana)苗高、生物量以及磷吸收效率等, 其原因是促进了根系生长和增加了表层土壤根系分布比例。Long等(2016)研究表明氮沉降可能会增强半干旱草原植物对磷的吸收, 且随着氮沉降的增加, 磷限制对于植物生长的限制不如预期那么重要。其次, 氮沉降还会影响凋落物的产量及其养分归还(陈美领等, 2016)。如张蔷等(2021)以杜鹃(Rhododendron simsii)灌丛为研究对象进行了为期两年的模拟氮沉降实验, 结果显示低氮、中氮和高氮处理样地的凋落物产量均高于对照样地; 在东北针叶林中也发现了类似的结果(Yan et al., 2018)。再者, 氮沉降还会影响凋落物分解速率, 有不少研究发现氮沉降会减少微生物以获取氮为目的的凋落物分解, 延缓了凋落物养分的归还, 可能会影响土壤磷的来源(Carreiro et al., 2000)。
氮沉降对土壤微生物组成和功能的影响也会影响土壤磷的转化, 如有研究发现氮添加降低了草地土壤溶磷细菌的丰度, 改变了溶磷细菌的群落组成, 最终导致有机磷矿化的增加和结合态无机磷溶解的减少(Widdig et al., 2019)。而与细菌相比, 真菌通常具有更强的溶磷功能(林启美等, 2001)。已有研究发现氮沉降改变了土壤真菌细菌比(F:B), 如氮添加后长白山温带森林有机层土壤的F:B显著升高(杨静怡等, 2020); 与之相反, 氮沉降降低了红椎(Castanopsis hystrix)人工幼林土壤的真菌生物量和F:B (洪丕征等, 2016)。也有研究发现氮添加没有改变F:B, 但随着氮添加水平升高丛枝菌根真菌的比例显著增加, 且长期的氮添加极大地增加了细菌应激指数, 并增强了和磷矿化有关的特定酶活性, 从而可能对植物的磷获取造成影响(Wang et al., 2018)。以上结果表明, 施氮水平、植被类型和季节变化等均可能影响土壤微生物对氮沉降的响应模式, 这一差异还可能改变土壤磷转化过程和土壤有效磷含量, 进而影响植物的磷吸收策略。
4.3 降水变化对土壤磷转化和植物磷吸收的影响
降水格局的改变是全球变化的重要内容, 会对植物生产力和土壤养分循环产生重大影响(Ruiz-Lozano et al., 1995)。首先, 降水的变化会影响土壤水分条件, 而土壤中水分含量又会影响土壤氧化还原条件, 从而对土壤中磷的吸附与释放产生影响, 改变土壤磷的有效性(张宝贵和李贵桐, 1998)。如Dijkstra等(2012)观察到土壤水分状况对于磷的有效性十分关键, 土壤有效磷含量随降水增加而减少, 植物中的磷含量也随土壤水分增加而增加(Hou et al., 2018)。而大量的降水控制实验发现, 隔离降水降低了植物的磷吸收, 主要体现在植物生长速率下降, 植物磷含量降低等方面(Munns, 1993; He & Dijkstra, 2014; Fan et al., 2021)。一方面, 干旱胁迫可能会限制植物光合能力和生长, 降低磷需求; 另一方面水分限制会引起植物根系形态和功能变化, 影响根系磷获取能力(Dunham & Nye, 1976)。如陈佳瑞等(2021)通过盆栽控水实验发现, 随着干旱的加剧, 紫丁香(Syringa oblata)、黄刺玫(Rosa xanthinal)、连翘(Forsythia suspensa) 3种灌木的叶片及茎的磷含量呈降低趋势, 认为可能与干旱影响植物磷吸收和在不同器官中的分配有关。另外, 水分是影响营养物质运输的重要介质, 降水增多也会对植物的磷吸收造成影响, 有研究发现增雨有利于水分对营养物质的运输, 可以提高植物的磷含量, 促进植株生长(张慧等, 2020)。
此外, 降水变化也会通过影响土壤微生物进而影响植物对磷的吸收。降水变化的扰动会影响土壤磷的可用性, 并可能导致植物和微生物对磷的吸收发生变化(Dijkstra et al., 2015)。如降水增加可以显著促进土壤微生物的生长(Nielsen & Ball, 2015), 而微生物的增加会促使植物和微生物之间的磷养分竞争加剧(Dijkstra et al., 2015)。许多研究都表明了土壤微生物生物量随着降水量的增加而增加(Cregger et al., 2012; Bell et al., 2014), 如增雨显著增加了内蒙古草原生态系统细菌丰度、Shannon-Wiener多样性指数、Chao 1指数和Faith系统发育多样性指数; Xu等(2022)发现降水增多还显著增加了丛枝菌根真菌丰富度。而植物与微生物对于磷的整体竞争随着干旱而减少, 因为干旱会抑制植物和微生物的生长, 两者对磷的需求减少(Dijkstra et al., 2015)。有研究发现水分与土壤微生物磷含量呈极显著的正相关关系, 水分减少抑制了微生物活性及其对磷的固持(贝昭贤等, 2018)。Kaisermann等(2017)发现干旱对细菌和真菌群落组成产生了遗留影响, 降低了同种土壤中的植物生长速率, 并对植物与土壤微生物养分竞争产生了影响。综上, 降水变化不仅通过影响土壤微生物数量和功能, 改变土壤磷周转过程和磷有效性, 还会通过影响植物生长来改变植物的磷需求。因此, 降水格局变化可能会改变磷的供需平衡, 影响植物磷吸收。
5 问题和展望
植物获取土壤磷的机制十分复杂, 特别是分子水平上植物如何感受低磷胁迫并作出响应的调控机制仍有待深入研究。虽然在农田生态系统中, 已对植物磷饥饿响应的基因表达、信号物质合成等做了有益探索, 但在森林生态系统中是否存在类似调控机制还缺乏研究。
植物与土壤微生物的相互作用, 如植物根系病害、豆科植物结瘤、菌根共生关系以及植物根系与微生物之间的非侵入性共生关系等均可能影响植物获取土壤磷的策略。目前已对拟南芥、水稻等植物的PHRs调控网络及其信号机制做了有益探索, 但尚不清楚不同类型植物的关键调控因子是否相同, 未来研究应继续验证和探索。
全球变化对植物磷获取策略的影响是一个综合多因素的过程, 但当前控制实验多关注单因子变化对磷转化的影响。亟需开展气温升高、氮沉降、水分变化等多因子交互实验, 揭示全球变化对植物磷获取策略及陆地生态系统磷循环的影响机制。
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Nutrient sensing and signalling in plants: potassium and phosphorus
Regulation of pyrroloquinoline quinone-dependent glucose dehydrogenase activity in the model rhizosphere-dwelling bacterium Pseudomonas putida KT2440
DOI:10.1128/AEM.00813-16
URL
[本文引用: 1]
\n Soil-dwelling microbes solubilize mineral phosphates by secreting gluconic acid, which is produced from glucose by a periplasmic glucose dehydrogenase (GDH) that requires pyrroloquinoline quinone (PQQ) as a redox coenzyme. While GDH-dependent phosphate solubilization has been observed in numerous bacteria, little is known concerning the mechanism by which this process is regulated. Here we use the model rhizosphere-dwelling bacterium\n Pseudomonas putida\n KT2440 to explore GDH activity and PQQ synthesis, as well as gene expression of the GDH-encoding gene (\n gcd\n ) and PQQ biosynthesis genes (\n pqq\n operon) while under different growth conditions. We also use reverse transcription-PCR to identify transcripts from the\n pqq\n operon to more accurately map the operon structure. GDH specific activity and PQQ levels vary according to growth condition, with the highest levels of both occurring when glucose is used as the sole carbon source and under conditions of low soluble phosphate. Under these conditions, however, PQQ levels limit\n in vitro\n phosphate solubilization. GDH specific activity data correlate well with\n gcd\n gene expression data, and the levels of expression of the\n pqqF\n and\n pqqB\n genes mirror the levels of PQQ synthesized, suggesting that one or both of these genes may serve to modulate PQQ levels according to the growth conditions. The\n pqq\n gene cluster (\n pqqFABCDEG\n ) encodes at least two independent transcripts, and expression of the\n pqqF\n gene appears to be under the control of an independent promoter and terminator.\n
Adsorption of a soil humic acid at the surface of goethite and its competitive interaction with phosphate
DOI:10.1016/j.geoderma.2006.10.011 URL [本文引用: 1]
Inorganic phosphate solubilization by two Penicillium species in solution culture and soil
DOI:10.1016/0038-0717(88)90058-2 URL [本文引用: 1]
Cloning of a mineral phosphate-solubilizing gene from Pseudomonas cepacia
DOI:10.1128/aem.61.3.972-978.1995
PMID:7540821
[本文引用: 1]
We have recently shown that the ability of some gram-negative bacteria to dissolve poorly soluble calcium phosphates (Mps+ phenotype) is the result of periplasmic oxidation of glucose to gluconic acid via the quinoprotein glucose dehydrogenase (GDH), a component of the direct oxidation pathway. Escherichia coli K-12 derivatives synthesize apo-GDH but not the cofactor pyrroloquinoline-quinone (PQQ) essential for formation of the holoenzyme. Therefore, in the absence of exogenous PQQ, these strains do not produce gluconic acid and are Mps-. Evidence is presented to show that expression of a single 396-base Pseudomonas cepacia open reading frame (designated gabY) in E. coli JM109 (a K-12 derivative) was sufficient to induce the Mps+ phenotype and production of gluconic acid. We present the nucleotide sequence of this open reading frame which coded for a protein (GabY) with a deduced M(r) of 14,235. Coupled transcription-translation of a plasmid (pSLY4 or pGAB1) carrying gabY resulted in production of a protein with an M(r) of 14,750. Disruption of the open reading frame of gabY via site-directed mutagenesis changed the phenotype to Mps- and eliminated gluconic acid production. The deduced amino acid sequence of gabY has no apparent homology with those of previously cloned direct oxidation pathway genes but does share regions highly homologous with the histidine permease system membrane-bound protein HisQ as well as other proteins in this family. In the presence of 1 microM exogenous PQQ, both JM109(pSLY4) and JM109(pGAB1) produced 10 times as much gluconic acid as was seen with either the plasmid or exogenous PQQ alone.(ABSTRACT TRUNCATED AT 250 WORDS)
Research progress of phosphate solubilizing microorganisms
解磷微生物的研究进展
Effects of simulated warming on soil phosphorus availability in subtropical Chinese fir plantation
模拟增温对中亚热带杉木人工林土壤磷有效性的影响
Soil microbial and nutrient responses to 7 years of seasonally altered precipitation in a Chihuahuan Desert grassland
DOI:10.1111/gcb.2014.20.issue-5 URL [本文引用: 1]
The down-regulation of Mt4-like genes by phosphate fertilization occurs systemically and involves phosphate translocation to the shoots
Mt4 is a cDNA representing a phosphate-starvation-inducible gene from Medicago truncatula that is down-regulated in roots in response to inorganic phosphate (Pi) fertilization and colonization by arbuscular mycorrhizal fungi. Split-root experiments revealed that the expression of the Mt4 gene in M. truncatula roots is down-regulated systemically by both Pi fertilization and colonization by arbuscular mycorrhizal fungi. A comparison of Pi levels in these tissues suggested that this systemic down-regulation is not caused by Pi accumulation. Using a 30-bp region of the Mt4 gene as a probe, Pi-starvation-inducible Mt4-like genes were detected in Arabidopsis and soybean (Glycine max L.), but not in corn (Zea mays L.). Analysis of the expression of the Mt4-like Arabidopsis gene, At4, in wild-type Arabidopsis and pho1, a mutant unable to load Pi into the xylem, suggests that Pi must first be translocated to the shoot for down-regulation to occur. The data from the pho1 and split-root studies are consistent with the presence of a translocatable shoot factor responsible for mediating the systemic down-regulation of Mt4-like genes in roots.
Root exudation of primary metabolites: mechanisms and their roles in plant responses to environmental stimuli
DOI:10.3389/fpls.2019.00157 [本文引用: 2]
Advance of plant phosphorus uptake improved by mycorrhiza fungi
菌根真菌促进植物磷吸收研究进展
Microbial enzyme shifts explain litter decay responses to simulated nitrogen deposition
DOI:10.1890/0012-9658(2000)081[2359:MESELD]2.0.CO;2 URL [本文引用: 1]
Root microbiota drive direct integration of phosphate stress and immunity
DOI:10.1038/nature21417 [本文引用: 1]
Population dynamics of Ranalisma rostratum, an endangered species growing in different habitats
不同生境内濒危植物长喙毛茛泽泻种群数量动态比较
Effects of drought stress on the stoichiometric characteristics in different organs of three shrub species
干旱胁迫对3种灌木不同器官化学计量特征的影响
DOI:10.13287/j.1001-9332.202101.002
[本文引用: 1]
采用盆栽控水试验,研究了3个水分处理,即田间持水量(FC)的(75±5)%、(55±5)%和(35±5)%,对丁香、黄刺梅、连翘的叶、茎、极细根(0~1 mm)、细根(1~2 mm)和粗根(>2 mm)化学计量特征的影响。结果表明: 3种灌木相同器官间氮(N)和磷(P)含量以及C∶N、C∶P、N∶P均存在显著差异。随着干旱胁迫的加剧,3种灌木各器官C含量总体上无显著变化;叶片N含量呈增加趋势,茎N含量逐渐下降,极细根和细根N含量均呈先上升后下降的趋势;叶片、茎P 含量呈降低趋势,极细根和细根P含量呈先增加后下降的趋势。3种灌木在干旱胁迫下叶C∶N呈降低趋势,叶和茎C∶P、N∶P均呈增加趋势。干旱胁迫对极细根C∶N和叶C∶P、N∶P影响最大,而对粗根C∶N、N∶P以及极细根C∶P的影响最小。土壤C、N含量与灌木各器官中C、N、P含量无显著相关性,但土壤P含量与叶和根系C、N、P含量均显著相关。土壤中相对缺乏的P是影响灌木器官化学计量特征的最重要因素。干旱对不同灌木不同器官化学计量的影响不同,叶和极细根的化学计量较其他器官对干旱胁迫更加敏感。干旱可能主要通过影响灌木对土壤中养分P的吸收和在不同器官中的分配,进而影响灌木不同器官的化学计量特征,尤其是与P相关的化学计量特征。
Influence of simulated nitrogen deposition on soil nitrogen mineralization rate under different forest stands
模拟氮沉降对温带不同森林类型土壤氮矿化速率的影响
Effect of nitrogen deposition on the soil phosphorus cycle in forest ecosystems: a review
氮沉降对森林土壤磷循环的影响
Advances in research on molecular mechanisms of phosphate-solubilizing microorganisms in soil
土壤微生物溶磷分子机理研究进展
Short-term effects of night warming and nitrogen addition on soil available nitrogen and microbial properties in subalpine coniferous forest, western Sichuan, China
DOI:10.3773/j.issn.1005-264x.2010.11.002
[本文引用: 1]
<p><em>Aims</em> The subalpine coniferous forest in eastern Qinghai-Tibet Plateau provides a natural laboratory for studying effects of climate change on terrestrial ecosystems. Research on responses of soil nitrogen availability and microbial properties to experimental warming and nitrogen addition can provide insights into their C resource/sink function under future climates. Methods We used an infrared heater combined with nitrogen addition to determine the short-term influences of two level of air temperature (ambient and warmed) and nitrogen addition (0 and 25 g N·m<sup>–2</sup>·a<sup>–1</sup>) on soil chemical properties, available nitrogen and microbial biomass. Important findings The warming manipulation increased mean air temperature and soil temperature at 5-cm depth by 1.93 and 4.19 °C, respectively, and the temperature increment was larger in summer and winter. Warming generally had no significant effects on soil pH, organic C, total N and microbial biomass, but it decreased soil ammonium nitrogen and increased nitrate nitrogen content. The warming effect was reduced with time. Nitrogen addition significantly increased available nitrogen and microbial biomass, but decreased soil pH, acidifying the soil. Compared with individually warming or adding nitrogen, the interaction of the two factors significantly increased organic C, available N and microbial biomass. Results suggest that soil nitrogen availability and microbial properties were sensitive to N status. Although soil nitrogen availability and microbial properties may adapt to temperature increase to some extent, the interaction of the warming and nitrogen addition significantly changed their response mode. Therefore, nitrogen deposition and multiple factors have interacting effects on the ecosystem and these should be further studied in this region.</p>
夜间增温和施氮对川西亚高山针叶林土壤有效氮和微生物特性的短期影响
DOI:10.3773/j.issn.1005-264x.2010.11.002
[本文引用: 1]
开展亚高山针叶林典型林地土壤有效氮和微生物特性对气候变化的响应研究, 对预测未来气候变化背景下亚高山针叶林生态系统C、N的源/汇功能具有重要意义。该文采用红外辐射加热器模拟增温结合外施氮肥的方法, 研究了川西亚高山针叶林下土壤化学特性、有效氮含量以及微生物生物量对夜间增温和施氮的短期响应。结果表明: 在模拟增温试验期间(2009年4月–2010年4月), 空气平均温度和5 cm土壤平均温度分别比对照提高了1.93和4.19 ℃, 增温幅度分别以夏季和冬季最为显著。增温对土壤pH值、有机碳、全氮和微生物生物量无显著影响。增温在试验前期降低了土壤NH<sub>4</sub><sup>+</sup>-N含量, 增加了NO<sub>3</sub><sup>–</sup>-N 含量, 其影响程度随着增温时间的延长而下降。施氮显著增加了有效氮和微生物生物量氮, 降低了土壤pH值, 使土壤表现出明显的酸化现象。与单独的增温和施氮处理相比, 增温和施氮联合处理对林下土壤的有效氮和微生物特性有显著的交互作用,显著增加了土壤的有机碳、有效氮及土壤微生物生物量氮含量, 并导致土壤进一步酸化。结果说明, 川西亚高山针叶林的土壤有效氮和微生物特性对土壤氮素状况的变化反应敏感, 而林下土壤有效氮和微生物特性对单独的温度升高表现出一定的适应性, 但更对增温和施氮双因素结合处理反应敏感且表现出不同的响应方式。因此, 该区域在未来全球变化下的氮沉降状况及气候变化的多因素协同效应值得长期深入的探讨。
Signaling network in sensing phosphate availability in plants
DOI:10.1146/arplant.2011.62.issue-1 URL [本文引用: 1]
Response of the soil microbial community to changes in precipitation in a semiarid ecosystem
DOI:10.1128/AEM.02050-12
PMID:23023755
[本文引用: 1]
Microbial communities regulate many belowground carbon cycling processes; thus, the impact of climate change on the structure and function of soil microbial communities could, in turn, impact the release or storage of carbon in soils. Here we used a large-scale precipitation manipulation (+18%, -50%, or ambient) in a piñon-juniper woodland (Pinus edulis-Juniperus monosperma) to investigate how changes in precipitation amounts altered soil microbial communities as well as what role seasonal variation in rainfall and plant composition played in the microbial community response. Seasonal variability in precipitation had a larger role in determining the composition of soil microbial communities in 2008 than the direct effect of the experimental precipitation treatments. Bacterial and fungal communities in the dry, relatively moisture-limited premonsoon season were compositionally distinct from communities in the monsoon season, when soil moisture levels and periodicity varied more widely across treatments. Fungal abundance in the drought plots during the dry premonsoon season was particularly low and was 4.7 times greater upon soil wet-up in the monsoon season, suggesting that soil fungi were water limited in the driest plots, which may result in a decrease in fungal degradation of carbon substrates. Additionally, we found that both bacterial and fungal communities beneath piñon pine and juniper were distinct, suggesting that microbial functions beneath these trees are different. We conclude that predicting the response of microbial communities to climate change is highly dependent on seasonal dynamics, background climatic variability, and the composition of the associated aboveground community.
Do nutrient limitation patterns shift from nitrogen toward phosphorus with increasing nitrogen deposition across the northeastern United States
DOI:10.1007/s10021-012-9550-2 URL [本文引用: 1]
Plant and microbial uptake of nitrogen and phosphorus affected by drought using 15N and 32P tracers
DOI:10.1016/j.soilbio.2014.12.021 URL [本文引用: 3]
Climate change alters stoichiometry of phosphorus and nitrogen in a semiarid grassland
DOI:10.1111/j.1469-8137.2012.04349.x
PMID:23005343
[本文引用: 1]
Nitrogen (N) and phosphorus (P) are essential nutrients for primary producers and decomposers in terrestrial ecosystems. Although climate change affects terrestrial N cycling with important feedbacks to plant productivity and carbon sequestration, the impacts of climate change on the relative availability of N with respect to P remain highly uncertain. In a semiarid grassland in Wyoming, USA, we studied the effects of atmospheric CO(2) enrichment (to 600 ppmv) and warming (1.5/3.0°C above ambient temperature during the day/night) on plant, microbial and available soil pools of N and P. Elevated CO(2) increased P availability to plants and microbes relative to that of N, whereas warming reduced P availability relative to N. Across years and treatments, plant N : P ratios varied between 5 and 18 and were inversely related to soil moisture. Our results indicate that soil moisture is important in controlling P supply from inorganic sources, causing reduced P relative to N availability during dry periods. Both wetter soil conditions under elevated CO(2) and drier conditions with warming can further alter N : P. Although warming may alleviate N constraints under elevated CO(2), warming and drought can exacerbate P constraints on plant growth and microbial activity in this semiarid grassland.© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.
Shifting carbon flow from roots into associated microbial communities in response to elevated atmospheric CO2
DOI:10.1073/pnas.0912421107
PMID:20534474
[本文引用: 1]
Rising atmospheric CO(2) levels are predicted to have major consequences on carbon cycling and the functioning of terrestrial ecosystems. Increased photosynthetic activity is expected, especially for C-3 plants, thereby influencing vegetation dynamics; however, little is known about the path of fixed carbon into soil-borne communities and resulting feedbacks on ecosystem function. Here, we examine how arbuscular mycorrhizal fungi (AMF) act as a major conduit in the transfer of carbon between plants and soil and how elevated atmospheric CO(2) modulates the belowground translocation pathway of plant-fixed carbon. Shifts in active AMF species under elevated atmospheric CO(2) conditions are coupled to changes within active rhizosphere bacterial and fungal communities. Thus, as opposed to simply increasing the activity of soil-borne microbes through enhanced rhizodeposition, elevated atmospheric CO(2) clearly evokes the emergence of distinct opportunistic plant-associated microbial communities. Analyses involving RNA-based stable isotope probing, neutral/phosphate lipid fatty acids stable isotope probing, community fingerprinting, and real-time PCR allowed us to trace plant-fixed carbon to the affected soil-borne microorganisms. Based on our data, we present a conceptual model in which plant-assimilated carbon is rapidly transferred to AMF, followed by a slower release from AMF to the bacterial and fungal populations well-adapted to the prevailing (myco-)rhizosphere conditions. This model provides a general framework for reappraising carbon-flow paths in soils, facilitating predictions of future interactions between rising atmospheric CO(2) concentrations and terrestrial ecosystems.
The influence of soil water content on the uptake of ions by roots. III. Phosphate, potassium, calcium and magnesium uptake and concentration gradients in soil
DOI:10.2307/2402270 URL [本文引用: 1]
Consortium of endophyte and rhizosphere phosphate solubilizing bacteria improves phosphorous use efficiency in wheat cultivars in phosphorus deficient soils
DOI:10.1016/j.rhisph.2020.100196 [本文引用: 1]
Decreased soil organic P fraction associated with ectomycorrhizal fungal activity to meet increased P demand under N application in a subtropical forest ecosystem
DOI:10.1007/s00374-017-1251-8 URL [本文引用: 1]
Long-term throughfall exclusion decreases soil organic phosphorus associated with reduced plant roots and soil microbial biomass in a subtropical forest
DOI:10.1016/j.geoderma.2021.115309 [本文引用: 1]
Unaltered soil microbial community composition, but decreased metabolic activity in a semiarid grassland after two years of passive experimental warming
DOI:10.1002/ece3.6862
PMID:33209291
[本文引用: 1]
Soil microbial communities regulate soil carbon feedbacks to climate warming through microbial respiration (i.e., metabolic rate). A thorough understanding of the responses of composition, biomass, and metabolic rate of soil microbial community to warming is crucial to predict soil carbon stocks in a future warmer climate. Therefore, we conducted a field manipulative experiment in a semiarid grassland on the Loess Plateau of China to evaluate the responses of the soil microbial community to increased temperature from April 2015 to December 2017. Soil temperature was 2.0°C higher relative to the ambient when open-top chambers (OTCs) were used. Warming did not affect microbial biomass or the composition of microbial functional groups. However, warming significantly decreased microbial respiration, directly resulting from soil pH decrease driven by the comediation of aboveground biomass increase, inorganic nitrogen increase, and moisture decrease. These findings highlight that the soil microbial community structure of semiarid grasslands resisted the short-term warming by 2°C, although its metabolic rate declined.© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Direct interaction between the arbuscular mycorrhizal fungus Glomus intraradices and different rhizosphere microorganisms
DOI:10.1046/j.1469-8137.1999.00366.x URL [本文引用: 1]
The transcriptional control of plant responses to phosphate limitation
DOI:10.1093/jxb/erh009
PMID:14718495
[本文引用: 1]
Plants have evolved an array of responses that adapt their growth to conditions of limited phosphate (Pi) supply. These involve biochemical and developmental changes that improve Pi acquisition and recycling, and protect against the stress of Pi starvation. The induction of these responses requires a sophisticated regulatory system that integrates information on external and internal plant Pi status and the details of this regulatory system are only just beginning to be elucidated. In this review, the current knowledge of this regulatory system is summarized, the hallmark of which is the central role of transcription factor PHR1 in the co-ordinated regulation of many phosphate-starvation-responsive genes. The role of hormonal signalling is also described, including auxins, ethylene and, particularly, cytokinins in the regulation of Pi-starvation responses.
Effects of warming and yak excreta addition on enzyme activity of marsh soil
增温和牦牛排泄物输入对沼泽土壤酶活性的影响
Auxin promotes Arabidopsis root growth by modulating gibberellin response
DOI:10.1038/nature01387 URL [本文引用: 1]
Inside the root microbiome: bacterial root endophytes and plant growth promotion
DOI:10.3732/ajb.1200572
PMID:23935113
[本文引用: 1]
Bacterial root endophytes reside in a vast number of plant species as part of their root microbiome, with some being shown to positively influence plant growth. Endophyte community structure (species diversity: richness and relative abundances) within the plant is dynamic and is influenced by abiotic and biotic factors such as soil conditions, biogeography, plant species, microbe-microbe interactions and plant-microbe interactions, both at local and larger scales. Plant-growth-promoting bacterial endophytes (PGPBEs) have been identified, but the predictive success at positively influencing plant growth in field conditions has been limited. Concurrent to the development of modern molecular techniques, the goal of predicting an organism's ability to promote plant growth can perhaps be realized by more thorough examination of endophyte community dynamics. This paper reviews the drivers of endophyte community structure relating to plant growth promotion, the mechanisms of plant growth promotion, and the current and future use of molecular techniques to study these communities.
The acquisition of phosphorus by Lupinus albus L. II. The effect of varying phosphorus supply and soil type on some characteristics of the soil/root interface
DOI:10.1007/BF02374725 URL [本文引用: 1]
Bacterial solubilization of mineral phosphates: historical perspective and future prospects
DOI:10.1017/S0889189300000886
URL
[本文引用: 1]
Maximum crop yields require sufficient phosphorus fertilization. Only phosphate in a soluble ionic form (Pi) is effective as a mineral nutrient. Current fertilizer technology supplies the soil solution with Pi via the application of large amounts of phosphate salts. Problems with this technology include energy-intensive production processes, the need for large scale mechanical application with associated environmental consequences, and reprecipitation of the phosphate into insoluble mineral complexes. It has been estimated that in some soils up to 75% of applied phosphate fertilizer may be lost to the plant because of mineral phase reprecipitation. Many approaches, ranging from cultural practices to biological inoculants such as mycorrhizal fungi, are being employed to enhance P-use efficiency. One area that is currently under-investigated is the ability of certain types of bacteria to solubilize mineral and organic phosphates. A review of the literature in the area of bacterial phosphate solubilization confirms that this trait is displayed by a wide range of bacteria. The phosphate starvation inducible (PSI) organic phosphate-solubilizing capability of E. coli is a component of a coordinately regulated gene system: the pho regulon. It has long been known that bacteria are also capable of solubilizing mineral phosphates such as hydroxyapatite. To date there has been no systematic study of the genetics of this phenomenon. Data from my laboratory indicate that the bacterial mineral phosphate-solubilizing (MPS) trait is regulated by the external level of Pi This conclusion is supported by results obtained from several types of molecular genetic studies. It is proposed that bacteria have mineral phosphate solubilizing (mps) genes. The potential agronomic applications of bacterial mineral and organic P solubilizing systems are discussed.
Phosphate transporter traffic facilitator is a plant-specific SEC12-related protein that enables the endoplasmic reticulum exit of a high-affinity phosphate transporter in Arabidopsis
DOI:10.1105/tpc.105.036640
URL
[本文引用: 1]
PHOSPHATE TRANSPORTER1 (PHT1) genes encode phosphate (Pi) transporters that play a fundamental role in Pi acquisition and remobilization in plants. Mutation of the Arabidopsis thaliana PHOSPHATE TRANSPORTER TRAFFIC FACILITATOR1 (PHF1) impairs Pi transport, resulting in the constitutive expression of many Pi starvation–induced genes, increased arsenate resistance, and reduced Pi accumulation. PHF1 expression was detected in all tissues, particularly in roots, flowers, and senescing leaves, and was induced by Pi starvation, thus mimicking the expression patterns of the whole PHT1 gene family. PHF1 was localized in endoplasmic reticulum (ER), and mutation of PHF1 resulted in ER retention and reduced accumulation of the plasma membrane PHT1;1 transporter. By contrast, the PIP2A plasma membrane protein was not mislocalized, and the secretion of Pi starvation–induced RNases was not affected in the mutant. PHF1 encodes a plant-specific protein structurally related to the SEC12 proteins of the early secretory pathway. However, PHF1 lacks most of the conserved residues in SEC12 proteins essential as guanine nucleotide exchange factors. Although it functions in early secretory trafficking, PHF1 likely evolved a novel mechanism accompanying functional specialization on Pi transporters. The identification of PHF1 reveals that plants are also endowed with accessory proteins specific for selected plasma membrane proteins, allowing their exit from the ER, and that these ER exit cofactors may have a phylum-specific origin.
Changes in gene expression in Arabidopsis shoots during phosphate starvation and the potential for developing smart plants
DOI:10.1104/pp.103.020941
PMID:12805589
[本文引用: 4]
Our aim was to generate and prove the concept of "smart" plants to monitor plant phosphorus (P) status in Arabidopsis. Smart plants can be genetically engineered by transformation with a construct containing the promoter of a gene up-regulated specifically by P starvation in an accessible tissue upstream of a marker gene such as beta-glucuronidase (GUS). First, using microarrays, we identified genes whose expression changed more than 2.5-fold in shoots of plants growing hydroponically when P, but not N or K, was withheld from the nutrient solution. The transient changes in gene expression occurring immediately (4 h) after P withdrawal were highly variable, and many nonspecific, shock-induced genes were up-regulated during this period. However, two common putative cis-regulatory elements (a PHO-like element and a TATA box-like element) were present significantly more often in the promoters of genes whose expression increased 4 h after the withdrawal of P compared with their general occurrence in the promoters of all genes represented on the microarray. Surprisingly, the expression of only four genes differed between shoots of P-starved and -replete plants 28 h after P was withdrawn. This lull in differential gene expression preceded the differential expression of a new group of 61 genes 100 h after withdrawing P. A literature survey indicated that the expression of many of these "late" genes responded specifically to P starvation. Shoots had reduced P after 100 h, but growth was unaffected. The expression of SQD1, a gene involved in the synthesis of sulfolipids, responded specifically to P starvation and was increased 100 h after withdrawing P. Leaves of Arabidopsis bearing a SQD1::GUS construct showed increased GUS activity after P withdrawal, which was detectable before P starvation limited growth. Hence, smart plants can monitor plant P status. Transferring this technology to crops would allow precision management of P fertilization, thereby maintaining yields while reducing costs, conserving natural resources, and preventing pollution.
Genetic responses to phosphorus deficiency
DOI:10.1093/aob/mch156
PMID:15292042
[本文引用: 1]
Phosphorus (P) is an essential macronutrient for plants. Plants take up P as phosphate (Pi) from the soil solution. Since little Pi is available in most soils, P fertilizers are applied to crops. However, the use of P fertilizers is unsustainable and may cause pollution. Consequently, there is a need to develop more P-use-efficient (PUE) crops and precise methods to monitor crop P-status.Manipulating the expression of genes to improve the PUE of crops could reduce their P fertilizer requirement. This has stimulated research towards the identification of genes and signalling cascades involved in plant responses to P deficiency. Genes that respond to P deficiency can be grouped into 'early' genes that respond rapidly and often non-specifically to P deficiency, or 'late' genes that impact on the morphology, physiology or metabolism of plants upon prolonged P deficiency.The use of micro-array technology has allowed researchers to catalogue the genetic responses of plants to P deficiency. Genes whose expression is altered by P deficiency include various transcription factors, which are thought to coordinate plant responses to P deficiency, and other genes involved in P acquisition and tissue P economy. Several common cis-regulatory elements have been identified in the promoters of these genes, suggesting that their expression might be coordinated. It is suggested that knowledge of the genes whose expression changes in response to P deficiency might allow the development of crops with improved PUE, and could be used in diagnostic techniques to monitor P deficiency in crops either directly using 'smart' indicator plants or indirectly through transcript profiling. The development of crops with improved PUE and the adoption of diagnostic technology could reduce production costs, minimize the use of a non-renewable resource, reduce pollution and enhance biodiversity.
Organic and inorganic soil phosphates and acid phosphatase activity in the rhizosphere of 80-year-old Norway spruce [Picea abies (L.) Karst.] trees
Ability and mechanism of Penicillium oxalicum HB1 solubilizing phosphates
草酸青霉菌HB1溶磷能力及作用机制
Drought effect on plant nitrogen and phosphorus: a meta-analysis
DOI:10.1111/nph.12952
PMID:25130263
[本文引用: 1]
Climate change scenarios forecast increased aridity in large areas worldwide with potentially important effects on nutrient availability and plant growth. Plant nitrogen and phosphorus concentrations (plant [N] and [P]) have been used to assess nutrient limitation, but a comprehensive understanding of drought stress on plant [N] and [P] remains elusive. We conducted a meta-analysis to examine responses of plant [N] and [P] to drought manipulation treatments and duration of drought stress. Drought stress showed negative effects on plant [N] (-3.73%) and plant [P] (-9.18%), and a positive effect on plant N:P (+ 6.98%). Drought stress had stronger negative effects on plant [N] and [P] in the short term (< 90 d) than in the long term (> 90 d). Drought treatments that included drying-rewetting cycles showed no effect on plant [N] and [P], while constant, prolonged, or intermittent drought stress had a negative effect on plant [P]. Our results suggest that negative effects on plant [N] and [P] are alleviated with extended duration of drought treatments and with drying-rewetting cycles. Availability of water, rather than of N and P, may be the main driver for reduced plant growth with increased long-term drought stress.© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
Effects of simulated nitrogen deposition on soil microbial biomass and community structure in a young plantation of Castanopsis hystrix
模拟氮沉降对红椎人工幼龄林土壤微生物生物量和微生物群落结构的影响
Effects of climate on soil phosphorus cycle and availability in natural terrestrial ecosystems
DOI:10.1111/gcb.14093
PMID:29450947
[本文引用: 1]
Climate is predicted to change over the 21st century. However, little is known about how climate change can affect soil phosphorus (P) cycle and availability in global terrestrial ecosystems, where P is a key limiting nutrient. With a global database of Hedley P fractions and key-associated physiochemical properties of 760 (seminatural) natural soils compiled from 96 published studies, this study evaluated how climate pattern affected soil P cycle and availability in global terrestrial ecosystems. Overall, soil available P, indexed by Hedley labile inorganic P fraction, significantly decreased with increasing mean annual temperature (MAT) and precipitation (MAP). Hypothesis-oriented path model analysis suggests that MAT negatively affected soil available P mainly by decreasing soil organic P and primary mineral P and increasing soil sand content. MAP negatively affected soil available P both directly and indirectly through decreasing soil primary mineral P; however, these negative effects were offset by the positive effects of MAP on soil organic P and fine soil particles, resulting in a relatively minor total MAP effect on soil available P. As aridity degree was mainly determined by MAP, aridity also had a relatively minor total effect on soil available P. These global patterns generally hold true irrespective of soil depth (≤10 cm or >10 cm) or site aridity index (≤1.0 or >1.0), and were also true for the low-sand (≤50%) soils. In contrast, available P of the high-sand (>50%) soils was positively affected by MAT and aridity and negatively affected by MAP. Our results suggest that temperature and precipitation have contrasting effects on soil P availability and can interact with soil particle size to control soil P availability.© 2018 John Wiley & Sons Ltd.
Soil organic phosphorus transformation during ecosystem development: a review
DOI:10.1007/s11104-017-3240-y URL [本文引用: 1]
Research summary on phosphate dissolution of phosphate solubilizing microorganisms
土壤解磷微生物及其解磷机制综述
Phosphate starvation root architecture and anthocyanin accumulation responses are modulated by the gibberellin-DELLA signaling pathway in Arabidopsis
Plant Physiology,
Effect of low-phosphorus stress on growth and physiological characteristics of new sainfoin lines at seedling stage
低磷胁迫对红豆草新品系幼苗生长及生理特性的影响
The impact of elevated carbon dioxide on the phosphorus nutrition of plants: a review
DOI:10.1093/aob/mcv088
PMID:26113632
[本文引用: 1]
Increasing attention is being focused on the influence of rapid increases in atmospheric CO2 concentration on nutrient cycling in ecosystems. An understanding of how elevated CO2 affects plant utilization and acquisition of phosphorus (P) will be critical for P management to maintain ecosystem sustainability in P-deficient regions.This review focuses on the impact of elevated CO2 on plant P demand, utilization in plants and P acquisition from soil. Several knowledge gaps on elevated CO2-P associations are highlighted.Significant increases in P demand by plants are likely to happen under elevated CO2 due to the stimulation of photosynthesis, and subsequent growth responses. Elevated CO2 alters P acquisition through changes in root morphology and increases in rooting depth. Moreover, the quantity and composition of root exudates are likely to change under elevated CO2, due to the changes in carbon fluxes along the glycolytic pathway and the tricarboxylic acid cycle. As a consequence, these root exudates may lead to P mobilization by the chelation of P from sparingly soluble P complexes, by the alteration of the biochemical environment and by changes to microbial activity in the rhizosphere. Future research on chemical, molecular, microbiological and physiological aspects is needed to improve understanding of how elevated CO2 might affect the use and acquisition of P by plants.© The Author 2015. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Phosphatase activity of extra-radical arbuscular mycorrhizal hyphae: a review
DOI:10.1023/A:1026582207192 URL [本文引用: 1]
Legacy effects of drought on plant-soil feedbacks and plant-plant interactions
DOI:10.1111/nph.14661
PMID:28621813
[本文引用: 1]
Interactions between aboveground and belowground biota have the potential to modify ecosystem responses to climate change, yet little is known about how drought influences plant-soil feedbacks with respect to microbial mediation of plant community dynamics. We tested the hypothesis that drought modifies plant-soil feedback with consequences for plant competition. We measured net pairwise plant-soil feedbacks for two grassland plant species grown in monoculture and competition in soils that had or had not been subjected to a previous drought; these were then exposed to a subsequent drought. To investigate the mechanisms involved, we assessed treatment responses of soil microbial communities and nutrient availability. We found that previous drought had a legacy effect on bacterial and fungal community composition that decreased plant growth in conspecific soils and had knock-on effects for plant competitive interactions. Moreover, plant and microbial responses to subsequent drought were dependent on a legacy effect of the previous drought on plant-soil interactions. We show that drought has lasting effects on belowground communities with consequences for plant-soil feedbacks and plant-plant interactions. This suggests that drought, which is predicted to increase in frequency with climate change, may change soil functioning and plant community composition via the modification of plant-soil feedbacks.© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.
Effect of sucrose on root morphology and substance accumulation at soybean seedling stage under low phosphorus stress
蔗糖对低磷胁迫条件下大豆苗期根系形态和物质积累的影响
How belowground interactions contribute to the coexistence of mycorrhizal and non-mycorrhizal species in severely phosphorus impoverished hyperdiverse ecosystems
DOI:10.1007/s11104-017-3427-2 URL [本文引用: 2]
Effect of phosphorus deficiency stress on root morphology and nutrient absorption of rice cultivars
With sandy culture, the difference of the root morphology, assimilation product distribution, mineral nutrient absorption and energy requirement of anion absorption were investigated among the low phosphorus tolerant rice (Oryza sativa L.) cultivars (Dld and Ltz3) and the low phosphorus sensitive cu
低磷胁迫对不同水稻品种根系形态和养分吸收的影响
以耐低磷品种大粒稻、莲塘早3号和低磷敏感品种沪占七、新三百粒为材料,采用砂培,测定常磷(10 mg/L)和低磷(0.5 mg/L)两个磷水平下,根系形态、同化物分配、矿质元素吸收、离子吸收耗能的品种差异。实验结果表明,低磷胁迫下,耐低磷品种的总根数、总根长、总根表面积、侧根长、侧根数及侧根密度均明显增加,而低磷敏感
Arbuscular mycorrhizal fungi promote the growth of Wedelia trilobata and the absorption of insoluble phosphorus
丛枝菌根真菌促进南美蟛蜞菊生长及对难溶磷的吸收
Capacity of some bacteria and fungi in dissolving phosphate rock
一些细菌和真菌的解磷能力及其机理初探
Regulation effect of soil P availability on mycorrhizal infection in relation to root architecture and P efficiency of Glycine max
磷有效性对大豆菌根侵染的调控及其与根构型、磷效率的关系
Cloning of an Erwinia herbicola gene necessary for gluconic acid production and enhanced mineral phosphate solubilization in Escherichia coli HB101: nucleotide sequence and probable involvement in biosynthesis of the coenzyme pyrroloquinoline quinone
Escherichia coli is capable of synthesizing the apo-glucose dehydrogenase enzyme (GDH) but not the cofactor pyrroloquinoline quinone (PQQ), which is essential for formation of the holoenzyme. Therefore, in the absence of exogenous PQQ, E. coli does not produce gluconic acid. Evidence is presented to show that the expression of an Erwinia herbicola gene in E. coli HB101(pMCG898) resulted in the production of gluconic acid, which, in turn, implied PQQ biosynthesis. Transposon mutagenesis showed that the essential gene or locus was within a 1.8-kb region of a 4.5-kb insert of the plasmid pMCG898. This 1.8-kb region contained only one apparent open reading frame. In this paper, we present the nucleotide sequence of this open reading frame, a 1,134-bp DNA fragment coding for a protein with an M(r) of 42,160. The deduced sequence of this protein had a high degree of homology with that of gene III (M(r), 43,600) of a PQQ synthase gene complex from Acinetobacter calcoaceticus previously identified by Goosen et al. (J. Bacteriol. 171:447-455, 1989). In minicell analysis, pMCG898 encoded a protein with an M(r) of 41,000. These data indicate that E. coli HB101(pMCG898) produced the GDH-PQQ holoenzyme, which, in turn, catalyzed the oxidation of glucose to gluconic acid in the periplasmic space. As a result of the gluconic acid production, E. coli HB101(pMCG898) showed an enhanced mineral phosphate-solubilizing phenotype due to acid dissolution of the hydroxyapatite substrate.
Impacts of arbuscular mycorrhizal fungi (AMF) on growth, N bio-fixation, and phosphorus uptake of legume crop
丛枝菌根真菌对豆科作物生长和生物固氮及磷素吸收的影响
DOI:10.13287/j.1001-9332.202105.022
[本文引用: 1]
本研究以大豆为材料,采用盆栽和田间试验,探讨丛枝菌根真菌(AMF)对豆科作物生长的影响。盆栽试验设置了接种(+AMF)和不接种(-AMF)丛枝菌根真菌处理,田间试验设置了AMF菌丝非限制与限制处理。盆栽试验结果表明: 接种AMF显著提高了大豆地上部生物量(16.5%)和大豆根瘤数(131.4%),地上部磷含量、磷吸收量、氮含量和氮吸收量也显著增加。田间试验中,AMF菌丝非限制处理下大豆的地上部生物量、根系生物量、根瘤数量分别比限制处理下显著提高了123.6%、61.5%和212.5%,地上部和根系磷吸收量、氮含量、氮吸收量均显著高于限制处理,大豆根际土壤速效氮和有效磷含量也均显著高于限制处理。本研究可为进一步认识豆科作物与AMF的共生关系及田间磷肥高效利用提供理论参考。
Nitrogen deposition promotes phosphorus uptake of plants in a semi-arid temperate grassland
DOI:10.1007/s11104-016-3022-y URL [本文引用: 1]
Isolation and identification of phosphorus solubilizing bacteria in mangrove soil and their characteristics
红树林土壤解磷菌的分离鉴定及解磷特性
Root phenes for enhanced soil exploration and phosphorus acquisition: tools for future crops
DOI:10.1104/pp.111.175414 PMID:21610180 [本文引用: 2]
Initial pH of medium affects organic acids production but do not affect phosphate solubilization
DOI:10.1590/S1517-838246246220131102
PMID:26273251
[本文引用: 1]
The pH of the culture medium directly influences the growth of microorganisms and the chemical processes that they perform. The aim of this study was to assess the influence of the initial pH of the culture medium on the production of 11 low-molecular-weight organic acids and on the solubilization of calcium phosphate by bacteria in growth medium (NBRIP). The following strains isolated from cowpea nodules were studied: UFLA03-08 (Rhizobium tropici), UFLA03-09 (Acinetobacter sp.), UFLA03-10 (Paenibacillus kribbensis), UFLA03-106 (Paenibacillus kribbensis) and UFLA03-116 (Paenibacillus sp.). The strains UFLA03-08, UFLA03-09, UFLA03-10 and UFLA03-106 solubilized Ca3(PO4)2 in liquid medium regardless of the initial pH, although without a significant difference between the treatments. The production of organic acids by these strains was assessed for all of the initial pH values investigated, and differences between the treatments were observed. Strains UFLA03-09 and UFLA03-10 produced the same acids at different initial pH values in the culture medium. There was no correlation between phosphorus solubilized from Ca3(PO4)2 in NBRIP liquid medium and the concentration of total organic acids at the different initial pH values. Therefore, the initial pH of the culture medium influences the production of organic acids by the strains UFLA03-08, UFLA03-09, UFLA03-10 and UFLA03-106 but it does not affect calcium phosphate solubilization.
The phosphorus composition of contrasting soils in pastoral, native and forest management in Otago, New Zealand: sequential extraction and 31P NMR
DOI:10.1016/j.geoderma.2005.01.020 URL [本文引用: 1]
Comparative aspects of cycling of organic C, N, S and P through soil organic matter
DOI:10.1016/0016-7061(81)90024-0 URL [本文引用: 1]
Elevated CO2 alters belowground exoenzyme activities in tussock tundra
DOI:10.1023/A:1004246720186 URL [本文引用: 1]
Physiological processes limiting plant growth in saline soils: some dogmas and hypotheses
A role for auxin redistribution in the responses of the root system architecture to phosphate starvation in Arabidopsis
DOI:10.1104/pp.105.060061
URL
[本文引用: 1]
The changes in root system architecture (RSA) triggered by phosphate (P) deprivation were studied in Arabidopsis (Arabidopsis thaliana) plants grown for 14 d on 1 mm or 3 μm P. Two different temporal phases were observed in the response of RSA to low P. First, lateral root (LR) development was promoted between days 7 and 11 after germination, but, after day 11, all root growth parameters were negatively affected, leading to a general reduction of primary root (PR) and LR lengths and of LR density. Low P availability had contrasting effects on various stages of LR development, with a marked inhibition of primordia initiation but a strong stimulation of activation of the initiated primordia. The involvement of auxin signaling in these morphological changes was investigated in wild-type plants treated with indole-3-acetic acid or 2,3,5-triiodobenzoic acid and in axr4-1, aux1-7, and eir1-1 mutants. Most effects of low P on RSA were dramatically modified in the mutants or hormone-treated wild-type plants. This shows that auxin plays a major role in the P starvation-induced changes of root development. From these data, we hypothesize that several aspects of the RSA response to low P are triggered by local modifications of auxin concentration. A model is proposed that postulates that P starvation results in (1) an overaccumulation of auxin in the apex of the PR and in young LRs, (2) an overaccumulation of auxin or a change in sensitivity to auxin in the lateral primordia, and (3) a decrease in auxin concentration in the lateral primordia initiation zone of the PR and in old laterals. Measurements of local changes in auxin concentrations induced by low P, either by direct quantification or by biosensor expression pattern (DR5::β-glucuronidase reporter gene), are in line with these hypotheses. Furthermore, the observation that low P availability mimicked the action of auxin in promoting LR development in the alf3 mutant confirmed that P starvation stimulates primordia emergence through increased accumulation of auxin or change in sensitivity to auxin in the primordia. Both the strong effect of 2,3,5-triiodobenzoic acid and the phenotype of the auxin-transport mutants (aux1, eir1) suggest that low P availability modifies local auxin concentrations within the root system through changes in auxin transport rather than auxin synthesis.
Impacts of altered precipitation regimes on soil communities and biogeochemistry in arid and semi-arid ecosystems
DOI:10.1111/gcb.12789
PMID:25363193
[本文引用: 1]
Altered precipitation patterns resulting from climate change will have particularly significant consequences in water-limited ecosystems, such as arid to semi-arid ecosystems, where discontinuous inputs of water control biological processes. Given that these ecosystems cover more than a third of Earth's terrestrial surface, it is important to understand how they respond to such alterations. Altered water availability may impact both aboveground and belowground communities and the interactions between these, with potential impacts on ecosystem functioning; however, most studies to date have focused exclusively on vegetation responses to altered precipitation regimes. To synthesize our understanding of potential climate change impacts on dryland ecosystems, we present here a review of current literature that reports the effects of precipitation events and altered precipitation regimes on belowground biota and biogeochemical cycling. Increased precipitation generally increases microbial biomass and fungal:bacterial ratio. Few studies report responses to reduced precipitation but the effects likely counter those of increased precipitation. Altered precipitation regimes have also been found to alter microbial community composition but broader generalizations are difficult to make. Changes in event size and frequency influences invertebrate activity and density with cascading impacts on the soil food web, which will likely impact carbon and nutrient pools. The long-term implications for biogeochemical cycling are inconclusive but several studies suggest that increased aridity may cause decoupling of carbon and nutrient cycling. We propose a new conceptual framework that incorporates hierarchical biotic responses to individual precipitation events more explicitly, including moderation of microbial activity and biomass by invertebrate grazing, and use this framework to make some predictions on impacts of altered precipitation regimes in terms of event size and frequency as well as mean annual precipitation. While our understanding of dryland ecosystems is improving, there is still a great need for longer term in situ manipulations of precipitation regime to test our model. © 2014 John Wiley & Sons Ltd.
Effects of simulated nitrogen deposition on growth and phosphorus efficiency of Pinus massoniana under low phosphorus stress
Atmospheric nitrogen (N) deposition dramatically raised in recent decades, resulting in increases of soil N availability and N/P ratio, which would impact plant growth and P efficiency under low P stress. Taking breeding population of Pinus massoniana as test materials, a pot experiment was conducted to simulate two P conditions, i. e., homogeneous low P availability vs. heterogeneous low P among soil layers, in combination with two N deposition levels on growth traits and P absorption and utilization efficiency of P. massoniana. Under the homogeneous low phosphorus condition, growth traits and P efficiency of P. massoniana were not significantly improved by simulated nitrogen deposition, but significant nitrogen x family interaction effect was detected, with the biomass of family 40x44 and 71x20 being increased, 36x29 and 73x23 being decreased. Under the heterogeneous low P condition, significant N effects on the seedling height, biomass and P absorption efficiency were observed, due to promoted root length and root distribution ratio of topsoil. In addition, the effects of simulated N deposition on growth and P efficiency of P. massoniana were relevant to the N/P ratio. Under the homogeneous low P condition, the N/P ratio of P. massoniana plant was 13.8, plants exhibited a low sensitivity to simulated N deposition, root secreted APase activity was increased but the plant growth was not promoted. In comparison, the plant N/P ratio was 9.7 under the heterogeneous low P condition, and the plant growth and P efficiency were significantly promoted, while no obvious change occurred in root secreted APase activity.
模拟氮沉降对低磷胁迫下马尾松生长和磷效率的影响
近年来大气N沉降日趋严重,导致森林土壤有效N含量增加,N/P发生改变,将会影响低P胁迫下林木的生长发育和P效率.本文以马尾松家系为研究对象,设置模拟N沉降与同质低P(介质表层与深层均缺P)、异质低P(介质表层P丰富、深层缺P)耦合的盆栽试验,研究N沉降对马尾松生长性状以及P吸收和利用效率的影响.结果表明: 同质低P下,模拟N沉降对马尾松生长性状和P效率影响较小,但存在显著的N×家系互作效应,家系40×44和71×20在N沉降后生物量增加,家系36×29和73×23生物量降低;异质低P下,模拟N沉降显著增加了马尾松苗高、生物量以及P吸收效率等,其原因是促进了根系生长和表层土壤中根系分布比例的增加.不同P环境下模拟N沉降对马尾松生长的影响,还与植株N/P有关.同质低P环境下,马尾松植株N/P为13.8,植株对N敏感性低,酸性磷酸酶活性增加,但未改善马尾松生长状况.在异质低P环境中,植株N/P为9.7,模拟N沉降显著增加了苗木生物量和P吸收效率,但未显著改变酸性磷酸酶活性.
Microbial products trigger amino acid exudation from plant roots
Plants naturally cycle amino acids across root cell plasma membranes, and any net efflux is termed exudation. The dominant ecological view is that microorganisms and roots passively compete for amino acids in the soil solution, yet the innate capacity of roots to recover amino acids present in ecologically relevant concentrations is unknown. We find that, in the absence of culturable microorganisms, the influx rates of 16 amino acids (each supplied at 2.5 microm) exceed efflux rates by 5% to 545% in roots of alfalfa (Medicago sativa), Medicago truncatula, maize (Zea mays), and wheat (Triticum aestivum). Several microbial products, which are produced by common soil microorganisms such as Pseudomonas bacteria and Fusarium fungi, significantly enhanced the net efflux (i.e. exudation) of amino acids from roots of these four plant species. In alfalfa, treating roots with 200 microm phenazine, 2,4-diacetylphloroglucinol, or zearalenone increased total net efflux of 16 amino acids 200% to 2,600% in 3 h. Data from (15)N tests suggest that 2,4-diacetylphloroglucinol blocks amino acid uptake, whereas zearalenone enhances efflux. Thus, amino acid exudation under normal conditions is a phenomenon that probably reflects both active manipulation and passive uptake by microorganisms, as well as diffusion and adsorption to soil, all of which help overcome the innate capacity of plant roots to reabsorb amino acids. The importance of identifying potential enhancers of root exudation lies in understanding that such compounds may represent regulatory linkages between the larger soil food web and the internal carbon metabolism of the plant.
Phosphorus nutrition of mycorrhizal trees
DOI:10.1093/treephys/tpq063
PMID:20631011
[本文引用: 1]
Globally, phosphorus (P) limits productivity of trees in many forests and plantations especially in highly weathered, acidic or calcareous profiles. Most trees form mycorrhizal associations which are prevalent in the organic and mineral soil horizons. This review critically examines mechanisms that enhance the acquisition of P by tree roots. Mycorrhizal roots have a greater capacity to take up phosphate (Pi) from the soil solution than non-mycorrhizal root tips. Factors that contribute to this include the extent of extraradical hyphal penetration of soil and the physiology and biochemistry of the fungal/soil and fungal/plant interfaces. Ectomycorrhizal (ECM) trees are likely to benefit from association with basidiomycetes that possess several high-affinity Pi transporters that are expressed in extraradical hyphae and whose expression is enhanced by P deficiency. To understand fully the role of these putative transporters in the symbiosis, data regarding their localization, Pi transport capacities and regulation are required. Some ECM fungi are able to effect release of Pi from insoluble mineral P through excretion of low-molecular-weight organic anions such as oxalate, but the relative contribution of insoluble P dissolution in situ remains to be quantified. How the production of oxalate is regulated by nitrogen remains a key question to be answered. Lastly, phosphatase release from mycorrhizas is likely to play a significant role in the acquisition of Pi from labile organic forms of P (Po). As labile forms of Po can constitute the major fraction of the total P in some tropical and temperate soils, a greater understanding of the forms of Po available to the phosphatases is warranted.
Biomass and productivity in Acacia crassicarpa plantation
厚荚相思人工林生物量和生产力的研究
Phosphate acquisition
Phosphorus is one of the major plant nutrients that is least available in the soil. Consequently, plants have developed numerous morphological, physiological, biochemical, and molecular adaptations to acquire phosphate (Pi). Enhanced ability to acquire Pi and altered gene expression are the hallmarks of plant adaptation to Pi deficiency. The intricate mechanisms involved in maintaining Pi homeostasis reflect the complexity of Pi acquisition and translocation in plants. Recent discoveries of multiple Pi transporters have opened up opportunities to study the molecular basis of Pi acquisition by plants. An increasing number of genes are now known to be activated under Pi starvation. Some of these genes may be involved in Pi acquisition, transfer, and signal transduction during Pi stress. This review provides an overview of plant adaptations leading to enhanced Pi acquisition, with special emphasis on recent developments in the molecular biology of Pi acquisition.
Costs of acquiring phosphorus by vascular land plants: patterns and implications for plant coexistence
DOI:10.1111/nph.14967
PMID:29292829
[本文引用: 1]
Content Summary 1420 I. Introduction 1421 II. Root adaptations that influence P acquisition 1422 III. Costs of P acquisition: general 1423 IV. Costs of P acquisition that are independent of soil P concentrations 1423 V. Costs of P acquisition that increase as soil P concentrations decline 1424 VI. Discussion and conclusions 1424 Acknowledgements 1425 References 1425 SUMMARY: We compare carbon (and hence energy) costs of the different modes of phosphorus (P) acquisition by vascular land plants. Phosphorus-acquisition modes are considered to be mechanisms of plants together with their root symbionts and structures such as cluster roots involved in mobilising or absorbing P. Phosphorus sources considered are soluble and insoluble inorganic and organic pools. Costs include operating the P-acquisition mechanisms, and resource requirements to construct and maintain them. For most modes, costs increase as the relevant soil P concentration declines. Costs can thus be divided into a component incurred irrespective of soil P concentration, and a component describing how quickly costs increase as the soil P concentration declines. Differences in sensitivity of costs to soil P concentration arise mainly from how economically mycorrhizal fungal hyphae or roots that explore the soil volume are constructed, and from costs of exudates that hydrolyse or mobilise insoluble P forms. In general, modes of acquisition requiring least carbon at high soil P concentrations experience a steeper increase in costs as soil P concentrations decline. The relationships between costs and concentrations suggest some reasons why different modes coexist, and why the mixture of acquisition modes differs between sites.© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.
Linkages of C:N:P stoichiometry and bacterial community in soil following afforestation of former farmland
DOI:10.1016/j.foreco.2016.06.004 URL [本文引用: 1]
Extracellular secretion of Aspergillus phytase from Arabidopsis roots enables plants to obtain phosphorus from phytate
DOI:10.1046/j.1365-313x.2001.00998.x URL [本文引用: 1]
Phosphorus cycling in deciduous forest soil differs between stands dominated by ecto- and arbuscular mycorrhizal trees
DOI:10.1111/nph.13720
PMID:26510093
[本文引用: 1]
Although much is known about how trees and their associated microbes influence nitrogen cycling in temperate forest soils, less is known about biotic controls over phosphorus (P) cycling. Given that mycorrhizal fungi are instrumental for P acquisition and that the two dominant associations - arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) fungi - possess different strategies for acquiring P, we hypothesized that P cycling would differ in stands dominated by trees associated with AM vs ECM fungi. We quantified soil solution P, microbial biomass P, and sequentially extracted inorganic and organic P pools from May to November in plots dominated by trees forming either AM or ECM associations in south-central Indiana, USA. Overall, fungal communities in AM and ECM plots were functionally different and soils exhibited fundamental differences in P cycling. Organic forms of P were more available in ECM plots than in AM plots. Yet inorganic P decreased and organic P accumulated over the growing season in both ECM and AM plots, resulting in increasingly P-limited microbial biomass. Collectively, our results suggest that P cycling in hardwood forests is strongly influenced by biotic processes in soil and that these are driven by plant-associated fungal communities. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
A conserved MYB transcription factor involved in phosphate starvation signaling both in vascular plants and in unicellular algae
DOI:10.1101/gad.204401
URL
[本文引用: 1]
Plants have evolved a number of adaptive responses to cope with growth in conditions of limited phosphate (Pi) supply involving biochemical, metabolic, and developmental changes. We prepared an EMS-mutagenized M2 population of an Arabidopsis thaliana transgenic line harboring a reporter gene specifically responsive to Pi starvation (AtIPS1∷GUS), and screened for mutants altered in Pi starvation regulation. One of the mutants,phr1 (phosphate starvation response 1), displayed reduced response of AtIPS1∷GUS to Pi starvation, and also had a broad range of Pi starvation responses impaired, including the responsiveness of various other Pi starvation-induced genes and metabolic responses, such as the increase in anthocyanin accumulation. PHR1 was positionally cloned and shown be related to the PHOSPHORUS STARVATION RESPONSE 1(PSR1) gene from Chlamydomonas reinhardtii. A GFP∷PHR1 protein fusion was localized in the nucleus independently of Pi status, as is the case for PSR1. PHR1 is expressed in Pi sufficient conditions and, in contrast to PSR1, is only weakly responsive to Pi starvation. PHR1, PSR1, and other members of the protein family share a MYB domain and a predicted coiled–coil (CC) domain, defining a subtype within the MYB superfamily, the MYB–CC family. Therefore, PHR1 was found to bind as a dimer to an imperfect palindromic sequence. PHR1-binding sequences are present in the promoter of Pi starvation-responsive structural genes, indicating that this protein acts downstream in the Pi starvation signaling pathway.
Warming and grazing increase mineralization of organic P in an alpine meadow ecosystem of Qinghai-Tibet Plateau, China
DOI:10.1007/s11104-012-1132-8 URL [本文引用: 2]
Effects of arbuscular-mycorrhizal glomus species on drought tolerance: physiological and nutritional plant responses
DOI:10.1128/aem.61.2.456-460.1995
PMID:16534929
[本文引用: 1]
The tolerance of lettuce plants (Lactuca sativa L. cv. Romana) to drought stress differed with the arbuscular-mycorrhizal fungal isolate with which the plants were associated. Seven fungal species belonging to the genus Glomus were studied for their ability to enhance the drought tolerance of lettuce plants. These fungi had different traits that affected the drought resistance of host plants. The ranking of arbuscular-mycorrhizal fungal effects on drought tolerance, based on the relative decreases in shoot dry weight, was as follows: Glomus deserticola > Glomus fasciculatum > Glomus mosseae > Glomus etunicatum > Glomus intraradices > Glomus caledonium > Glomus occultum. In this comparative study specific mycorrhizal fungi had consistent effects on plant growth, mineral uptake, the CO(inf2) exchange rate, water use efficiency, transpiration, stomatal conductance, photosynthetic phosphorus use efficiency, and proline accumulation under either well-watered or drought-stressed conditions. The ability of the isolates to maintain plant growth effectively under water stress conditions was related to higher transpiration rates, levels of leaf conductance, and proline, N, and P contents. Differences in proline accumulation in leaves among the fungal symbioses suggested that the fungi were able to induce different degrees of osmotic adjustment. The detrimental effects of drought were not related to decreases in photosynthesis or water use efficiency. Neither of these parameters was related to P nutrition. The differences in P and K acquisition, transpiration, and stomatal conductance were related to the mycorrhizal efficiencies of the different fungi. Our observations revealed the propensities of different Glomus species to assert their protective effects during plant water stress. The greater effectiveness of G. deserticola in improving water deficit tolerance was associated with the lowest level of growth reduction (9%) under stress conditions. The growth of plants colonized by G. occultum was reduced by 70% after a progressive drought stress period. In general, the different protective effects of the mycorrhizal isolates were not associated with colonizing ability. Nevertheless, G. deserticola was the most efficient fungus and exhibited the highest levels of mycorrhizal colonization, as well as the greatest stimulation of physiological parameters.
Isolation and evaluation of rock phosphate solubilizing fungi as potential biofertilizer
Biological nitrogen fixation in non-legume plants
DOI:10.1093/aob/mct048
PMID:23478942
[本文引用: 1]
Nitrogen is an essential nutrient in plant growth. The ability of a plant to supply all or part of its requirements from biological nitrogen fixation (BNF) thanks to interactions with endosymbiotic, associative and endophytic symbionts, confers a great competitive advantage over non-nitrogen-fixing plants.Because BNF in legumes is well documented, this review focuses on BNF in non-legume plants. Despite the phylogenic and ecological diversity among diazotrophic bacteria and their hosts, tightly regulated communication is always necessary between the microorganisms and the host plant to achieve a successful interaction. Ongoing research efforts to improve knowledge of the molecular mechanisms underlying these original relationships and some common strategies leading to a successful relationship between the nitrogen-fixing microorganisms and their hosts are presented.Understanding the molecular mechanism of BNF outside the legume-rhizobium symbiosis could have important agronomic implications and enable the use of N-fertilizers to be reduced or even avoided. Indeed, in the short term, improved understanding could lead to more sustainable exploitation of the biodiversity of nitrogen-fixing organisms and, in the longer term, to the transfer of endosymbiotic nitrogen-fixation capacities to major non-legume crops.
A phosphate starvation response-centered network regulates mycorrhizal symbiosis
DOI:10.1016/j.cell.2021.09.030
PMID:34644527
[本文引用: 1]
To secure phosphorus (P) from soil, most land plants use a direct phosphate uptake pathway via root hairs and epidermis and an indirect phosphate uptake pathway via mycorrhizal symbiosis. The interaction between these two pathways is unclear. Here, we mapped a network between transcription factors and mycorrhizal symbiosis-related genes using Y1H. Intriguingly, this gene regulatory network is governed by the conserved P-sensing pathway, centered on phosphate starvation response (PHR) transcription factors. PHRs are required for mycorrhizal symbiosis and regulate symbiosis-related genes via the P1BS motif. SPX-domain proteins suppress OsPHR2-mediated induction of symbiosis-related genes and inhibit mycorrhizal infection. In contrast, plants overexpressing OsPHR2 show improved mycorrhizal infection and are partially resistant to P-mediated inhibition of symbiosis. Functional analyses of network nodes revealed co-regulation of hormonal signaling and mycorrhizal symbiosis. This network deciphers extensive regulation of mycorrhizal symbiosis by endogenous and exogenous signals and highlights co-option of the P-sensing pathway for mycorrhizal symbiosis.Copyright © 2021 Elsevier Inc. All rights reserved.
Determinate root growth and meristem maintenance in angiosperms
DOI:10.1093/aob/mcm251
PMID:17954472
[本文引用: 1]
The difference between indeterminate and determinate growth in plants consists of the presence or absence of an active meristem in the fully developed organ. Determinate root growth implies that the root apical meristem (RAM) becomes exhausted. As a consequence, all cells in the root tip differentiate. This type of growth is widely found in roots of many angiosperm taxa and might have evolved as a developmental adaptation to water deficit (in desert Cactaceae), or low mineral content in the soil (proteoid roots in various taxa).This review considers the mechanisms of determinate root growth to better understand how the RAM is maintained, how it functions, and the cellular and genetic bases of these processes. The role of the quiescent centre in RAM maintenance and exhaustion will be analysed. During root ageing, the RAM becomes smaller and its organization changes; however, it remains unknown whether every root is truly determinate in the sense that its RAM becomes exhausted before senescence. We define two types of determinate growth: constitutive where determinacy is a natural part of root development; and non-constitutive where determinacy is induced usually by an environmental factor. Determinate root growth is proposed to include two phases: the indeterminate growth phase, when the RAM continuously produces new cells; and the termination growth phase, when cell production gradually decreases and eventually ceases. Finally, new concepts regarding stem cells and a stem cell niche are discussed to help comprehend how the meristem is maintained in a broad taxonomic context.
Phosphate transport in plants
DOI:10.1023/A:1022376332180 URL [本文引用: 1]
Functional diversity in arbuscular mycorrhizal (AM) symbioses: the contribution of the mycorrhizal P uptake pathway is not correlated with mycorrhizal responses in growth or total P uptake
DOI:10.1111/nph.2004.162.issue-2 URL [本文引用: 2]
Differential expression of the LePS2 phosphatase gene family in response to phosphate availability, pathogen infection and during development
DOI:10.1034/j.1399-3054.2003.00091.x URL [本文引用: 1]
Research progress of phosphate dissolving microorganisms and their promotion effect on plant growth
土壤溶磷微生物及其对植物促生作用研究进展
溶磷微生物可使土壤中难溶性或不溶性磷转化成易于被植物吸收利用的磷,从而提高土壤供磷水平,此过程是农田磷源可再生利用的有效途径之一。对溶磷微生物的种类、在土壤中的生态分布特征、溶磷机制、对植物的促生作用及其机制进行了综述,并根据目前溶磷微生物的利用现状,对今后该领域的研究方向进行了展望。
Interactive effects of phosphorus deficiency and exogenous auxin on root morphological and physiological traits in white lupin (Lupinus albus L.)
DOI:10.1007/s11427-013-4461-9 URL [本文引用: 1]
Screening of a highly efficient phosphate-solubilizing bacterium and validation of its phosphate-solubilizing effect
一株高效解磷菌的筛选及其解磷效果验证
DOI:10.13560/j.cnki.biotech.bull.1985.2019-0969
[本文引用: 1]
为开发新型、安全及高效的解磷菌肥,从农田中分离筛选出多株菌,通过解磷能力验证,最终选出解磷能力最强的X-P18菌株,经16S rDNA分析鉴定为贝莱斯芽孢杆菌。同时,通过优化X-P18菌株液体发酵培养条件并测定发酵液中小分子有机酸种类,初步探究菌株的解磷机理。将该解磷菌施用于黑叶葵扇白菜盆栽,并对小白菜的基本指标进行分析。结果表明,X-P18菌株在无机磷液体培养基中,溶磷量为495.4 mg/L,具有分泌乙酸及其他少量小分子有机酸的特性。培养基初始接种量为1%、pH8.0、碳源为甘露糖、氮源为硫酸铵、培养温度为30℃时,X-P18菌株的解磷能力最佳,溶磷量为582.4 mg/L,比优化前高出17.6%。在X-P18菌剂添加量为2×10<sup>9 </sup>CFU/盆时,对黑叶葵扇白菜的促生效果最明显,鲜重增加65.5%,叶片全磷含量增加46.9%。
Research advances in effect of phosphate-solubilizing microorganisms on soil phosphorus resource utilization
解磷微生物对土壤磷资源利用影响的研究进展
Characterization and sequence of PhoC, the principal phosphate-irrepressible acid phosphatase of Morganella morganii
DOI:10.1099/00221287-140-6-1341 URL [本文引用: 1]
Short on phosphate: plant surveillance and countermeasures
Metabolism depends on inorganic phosphate (P(i)) as reactant, allosteric effector and regulatory moiety in covalent protein modification. To cope with P(i) shortage (a common situation in many ecosystems), plants activate a set of adaptive responses to enhance P(i) recycling and acquisition by reprogramming metabolism and restructuring root system architecture. The physiology of P(i) starvation responses has become well understood, and so current research focuses on the initial molecular events that sense, transmit and integrate information about external and internal P(i) status. Recent studies have provided evidence for P(i) as a signaling molecule and initial insight into the coordination of P(i) deficiency responses at the cellular and molecular level.
Transcripts of MYB-like genes respond to phosphorous and nitrogen deprivation in Arabidopsis
DOI:10.1007/s00425-004-1305-7 URL [本文引用: 1]
Influence of arbuscular mycorrhizal mycelial exudates on soil bacterial growth and community structure
DOI:10.1111/j.1574-6941.2007.00337.x
PMID:17535297
[本文引用: 1]
Plant root systems colonized by arbuscular mycorrhizal (AM) fungi have previously been shown to influence soil bacterial populations; however, the direct influence of the AM extraradical mycelium itself on bacterial growth and community composition is not well understood. In this study, we investigated the effects of exudates produced by AM extraradical mycelia on the growth and development of an extracted soil bacterial community in vitro. The chemical composition of the mycelial exudates was analysed using proton nuclear magnetic resonance spectrometry. Following the addition of exudates to a bacterial community extracted from soil, bacterial growth and vitality were determined using a bacterial vitality stain and fluorescence microscopy. Changes in community composition were also analysed at various times over the course of 3 days by terminal restriction fragment length polymorphism analysis, in combination with cloning and sequencing of 16S rRNA genes. Mycelial exudates increased bacterial growth and vitality and changed bacterial community composition. Several Gammaproteobacteria, including a taxon within the Enterobacteriaceae, increased in frequency of occurrence in response to AM mycelial exudates. This study is the first attempt to identify carbohydrates from the extraradical mycelium of an AM fungus, and demonstrates the direct effects of mycelial exudates on a soil bacterial community.
Regulation of the pho regulon in Escherichia coli K-12//Schaechter M
Potential contribution of lysed bacterial cells to phosphorus solubilisation in two rewetted Australian pasture soils
DOI:10.1016/S0038-0717(02)00244-4 URL [本文引用: 1]
Warming and disturbance alter soil microbiome diversity and function in a northern forest ecotone
DOI:10.1093/femsec/fiaa108 [本文引用: 1]
Phosphorus acquisition and use: critical adaptations by plants for securing a nonrenewable resource
DOI:10.1046/j.1469-8137.2003.00695.x
PMID:33873400
[本文引用: 1]
Phosphorus (P) is limiting for crop yield on > 30% of the world's arable land and, by some estimates, world resources of inexpensive P may be depleted by 2050. Improvement of P acquisition and use by plants is critical for economic, humanitarian and environmental reasons. Plants have evolved a diverse array of strategies to obtain adequate P under limiting conditions, including modifications to root architecture, carbon metabolism and membrane structure, exudation of low molecular weight organic acids, protons and enzymes, and enhanced expression of the numerous genes involved in low-P adaptation. These adaptations may be less pronounced in mycorrhizal-associated plants. The formation of cluster roots under P-stress by the nonmycorrhizal species white lupin (Lupinus albus), and the accompanying biochemical changes exemplify many of the plant adaptations that enhance P acquisition and use. Physiological, biochemical, and molecular studies of white lupin and other species response to P-deficiency have identified targets that may be useful for plant improvement. Genomic approaches involving identification of expressed sequence tags (ESTs) found under low-P stress may also yield target sites for plant improvement. Interdisciplinary studies uniting plant breeding, biochemistry, soil science, and genetics under the large umbrella of genomics are prerequisite for rapid progress in improving nutrient acquisition and use in plants. Contents I. Introduction 424 II. The phosphorus conundrum 424 III. Adaptations to low P 424 IV. Uptake of P 424 V. P deficiency alters root development and function 426 VI. P deficiency modifies carbon metabolism 431 VII. Acid phosphatase 436 VIII. Genetic regulation of P responsive genes 437 IX. Improving P acquisition 439 X. Synopsis 440.
Endophyte-promoted phosphorus solubilization in Populus
DOI:10.3389/fpls.2020.567918 [本文引用: 1]
Responses of soil microbial community to continuous experimental nitrogen additions for 13 years in a nitrogen-rich tropical forest
DOI:10.1016/j.soilbio.2018.03.009 URL [本文引用: 1]
Warming reduces the production of a major annual forage crop on the Tibetan Plateau
DOI:10.1016/j.scitotenv.2021.149211 [本文引用: 1]
Effects of short-term and long-term warming on soil nutrients, microbial biomass and enzyme activities in an alpine meadow on the Qinghai-Tibet Plateau of China
DOI:10.1016/j.soilbio.2014.05.014 URL
Rapid induction of regulatory and transporter genes in response to phosphorus, potassium, and iron deficiencies in tomato roots: evidence for cross talk and root/rhizosphere-mediated signals
DOI:10.1104/pp.008854
URL
[本文引用: 4]
Mineral nutrient deficiencies constitute major limitations for plant growth on agricultural soils around the world. To identify genes that possibly play roles in plant mineral nutrition, we recently generated a high-density array consisting of 1,280 genes from tomato (Lycopersicon esculentum) roots for expression profiling in nitrogen (N) nutrition. In the current study, we used the same array to search for genes induced by phosphorus (P), potassium (K+), and iron (Fe) deficiencies. RNA gel-blot analysis was conducted to study the time-dependent kinetics for expression of these genes in response to withholding P, K, or Fe. Genes previously not associated with P, K, and Fe nutrition were identified, such as transcription factor, mitogen-activated protein (MAP) kinase, MAP kinase kinase, and 14-3-3 proteins. Many of these genes were induced within 1 h after withholding the specific nutrient from roots of intact plants; thus, RNA gel-blot analysis was repeated for specific genes (transcription factor and MAP kinase) in roots of decapitated plants to investigate the tissue-specific location of the signal triggering gene induction. Both genes were induced just as rapidly in decapitated plants, suggesting that the rapid response to the absence of P, K, or Fe in the root-bathing medium is triggered either by a root-localized signal or because of root sensing of the mineral environment surrounding the roots. We also show that expression of Pi, K, and Fe transporter genes were up-regulated by all three treatments, suggesting coordination and coregulation of the uptake of these three essential mineral nutrients.
Effects of climate warming on the key process and index of black soil carbon and nitrogen cycle during freezing period
DOI:10.1021/es702565h URL [本文引用: 1]
气候变暖对冻结期黑土碳氮循环关键过程及指标的影响
Rice SPX1 and SPX2 inhibit phosphate starvation responses through interacting with PHR2 in a phosphate-dependent manner
Transcriptomic analysis of metabolic changes by phosphorus stress in rice plant roots
Phosphate-solubilizing bacteria associated with proteoid roots of seedlings of waratah [Telopea speciosissima (Sm.) R. Br.]
DOI:10.1111/j.1469-8137.1994.tb02995.x
PMID:33874565
[本文引用: 1]
Bacteria isolated from proteoid roots and non-proteoid lateral roots of Telopea speciosissima (Sm.) R.Br. seedlings were able to acidify the medium and solubilize calcium phosphates when grown in culture in the presence of ammonium salts and an appropriate carbon source. In general this activity was not detected when NO was substituted for NH, and it is proposed that protons were secreted in exchange for ammonium ions. Cation exchange between these protons and calcium in the medium is a possible cause of the calcium phosphate solubilizing activity. The relevance of these data to phosphate-solubilizing activity in natural environments is discussed.
Nitrogen and phosphorus additions alter the abundance of phosphorus-solubilizing bacteria and phosphatase activity in grassland soils
DOI:10.3389/fenvs.2019.00185 [本文引用: 1]
Phosphate starvation triggers distinct alterations of genome expression in Arabidopsis roots and leaves
DOI:10.1104/pp.103.021022
URL
[本文引用: 1]
Arabidopsis genome expression pattern changes in response to phosphate (Pi) starvation were examined during a 3-d period after removal of Pi from the growth medium. Available Pi concentration was decreased after the first 24 h of Pi starvation in roots by about 22%, followed by a slow recovery during the 2nd and 3rd d after Pi starvation, but no significant change was observed in leaves within the 3 d of Pi starvation. Microarray analysis revealed that more than 1,800 of the 6,172 genes present in the array were regulated by 2-fold or more within 72 h from the onset of Pi starvation. Analysis of these Pi starvation-responsive genes shows that they belong to wide range of functional categories. Many genes for photosynthesis and nitrogen assimilation were down-regulated. A complex set of metabolic adaptations appears to occur during Pi starvation. More than 100 genes each for transcription factors and cell-signaling proteins were regulated in response to Pi starvation, implying major regulatory changes in cellular growth and development. A significant fraction of those regulatory genes exhibited distinct or even contrasting expression in leaves and roots in response to Pi starvation, supporting the idea that distinct Pi starvation response strategies are used for different plant organs in response to a shortage of Pi in the growth medium.
Status quo and prospects of phosphate-soluble microorganisms
解磷微生物研究现状及展望
Research progress on the root adaptation mechanism of plants under low phosphorus stress
植物耐低磷胁迫的根系适应性机制研究进展
Climate warming promotes deterministic assembly of arbuscular mycorrhizal fungal communities
DOI:10.1111/gcb.v28.3 URL [本文引用: 1]
Sequestration of atmospheric CO2 in boreal forest carbon pools in northeastern China: effects of nitrogen deposition
DOI:10.1016/j.agrformet.2017.09.015 URL [本文引用: 1]
Distribution and storage of soil organic carbon across the desert grasslands in the southeastern fringe of the Tengger Desert, China
DOI:10.17521/cjpe.2017.0068 URL [本文引用: 1]
腾格里沙漠东南缘荒漠草地不同群落类型土壤有机碳分布及储量特征
DOI:10.17521/cjpe.2017.0068
[本文引用: 1]
荒漠生态系统环境的复杂性及其高度的空间异质性是土壤碳储量估算结果不确定性的重要因素。通过调查取样和室内分析, 研究了腾格里沙漠东南缘10种主要荒漠草地群落各土壤层(0-5、5-10、10-20、20-30、30-50、50-70和70-100 cm土层)的土壤有机碳(SOC)含量、垂直分布特征和影响因素, 以及0-5、0-20、0-50和0-100 cm土层剖面的土壤有机碳密度(SOCD)。结果显示, 灌木群落是SOC含量的重要影响因素, 各土层SOC含量在10种群落间均存在显著差异。从土壤表层到深层, SOC含量主要表现为由高至低和先升高后降低两种特征。SOC含量与黏粉粒含量、全氮(N)、全磷(P)和电导率呈显著的正相关关系, 与砂粒含量呈显著的负相关关系。各群落之间, 0-5、0-20、0-50和0-100 cm土层剖面的SOCD均存在显著差异, 其平均值分别为0.118、0.478、1.159和1.936 kg·m <sup>-2</sup>。这一结果低于全球和全国草地SOCD的平均值, 利用全国数据或该研究的平均值可能会高估或低估区域土壤有机碳储量, 因此利用不同群落的SOCD能增加荒漠地区土壤有机碳储量估算的确定性。
Effects of nitrogen and phosphorus addition on soil microbial community and amino sugar in a temperate forest on Changbai Mountain, northeast China
DOI:10.13287/j.1001-9332.202006.007
[本文引用: 1]
We carried out an experiment including nitrogen addition (N, 50 kg N·hm<sup>-2</sup>·a<sup>-1</sup>), phosphorus addition (P, 25 kg P·hm<sup>-2</sup>·a<sup>-1</sup>) and both nitrogen and phosphorus addition (NP, 50 kg N·hm<sup>-2</sup>·a<sup>-1</sup>+25 kg P·hm<sup>-2</sup>·a<sup>-1</sup>) in a natural Korean pine broad-leaved mixed forest on Changbai Mountain to examine the effects of single and combined N and P additions on soil microbial community composition and amino sugar. The results showed that N and P addition significantly reduced total microbial biomass by 19.5% and 24.6% in the organic layer of soil, while P addition significantly reduced the biomass of bacteria and fungi by 23.8% and 19.3%, respectively. In the mineral layer, N, P and NP addition significantly increased total microbial biomass by 94.8%, 230.9%, and 115.0% respectively, while the biomass of bacteria and fungi were significantly increased under all the treatments. The fungi to bacteria ratio (F/B) was significantly increased in the organic layer by N addition, while was decreased in the mineral layer soil by NP addition. The Gram-positive bacteria to Gram-negative bacteria ratio showed positive response to N, P and NP addition. Soil amino sugars responded differently to different treatments. N, P and NP addition significantly decreased glucosamine content by 41.3%, 48.8% and 36.4% in the organic layer, while N and NP addition increased muramic acid content by 43.0% and 71.1%, respectively. The contents of glucosamine and muramic acid in the mineral layer did not change significantly in response to N addition but increased significantly in response to both P addition and NP addition. The glucosamine to muramic acid ratio in the organic layer significantly decreased under fertilization treatments, indicating that N and P addition increased the relative contribution of bacteria to soil organic carbon accumulation. The changes in soil amino sugar contents were closely related to the change in microbial community composition after N and P addition, both of which were affected by changes in soil chemical properties.
氮磷添加对长白山温带森林土壤微生物群落组成和氨基糖的影响
DOI:10.13287/j.1001-9332.202006.007
[本文引用: 1]
在长白山阔叶红松林中设置氮添加(N,50 kg N·hm<sup>-2</sup>·a<sup>-1</sup>)、磷添加(P,25 kg P·hm<sup>-2</sup>·a<sup>-1</sup>)和氮磷添加(NP,50 kg N·hm<sup>-2</sup>·a<sup>-1</sup>+25 kg P·hm<sup>-2</sup>·a<sup>-1</sup>)试验,分析氮磷添加对有机层和矿质层土壤微生物群落组成和氨基糖的影响。结果表明: 在有机层土壤中,N、P添加使总微生物生物量显著降低19.5%和24.6%,P添加还使细菌和真菌生物量分别显著降低23.8%和19.3%;在矿质层土壤中,N、P和NP添加使总微生物生物量显著增加94.8%、230.9%和115.0%,细菌和真菌生物量在施肥处理下显著增加。N添加下有机层土壤真菌与细菌生物量比值(F/B)显著增大,而NP添加使矿质层土壤F/B显著减小。革兰氏阳性菌与革兰氏阴性菌比对N、P和NP添加有显著正响应。土壤氨基糖对不同处理的响应不同。在有机层土壤中,N、P和NP添加使氨基葡萄糖含量分别减少41.3%、48.8%和36.4%,而N和NP添加分别使胞壁酸含量显著增加43.0%和71.1%;在矿质层土壤中,氨基葡萄糖和胞壁酸含量在N添加下无显著变化,而在P和NP添加下显著增加。在施肥处理下,有机层土壤中氨基葡萄糖与胞壁酸比值显著减小,表明N、P添加增加了细菌对土壤有机碳积累的相对贡献。N、P添加后土壤氨基糖含量的变化与微生物群落组成的变化密切相关,且二者均受到土壤化学性质变化的影响。
Evaluating P availability influenced by warming and N deposition in a subtropical forest soil: a bioassay mesocosm experiment
DOI:10.1007/s11104-019-04246-z [本文引用: 2]
Root exudates and their ecological consequences in forest ecosystems: problems and perspective
DOI:10.17521/cjpe.2018.0156 URL [本文引用: 1]
森林根系分泌物生态学研究: 问题与展望
DOI:10.17521/cjpe.2018.0156
[本文引用: 1]
植物根际过程与调控机理研究已成为当前土壤学最活跃、最敏感的研究领域, 而根系分泌物作为根系-土壤-微生物界面物质能量交换和信息传递的重要媒介物质, 是构成根际微生态系统活力与功能特征的内在驱动因素, 是根际概念与根际过程存在的重要前提和基础。然而, 由于传统的根际过程研究更强调以实际生产问题为导向, 加之农作物生长周期较短、操作便利等诸多因素, 以往对植物根系分泌物研究主要聚焦在农业生态系统, 而有关根系分泌物在森林生态系统中的重要作用与调控机理研究甚少, 认识相对零散和片段化。基于此, 该文结合作者实际研究工作中的主要成果和该领域国际前沿动态, 综述了森林根系分泌物的生态重要性, 重点论述了目前森林根系分泌物生态学研究中存在的主要问题与不足, 在此基础上展望了未来森林根系分泌物生态学研究中值得关注的重点方向和研究内容。
Combination of nitrogen and phosphorus fertilization enhance ecosystem carbon sequestration in a nitrogen-limited temperate plantation of northern China
DOI:10.1016/j.foreco.2015.01.004 URL [本文引用: 1]
Roles of soil organisms on the enhancement of plant availability of soil phosphorus
土壤生物在土壤磷有效化中的作用
Adaptable effects of phosphorus stress on different genotypes of faba-bean
不同基因型春蚕豆对磷胁迫的适应性反应
A study on active soil organic carbon and available nutrients for protected soil under long-term fertilization
长期定位施肥条件下土壤活性有机碳变化及其与土壤速效养分的相关性
Effects of AMF inoculation, short-term warming and increasing precipitation on nitrogen and phosphorus absorption of plant
AMF及短期增温增雨互作对植物吸收氮磷功能的影响
DOI:10.11733/j.issn.1007-0435.2020.04.021
[本文引用: 1]
为研究气候变化是否影响植物-真菌共生关系及植物的吸收功能,本试验以青藏草原优势植物垂穗披碱草(Elymus nutans)和草地早熟禾(Poa pratensis)为研究对象,选取管柄囊霉属(Funneliformis)丛枝菌根真菌(Arbuscular mycorrizal fungi,AMF)进行室内控制试验。结果表明:增温促进了AMF和植物的共生,接种AMF及短期增温增雨互作对植物吸收氮磷功能的影响并不一致。对垂穗披碱草而言,增温增雨对其生长的土壤pH、全氮和有效磷含量均无显著影响,增雨促进了植物磷吸收(PPPPP<0.05)。
Effects of nitrogen deposition and increased precipitation on soil phosphorus dynamics in a temperate forest
DOI:10.1016/j.geoderma.2020.114650
The arbuscular mycorrhizal fungus Rhizophagus irregularis MUCL 43194 induces the gene expression of citrate synthase in the tricarboxylic acid cycle of the phosphate-solubilizing bacterium Rahnella aquatilis HX2
DOI:10.1007/s00572-018-0871-7
PMID:30368606
[本文引用: 1]
An increasing number of studies have demonstrated that arbuscular mycorrhizal fungi can cooperate with other soil microorganisms, e.g., bacteria, which develop near or on the surface of the extraradical hyphae where they perform multiple functions. However, the mechanisms involved in this privileged relationship are still poorly known. In the present study, we investigated how the arbuscular mycorrhizal fungus Rhizophagus irregularis MUCL 43194 influences the three pace-making enzymes (i.e., citrate synthase, isocitrate dehydrogenase, and α-oxoglutarate dehydrogenase) of the tricarboxylic acid (TCA) cycle in the phosphate-solubilizing bacterium Rahnella aquatilis HX2. The study was conducted under strict in vitro culture conditions and analysis made at the transcriptional level. Results showed that R. irregularis induced the expression of the gene-encoding citrate synthase (gltA), the pace-making enzyme involved in the first step of the TCA cycle, in R. aquatilis at all time points of observation (i.e., 1, 6, 12, 24, 48, and 72 h). The expression of the gene-encoding isocitrate dehydrogenase (icd) significantly decreased at 6, 12, 24, 48, and 72 h and the expression of the gene-encoding α-oxoglutarate dehydrogenase E1 component (kgdhc) significantly increased at 1, 6, and 48 h. The above results suggested that R. irregularis may influence the level of adenosine triphosphate production in R. aquatilis and thus the metabolism of the bacterium by stimulating the expression of gltA involved in the TCA cycle. Our results suggest a fine-tuned dialog between R. irregularis MUCL 43194 and R. aquatilis HX2 and emphasize the complexity of the interactions that might take place at the hyphal surface of arbuscular mycorrhizal fungi hosting communities of microbes.
Effects of warming and nitrogen deposition on the coupling mechanism between soil nitrogen and phosphorus in Songnen Meadow Steppe, northeastern China
DOI:10.1016/j.soilbio.2013.05.015 URL [本文引用: 1]
Effects of nitrogen addition on litter production and leaf decomposition in Rhododendron simsii shrubland in the mid-subtropical of China
氮添加对中亚热带杜鹃灌丛凋落物生产和叶分解的影响
Comparison of the responses between maize and faba bean to low phosphorus stress in the field
田间玉米和蚕豆对低磷胁迫响应的差异比较
In situ root phenotypes of cotton seedlings under phosphorus stress revealed through RhizoPot
Effects of environmental stress on characteristics of low molecular weight organic acids secreted by macrophyte roots
DOI:10.18307/2020.0215 URL [本文引用: 2]
环境胁迫对水生植物根系分泌小分子量有机酸(LMWOAs)的影响特征
Effects of warming on soil microbial communities of a subtropical monsoon evergreen broad-leaved forest in southern China
增温对南亚热带季风常绿阔叶林土壤微生物群落的影响
DOI:10.16258/j.cnki.1674-5906.2019.05.004
[本文引用: 1]
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Mobilization of soil phosphate after 8 years of warming is linked to plant phosphorus-acquisition strategies in an alpine meadow on the Qinghai-Tibetan Plateau
DOI:10.1111/gcb.15914
PMID:34606141
[本文引用: 1]
Phosphorus (P) is essential for productivity of alpine grassland ecosystems, which are sensitive to global warming. We tested the hypotheses that (1) mobilized 'calcium-bound inorganic P' (Ca-P ) is a major source of plant-available P in alpine meadows with alkaline soils after long-term warming, (2) mobilization of Ca-P is linked to effective plant carboxylate-releasing P-acquisition strategies under warming, and (3) the mobilization is also related to plant nitrogen (N)-acquisition. We conducted an eight-year warming experiment in an alpine meadow (4,635 m above sea level) on the Qinghai-Tibetan Plateau. A significant increase in P concentration in both aboveground and belowground biomass indicates an increased mobilization and assimilation of P by plants under warming. We observed a significant decrease in Ca-P, no change in moderately-labile organic P, and an increase in highly-resistant organic P after warming. There was no increase in phosphatase activities. Our results indicate that Ca-P, rather than organic P was the major source of plant-available P for alpine meadows under warming. Higher leaf manganese concentrations of sedges and forbs after warming indicate that carboxylates released by these plants are a key mechanism of Ca-P mobilization. The insignificant increase in Rhizobiales after warming and the very small cover of legumes show a minor role of N-acquisition strategies in solubilizing phosphate. The insignificant change in relative abundance of mycorrhizal fungi and bacteria related to P cycling after warming shows a small contribution of microorganisms to Ca-P mobilization. The significant increase in leaf N and P concentrations and N:P ratio of grasses and no change in sedge leaf N:P ratio reflect distinct responses of plant nutrient status to warming due to differences in P-acquisition strategies. We highlight the important effects of belowground P-acquisition strategies, especially plant carboxylate-releasing P-acquisition strategies on responses of plants to global changes in alpine meadows.This article is protected by copyright. All rights reserved.
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