大气CO_2浓度升高对拟南芥根毛发育与养分吸收的影响及根系对养分的响应机理
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摘要
大气CO浓度持续升高给人类生活和农业生产带来的深刻影响已经引起全球的广泛关注。CO2作为植物光合作用的原料之一,其浓度升高被多数研究证明对植物的生长和养分代谢产生了深刻的影响。CO2浓度升高会促进植物碳水化合物的积累,从而促进植物的生长,而这种持续生长需要更多的养分和水分供应。根系作为植物吸收养分和水分的重要器官,其生长发育在植物响应CO2浓度升高过程中扮演着重要的角色。根毛是根部特化的一种表皮细胞,在吸收土壤水分及养分,尤其是在吸收诸如磷(P),铁(Fe)等难移动营养元素时发挥着重要作用。同时,根毛的生长又容易受到外界生物和非生物因素的影响,尤其是土壤养分的供应水平。本研究选用模式植物拟南芥(Arabidopsis thaliana (L.) Heynh)作为主要研究材料,采用水培试验和可控培养箱试验,通过根系扫描、荧光显微镜、透射电镜和激光共聚焦镜观察、生理学、药理学以及分子生物学等方法与手段,开展了CO2浓度升高对拟南芥根毛生长发育的影响及其机理、CO2浓度升高对不同供氮形态下低供磷拟南芥磷吸收利用的影响及其机理以及Mg对拟南芥根毛生长发育的影响及其机理等研究,主要研究结果如下：
1.CO2浓度升高对拟南芥根毛生长发育的影响及其机理
与生长在正常CO2浓度(350gL L-1)条件下的拟南芥根毛相比,生长在升高CO2浓度(800μL L-1)条件下的拟南芥根毛长度和密度显著增加,而且‘H’型细胞(生毛细胞)数目也显著增多。CO2浓度升高提高了植物根系中生长素水平。在升高CO2浓度条件下,加入外源生长素极性输入阻断剂1-NOA或者生长素极性输出阻断剂NPA抑制了CO2浓度升高诱导的根毛形成。反之,在正常CO2浓度条件下,加入外源生长素类似物NAA则显著促进了根毛的形成。而且,CO2浓度升高处理下,生长素响应不敏感突变体axrl-3和axr4-1以及生长素极性运输突变体auxl-7和pin1-1根毛的长度和密度与生长在正常CO2浓度条件下的根毛相比没有显著差异。CO2浓度升高和NAA处理显著提高了根毛细胞分化和伸长促进基因CPC, TRY和ROP2的表达,同时降低了根毛分化抑制基因WER, GL2, GL3和TTG1的表达,但是,NPA处理得到与之相反的结果。总之,本研究发现CO2浓度升高通过生长素信号途径影响其下游的根毛细胞特化和伸长基因,从而促进根毛的生长发育。
2.CO2浓度升高对不同供氮形态下低供磷拟南芥磷吸收利用的影响及其机理
磷营养是植物响应大气CO2浓度升高的一个关键因子,然而,目前关于C02浓度升高对不同供N形态下植物P吸收利用的影响还不清楚。本研究选取5周龄拟南芥,在不同供N形态(NO3-和NH4+,2mM)和低P(0.5μM)条件下,分别在正常C02浓度(350μL L-1)和升高C02浓度(800μL L-1)条件下培养1周。研究结果表明,在供硝时,生长在正常CO2浓度下的拟南芥新叶呈暗绿色,而在升高C02浓度下,叶片生长正常呈嫩绿色。不同的是,在供铵情况下,无论是生长在正常C02还是升高C02浓度下,拟南芥叶片均呈暗绿色。同时,C02浓度升高降低了供硝拟南芥叶片中叶绿素和花青素的含量,而对供铵拟南芥叶片中的叶绿素和花青素含量没有影响。进一步研究结果显示,CO2浓度升高均促进了两种供N条件下植物的生长,但是降低了供铵植物每单位根重P吸收量和地上部P含量。与此相反,供硝植物的每单位根重P吸收量和植物组织中P含量均随CO2浓度升高而增加。而当P供应充足(0.5mM),其他处理条件与低P处理相同的情况下,发现C02浓度升高同样促进了两种供N条件下植物的生长,但是却降低了其体内P含量和P吸收量,且对叶片叶绿素和花青素含量没有显著影响。同时,CO2浓度升高增加了低P条件下供硝植物根冠比、根系总面积、酸性磷酸酶活性以及P吸收、运输、循环和再分配关键基因的表达。此外,与正常CO2浓度条件下相比,生长在高浓度CO2条件下的供硝植物根系中NO含量明显升高,而供铵植物根系NO含量却显著降低。升高CO2浓度条件下,加入外源NO清除剂cPTIO则抑制了两种供N条件下拟南芥根系对P的吸收。综合以上结果,本研究得出升高CO2浓度和硝态氮互作有助于植物从低P环境中吸收和利用P,从而缓解植物的缺磷症状,而在这一过程中,NO起到了很重要的介导调控作用。
3.Mg对拟南芥根毛生长发育的影响及其机理
根毛是植物根系吸收养分和水分的一个重要器官,根毛的生长容易受到土壤环境,尤其是养分水平的影响。Mg是植物所需的中量元素之一,但是目前有关Mg对植物根毛生长发育的影响及参与其中的生理和分子机制还不清楚。
本研究利用模式植物拟南芥开展了Mg对其根毛生长的影响。选择拟南芥根毛生长较旺盛的时期(第5周)将其在7个不同供Mg水平(MgSO4,浓度从0.5至10000μM)下培养7d,观察其根毛生长变化。取最低Mg浓度(0.5μM M1),和最高Mg浓度(10000M7)处理下分别生长4d和7d的根系以及在正常Mg浓度(500μM M3)生长4d根系进行RNA全转录组分析,并将RNA转录组分析的数据已经上传至NCBI, GEO编号为GSE42899。
生长在不同Mg浓度下的植物根毛在前4天并没有显示出生长差异,而从第5天开始根毛生长显示出处理间差异。处理7d后,统计检验结果显示供Mg水平越低,拟南芥根毛生长越旺盛,随着Mg浓度的提高,根毛逐渐减少。生长在低Mg下的拟南芥根毛长度和密度均显著增加,皮层细胞体积变小,‘H，型细胞(生毛细胞)数目增多且分化为成熟根毛的比例提高。而生长在高Mg(5000μM)下的拟南芥根毛稀疏且大部分根毛突体未伸长便停止生长。当供Mg浓度达到10000rtM时,根毛的生长完全被抑制,且皮层细胞体积变大,‘H，型细胞数目显著减少,部分根毛细胞壁瓦解,细胞质外流。拟南芥地上部和根系中总Ca浓度和根尖ROS水平随Mg浓度的增加而降低,生长在低Mg和正常Mg浓度下的拟南芥根毛突体和根毛细胞尖端均形成胞质Ca2+(cCa2+)梯度和ROS富集,而且。与此相反,生长在高浓度下的拟南芥根毛细胞起始和伸长部位未观察到cCa2+梯度和ROS富集。在低Mg处理下,加入外源BAPTA(Ca2+整合剂)或DPI(NADPH氧化酶抑制剂)会抑制低Mg对根毛生长的促进效应,同时根毛细胞尖端cCa2+梯度消失。反之,在高Mg浓度条件下,加入外源CaCl2或者ROS产生剂PMS时,被高Mg抑制的根毛恢复生长,虽然恢复程度随着Mg浓度的升高略有降低,同时根毛细胞伸长部位形成(Ca2+)c梯度和ROS。此外,不同供Mg处理对根毛缺陷突变体rhd2-1的根毛生长没有显著影响。
RNA转录组测序数据差异表达分析结果显示共有557个基因在不同处理条件下存在差异表达(FDR<0.05),进一步对不同Mg处理条件之间差异表达基因的GO富集结果显示不同Mg处理对拟南芥根系基因变化涉及的生物功能主要包括‘胁迫响应’、‘氧化还原反应’、‘离子转运’和‘细胞壁组织’。其中“胁迫响应”出现在所有的处理条件下,而“离子转运”主要发生在L7(0.5μM Mg处理7d)处理下,而“氧化还原反应”则共同出现在L7和H7(10000μMMg处理7d)。此外,决定根毛形态和伸长的‘H’基因均在L7下高表达,H7下低表达。值得一提的是,“细胞壁组织”相关基因特异性在H7处理下富集,而且这些基因的表达均被H7严重抑制,表明高Mg对拟南芥根毛生长的抑制与细胞壁的合成和定位缺陷有关。
综合以上,本研究得出Mg浓度会显著影响拟南芥根毛的生长发育,表现为Mg缺乏促进根毛的生长,而Mg过量抑制根毛的生长。低Mg提高了根系ROS和胞质Ca2+浓度并促进其在根毛尖富集,这些信号促进了根毛细胞分化和伸长基因,尤其是根毛伸长基因的表达从而促进根毛的生长。而长时期高Mg处理降低了根系的ROS和胞质Ca2+浓度,进而抑制根毛发育基因,特别是细胞壁合成和定位的关键基因,导致根毛形成受阻。而这一过程与植物细胞内Ca2+和ROS信号调控作用密切相关。
1. Development of root hairs in Arabidopsis thaliana (L.) Heynh. under elevated CO2
Root hairs may play a critical role in nutrient acquisition of plants grown under elevated CO2. This study investigated how elevated CO2enhanced the development of root hairs in Arabidopsis thaliana (L.) Heynh. The plants under elevated CO2(800μL L-1) had denser and longer root hairs, and more H-positioned cells in root epidermis than those under ambient CO2(350μL L-1). The elevated CO2increased auxin production in roots. Under elevated CO2, application of either1-naphthoxyacetic acid (1-NOA) or N-1-naphthylphthalamic acid (NPA) blocked the enhanced development of root hairs. The opposite was true when the plants under ambient CO2were treated with1-naphthylacetic acid (NAA), an auxin analogue. Furthermore, the elevated CO2did not enhance the development of root hairs in auxin-response mutants, axrl-3, and auxin-transporter mutants, axr4-1, auxl-7and pinl-1. Both elevated CO2and NAA application increased expressions of caprice, triptychon and rho-related protein from plants2, and decreased expressions of werewolf, GLABRA2, GLABRA3and the transparent testa glabra1, genes related to root-hair development, while1-NOA and NPA application had an opposite effect. Our study suggests that elevated CO2enhanced the development of root hairs in Arabidopsis via the well-characterized auxin signalling and transport that modulate the initiation of root hairs and the expression of its specific genes.
2. Effect of elevated CO2on phosphorus nutrition of phosphate-deficient Arabidopsis thaliana (L.) Heynh. under different nitrogen forms
Phosphorus (P) nutrition is always a key issue regarding plants responses to elevated CO2. Yet it is unclear of how elevated CO2affect P uptake under different nitrogen (N) forms. This study investigated the influence of elevated CO2(800μL L-1) on P uptake and utilization by Arabidopsis grown in pH-buffered phosphate (Pi)-deficient (0.5μM) hydroponic culture supplying with2mM nitrate (NO3-) or ammonium (NH4+). After7-d treatment, elevated CO2enhanced biomass production of both NO3-and NH4+-fed plants but decreased Pi amount absorbed by per weight of roots and Pi concentration in shoots of plants supplied with NH4+. In comparison, elevated CO2increased amount of Pi absorbed by per weight of roots as well as Pi concentration in plants and alleviated P-deficiency-induced symptoms of plants supplied with NO3-. Elevated CO2also increased root/shoot ratio, total root surface area and acid phosphatase activity, and enhanced expressions of genes or transcriptional factors involving in P uptake, allocation and remobilization in Pi-deficient plants. Furthermore, elevated CO2increased the NO level in roots of NO3--fed plants but decreased it in NH4+-fed plants. NO scavenger cPTIO inhibited plant P acquisition by roots under elevated CO2. Considering all of these findings, this study concludes that a combination of elevated CO2and NO3-nutrition can induce a set of plant adaptive strategies to improve P status from Pi-deficient soluble sources and NO may be a signaling molecule that controls these processes.
3. Regulation of root hair development in Arabidopsis thaliana (L.) Heynh. by magnesium
Root hairs are frequently reported to be plastic in response to nutrient supply in most plants, but relatively little is known about their development along with magnesium (Mg) availability, and evidence is also scarce about the signals involved in this process. Here, we characterized the response of root hair development to Mg availability in Arabidopsis thaliana grown in hydroponic culture with varied Mg concentrations, ranging from0.5to10000μM. Both density and length of root hairs decreased logarithmically in response to increasing Mg concentrations within that range, which correlated with the initiation of new trichoblast files and likelihood of trichoblasts to form hairs. Low-Mg availability resulted in greater concentrations of reactive oxygen species (ROS) and Ca2+in the roots and displayed a stronger tip-focused gradient of ROS and cytosolic Ca2+during initiation and elongation of root hairs, and this gradient could be eliminated by DPI or BAPTA drugs. Application of either BAPTA or DPI to low Mg treatment blocked the enhanced development of root hairs. The opposite was true when the plants under high Mg were supplied with Ca2+or PMS. Furthermore, root hair development of atrbohC/root hair defective2-1(rhd2-1) mutant was not affected by Mg availability. Whole-genome transcriptome data indentified a total of557differentially expressed genes, and the maximum differential expressed genes involved in'stress response','oxidation reduction','ion transport' and 'cell wall organization'. A greater fraction of genes expressed at later developmental stages as well as hair cell genes involving cell wall organization were up-regulated by L7(0.5μM Mg for7d) and down-regulated by H7(10000μM Mg for7d). No change in expression of genes encoding Ca2+and Mg uptake. Taking all of these findings together, a distinct and previously poorly characterized response of Arabidopsis root hair development to Mg availability is presented where ROS and Ca2+were the signaling molecules that control this process.
1.CO2浓度升高对拟南芥根毛生长发育的影响及其机理
与生长在正常CO2浓度(350gL L-1)条件下的拟南芥根毛相比,生长在升高CO2浓度(800μL L-1)条件下的拟南芥根毛长度和密度显著增加,而且‘H’型细胞(生毛细胞)数目也显著增多。CO2浓度升高提高了植物根系中生长素水平。在升高CO2浓度条件下,加入外源生长素极性输入阻断剂1-NOA或者生长素极性输出阻断剂NPA抑制了CO2浓度升高诱导的根毛形成。反之,在正常CO2浓度条件下,加入外源生长素类似物NAA则显著促进了根毛的形成。而且,CO2浓度升高处理下,生长素响应不敏感突变体axrl-3和axr4-1以及生长素极性运输突变体auxl-7和pin1-1根毛的长度和密度与生长在正常CO2浓度条件下的根毛相比没有显著差异。CO2浓度升高和NAA处理显著提高了根毛细胞分化和伸长促进基因CPC, TRY和ROP2的表达,同时降低了根毛分化抑制基因WER, GL2, GL3和TTG1的表达,但是,NPA处理得到与之相反的结果。总之,本研究发现CO2浓度升高通过生长素信号途径影响其下游的根毛细胞特化和伸长基因,从而促进根毛的生长发育。
2.CO2浓度升高对不同供氮形态下低供磷拟南芥磷吸收利用的影响及其机理
磷营养是植物响应大气CO2浓度升高的一个关键因子,然而,目前关于C02浓度升高对不同供N形态下植物P吸收利用的影响还不清楚。本研究选取5周龄拟南芥,在不同供N形态(NO3-和NH4+,2mM)和低P(0.5μM)条件下,分别在正常C02浓度(350μL L-1)和升高C02浓度(800μL L-1)条件下培养1周。研究结果表明,在供硝时,生长在正常CO2浓度下的拟南芥新叶呈暗绿色,而在升高C02浓度下,叶片生长正常呈嫩绿色。不同的是,在供铵情况下,无论是生长在正常C02还是升高C02浓度下,拟南芥叶片均呈暗绿色。同时,C02浓度升高降低了供硝拟南芥叶片中叶绿素和花青素的含量,而对供铵拟南芥叶片中的叶绿素和花青素含量没有影响。进一步研究结果显示,CO2浓度升高均促进了两种供N条件下植物的生长,但是降低了供铵植物每单位根重P吸收量和地上部P含量。与此相反,供硝植物的每单位根重P吸收量和植物组织中P含量均随CO2浓度升高而增加。而当P供应充足(0.5mM),其他处理条件与低P处理相同的情况下,发现C02浓度升高同样促进了两种供N条件下植物的生长,但是却降低了其体内P含量和P吸收量,且对叶片叶绿素和花青素含量没有显著影响。同时,CO2浓度升高增加了低P条件下供硝植物根冠比、根系总面积、酸性磷酸酶活性以及P吸收、运输、循环和再分配关键基因的表达。此外,与正常CO2浓度条件下相比,生长在高浓度CO2条件下的供硝植物根系中NO含量明显升高,而供铵植物根系NO含量却显著降低。升高CO2浓度条件下,加入外源NO清除剂cPTIO则抑制了两种供N条件下拟南芥根系对P的吸收。综合以上结果,本研究得出升高CO2浓度和硝态氮互作有助于植物从低P环境中吸收和利用P,从而缓解植物的缺磷症状,而在这一过程中,NO起到了很重要的介导调控作用。
3.Mg对拟南芥根毛生长发育的影响及其机理
根毛是植物根系吸收养分和水分的一个重要器官,根毛的生长容易受到土壤环境,尤其是养分水平的影响。Mg是植物所需的中量元素之一,但是目前有关Mg对植物根毛生长发育的影响及参与其中的生理和分子机制还不清楚。
本研究利用模式植物拟南芥开展了Mg对其根毛生长的影响。选择拟南芥根毛生长较旺盛的时期(第5周)将其在7个不同供Mg水平(MgSO4,浓度从0.5至10000μM)下培养7d,观察其根毛生长变化。取最低Mg浓度(0.5μM M1),和最高Mg浓度(10000M7)处理下分别生长4d和7d的根系以及在正常Mg浓度(500μM M3)生长4d根系进行RNA全转录组分析,并将RNA转录组分析的数据已经上传至NCBI, GEO编号为GSE42899。
生长在不同Mg浓度下的植物根毛在前4天并没有显示出生长差异,而从第5天开始根毛生长显示出处理间差异。处理7d后,统计检验结果显示供Mg水平越低,拟南芥根毛生长越旺盛,随着Mg浓度的提高,根毛逐渐减少。生长在低Mg下的拟南芥根毛长度和密度均显著增加,皮层细胞体积变小,‘H，型细胞(生毛细胞)数目增多且分化为成熟根毛的比例提高。而生长在高Mg(5000μM)下的拟南芥根毛稀疏且大部分根毛突体未伸长便停止生长。当供Mg浓度达到10000rtM时,根毛的生长完全被抑制,且皮层细胞体积变大,‘H，型细胞数目显著减少,部分根毛细胞壁瓦解,细胞质外流。拟南芥地上部和根系中总Ca浓度和根尖ROS水平随Mg浓度的增加而降低,生长在低Mg和正常Mg浓度下的拟南芥根毛突体和根毛细胞尖端均形成胞质Ca2+(cCa2+)梯度和ROS富集,而且。与此相反,生长在高浓度下的拟南芥根毛细胞起始和伸长部位未观察到cCa2+梯度和ROS富集。在低Mg处理下,加入外源BAPTA(Ca2+整合剂)或DPI(NADPH氧化酶抑制剂)会抑制低Mg对根毛生长的促进效应,同时根毛细胞尖端cCa2+梯度消失。反之,在高Mg浓度条件下,加入外源CaCl2或者ROS产生剂PMS时,被高Mg抑制的根毛恢复生长,虽然恢复程度随着Mg浓度的升高略有降低,同时根毛细胞伸长部位形成(Ca2+)c梯度和ROS。此外,不同供Mg处理对根毛缺陷突变体rhd2-1的根毛生长没有显著影响。
RNA转录组测序数据差异表达分析结果显示共有557个基因在不同处理条件下存在差异表达(FDR<0.05),进一步对不同Mg处理条件之间差异表达基因的GO富集结果显示不同Mg处理对拟南芥根系基因变化涉及的生物功能主要包括‘胁迫响应’、‘氧化还原反应’、‘离子转运’和‘细胞壁组织’。其中“胁迫响应”出现在所有的处理条件下,而“离子转运”主要发生在L7(0.5μM Mg处理7d)处理下,而“氧化还原反应”则共同出现在L7和H7(10000μMMg处理7d)。此外,决定根毛形态和伸长的‘H’基因均在L7下高表达,H7下低表达。值得一提的是,“细胞壁组织”相关基因特异性在H7处理下富集,而且这些基因的表达均被H7严重抑制,表明高Mg对拟南芥根毛生长的抑制与细胞壁的合成和定位缺陷有关。
综合以上,本研究得出Mg浓度会显著影响拟南芥根毛的生长发育,表现为Mg缺乏促进根毛的生长,而Mg过量抑制根毛的生长。低Mg提高了根系ROS和胞质Ca2+浓度并促进其在根毛尖富集,这些信号促进了根毛细胞分化和伸长基因,尤其是根毛伸长基因的表达从而促进根毛的生长。而长时期高Mg处理降低了根系的ROS和胞质Ca2+浓度,进而抑制根毛发育基因,特别是细胞壁合成和定位的关键基因,导致根毛形成受阻。而这一过程与植物细胞内Ca2+和ROS信号调控作用密切相关。
1. Development of root hairs in Arabidopsis thaliana (L.) Heynh. under elevated CO2
Root hairs may play a critical role in nutrient acquisition of plants grown under elevated CO2. This study investigated how elevated CO2enhanced the development of root hairs in Arabidopsis thaliana (L.) Heynh. The plants under elevated CO2(800μL L-1) had denser and longer root hairs, and more H-positioned cells in root epidermis than those under ambient CO2(350μL L-1). The elevated CO2increased auxin production in roots. Under elevated CO2, application of either1-naphthoxyacetic acid (1-NOA) or N-1-naphthylphthalamic acid (NPA) blocked the enhanced development of root hairs. The opposite was true when the plants under ambient CO2were treated with1-naphthylacetic acid (NAA), an auxin analogue. Furthermore, the elevated CO2did not enhance the development of root hairs in auxin-response mutants, axrl-3, and auxin-transporter mutants, axr4-1, auxl-7and pinl-1. Both elevated CO2and NAA application increased expressions of caprice, triptychon and rho-related protein from plants2, and decreased expressions of werewolf, GLABRA2, GLABRA3and the transparent testa glabra1, genes related to root-hair development, while1-NOA and NPA application had an opposite effect. Our study suggests that elevated CO2enhanced the development of root hairs in Arabidopsis via the well-characterized auxin signalling and transport that modulate the initiation of root hairs and the expression of its specific genes.
2. Effect of elevated CO2on phosphorus nutrition of phosphate-deficient Arabidopsis thaliana (L.) Heynh. under different nitrogen forms
Phosphorus (P) nutrition is always a key issue regarding plants responses to elevated CO2. Yet it is unclear of how elevated CO2affect P uptake under different nitrogen (N) forms. This study investigated the influence of elevated CO2(800μL L-1) on P uptake and utilization by Arabidopsis grown in pH-buffered phosphate (Pi)-deficient (0.5μM) hydroponic culture supplying with2mM nitrate (NO3-) or ammonium (NH4+). After7-d treatment, elevated CO2enhanced biomass production of both NO3-and NH4+-fed plants but decreased Pi amount absorbed by per weight of roots and Pi concentration in shoots of plants supplied with NH4+. In comparison, elevated CO2increased amount of Pi absorbed by per weight of roots as well as Pi concentration in plants and alleviated P-deficiency-induced symptoms of plants supplied with NO3-. Elevated CO2also increased root/shoot ratio, total root surface area and acid phosphatase activity, and enhanced expressions of genes or transcriptional factors involving in P uptake, allocation and remobilization in Pi-deficient plants. Furthermore, elevated CO2increased the NO level in roots of NO3--fed plants but decreased it in NH4+-fed plants. NO scavenger cPTIO inhibited plant P acquisition by roots under elevated CO2. Considering all of these findings, this study concludes that a combination of elevated CO2and NO3-nutrition can induce a set of plant adaptive strategies to improve P status from Pi-deficient soluble sources and NO may be a signaling molecule that controls these processes.
3. Regulation of root hair development in Arabidopsis thaliana (L.) Heynh. by magnesium
Root hairs are frequently reported to be plastic in response to nutrient supply in most plants, but relatively little is known about their development along with magnesium (Mg) availability, and evidence is also scarce about the signals involved in this process. Here, we characterized the response of root hair development to Mg availability in Arabidopsis thaliana grown in hydroponic culture with varied Mg concentrations, ranging from0.5to10000μM. Both density and length of root hairs decreased logarithmically in response to increasing Mg concentrations within that range, which correlated with the initiation of new trichoblast files and likelihood of trichoblasts to form hairs. Low-Mg availability resulted in greater concentrations of reactive oxygen species (ROS) and Ca2+in the roots and displayed a stronger tip-focused gradient of ROS and cytosolic Ca2+during initiation and elongation of root hairs, and this gradient could be eliminated by DPI or BAPTA drugs. Application of either BAPTA or DPI to low Mg treatment blocked the enhanced development of root hairs. The opposite was true when the plants under high Mg were supplied with Ca2+or PMS. Furthermore, root hair development of atrbohC/root hair defective2-1(rhd2-1) mutant was not affected by Mg availability. Whole-genome transcriptome data indentified a total of557differentially expressed genes, and the maximum differential expressed genes involved in'stress response','oxidation reduction','ion transport' and 'cell wall organization'. A greater fraction of genes expressed at later developmental stages as well as hair cell genes involving cell wall organization were up-regulated by L7(0.5μM Mg for7d) and down-regulated by H7(10000μM Mg for7d). No change in expression of genes encoding Ca2+and Mg uptake. Taking all of these findings together, a distinct and previously poorly characterized response of Arabidopsis root hair development to Mg availability is presented where ROS and Ca2+were the signaling molecules that control this process.
引文
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