,, 赵继浩, 李金融, 钱必长, 刘兆新, 高芳, 杨东清,, 李向东,山东农业大学农学院/作物生物学国家重点实验室,山东泰安 271018Effects of Exogenous 6-BA on Root Growth and Pod Yield of Flooded Peanut at Different Growth Stages
LI Ying,, ZHAO JiHao, LI JinRong, QIAN BiChang, LIU ZhaoXin, GAO Fang, YANG DongQing,, LI XiangDong,College of Agriculture, Shandong Agricultural University/State Key Laboratory of Crop Biology, Taian 271018, Shandong通讯作者:
责任编辑: 杨鑫浩
收稿日期:2019-12-30接受日期:2020-02-26网络出版日期:2020-09-16
| 基金资助: |
Received:2019-12-30Accepted:2020-02-26Online:2020-09-16
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李颖, 赵继浩, 李金融, 钱必长, 刘兆新, 高芳, 杨东清, 李向东. 外源6-BA对不同生育时期淹水花生根系生长和荚果产量的影响[J]. 中国农业科学, 2020, 53(18): 3665-3678 doi:10.3864/j.issn.0578-1752.2020.18.004
LI Ying, ZHAO JiHao, LI JinRong, QIAN BiChang, LIU ZhaoXin, GAO Fang, YANG DongQing, LI XiangDong.
0 引言
【研究意义】黄淮海地区是我国重要的花生生产区,该区域属温带大陆性季风气候,夏季降水量多且集中,易导致农田土壤积水[1,2],对花生生产造成很大的影响。因此,研究不同生育时期淹水对花生生长发育的影响和栽培调控的作用机制,对花生品种选育、栽培管理等具有重要理论参考意义。【前人研究进展】根系是植物吸收养分的主要器官,也是植物激素等物质合成的器官[3],其生长分布与功能对地上部植株发育和产量形成有重要调节作用[4,5]。淹水条件下,根系缺氧,其细胞呼吸代谢方式发生转变,表现为有氧呼吸锐降,无氧呼吸增强[6,7]。前人研究发现植物淹水后根系乳酸脱氢酶（LDH）、乙醇脱氢酶（ADH）活性增加,苹果酸脱氢酶（MDH）活性降低[8,9]。植物内源激素脱落酸、细胞分裂素等作为植物体内的痕量信号分子,对于调节植物的生长发育过程和环境的应答具有十分重要的意义[10]。淹水环境下叶片ABA含量会明显升高,致使气孔开度减小或关闭[11],影响作物的光合作用,进而影响干物质积累,最终降低产量[12]。玉米素核苷（ZR）在逆境中起到从地下到地上的信息介质作用[13]。通过对河竹的研究发现,淹水显著降低了根系ZR含量,提高了ABA含量[14]。6-苄基腺嘌呤（6-BA）是一种人工合成的细胞分裂素。REN等[15]研究发现外源6-BA增加了淹水胁迫下的夏玉米叶片叶绿素含量,提高了光合性能,延缓了叶片衰老,有效地缓解了涝渍对夏玉米的不利影响。【本研究切入点】淹水胁迫对作物发育影响的研究已有报道,但淹水对不同生育期花生根系生长及6-BA对淹水的缓解作用尚缺乏深入研究。【拟解决的关键问题】本研究选用山花108为材料,研究外源细胞分裂素对不同生育时期淹水胁迫下花生根系干重、根长密度、根系呼吸酶活性及根系内源激素的影响,探讨淹水逆境下外源6-BA对不同生育时期根系生长的调控作用,深入认识激素在淹水逆境下调控根系发育及产量形成的作用机制,为花生生长的化学调控提供理论依据。1 材料与方法
1.1 试验设计
试验于2018—2019年在山东农业大学农学试验农场和作物生物学国家重点实验室进行。试验采用土柱栽培,供试土壤基本养分状况见表1。Table 1
表1
表1试验土壤基本养分状况
Table 1
| 年份Year | 有机质Organic matter (g·kg-1) | 全氮Total nitrogen (g·kg-1) | 速效磷Available P (mg·kg-1) | 速效钾Available K (mg·kg-1) | pH |
|---|---|---|---|---|---|
| 2018 | 16.68 | 0.768 | 76.53 | 108.27 | 7.6 |
| 2019 | 15.83 | 0.735 | 73.38 | 101.25 | 7.4 |
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花生品种选用山花108,种植于直径32 cm,高100 cm的PVC管中。每土柱施6.04 g复合肥（N、P2O5、K2O含量均为15%）。播种前进行灌水沉实,使土柱与大田状况尽可能一致。2年试验均于4月27日播种,每柱播3粒种子,出苗后留生长一致的花生2株,9月8日收获。其他如除草、除虫等田间管理同一般高产大田。在花生V3、R3、R5、R7[16]4个时期进行淹水处理（水面高于土面3 cm）,淹水持续10 d。淹水结束后,每天18：00后叶片喷施15 mg·L-16-BA,用量为250 mL·m-2,连续喷5 d。为使激素能更好地附着于叶片,激素溶液中加入0.5% Tween-20。试验设9个处理组合,分别为正常水分管理（生育期内保持田间持水量60%—70%）（CK）、苗期淹水处理（V3-W）、苗期淹水后喷施6-BA处理（V3-S）、花针期淹水处理（R3-W）、花针期淹水后喷施6-BA处理（R3-S）、结荚期淹水处理（R5-W）、结荚期淹水后喷施6-BA处理（R5-S）、饱果期淹水处理（R7-W）、饱果期淹水后喷施6-BA处理（R7-S）,每次取样3次重复。
1.2 测定项目与方法
1.2.1 取样方法 分别在V3时期淹水处理后0、5、10、15、20、60、80、100 d;R3时期淹水处理后0、5、10、15、20、40、60、80 d;R5时期淹水处理后0、5、10、15、20、40 d;R7时期淹水处理后0、5、10、15 d取样,每次取6柱,分为两部分,一部分用于测定植株干物质积累和根系形态分析;另一部分用去离子水冲洗,吸水纸吸干后保存于-80℃冰箱中,用于测定根系呼吸酶活性和内源激素含量。1.2.2 根系形态测定分析和干物质积累测定 采用扫描系统（Epson Perfection V800 Photo）对根系进行扫描。扫描时将根系放入特制的透明托盘内,加入3—5 ml水以避免根系分支的互相缠绕。扫描后采用分析程序（Win Rhizo Pro Vision 5.0 a）对图像进行分析,得到根系长度,计算根长密度。扫描后的根系在105℃中杀青0.5 h, 65℃烘箱中烘干48 h至恒重后测定植株根系干物质量。
1.2.3 根系无氧呼吸酶活性的测定 选取长3— 5 cm、直径1.5 mm左右的生长根,称取0.5 g左右,放入预冷研钵中,加入2 mL pH 6.8的50 mmol·L-1 Tris-HCl提取液（含5 mmol·L-1 MgCl2、5 mmol·L-1 β-巯基乙醇、体积分数为15%甘油、l mmol·L-1 EDTA、1 mmol·L-1EGTA和0.1 mmol·L-1苯甲基磺酰氟）,冰浴研磨,于4℃12 000×g离心20 min,上清液即为酶液。
参考WATER等[17]方法测定乙醇脱氢酶（ADH）活性。取2.85 mL pH 8.0的150 mmol·L-1 Tris-HCl反应液（含0.3 mmol·L-1 NADH）和50 μL酶液混匀,然后加入30 μL 95%乙醇启动反应,于340 nm处测定吸光度值的变化。
依据BERGMEGER[18]方法测定乳酸脱氢酶（LDH）活性。取3 mL pH 7.0的0.1 mol·L-1 磷酸缓冲液（含4 μmol·L-1 NADH、0.2 mol·L-1 丙酮酸）,用0.15 mL酶提取液启动反应,于340 nm处测定吸光度值的变化。
1.2.4 根系有氧呼吸酶活性的测定 选取长3—5 cm、直径1.5 mm左右的生长根,称取0.5 g左右,放入预冷研钵中,加入2 mL pH 8.0的50 mmol·L-1 Tris-HCl提取液（含5 mmol·L-1 MgCl2、25 mmol·L-1 β-巯基乙醇、5 mmol·L-1 EDTA）,冰浴研磨后,于4℃12 000×g离心20 min,上清液即为酶液。
参照薛应龙[19]的方法测定苹果酸脱氢酶（MDH）活性。取880 μL pH 8.0的25 mmol·L-1Tris-HCl（含1 mmol·L-1 EDTA、0.2 mmol·L-1 NADH、20 μL酶液）,用100 μL的0.5 mmol·L-1草酰乙酸启动反应,于340 nm处测定吸光度的变化。
1.2.5 根系内源激素含量测定 采用酶联免疫法测定根系内源ZR和ABA的含量[20]。液氮研磨样品成粉末状,称取0.2 g,加2 mL样品提取液（80%甲醇,内含1 mmol·L-1二叔丁基对甲苯酚）,并在4℃下保存过夜,10 000×g离心20 min,取上清液。沉淀继续用1 mL提取液,提取2次,合并上清液。上清液过Sep-Pak C-18固相萃取柱（Waters,USA）。具体步骤是：100%甲醇活化→纯水洗柱→上样→纯水冲洗→80%甲醇洗脱。将洗脱液转入5 mL离心管,氮气吹干,用1 mL PBS（0.01 mol·L-1,pH7.4）复溶,然后用ZR和ABA酶联免疫试剂盒（南京草本源提供）测定ZR和ABA含量。
1.2.6 净光合速率测定 采用Li-6400便携式光合仪（Li-Cor Inc,USA）测定主茎倒三叶的净光合速率（Pn,μmol CO2·m-2·s-1）,各处理选取5株长势一致的植株。
1.2.7 SPAD值测定 采用MultiSpeQ多功能植物测量仪（Photosynp,USA）测定主茎倒三叶SPAD值,测定时每处理选取10个有代表性的叶片。
1.2.8 花生产量测定 收获时测定单株产量。每处理取10株考察单株结果数,待花生荚果晒干至恒重后测定单株产量。
1.3 数据统计分析
采用Microsoft Excel 2007与SigmaPlot 12.0进行数据处理与作图,用DPS10.0统计分析软件对试验数据进行分析,差异显著性检验采用LSD法。由于2年趋势相近,文中数据为2018与2019年数据平均值。2 结果
2.1 外源6-BA对淹水胁迫下根系生长发育的影响
2.1.1 外源6-BA对淹水胁迫下不同土层根长密度的影响 图1是不同生育时期淹水结束后5 d的根长密度。可以看出,与CK相比,各生育时期淹水均降低了根长密度,V3时期淹水对根系生长影响最大,表现为20—40 cm土层无根系。与淹水处理相比,淹水后喷施6-BA显著增加了V3、R3和R5时期的根长密度。2018与2019生长季,V3-S处理0—20 cm土层平均增加了21.75%;R3-S处理20—40 cm土层平均增加了11.35%;R5-S处理40—60 cm土层增加了22.56%。但饱果期淹水后喷施6-BA对根长密度无显著影响。图1
新窗口打开|下载原图ZIP|生成PPT图1外源6-BA对不同生育时期淹水胁迫下花生根长密度的影响
A,B,C,D表示V3,R3,R5,R7时期淹水处理。下同
Fig. 1Effects of exogenous 6-BA on root length density in peanut under water-logging stress at different growth stages
A, B, C, and D represent flooding during the V3, R3, R5 and R7 periods, respectively. The same as below
2.1.2 外源6-BA对淹水胁迫下不同土层根系干重的影响 图2是不同生育时期淹水结束后5 d的根系干重。可以看出,各生育时期淹水均抑制了根系的生长,降低了根系的干重。2018与2019生长季,V3-W处理20—40 cm土层根系干重为零;而R3-W处理40—60 cm土层的根系干重平均降低了39.33%;R5-W处理40—60 cm土层的根系干重平均降低了15.93%。淹水后喷施6-BA后增加了V3-S、R3-S、R5-S处理的根系干重,其中V3-S处理0—20 cm土层根系干重较V3-W处理增加了4.72%,R3-S处理20—40 cm土层根系干重较R3-W处理增加了8.42%。饱果期淹水及喷施6-BA对根系干重无显著影响。
图2
新窗口打开|下载原图ZIP|生成PPT图2外源6-BA对不同生育时期淹水胁迫下花生根系干重的影响
Fig. 2Effects of exogenous 6-BA on dry root weight in peanut under water-logging stress at different growth stages
2.2 外源6-BA对淹水胁迫下根系呼吸酶活性的影响
2.2.1 外源6-BA对淹水胁迫下根系乙醇脱氢酶（ADH）活性变化的影响 由图3可以看出,不同生育时期淹水均提高了根系ADH活性。淹水结束后0 d,V3-W、R3-W、R5-W与R7-W 2年较CK平均增加了12.50倍、8.41倍、6.74倍与6.16倍。其中V3-W于淹水结束后0—20 d内逐渐恢复到CK水平;R3-W处理于淹水结束后20—40 d恢复到CK水平;而R5-W处理于淹水结束后40 d逐渐恢复到与CK差异较小。淹水后喷施6-BA增加了根系ADH活性,且以花针期喷施效果最为明显。图3
新窗口打开|下载原图ZIP|生成PPT图3外源6-BA对不同生育时期淹水胁迫下花生根系乙醇脱氢酶活性变化的影响
Fig. 3Effects of exogenous 6-BA on ADH activity changes in peanut root under water-logging stress at different growth stages
2.2.2 外源6-BA对淹水胁迫下根系乳酸脱氢酶（LDH）活性变化的影响 由图4可知,不同生育时期淹水显著增加了根系LDH活性。且随着淹水时期的延后,增加幅度逐渐降低。淹水结束后,V3-W处理2年较CK平均增加了18.99倍,而R7-W处理增加了6.66倍。且V3-W处理于淹水结束后0—20 d内逐渐恢复到CK水平;R5-W处理淹水结束后20—40 d恢复到与CK差异较小。淹水后喷施6-BA缓解了LDH活性的升高,尤其降低了R3-S处理LDH的活性,喷施10 d后,LDH活性比R3-W处理下降了7.40%。
图4
新窗口打开|下载原图ZIP|生成PPT图4外源6-BA对不同生育时期淹水胁迫下花生根系乳酸脱氢酶活性变化的影响
Fig. 4Effects of exogenous 6-BA on LDH activity changes in peanut root under water-logging stress at different growth stages
2.2.3 外源6-BA对淹水胁迫下根系苹果酸脱氢酶（MDH）活性变化的影响 由图5可知,不同生育时期淹水均降低了根系苹果酸脱氢酶（MDH）的活性。淹水结束后0 d,V3-W、R3-W、R5-W与R7-W处理2年较CK降低了65.16%、57.15%、49.25%、44.16%。且V3-W于淹水结束后0—20 d内逐渐恢复到CK水平;R3-W在淹水结束后15—20 d恢复到CK水平。淹水后喷施6-BA缓解了MDH活性的下降,在喷施激素10 d后,V3-S处理较V3-W处理2年平均增加了8.94%;R3-S处理增加了8.97%。饱果期淹水后喷施6-BA对MDH活性无显著影响。
图5
新窗口打开|下载原图ZIP|生成PPT图5外源6-BA对不同生育时期淹水胁迫下花生根系苹果酸脱氢酶活性变化的影响
Fig. 5Effects of exogenous 6-BA on MDH activity changes in peanut root under water-logging stress at different growth stages
2.3 外源6-BA对淹水胁迫下根源激素变化的影响
2.3.1 外源6-BA对淹水胁迫下根源ZR变化的影响 随着生育时期的推进,根系中ZR含量表现为先升后降,在结荚期达到最大（图6）。淹水结束后,V3-W、R3-W、R5-W与R7-W处理2年平均降低了16.84%、15.61%、15.35%、6.58%。胁迫解除后,V3-W与R3-W处理ZR含量增加并逐渐恢复到CK水平。淹水后喷施6-BA增加了根系ZR含量,增加幅度表现为R3-S>V3-S>R5-S>R7-S,分别较各淹水处理增加了5.47%、4.51%、4.31%、1.78%。图6
新窗口打开|下载原图ZIP|生成PPT图6外源6-BA对不同生育时期淹水胁迫下花生根系ZR含量变化的影响
Fig. 6Effects of exogenous 6-BA on ZR content changes in peanut root under water-logging stress at different growth stages
2.3.2 外源6-BA对淹水胁迫下根源ABA变化的影响 2018与2019年,根系ABA含量呈现先降后升的趋势,并于结荚期达到最低值（图7）。与CK相比,不同生育时期淹水均增加了根系ABA含量,V3-W处理2年平均增加了30.84%;R3-W处理增加了21.35%;R5-W处理增加了17.21%;R7-W处理增加了11.91%。且V3-W与R3-W处理分别于淹水结束后20—60 d与40—60 d内逐渐恢复到CK水平。淹水后喷施6-BA降低了ABA含量,以R3-S处理喷施效果明显,2个生长季平均比R3-W处理降低了5.66%。
图7
新窗口打开|下载原图ZIP|生成PPT图7外源6-BA对不同生育时期淹水胁迫下花生根系ABA含量变化的影响
Fig. 7Effects of exogenous 6-BA on ABA content changes in peanut root under water-logging stress at different growth stages
2.4 花生根系内源激素与根系呼吸酶的关系
由表2相关分析结果表明,根系脱落酸（ABA）含量与乳酸脱氢酶（LDH）活性呈极显著正相关关系,与乙醇脱氢酶（ADH）、苹果酸脱氢酶（MDH）活性相关性不显著。根系玉米素核苷（ZR）含量与苹果酸脱氢酶（MDH）活性呈极显著正相关关系,与乙醇脱氢酶（ADH）、乳酸脱氢酶（LDH）活性无显著性相关关系。Table 2
表2
表2花生根系内源激素与根系呼吸酶的相关性
Table 2
| 玉米素核苷ZR | 脱落酸ABA | 乙醇脱氢酶ADH | 乳酸脱氢酶LDH | |
|---|---|---|---|---|
| 脱落酸 ABA | 0.397 | |||
| 乙醇脱氢酶 ADH | 0.286 | 0.480 | ||
| 乳酸脱氢酶 LDH | 0.305 | 0.685** | 0.898** | |
| 苹果酸脱氢酶 MDH | 0.705** | 0.006 | -0.225 | -0.265 |
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2.5 外源6-BA对淹水胁迫下叶片光合特性的影响
2.5.1 外源6-BA对淹水胁迫下叶片净光合速率（Pn）变化的影响 由图8可以看出,叶片净光合速率在花生生育期内呈单峰变化,在结荚期达到最大值。不同生育时期淹水后花生净光合速率下降。V3-W、R3-W、R5-W与R7-W处理Pn 2年平均降低了17.98%、19.74%、25.57%与14.32%。淹水结束后,V3-W与R3-W处理净光合速率逐渐恢复到CK水平。喷施6-BA后提高了主茎倒三叶的Pn,其中以R3-S处理增加效果显著,比R3-W处理增加了7.92%。饱果期淹水后喷施6-BA对Pn无显著影响。图8
新窗口打开|下载原图ZIP|生成PPT图8外源6-BA对不同生育时期淹水胁迫下花生叶片Pn变化的影响
Fig. 8Effects of exogenous 6-BA on Pn changes in peanut leaves under water-logging stress at different growth stages
2.5.2 外源6-BA对淹水胁迫下叶片SPAD值变化的影响 由图9可以看出,淹水使花生主茎倒三叶的SPAD值降低。降低幅度表现为R5-W>R3-W>V3-W>R7-W。喷施6-BA后,SPAD值提高,增加幅度表现为R3-S>R5-S>V3-S>R7-S。淹水结束后0 d,2018与2019生长季R5-W处理SPAD值较CK平均降低了37.05%。喷施6-BA后5 d,R3-S处理较R3-W平均增加了7.98%。R3-W与R3-S处理于胁迫解除后60—80 d逐渐恢复。R5-W与R5-S处理在淹水解除后一直低于CK。而R7-W与R7-S处理SPAD值于淹水结束后15 d内逐渐降低。
图9
新窗口打开|下载原图ZIP|生成PPT图9外源6-BA对不同生育时期淹水胁迫下花生叶片SPAD值变化的影响
Fig. 9Effects of exogenous 6-BA on the SPAD value of peanut leaves under water-logging stress at different growth stages
2.6 外源6-BA对淹水胁迫下荚果产量的影响
由表3可知,不同生育时期淹水显著降低了花生的单株结果数、单株产量、出仁率。与CK相比,R5时期淹水减产幅度最大,R5-W处理的单株产量2年度分别降低30.43%（2018）与25.75%（2019）。喷施6-BA后,不同生育时期喷施外源6-BA提高了花生单株产量,但增加幅度不同,以花针期喷施增产幅度最大。与R3-W处理相比,R3-S处理的单株产量2年分别提高了5.68%（2018）与8.10%（2019）,这是由于外源6-BA显著提高了单株结果数与双仁果率。与R7-W处理相比,饱果期喷施6-BA对单株结果数及百果重无显著影响。Table 3
表3
表3外源6-BA对不同生育时期淹水胁迫下花生产量及产量构成因素的影响
Table 3
| 年份 Year | 处理 Treatment | 单株结果数 Pods number per plant | 百果重 Weight per 100 kernel (g) | 双果仁率 Double kernel rate (%) | 单株产量 Pod mass per plant (g) | 出仁率 Kernel rate (%) |
|---|---|---|---|---|---|---|
| 2018 | CK | 20.47a | 135.49a | 66.30a | 27.05a | 68.46a |
| V3-W | 18.33d | 129.85abc | 61.77bc | 23.80c | 67.61a | |
| V3-S | 20.39bc | 133.27a | 62.98ab | 25.06b | 67.93a | |
| R3-W | 16.67e | 126.51bcd | 55.70d | 22.17d | 66.22abc | |
| R3-S | 17.33e | 131.92ab | 58.28cd | 23.43c | 65.87abc | |
| R5-W | 14.17g | 122.24d | 55.48d | 18.82e | 68.34a | |
| R5-S | 15.67f | 124.92cd | 55.79d | 19.62e | 67.13ab | |
| R7-W | 19.67c | 129.35abc | 64.66ab | 24.70b | 62.05c | |
| R7-S | 20.33bc | 130.17abc | 64.95ab | 25.15b | 62.49bc | |
| 2019 | CK | 18.20a | 148.85a | 65.40a | 27.21a | 67.55a |
| V3-W | 17.83a | 146.84ab | 63.65ab | 26.17a | 66.95a | |
| V3-S | 18.00a | 150.28a | 64.44a | 27.05a | 67.52a | |
| R3-W | 15.33abc | 137.55bc | 55.51c | 22.60bc | 64.90abc | |
| R3-S | 16.83ab | 145.51ab | 58.50bc | 24.43ab | 65.42ab | |
| R5-W | 14.83c | 131.11c | 57.97bc | 19.39c | 59.07d | |
| R5-S | 15.25bc | 136.90bc | 58.54bc | 20.88c | 61.71bcd | |
| R7-W | 17.25a | 147.02ab | 56.53c | 25.37ab | 60.83cd | |
| R7-S | 17.33a | 148.00a | 56.73c | 25.73ab | 61.05bcd |
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3 讨论
3.1 外源6-BA对不同生育时期淹水胁迫下花生根系呼吸特性的影响
淹水导致可用的O2含量降低,根系从有氧呼吸转化无氧呼吸,表现为无氧呼吸酶ADH与LDH活性增强[21],有氧呼吸酶MDH活性降低[22],从而可以促进乙醇发酵与乳酸发酵,提高NADH的再氧化过程,并维持适当的ATP水平[23]。僧珊珊等[24]研究发现淹水条件下,与耐涝品种相比,不耐涝玉米品种根系的ADH与LDH活性显著增加。另外,张凤等[25]的研究发现,不同生育时期淹水,花生根系的ADH活性差异显著。本研究结果表明不同生育时期淹水均抑制了花生根系MDH活性,同时诱导了ADH、LDH活性的上升;不同生育时期相比,苗期酶活性增加幅度最大,表明淹水对根系生育前期的呼吸代谢影响较大。淹水胁迫解除后,ADH、LDH活性逐渐恢复到对照水平。淹水后喷施外源6-BA显著提高了根系MDH与ADH活性,降低了LDH活性,且淹水胁迫解除后缩短了恢复期;各生育时期喷施6-BA相比,花针期喷施6-BA处理的ADH与MDH活性变化幅度最大,这表明淹水后施用外源6-BA可改善根系的呼吸作用;而ADH活性的提高可以促进乙醛转化为乙醇,降低其对根系细胞的毒性,缓解其对植株的伤害[26]。前人研究表明,LDH作为催化乳酸发酵的关键酶,其催化产生的乳酸引起胞质酸化是引起植株死亡的主要原因之一[27]。外源6-BA的施用减缓了其活性的升高,抑制了胞质的酸化程度,降低了根尖细胞的损伤,从而提高了花生的抗涝性。3.2 外源6-BA对不同生育时期淹水胁迫下花生根系内源激素的影响
ABA和CTK是两种重要的激素信号物质,在逆境胁迫中具有重要的作用[28]。前人研究发现,在涝渍胁迫下,内源性ABA含量迅速增加,通过合成第二信使（Ca2+、ROS、NO）来调节气孔开闭[29]。WANG等[30]发现叶片气孔关闭不仅仅受ABA的影响,而是ABA含量增加和ZR含量下降共同作用的结果。ZR在根中催化合成,通过木质部转运到枝条中以协调气孔开闭[31]。本研究发现各生育时期淹水均增加了花生根系中ABA的含量,降低了ZR的含量。这是由于淹水促进了细胞分裂素氧化酶基因的表达,提高了氧化酶活性,不利于ZR积累[32]。前人研究发现,ABA通过延缓细胞伸长来抑制根系的生长,而此过程的实现需要抑制CTK信号的转导[33]。本研究发现,不同生育时期淹水抑制了根系生长,降低了根系长度。ABA抑制根的生长与NADPH氧化酶的活化和ROS的积累有关[34]。胁迫后喷施外源细胞分裂素可以提高内源CTK的含量[35]。刘敬然等认为外源6-BA可以显著提高棉花CTK含量,降低ABA含量[36]。本研究发现,淹水后喷施6-BA降低了根系ABA含量,提高了ZR含量。与淹水处理相比,淹水后喷施6-BA增加了根系长度,提高了根系干重,表明外源6-BA一方面降低了内源ABA对根细胞伸长的抑制作用,促进根系的生长;另一方面提高了叶片的光合速率,促进了光合产物向根系的转运,进而提高根系生物量。研究发现,缺氧条件下低能量信号传导与植物激素信号传导途径密切相关[37]。而淹水缺氧条件下,根系产生ATP的主要机制为糖酵解与发酵。本试验相关性分析表明,ABA含量与LDH活性呈极显著正相关关系,ZR含量与MDH活性呈极显著正相关关系,这表明内源ZR和ABA含量变化在根系呼吸中起重要作用。喷施外源细胞分裂素通过调节内源ZR、ABA含量,进而调控LDH和MDH活性。3.3 外源6-BA对不同生育时期淹水胁迫下花生产量及其构成因素的影响
淹水影响叶片的光合作用,光合同化生产能力减弱,进而导致干物质积累量减少,影响籽粒充实,最终导致产量下降[38,39]。前人研究表明,淹水后小麦旗叶的叶绿素含量显著下降[40]。叶绿素是植物叶片光合作用的物质基础,其含量高低能反映植物的生长状况和叶片的光合能力。本研究表明,不同生育时期淹水后主茎倒三叶的SPAD值显著降低（图9）,分析原因为淹水胁迫影响了花生的呼吸代谢,细胞内积累较多的有害物质（氧自由基、乙醛、乳酸等）,破坏了叶绿体的结构,加速了叶绿体的分解,最终导致叶绿素含量下降[41,42]。花生淹水后地上部叶片光合速率显著降低（图8）,进而影响花生的干物质积累与分配,降低了花生的单株结果数与百果重,最终降低花生产量。本研究结果表明,R5时期淹水显著降低了花生的单株结果数,单株产量2年平均降低29.58%。研究发现涝后喷施6-BA能够提高作物的叶面积指数[15],增加叶绿素含量[43],从而提高作物光合生产能力,最终提高产量。本研究表明,淹水后喷施6-BA可显著提高花生单株产量,且以R3时期处理效果显著,产量比淹水处理平均提高6.89%,这表明6-BA对花生花针期淹水的调控效应最明显。4 结论
淹水通过增加根系内源ABA含量,降低ZR含量,一方面降低了根系MDH活性,提高了根系ADH与LDH活性;另一方面降低了叶片的叶绿素含量与光合速率,减少了光合产物的积累,产量降低。淹水后喷施外源6-BA降低了根系内源ABA含量,提高了ZR含量,一方面提高了根系ADH、MDH活性,降低了LDH活性,改善根系的呼吸性能,促进根系生长;另一方面提高了叶片叶绿素含量和光合速率,促进干物质的积累与分配,提高了单株结果数与单株产量,且不同生育时期相比,花针期喷施单株产量增加效果明显。参考文献 原文顺序
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被引期刊影响因子
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为了引导和深化黄淮海区域花生生产的区域分工、高效合理利用农业资源和提升花生产品质量, 基于GIS技术和数理统计方法, 对黄淮海区域花生生产与品质特征进行了研究。结果表明: 该区花生生产技术先进, 单产水平高, 出口基础好, 加工业已见雏形, 种植规模与产量在全国占有举足轻重的地位, 是稳定我国植物油市场的重要区域; 受生态条件所致, 生产上应用的品种主要是普通型花生, 属食用中蛋白、油用高脂肪品质类型, 且油亚比值(O/L)普遍偏低, 黄曲霉毒素、丁酰肼和重金属含量超标已成为花生出口的主要障碍; 在空间分布上, 花生蛋白质含量东高西低、南高北低、丘陵区高于平原区, 脂肪含量北高南低、丘陵区高于平原区, O/L比值呈现由西向东、由北向南增加的趋势。气候与土壤肥力条件是影响花生蛋白质、脂肪含量及O/L比值高低和空间分布的主导因子。
URL [本文引用: 1]
为了引导和深化黄淮海区域花生生产的区域分工、高效合理利用农业资源和提升花生产品质量, 基于GIS技术和数理统计方法, 对黄淮海区域花生生产与品质特征进行了研究。结果表明: 该区花生生产技术先进, 单产水平高, 出口基础好, 加工业已见雏形, 种植规模与产量在全国占有举足轻重的地位, 是稳定我国植物油市场的重要区域; 受生态条件所致, 生产上应用的品种主要是普通型花生, 属食用中蛋白、油用高脂肪品质类型, 且油亚比值(O/L)普遍偏低, 黄曲霉毒素、丁酰肼和重金属含量超标已成为花生出口的主要障碍; 在空间分布上, 花生蛋白质含量东高西低、南高北低、丘陵区高于平原区, 脂肪含量北高南低、丘陵区高于平原区, O/L比值呈现由西向东、由北向南增加的趋势。气候与土壤肥力条件是影响花生蛋白质、脂肪含量及O/L比值高低和空间分布的主导因子。
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Plant adaptive potential is critically dependent upon efficient communication and co-ordination of resource allocation and signalling between above- and below-ground plant parts. Plant roots act as gatekeepers that sense and encode information about soil physical, chemical and biological factors, converting them into a sophisticated network of signals propagated both within the root itself, and also between the root and shoot, to optimise plant performance for a specific set of conditions. In return, plant roots receive and decode reciprocal information coming from the shoot. The communication modes are highly diverse and include a broad range of physical (electric and hydraulic signals, propagating Ca2+ and ROS waves), chemical (assimilates, hormones, peptides and nutrients), and molecular (proteins and RNA) signals. Further, different signalling systems operate at very different timescales. It remains unclear whether some of these signalling systems operate in a priming mode(s), whereas others deliver more specific information about the nature of the signal, or whether they carry the same 'weight'. This review summarises the current knowledge of the above signalling mechanisms, and reveals their hierarchy, and highlights the importance of integration of these signalling components, to enable optimal plant functioning in a dynamic environment.
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Alcohol dehydrogenase (Adh) is the key enzyme in alcohol fermentation. We analyzed Adh expression in order to clarify the role of Adh of soybeans (Glycine max) to flooding stress. Proteome analysis confirmed that expression of Adh is significantly upregulated in 4-day-old soybean seedlings subjected to 2 days of flooding. Southern hybridization analysis and soybean genome database search revealed that soybean has at least 6 Adh genes. The GmAdh2 gene that responded to flooding was isolated from soybean cultivar Enrei. Adh2 expression was markedly increased 6 h after flooding and decreased 24 h after floodwater drainage. In situ hybridization and Western blot indicated that flooding strongly induces Adh2 expression in RNA and protein levels in the root apical meristem. Osmotic, cold, or drought stress did not induce expression of Adh2. These results indicate that Adh2 is a flooding-response specific soybean gene expressed in root tissue.
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研究了水分胁迫对花生幼苗叶片内源激素含量的影响。结果表明，GA3含量先升后降，出现了两个高峰值，ABA含量则逐渐升高，IAA和ZR先降低后升高。随着胁迫时间的延长，叶片中ABA/IAA的值先升后降，ABA/ZR逐渐升高。
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研究了水分胁迫对花生幼苗叶片内源激素含量的影响。结果表明，GA3含量先升后降，出现了两个高峰值，ABA含量则逐渐升高，IAA和ZR先降低后升高。随着胁迫时间的延长，叶片中ABA/IAA的值先升后降，ABA/ZR逐渐升高。
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选用登海605和郑单958为试验材料，在大田条件下设置淹水时期(三叶期、拔节期、花后10 d)和淹水持续时间(淹水3 d和6 d)处理研究淹水对夏玉米光合特性的影响。结果表明，淹水胁迫后叶面积指数和叶绿素含量显著下降，净光合速率(Pn)、气孔导度(Gs)、细胞间隙CO2浓度(Ci)显著降低，三叶期淹水6 d对其影响最显著，开花期郑单958和登海605功能叶片的Pn、Gs、Ci较对照的分别下降21.24%、33.65%、16.49%和24.50%、32.31%、10.99%。淹水后叶片的Fv/Fm、Fm/Fo、ΦPSII显著下降，三叶期淹水6 d后影响最显著，登海605和郑单958分别下降16.86%、17.12%、11.67%和13.58%、21.40%、22.52%，这是导致夏玉米产量显著下降的主要光合生理机制，三叶期淹水6 d产量降幅最大，登海605和郑单958较对照分别下降41.51%和40.96%。三叶期淹水造成的影响最大，拔节期淹水次之，开花后10 d淹水造成的影响较小，其影响随淹水持续时间的延长而加剧。
DOI:10.3724/SP.J.1006.2015.00329URL [本文引用: 2]
选用登海605和郑单958为试验材料，在大田条件下设置淹水时期(三叶期、拔节期、花后10 d)和淹水持续时间(淹水3 d和6 d)处理研究淹水对夏玉米光合特性的影响。结果表明，淹水胁迫后叶面积指数和叶绿素含量显著下降，净光合速率(Pn)、气孔导度(Gs)、细胞间隙CO2浓度(Ci)显著降低，三叶期淹水6 d对其影响最显著，开花期郑单958和登海605功能叶片的Pn、Gs、Ci较对照的分别下降21.24%、33.65%、16.49%和24.50%、32.31%、10.99%。淹水后叶片的Fv/Fm、Fm/Fo、ΦPSII显著下降，三叶期淹水6 d后影响最显著，登海605和郑单958分别下降16.86%、17.12%、11.67%和13.58%、21.40%、22.52%，这是导致夏玉米产量显著下降的主要光合生理机制，三叶期淹水6 d产量降幅最大，登海605和郑单958较对照分别下降41.51%和40.96%。三叶期淹水造成的影响最大，拔节期淹水次之，开花后10 d淹水造成的影响较小，其影响随淹水持续时间的延长而加剧。
DOI:10.3146/i0095-3679-9-1-11URL [本文引用: 1]
DOI:10.1093/jxb/42.11.1437URL [本文引用: 1]
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灌浆初期籽粒中ABA(脱落酸)含量(ng g-1FW)和ABA与GA(赤霉素)的比值，强势粒高于弱势粒，籽粒充实度好的组合高于籽粒充实度差的组合。开花后2天，喷施外源ABA，籽粒中ABA含量和ABA/GA增大，喷施外源GA后籽粒中GA含量增加，ABA/GA减小。低浓度(15mg/L)ABA处理后，灌浆初期籽粒中ADPG焦磷酸酶和淀粉合成酶活性及淀粉含量增加，
URL [本文引用: 1]
灌浆初期籽粒中ABA(脱落酸)含量(ng g-1FW)和ABA与GA(赤霉素)的比值，强势粒高于弱势粒，籽粒充实度好的组合高于籽粒充实度差的组合。开花后2天，喷施外源ABA，籽粒中ABA含量和ABA/GA增大，喷施外源GA后籽粒中GA含量增加，ABA/GA减小。低浓度(15mg/L)ABA处理后，灌浆初期籽粒中ADPG焦磷酸酶和淀粉合成酶活性及淀粉含量增加，
DOI:10.1104/pp.116.4.1573URLPMID:9536076 [本文引用: 1]
The goal of this research was to resolve the hypoxic and anoxic responses of maize (Zea mays) sucrose (Suc) synthases known to differ in their sugar regulation. The two maize Suc synthase genes, Sus1 and Sh1, both respond to sugar and O2, and recent work suggests commonalities between these signaling systems. Maize seedlings (NK508 hybrid, W22 inbred, and an isogenic sh1-null mutant) were exposed to anoxic, hypoxic, and aerobic conditions (0, 3, and 21% O2, respectively), when primary roots had reached approximately 5 cm. One-centimeter tips were excised for analysis during the 48-h treatments. At the mRNA level, Sus1 was rapidly up-regulated by hypoxia (approximately 5-fold in 6 h), whereas anoxia had less effect. In contrast, Sh1 mRNA abundance increased strongly under anoxia (approximately 5-fold in 24 h) and was much less affected by hypoxia. At the enzyme level, total Suc synthase activity rose rapidly under hypoxia but showed little significant change during anoxia. The contributions of SUS1 and SH1 activities to these responses were dissected over time by comparing the sh1-null mutant with the isogenic wild type (Sus+, Sh1+). Sh1-dependent activity contributed most markedly to a rapid protein-level response consistently observed in the first 3 h, and, subsequently, to a long-term change mediated at the level of mRNA accumulation at 48 h. A complementary midterm rise in SUS1 activity varied in duration with genetic background. These data highlight the involvement of distinctly different genes and probable signal mechanisms under hypoxia and anoxia, and together with earlier work, show parallel induction of
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以美早/东北山樱桃(Prunus serrulataG.Don)和美早/马哈利(P. mahaleb L.)为试材，研究了淹水对甜樱桃根系（生长根和褐色木质根）呼吸强度和呼吸酶活性的影响.结果表明：淹水过程中，两种甜樱桃砧木生长根和褐色木质根呼吸强度均呈下降趋势，生长根降幅更大；东北山樱桃生长根和褐色木质根呼吸强度降幅分别是马哈利的1.47和1.36倍.丙酮酸脱羧酶(PDC)、乳酸脱氢酶(LDH)活性在两类根系中均呈先升后降趋势，乙醇脱氢酶(ADH)活性在生长根中亦先升后降，而在褐色木质根中为上升趋势，3种酶活性变化幅度表现为生长根大于褐色木质根；东北山樱桃ADH和LDH活性增幅大于马哈利，但PDC活性则相反.两类根系苹果酸脱氢酶(MDH)活性均下降,且生长根降幅大于褐色木质根；东北山樱桃MDH活性降幅大于马哈利.说明生长根对淹水的敏感性强于褐色木质根；与马哈利相比，东北山樱桃对淹水更敏感.
URL [本文引用: 1]
以美早/东北山樱桃(Prunus serrulataG.Don)和美早/马哈利(P. mahaleb L.)为试材，研究了淹水对甜樱桃根系（生长根和褐色木质根）呼吸强度和呼吸酶活性的影响.结果表明：淹水过程中，两种甜樱桃砧木生长根和褐色木质根呼吸强度均呈下降趋势，生长根降幅更大；东北山樱桃生长根和褐色木质根呼吸强度降幅分别是马哈利的1.47和1.36倍.丙酮酸脱羧酶(PDC)、乳酸脱氢酶(LDH)活性在两类根系中均呈先升后降趋势，乙醇脱氢酶(ADH)活性在生长根中亦先升后降，而在褐色木质根中为上升趋势，3种酶活性变化幅度表现为生长根大于褐色木质根；东北山樱桃ADH和LDH活性增幅大于马哈利，但PDC活性则相反.两类根系苹果酸脱氢酶(MDH)活性均下降,且生长根降幅大于褐色木质根；东北山樱桃MDH活性降幅大于马哈利.说明生长根对淹水的敏感性强于褐色木质根；与马哈利相比，东北山樱桃对淹水更敏感.
DOI:10.1146/annurev-arplant-043014-114813URL [本文引用: 1]
DOI:10.3864/j.issn.0578-1752.2012.20.003URL [本文引用: 1]
【目的】研究不同玉米品种幼苗根显微结构及呼吸代谢对淹水胁迫的响应差异。【方法】以耐涝性不同的玉米品种浚单20（耐涝性强）和登海662（耐涝性弱）为试材，设CK、淹水2、4、6、8 d共5个处理，比较淹水胁迫下不同玉米品种苗期根结构及呼吸代谢差异。【结果】淹水后，两品种幼苗的根干重、根总长度、根系活力均明显下降，随着淹水天数增加，下降幅度增大，两品种间表现为登海662各指标下降幅度显著高于浚单20。其中淹水8 d时，登海662和浚单20根干重分别比对照降低29.7%和12.1%，根系活力分别比对照降低30.6%和8.7%。淹水增加了两品种玉米根气腔面积和根孔积率，平均比对照增加3.9倍和2.8倍，且随着淹水时间延长而增大。淹水使玉米根系的乙醇脱氢酶（ADH）、丙酮酸脱羧酶（PDC）和乳酸脱氢酶（LDH）活性比对照平均增加27.7%、55.0%和2.6倍，其中登海662的增加幅度大于浚单20，且差异显著。【结论】不同玉米品种根系对淹水胁迫的响应不同，耐涝品种具有完整根结构、较高的根系活力、大面积较发达的通气组织、适度的无氧呼吸代谢，因而能够更好地适应淹水胁迫，提高干物质生产能力。
DOI:10.3864/j.issn.0578-1752.2012.20.003URL [本文引用: 1]
【目的】研究不同玉米品种幼苗根显微结构及呼吸代谢对淹水胁迫的响应差异。【方法】以耐涝性不同的玉米品种浚单20（耐涝性强）和登海662（耐涝性弱）为试材，设CK、淹水2、4、6、8 d共5个处理，比较淹水胁迫下不同玉米品种苗期根结构及呼吸代谢差异。【结果】淹水后，两品种幼苗的根干重、根总长度、根系活力均明显下降，随着淹水天数增加，下降幅度增大，两品种间表现为登海662各指标下降幅度显著高于浚单20。其中淹水8 d时，登海662和浚单20根干重分别比对照降低29.7%和12.1%，根系活力分别比对照降低30.6%和8.7%。淹水增加了两品种玉米根气腔面积和根孔积率，平均比对照增加3.9倍和2.8倍，且随着淹水时间延长而增大。淹水使玉米根系的乙醇脱氢酶（ADH）、丙酮酸脱羧酶（PDC）和乳酸脱氢酶（LDH）活性比对照平均增加27.7%、55.0%和2.6倍，其中登海662的增加幅度大于浚单20，且差异显著。【结论】不同玉米品种根系对淹水胁迫的响应不同，耐涝品种具有完整根结构、较高的根系活力、大面积较发达的通气组织、适度的无氧呼吸代谢，因而能够更好地适应淹水胁迫，提高干物质生产能力。
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URL [本文引用: 1]
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淹水胁迫引起弱光环境，使气体扩散受限，叶片细胞膜脂过氧化加剧，体内保护酶系统受损，叶绿素降解，丙二醛含量积累，光合速率下降。为了适应淹水环境，植物通过生理生化机制的调节来保证淹水条件下的生命活动。如细胞通过调节渗透物质的含量来保持渗透势的平衡；细胞内各种抗氧化酶活性增加，以清除自由基，避免或者减轻细胞受到伤害；改变代谢途径和激素调节以保持能量储备和低的代谢速率。本文综述了淹水胁迫对细胞膜系统及功能、植物光合作用、植物呼吸、激素、生理代谢、基因调控的影响和淹水结束后植物的生理生态学变化，介绍了植物适应淹水胁迫的机制，并指出植物耐淹响应的分子机理，环境因素对淹没植物的影响，森林淹水胁迫的定位观测是今后需要研究的方面。
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淹水胁迫引起弱光环境，使气体扩散受限，叶片细胞膜脂过氧化加剧，体内保护酶系统受损，叶绿素降解，丙二醛含量积累，光合速率下降。为了适应淹水环境，植物通过生理生化机制的调节来保证淹水条件下的生命活动。如细胞通过调节渗透物质的含量来保持渗透势的平衡；细胞内各种抗氧化酶活性增加，以清除自由基，避免或者减轻细胞受到伤害；改变代谢途径和激素调节以保持能量储备和低的代谢速率。本文综述了淹水胁迫对细胞膜系统及功能、植物光合作用、植物呼吸、激素、生理代谢、基因调控的影响和淹水结束后植物的生理生态学变化，介绍了植物适应淹水胁迫的机制，并指出植物耐淹响应的分子机理，环境因素对淹没植物的影响，森林淹水胁迫的定位观测是今后需要研究的方面。
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为了解涝渍条件下不同树种的耐涝性和适应性，通过田间模拟试验，分析了涝渍胁迫对1年生落羽杉、美国山核桃和乌桕实生苗生长及能量代谢酶的影响.试验分为对照、渍水和淹水3个处理，处理时间为60 d.结果表明：在渍水和淹水条件下，3种树种的相对生长率均表现为落羽杉>美国山核桃>乌桕，落羽杉耐涝性最强，乌桕耐涝性最弱.涝渍条件下，3种树种的根冠比显著增加，更多的光合产物被分配到根系.3树种乙醇脱氢酶和乳酸脱氢酶活性显著升高，其中耐涝性强的落羽杉增加幅度不大，但一直维持在较高水平；而乌桕和美国山核桃在处理初期增幅较大，在处理后期呈下降趋势.3种树种的苹果酸脱氢酶、磷酸己糖异构酶和葡萄糖-6-磷酸脱氢酶-6-磷酸葡萄糖酸脱氢酶的活性均低于对照，其中落羽杉降幅最低，淹水条件下分别下降35.6%、21.0%和22.7%.耐涝性强的树种能够通过自身的调节，维持各种能量代谢途径的强度，为树种在低氧条件下的生命活动提供能量，进而维持一定的生长量.
URL [本文引用: 1]
为了解涝渍条件下不同树种的耐涝性和适应性，通过田间模拟试验，分析了涝渍胁迫对1年生落羽杉、美国山核桃和乌桕实生苗生长及能量代谢酶的影响.试验分为对照、渍水和淹水3个处理，处理时间为60 d.结果表明：在渍水和淹水条件下，3种树种的相对生长率均表现为落羽杉>美国山核桃>乌桕，落羽杉耐涝性最强，乌桕耐涝性最弱.涝渍条件下，3种树种的根冠比显著增加，更多的光合产物被分配到根系.3树种乙醇脱氢酶和乳酸脱氢酶活性显著升高，其中耐涝性强的落羽杉增加幅度不大，但一直维持在较高水平；而乌桕和美国山核桃在处理初期增幅较大，在处理后期呈下降趋势.3种树种的苹果酸脱氢酶、磷酸己糖异构酶和葡萄糖-6-磷酸脱氢酶-6-磷酸葡萄糖酸脱氢酶的活性均低于对照，其中落羽杉降幅最低，淹水条件下分别下降35.6%、21.0%和22.7%.耐涝性强的树种能够通过自身的调节，维持各种能量代谢途径的强度，为树种在低氧条件下的生命活动提供能量，进而维持一定的生长量.
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DOI:10.1016/j.phytochem.2014.03.015URLPMID:24713571 [本文引用: 1]
Nitric oxide (NO) is a free radical gas involved in a myriad of plant physiological processes including immune responses. How NO mediates its biological effects in plant facing microbial pathogen attack is an unresolved question. Insights into the molecular mechanisms by which it propagates signals reveal the contribution of this simple gas in complex signaling pathways shared with reactive oxygen species (ROS) and the second messenger Ca(2+). Understanding of the subtle cross-talks operating between these signals was greatly improved by the recent identification and the functional analysis of proteins regulated through S-nitrosylation, a major NO-dependent post-translational protein modification. Overall, these findings suggest that NO is probably an important component of the mechanism coordinating and regulating Ca(2+) and ROS signaling in plant immunity.
DOI:10.1016/0005-1098(94)90030-2URL [本文引用: 1]
DOI:10.1007/s00344-005-0103-1URL [本文引用: 1]
In this article we review evidence for a variety of long-distance signaling pathways involving hormones and nutrient ions moving in the xylem sap. We argue that ABA has a central role to play, at least in root-to-shoot drought stress signaling and the regulation of functioning, growth, and development of plants in drying soil. We also stress the importance of changes in the pH of the leaf cell apoplast as influenced both by edaphic and climatic variation, as a regulator of shoot growth and functioning, and we show how changes in xylem and apoplastic pH can affect the way in which ABA regulates stomatal behavior and growth. The sensitivity to drought of the pH/ABA sensing and signaling mechanism is emphasized. This allows regulation of plant growth, development and functioning, and particularly shoot water status, as distinct from stress lesions in growth and other processes as a reaction to perturbations such as soil drying.
DOI:10.1016/j.chemosphere.2006.10.007URLPMID:17123580 [本文引用: 1]
The effects of multiple heavy metal stress on the activity of antioxidative enzymes and lipid peroxidation were studied in leaves and roots of two mangrove plants, Kandelia candel and Bruguiera gymnorrhiza, grown under control (10 per thousand NaCl nutrient solution) or five levels of multiple heavy metal stress (10 per thousand NaCl nutrient solution containing different concentration of Pb2+, Cd2+, and Hg2+). Leaves and roots of control and heavy metal-stressed plants were harvested after two months. In leaves of heavy metal-stressed plants superoxide dismutase (SOD) and peroxidase (POD) activities fluctuated in different stress levels compared to the control, while catalase (CAT) activity increased with stress levels in K. candel, but remained unchanged in leaves of B. gymnorrhiza. In comparison with the control, the dynamic tendency of SOD, CAT, and POD activities in roots of heavy metal-stressed plants all ascended, and then declined. The increase in enzyme activities demonstrated that K. candel is more tolerant to heavy metals than B. gymnorrhiza. Lipid peroxidation was enhanced only in leaves of heavy metal-stressed B. gymnorrhiza. These results indicate that in heavy-metal stress antioxidative activities may play an important role in K. candel and B. gymnorrhiza and that cell membrane in leaves and roots of K. candel have greater stability than those of B. gymnorrhiza. For pollution monitoring purposes, POD activity in roots and leaves maybe serve as a biomarker of heavy metal stress in K. candel, while lipid peroxidation maybe serve as biomarker in B. gymnorrhiza.
DOI:10.1080/15592324.2019.1578632URLPMID:30741075 [本文引用: 1]
Cell elongation, which plays an important role in root penetration into the soil, responds to a variety of environmental factors. A previous study demonstrated that abscisic acid, a phytohormone involved in stress responses, inhibits root growth by delaying the onset of cell elongation. In contrast, we recently reported that cytokinins promote elongation of root cells by enhancing actin bundling. However, the control of root cell elongation through the interaction between abscisic acid and cytokinin signaling has not yet been uncovered. Here, we show that abscisic acid-induced delay in cell elongation requires inhibition of cytokinin signaling; further, stress is signaled to cell elongation by the pathway mediated by B-type ARABIDOPSIS RESPONSE REGULATOR 2 (ARR2), which retards root growth.
DOI:10.1093/jxb/ert228URLPMID:23963673 [本文引用: 1]
Reactive oxygen species (ROS) originating from the NADPH oxidases AtrbohD and AtrbohF play an important role in abscisic acid (ABA)-inhibited primary root growth in Arabidopsis. However, the mechanisms underlying this process remain elusive. In this study, the double mutant atrbohD1/F1 and atrbohD2/F2, in which both AtrbohD and AtrbohF were disrupted, were less sensitive to ABA suppression of root cell elongation than wild-type (WT) plants. Furthermore, the double mutants showed impaired ABA responses in roots, including ROS generation, cytosolic Ca(2+) increases, and activation of plasma membrane Ca(2+)-permeable channels compared with WT. Exogenous H2O2 can activate the Ca(2+) currents in roots of atrbohD1/F1. In addition, exogenous application of the auxin transport inhibitor naphthylphthalamic acid effectively promoted ABA inhibition of root growth of the mutants relative to that of WT. The ABA-induced decreases in auxin sensitivity of the root tips were more pronounced in WT than in atrbohD1/F1. These findings suggest that both AtrbohD and AtrbohF are essential for ABA-promoted ROS production in roots. ROS activate Ca(2+) signalling and reduce auxin sensitivity of roots, thus positively regulating ABA-inhibited primary root growth in Arabidopsis.
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DOI:10.3724/SP.J.1006.2013.01078URL [本文引用: 1]
大田播种期(4月25日和5月25日)和植物生长调节剂(6-BA和ABA)试验于2006年在江苏南京(长江流域下游棉区)和河南安阳(黄河流域黄淮棉区)进行，以科棉1号和美棉33B品种为材料，研究外源6-BA和ABA对不同播种期棉铃对位叶光合产物形成与运转的影响及其与产量品质的关系。结果表明: (1) 6-BA显著提高棉铃对位叶蔗糖含量和转化率及淀粉含量，而ABA则主要调节棉铃对位叶内源激素平衡。(2)不同播种期条件下(4月25日和5月25日)，外源6-BA均可使棉花单铃重增加、纤维品质提高，外源ABA在晚播(5月25日，南京试点和安阳试点棉铃发育期MDTmin分别为20.9℃和16.5℃)条件下可使棉花产量(安阳试点)和纤维主要品质指标下降幅度减小；外源6-BA和ABA对棉籽主要品质性状的作用均不显著。(3)晚播条件下外源6-BA和ABA均可提高相对棉花主要品质指标，但两者作用机制不同，外源6-BA主要是提高棉铃对位叶光合产物含量和蔗糖转化率，而ABA则主要是诱导棉株抗逆性。
DOI:10.3724/SP.J.1006.2013.01078URL [本文引用: 1]
大田播种期(4月25日和5月25日)和植物生长调节剂(6-BA和ABA)试验于2006年在江苏南京(长江流域下游棉区)和河南安阳(黄河流域黄淮棉区)进行，以科棉1号和美棉33B品种为材料，研究外源6-BA和ABA对不同播种期棉铃对位叶光合产物形成与运转的影响及其与产量品质的关系。结果表明: (1) 6-BA显著提高棉铃对位叶蔗糖含量和转化率及淀粉含量，而ABA则主要调节棉铃对位叶内源激素平衡。(2)不同播种期条件下(4月25日和5月25日)，外源6-BA均可使棉花单铃重增加、纤维品质提高，外源ABA在晚播(5月25日，南京试点和安阳试点棉铃发育期MDTmin分别为20.9℃和16.5℃)条件下可使棉花产量(安阳试点)和纤维主要品质指标下降幅度减小；外源6-BA和ABA对棉籽主要品质性状的作用均不显著。(3)晚播条件下外源6-BA和ABA均可提高相对棉花主要品质指标，但两者作用机制不同，外源6-BA主要是提高棉铃对位叶光合产物含量和蔗糖转化率，而ABA则主要是诱导棉株抗逆性。
DOI:10.1104/pp.17.01232URLPMID:29097391 [本文引用: 1]
Flooding is detrimental for nearly all higher plants, including crops. The compound stress elicited by slow gas exchange and low light levels under water is responsible for both a carbon and an energy crisis ultimately leading to plant death. The endogenous concentrations of four gaseous compounds, oxygen, carbon dioxide, ethylene, and nitric oxide, change during the submergence of plant organs in water. These gases play a pivotal role in signal transduction cascades, leading to adaptive processes such as metabolic adjustments and anatomical features. Of these gases, ethylene is seen as the most consistent, pervasive, and reliable signal of early flooding stress, most likely in tight interaction with the other gases. The production of reactive oxygen species (ROS) in plant cells during flooding and directly after subsidence, during which the plant is confronted with high light and oxygen levels, is characteristic for this abiotic stress. Low, well-controlled levels of ROS are essential for adaptive signaling pathways, in interaction with the other gaseous flooding signals. On the other hand, excessive uncontrolled bursts of ROS can be highly damaging for plants. Therefore, a fine-tuned balance is important, with a major role for ROS production and scavenging. Our understanding of the temporal dynamics of the four gases and ROS is basal, whereas it is likely that they form a signature readout of prevailing flooding conditions and subsequent adaptive responses.
DOI:10.1093/jxb/ers148URL [本文引用: 1]
Here we highlight how both the root and shoot environment impact on whole plant hormone balance, particularly under stresses such as soil drying, and relate hormone ratios and relative abundances to processes influencing plant performance and yield under both mild and more severe stress. We discuss evidence (i) that abscisic acid (ABA) and ethylene act antagonistically on grain-filling rate amongst other yield-impacting processes; (ii) that ABA's effectiveness as an agent of stomatal closure can be modulated by coincident ethylene or cytokinin accumulation; and (iii) that enhanced cytokinin production can increase growth and yield by improving foliar stay-green indices under stress, and by improving processes that impact grain-filling and number, and that this can be the result of altered relative abundances of cytokinin and ABA (and other hormones). We describe evidence and novel processes whereby these phenomena are/could be amenable to manipulation through genetic and management routes, such that plant performance and yield can be improved. We explore the possibility that a range of ABA-ethylene and ABA-cytokinin relative abundances could represent targets for breeding/managing for yield resilience under a spectrum of stress levels between severe and mild, and could circumvent some of the pitfalls so far encountered in the massive research effort towards breeding for increases in the complex trait of yield.
DOI:10.3724/SP.J.1006.2019.84160URL [本文引用: 1]
淹水胁迫是作物生长发育过程中遭受的主要非生物胁迫之一, 探究提高绿豆耐淹性的机制对绿豆抗涝栽培具有重要意义。本文在2017—2018年以耐淹性不同的绿豆品种绿丰2号和绿丰5号为试验材料, 采用盆栽方式探究了烯效唑(S3307)对淹水胁迫下绿豆叶片生理、光合作用及产量的影响。结果表明, 在不同生育时期淹水胁迫下, 绿豆叶片的叶绿素含量(SPAD)及光合特性参数均显著下降, 丙二醛(MDA)含量显著增加, 始花期(R1期)淹水胁迫下绿豆的减产率为24.70%~33.63%, 始粒期(R5期)减产率为18.07%~28.87%。2个绿豆品种均表现为R1期受淹水胁迫危害程度大于R5期, 绿丰2号耐淹性强于绿丰5号。喷施S3307后显著提高淹水胁迫下绿豆叶片的SPAD、净光合速率(Pn)、蒸腾速率(Tr)和气孔导度(Gs), 并显著降低了MDA含量。绿豆在R1期淹水胁迫下的缓解率为28.91%~52.34%, R5期缓解率为13.77%~27.36%。表明叶面喷施S3307可有效提高淹水胁迫下绿豆叶片的生理功能及光合能力, 进而降低减产幅度, 但不同淹水时期和绿豆品种对S3307的调控响应存在差异。
DOI:10.3724/SP.J.1006.2019.84160URL [本文引用: 1]
淹水胁迫是作物生长发育过程中遭受的主要非生物胁迫之一, 探究提高绿豆耐淹性的机制对绿豆抗涝栽培具有重要意义。本文在2017—2018年以耐淹性不同的绿豆品种绿丰2号和绿丰5号为试验材料, 采用盆栽方式探究了烯效唑(S3307)对淹水胁迫下绿豆叶片生理、光合作用及产量的影响。结果表明, 在不同生育时期淹水胁迫下, 绿豆叶片的叶绿素含量(SPAD)及光合特性参数均显著下降, 丙二醛(MDA)含量显著增加, 始花期(R1期)淹水胁迫下绿豆的减产率为24.70%~33.63%, 始粒期(R5期)减产率为18.07%~28.87%。2个绿豆品种均表现为R1期受淹水胁迫危害程度大于R5期, 绿丰2号耐淹性强于绿丰5号。喷施S3307后显著提高淹水胁迫下绿豆叶片的SPAD、净光合速率(Pn)、蒸腾速率(Tr)和气孔导度(Gs), 并显著降低了MDA含量。绿豆在R1期淹水胁迫下的缓解率为28.91%~52.34%, R5期缓解率为13.77%~27.36%。表明叶面喷施S3307可有效提高淹水胁迫下绿豆叶片的生理功能及光合能力, 进而降低减产幅度, 但不同淹水时期和绿豆品种对S3307的调控响应存在差异。
DOI:10.1016/j.fcr.2012.09.006URL [本文引用: 1]
Waterlogging may be involved in the grain-filling disorder known as 'abnormal early ripening' (AER) in wheat (Triticum aestivum) in western Japan. This study carried out over two consecutive cropping seasons (2009/10 and 2010/11), examines whether or not wheat plants exposed to waterlogging at jointing exhibit early leaf senescence during grain filling. The effects of a single waterlogging at jointing (WLjoint) or after anthesis (WLanthe), and of a double waterlogging at the same two stages (WW) were compared in three Japanese wheat cultivars. Waterlogging for 21 days was imposed in all cases except for after anthesis in 2010/11 when the treatment was for only 11 days. In both cropping seasons, grain yield in the WLjoint plants was 10-15% lower than that in the well-drained plants. The lower yields were due to smaller grain weights, rather than to decreases in spike number or grain number per spike. The WLanthe treatment also reduced yield by 15%, although the duration of stress in the WLanthe plots was 10 days shorter than in the WLjoint, plots. This indicates that the WLanthe treatment had a more negative impact on yield. Plants in the WW plots exhibited reductions in grain yield of from 22 to 35% depending on the cultivar. The smaller grain weights of waterlogged plants was a result of both a 1-5 days shorter grain-filling period and also lower grain growth rates in the later stages of filling. Plants in the WLjoint and WW plots did not show reductions in leaf greenness, stomatal conductance or leaf water content during or after waterlogging at jointing. However, the plants in the WLjoint and WW plots,and also in the WLanthe plots, suffered reductions in these parameters about 14 days after anthesis. At the same time, the plants in the WW plots showed the most marked leaf senescence and the lowest leaf water contents (0.39 g g(-1) in the WW plots vs 0.85 g g(-1) in the well-drained plots). At maturity, concentrations of water-soluble carbohydrates (WSCs) in culms were highest in the WW plots, probably indicating that remobilization of the WSCs from the culms to the grains was reduced. These results indicate that root injury by waterlogging at jointing induces rapid leaf senescence in the grain filling period. Comparisons among the WLjoint, WLanthe and WW plots indicate that in wheat, root sensitivity to waterlogging rises markedly during the post-anthesis period. (c) 2012 Elsevier B.V.
DOI:10.1016/j.tplants.2019.02.007URLPMID:30857921 [本文引用: 1]
Increasing flooding events have detrimentally impacted food security amid a growing global population. Complete submergence of plants represents the most severe flooding stress and studies have identified underwater responses to low oxygen and light availability. However, knowledge on plant responses during the post-submergence phase is limited. It is important to consider how plants can resume vegetative growth after enduring submergence and post-submergence stress. This review highlights current knowledge on physiological and molecular adaptations following desubmergence. Interplays of reactive oxygen species (ROS), energy depletion, photoinhibition, desiccation stress, and hormonal signaling have been characterized as components of the post-submergence stress response. Active elucidation of key genes and traits enhancing post-submergence adaptations is highly relevant for the improvement of submergence tolerance and ultimately crop yield.
URL [本文引用: 1]
采用雾培植株根际通CO2处理方式，研究了开花结果期根际CO2浓度升高对网纹甜瓜光合作用及产量和品质的影响.结果表明：在网纹甜瓜果实发育期间，与350 μL·L-1（对照）处理相比，根际2500和5000 μL CO2·L-1处理的叶片光合色素含量、净光合速率（Pn）、气孔导度（gs）、胞间CO2浓度（Ci）及PSⅡ最大光化学效率（Fv/Fm）均不同程度降低，而气孔限制值（Ls）显著提高，且5000 μL CO2·L-1处理的变化幅度高于2500 μL CO2·L-1处理；单株产量、果实中维生素C和可溶性糖含量显著降低，有机酸含量显著提高.可见，网纹甜瓜果实发育期间根际CO2浓度超过2500 μL·L-1时，其光合作用及果实发育会受到显著抑制，从而导致产量和品质降低.
URL [本文引用: 1]
采用雾培植株根际通CO2处理方式，研究了开花结果期根际CO2浓度升高对网纹甜瓜光合作用及产量和品质的影响.结果表明：在网纹甜瓜果实发育期间，与350 μL·L-1（对照）处理相比，根际2500和5000 μL CO2·L-1处理的叶片光合色素含量、净光合速率（Pn）、气孔导度（gs）、胞间CO2浓度（Ci）及PSⅡ最大光化学效率（Fv/Fm）均不同程度降低，而气孔限制值（Ls）显著提高，且5000 μL CO2·L-1处理的变化幅度高于2500 μL CO2·L-1处理；单株产量、果实中维生素C和可溶性糖含量显著降低，有机酸含量显著提高.可见，网纹甜瓜果实发育期间根际CO2浓度超过2500 μL·L-1时，其光合作用及果实发育会受到显著抑制，从而导致产量和品质降低.
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