花铃期渍水对棉铃对位叶蔗糖代谢及铃重的影响

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杨长琴, 刘瑞显, 张国伟, 徐立华, 周治国. 花铃期渍水对棉铃对位叶蔗糖代谢及铃重的影响. 作物学报, 2014, 40(5): 908-914[YANG Chang-Qin, LIU Rui-Xian, ZHANG Guo-Wei, XU Li-Hua, ZHOU Zhi-Guo. Effects of Waterlogging on Sucrose Metabolism of the Subtending Leaf of Cotton Boll and Boll Weight during Flowering and Boll-forming Stage. 作物学报, 2014, 40(5): 908-914] 复制到剪切板

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花铃期渍水对棉铃对位叶蔗糖代谢及铃重的影响

杨长琴^1,², 刘瑞显², 张国伟², 徐立华², 周治国^1,^*

¹ 南京农业大学农学院 / 农业部南方作物生理生态重点开放实验室, 江苏南京210095

²江苏省农业科学院经济作物研究所, 江苏南京210014

^*通讯作者(Corresponding author): 周治国, E-mail:giscott@njau.edu.cn, Tel: 025-84396813

第一作者联系方式: E-mail:ychq2003@qq.com

收稿日期:2014-01-12

基金:本研究由国家现代农业产业技术体系建设专项(CARS-18-20)和国家公益性行业(农业)科研专项资金项目(200903003)资助;

摘要

试验以美棉33B为材料, 于2007年和2009年在江苏南京(32º02′N, 118º50′E)江苏省农业科学院防雨棚的水泥池中进行。以正常灌水(土壤相对含水量为75%±5%)为对照, 于棉花中部(6~8)果枝开花时进行渍水7 d和14 d (保持地面2~3 cm的水层)处理。研究花铃期渍水对棉铃对位叶蔗糖代谢及铃重形成的影响。结果表明, 花铃期渍水条件下棉铃对位叶可溶性糖、蔗糖、淀粉含量增加, 蔗糖/淀粉比值下降, 且随渍水持续期的延长变化幅度增大, 表明花铃期渍水致使叶片蔗糖外运受阻且随渍水时间的延长外运受阻加重; 棉铃对位叶蔗糖磷酸合酶、蔗糖合酶和酸性转化酶活性增强, 且随渍水持续期的延长和果枝部位的上升而增大; 棉铃对位叶可溶性糖、蔗糖和淀粉含量与蔗糖磷酸合酶、蔗糖合酶及酸性转化酶活性显著正相关(P<0.05), 表明渍水条件下蔗糖代谢被激活但并未促进叶片蔗糖的外运, 铃重随果枝部位上升和渍水持续期的延长下降幅度增大。这是渍水条件下铃重下降的重要原因。

关键词: 棉花(Gossypium hirsutum L.); 花铃期; 渍水; 叶片蔗糖代谢; 铃重

Effects of Waterlogging on Sucrose Metabolism of the Subtending Leaf of Cotton Boll and Boll Weight during Flowering and Boll-forming Stage

YANG Chang-Qin^1,², LIU Rui-Xian², ZHANG Guo-Wei², XU Li-Hua², ZHOU Zhi-Guo^1,^*

¹Key Laboratory of Crop Physiology & Ecology in Southern China, Ministry of Agriculture / College of Agriculture, Nanjing Agricultural University, Nanjing 210095, China

²Institute of Industrial Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

Fund:

Abstract

Two year experiments were conducted at experimental station of Jiangsu Academy of Agricultural Sciences of Nanjing City with waterlogging treatments for seven days and fourteen days in pools with rainproof shelter at anthesis of the 6th or 8th fruiting branches of cotton NuCOTN 33B, taking well water conditions as control. The results showed that contents of soluble sugar, sucrose and starch of subtending leaves of bolls increased and the ratio of sucrose content to starch content decreased under waterlogging as compared with control and the increase ranges rised with lasting waterlogging duration, indicating that the sucrose exportation of subtending leaves was inhibited and became worse with lasting waterlogging duration. The activities of sucrose phosphate synthase (SPS), sucrose synthase (SS), and acid invertase (AI) increased and got higher with the rise in fruit branch position and waterlogging duration. The contents of soluble sugar, sucrose and starch were positively and significantly correlated with activities of SPS, SS, and AI (P<0.05), indicating that the increased activities of SPS, SS, and AI do not facilitate sucrose transiting out. As a result, the boll weight declined under waterlogging with the rise in fruit branch position and waterlogging duration. Therefore, the increased enzyme activity in sucrose metabolism and the impaired sucrose efflux are the main causes of boll weight decrease under waterlogging.

Keyword: Cotton (Gossypium hirsutum L.); Flowering and boll-forming stage; Waterlogging; Leaves sucrose metabolism; Boll weight

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受季节性降雨和灾害性天气的影响, 在棉花生长季节时有渍水发生, 严重影响棉花的生长及产量。Range等^{[ 1]}和郭文琦等^{[ 2, 3]}曾报道渍水对棉花生长影响主要集中在光合性能、干物质累积、产量及根系生长等方面。对其他植物的研究发现, 碳水化合物在植物响应逆境胁迫的过程中具有重要意义, 尤其非结构性碳水化合物(如蔗糖和淀粉)的代谢在很大程度上影响着植株的生长以及对环境的响应^{[ 4, 5, 6]}。棉铃发育所需养分主要来源于其棉铃对位叶的光合作用, 而蔗糖是光合作用的主要产物和运输形式, 因此, 开展花铃期渍水对棉铃对位叶蔗糖代谢生理的研究具重要意义。

Malik等^{[ 7]}、Chen等^{[ 8]}和Liao等^{[ 9]}研究表明, 淹水胁迫下大多数不耐淹植物光合产物运输到根和竞争性库器官的量减少, 导致叶片中可溶性糖和淀粉积累, 也有研究认为植物在渍水条件下由于缺氧代谢获取的能量效率较低, 必须消耗大量的碳水化合物, 叶片非结构性糖含量降低甚至糖分迅速耗尽^{[ 10, 11]}。造成上述分歧的原因与所用材料和处理方式等有关, 但不同研究的渍水条件下叶片碳水化合物含量的变化最终均抑制植物光合和生长。水分胁迫对植物蔗糖代谢的影响是通过影响蔗糖代谢相关酶的活性来实现的。一般认为, 植物叶片中蔗糖磷酸合酶是蔗糖合成的关键限速酶^{[ 12]}, 其活性与蔗糖含量密切相关^{[ 13, 14]}; 蔗糖合酶既可催化蔗糖合成又可催化蔗糖分解, 植物光合组织中的蔗糖合酶催化蔗糖合成的活性较高^{[ 15, 16]}; 叶片中酸性转化酶是蔗糖降解主要酶^{[ 17]}。叶片中蔗糖磷酸合酶、蔗糖合酶和酸性转化酶协调作用, 有利于形成蔗糖浓度梯度, 加快蔗糖从韧皮部筛管“卸载”到库器官。

棉铃重是棉花产量构成的重要因子。棉铃对位叶作为棉铃养分的主要供应者, 其生理特性直接影响棉铃发育所需养分的合成和转运^{[ 18]}。张文静等^{[ 19]}研究表明棉铃对位叶的生理特性存在基因型差异且与铃重的形成关系密切, 棉铃对位叶C/N适宜、内源保护酶活性高有利于铃重提高^{[ 20]}, 最近研究发现, 外源6-BA显著提高棉铃对位叶蔗糖含量和转化率及淀粉含量, 促进光合产物向棉铃的运输, 从而提高铃重^{[ 21]}。但未见有关花铃期渍水如何影响棉铃对位叶蔗糖代谢及铃重的报道。本试验在池栽条件下研究花铃期渍水对棉花不同果枝部位叶片碳水化合物动态变化、蔗糖代谢及其对铃重的影响, 以期明确花铃期渍水影响棉花铃重的叶片蔗糖代谢生理。

1 材料与方法

1.1 试验设计

试验于2007年和2009年在江苏南京(32°02′N、118°50′E, 长江流域下游棉区)江苏省农业科学院防雨棚水泥池进行, 供试土壤为黄棕壤土。2007年和2009年土壤分别含有机质15.3 g kg^-1和14.3 g kg^-1, 全氮0.8 g kg^-1和0.9 g kg^-1, 碱解氮63.4 mg kg^-1和52.6 mg kg^-1, 速效磷29.9 mg kg^-1和23.8 mg kg^-1, 速效钾107.5 mg kg^-1和93.2 mg kg^-1。供试棉花品种为美棉33B。2007年于4月17日播种, 5月17日移栽; 2009年于4月25日播种, 5月22日移栽。两年种植密度分别为每公顷3.6万株和3.9万株, 小区面积4.5 m × 3.3 m, 4个重复。施纯氮225 kg hm^-2、过磷酸钙750 kg hm^-2、氯化钾225 kg hm^-2, 基肥和初花肥各占50%。其他措施按高产要求进行。

于中部6~8台果枝进入开花期(2007年7月18日; 2009年7月24日)开始水分处理。2007年设2个处理, 即对照和渍水14 d。对照全生育期土壤相对含水量(土壤相对含水量=土壤含水量/田间持水量)保持在(75±5)%; 渍水14 d处理保持地面2~3 cm的水层, 持续时间为14 d, 渍水结束后土壤相对含水量自然减少至(75±5)%。2009年设3个处理, 即对照、渍水7 d和渍水14 d, 处理方式同2007年。

1.2 测定内容与方法

1.2.1 碳水化合物含量与酶活性试验于棉株2~4 (LFB)、6~8 (MFB)、10~12 (UFB)果枝第一、二果节开花时, 挂牌标记当日所开白花。从花后10 d开始每7 d取生长发育一致棉铃8~10个及其对位叶片(每天上午9:00~10:00取样, 棉铃作其他测定), 一部分叶片用液氮速冻并在-30℃保存测定酶活性; 另一部分叶片105℃杀青、80℃烘干, 粉碎过筛后作碳水化合物含量测定。用蒽酮比色法^{[ 22]}测定可溶性总糖和淀粉含量; 用间苯二酚比色法^{[ 22]}测定蔗糖含量。用果糖和UDPG比色法^{[ 23]}测定蔗糖磷酸合酶和蔗糖合酶(合成方向)活性; 用DNS显色法^{[ 22]}测定酸性转化酶活性。

1.2.2 铃重吐絮期按果枝部位收标记的正常成熟棉铃20个, 晒干后称重。

2 结果与分析

2.1 铃重变化

由表1可见, 花铃期渍水条件下不同果枝部位棉铃铃重显著降低。渍水7 d处理下部、中部和上部果枝铃重分别下降7.10%、7.99%和19.55% (2009年); 渍水14 d处理则分别下降13.80%、14.06%和27.81% (2007年)和13.33%、12.80%和29.03% (2009

表1 花铃期渍水对棉花铃重的影响(2007年和2009年) Table 1 Effects of waterlogging on boll weight during flowering and boll-forming stage in 2007 and 2009 (g boll^-1)

年)。表明不同果枝部位间, 花铃期渍水对铃重的影响上部大于下部和中部; 渍水处理间随渍水持续期的延长铃重下降幅度增大。

2.2 棉铃对位叶碳水化合物含量

2.2.1 可溶性糖含量

图1表明, 花铃期渍水条件下棉铃对位叶可溶性糖含量大幅度增加, 其峰值出现时期随果枝部位上升提前。2009年渍水7 d处理下部、中部和上部果枝棉铃对位叶可溶性糖含量峰值分别出现在花后31、31和17 d; 而渍水14 d处理分别出现在花后31、31和24 d。渍水7 d处理的下部和中部果枝分别于花后24 d和17 d可溶性糖含量较高, 而这正是水分处理结束后1周时间, 表明棉铃对位叶可溶性糖含量变化是响应土壤水分变化的。其后不同果枝部位以渍水14 d处理棉铃对位叶可溶性糖含量较高, 表明随渍水持续时间的延长, 棉铃对位叶增加可溶性糖含量以适应土壤水分变化。花后38 d渍水14 d处理不同果枝部位棉铃对位叶可溶性糖含量均降低, 表明叶片中可利用性糖含量降低。

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图1 花铃期渍水对棉铃对位叶碳水化合物含量的影响(2007年和2009年)LFB、MFB和UFB分别代表下部、中部和上部果枝。Fig. 1 Effects of waterlogging on carbohydrate contents of subtending leaves of bolls during flowering and boll-forming
stage in 2007 and 2009LFB, MFB, and UFB stand for low fruiting branch, middle fruiting branch, and upper fruiting branch, respectively.

2.2.2 蔗糖含量

与可溶性糖相似(图1), 蔗糖峰值期含量以渍水14 d处理较高。花后38 d渍水14 d处理中、上部果枝棉铃对位叶蔗糖含量降低, 表明叶片可供外运蔗糖减少。

2.2.3 淀粉含量

由图1可见, 花铃期渍水条件下棉铃对位叶淀粉含量大幅度高于对照, 且中部和上部果枝增加幅度大于下部果枝, 淀粉含量峰值出现的时间随果枝部位上升提前, 如2009年渍水7 d处理下部、中部和上部果枝棉铃对位叶淀粉含量峰值分别出现在花后31、24和17 d; 而渍水14 d处理分别出现在花后24、24和17 d。渍水处理间相比, 中部和上部果枝棉铃对位叶淀粉含量以渍水14 d处理较高, 尤其在花后后期仍保持较高水平。表明随渍水持续期的延长, 渍水对棉铃对位叶蔗糖外运抑制增强。

2.3 棉铃对位叶蔗糖代谢相关酶活性

2.3.1 蔗糖磷酸合酶活性图2表明, 花铃期渍水条件下棉铃对位叶蔗糖磷酸合酶的活性增加, 随着果枝部位的上升, 渍水处理与对照棉铃对位叶蔗糖磷酸合酶活性差异变大, 且其活性峰值出现时期提前。2009年渍水处理下部、中部和上部果枝棉铃对位叶蔗糖磷酸合酶活性峰值分别出现在花后31、24和17 d。渍水处理间相比, 以渍水14 d处理棉铃对位叶蔗糖磷酸合酶活性较高。表明随渍水时间的延长, 棉铃对位叶通过蔗糖磷酸合酶活性的提高以适应土壤水分变化。

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图2 花铃期渍水对棉铃对位叶蔗糖代谢相关酶活性的影响(2007年和2009年)LFB、MFB和UFB分别代表下部、中部和上部果枝。Fig. 2 Effects of waterlogging on sucrose metabolism enzyme activity of subtending leaves of bolls during flowering and boll-forming stage in 2007 and 2009LFB, MFB, and UFB stand for low fruiting branch, middle fruiting branch, and upper fruiting branch, respectively.

2.3.2 蔗糖合酶活性图2表明, 花铃期渍水条件下棉铃对位叶蔗糖合酶活性动态与蔗糖磷酸合酶活性动态相似, 渍水处理间相比, 以渍水14 d处理的酶活性较高。

2.3.3 酸性转化酶活性图2表明, 渍水7 d处理花后17 d后棉铃对位叶酸性转化酶活性动态与对照相似但高于对照。渍水14 d处理下部和中部果枝花后10 d棉铃对位叶酸性转化酶活性略低, 其后大幅上升, 因而有利于花后初期叶片蔗糖外运但后期可供外运蔗糖减少。随花后天数增加和果枝部位上升, 渍水14 d处理酸性转化酶活性与对照及渍水7 d处理间差异变大, 如2009年渍水14 d处理下部、中部和上部果枝花后31 d棉铃对位叶酸性转化酶活性分别是对照的1.43、1.59和2.94倍; 是渍水7 d处理的1.23、1.37和1.91倍。表明渍水时间延长对棉铃对位叶酸性转化酶活性影响加大, 影响叶片可供外运的蔗糖水平。

2.4 蔗糖代谢相关物质和酶活的关系

花铃期渍水条件下棉铃对位叶中碳水化合物含量、蔗糖代谢酶活性均大幅度增加。相关分析发现(表2), 对照棉铃对位叶可溶性糖、蔗糖和淀粉含量与蔗糖磷酸合酶呈极显著正相关( P<0.01)、与蔗糖合酶活性呈正相关但未达显著水平、与酸性转化酶活性则呈显著负相关( P<0.05); 渍水处理未改变棉铃对位叶可溶性糖、蔗糖和淀粉含量与蔗糖磷酸合酶、蔗糖合酶的关系, 且与蔗糖合酶活性的相关达极显著水平( P<0.01), 但棉铃对位叶蔗糖、可溶性糖和淀粉含量与酸性转化酶活性呈显著正相关( P<0.05)。表明对照棉铃对位叶酸性转化酶活性的增加促进了蔗糖的降解外运, 而渍水处理棉铃对位叶酸性转化酶活性的增加促进了蔗糖降解的同时叶片中碳水化合物含量并未降低, 即酸性转化酶活性的增加并未促进叶片蔗糖外运。

表2 棉铃对位叶碳水化合物含量与蔗糖代谢相关酶活性相关关系 Table 2 Relationship between carbohydrate contents and enzyme activities related to sucrose metabolism of subtending
leaves of bolls

3 讨论

棉铃对位叶碳水化合物是光合作用的产物, 蔗糖则是光合作用的主要产物, 也是向库器官长距离运输的主要形式。叶片中蔗糖的水平取决于光合速率、蔗糖代谢和蔗糖外运的协调作用^{[ 24]}。郭文琦等^{[ 25]}盆栽试验结果表明, 渍水条件下棉株光合速率下降, 光合产物减少, 致使铃数减少、铃重降低。但有关棉铃对位叶-棉铃蔗糖代谢及运转的生理机制缺乏研究。本文研究发现, 花铃期渍水条件下棉铃对位叶可溶性糖、蔗糖和淀粉含量均大幅度增加, 与小麦^{[ 7]}、宽叶独行菜( Lepidium latifolium)^{[ 8]}等大多数不耐渍水物种具有相同的特征。利容千等^{[ 11]}认为, 由于植物在渍水条件下缺氧代谢获取能量的效率较低, 必须消耗大量的碳水化合物, 因此, 渍水条件下叶片碳水化合物的累积可能是其耐渍水环境的重要因素^{[ 4]}。Albrecht等^{[ 26]}和Islam等^{[ 27]}研究还表明, 渍水条件下叶片碳水化合累积是碳水化合物向库器官和根系运输受阻的结果, 因此, 棉铃对位叶碳水化合物累积不利于库器官-棉铃的发育。叶片蔗糖代谢变化对棉铃发育的影响一方面致使铃重降低, 就不同果枝部位而言, 渍水处理棉铃对位叶碳水化合物增加的幅度以中部和上部果枝大于下部果枝、且随渍水持续期延长而增加, 而铃重随果枝部位上升和渍水持续期的延长降幅增大; 另一方面致使棉铃脱落、铃数减少, 如2009年对照下部、中部和上部果枝成铃数分别为每株6.0、4.7和2.7个, 渍水7 d不同果枝部位成铃数分别降低了12.3%、42.4%和34.0%、渍水14 d分别降低了17.4%、35.2%和65.4%, 随果枝部位上升, 铃数降幅增大。

叶片碳水化合物水平受蔗糖代谢相关酶的调节。Yang等^{[ 28]}和王维等^{[ 29]}研究表明, 轻度、适度的土壤水分胁迫可诱导水稻茎部蔗糖磷酸合酶活性升高而酸性转化酶下降, 利于贮藏碳水化合物的输出, Hirano等^{[ 30]}研究发现淹水下水稻上层叶片的蔗糖磷酸合酶活性被高度激活, Harada等^{[ 31]}发现厌氧条件下水池草芽蔗糖磷酸合酶、蔗糖合酶和酸性转化酶活性均增加。本试验中渍水处理棉铃对位叶蔗糖磷酸合酶和蔗糖合酶活性大幅度增加, 酸性转化酶活性在花后10 d低, 其后也大幅度增加。表明渍水条件下棉铃对位叶蔗糖代谢较活跃, 蔗糖合成和降解代谢同时进行^{[ 32]}。相关分析发现, 渍水处理棉铃对位叶可溶性糖、蔗糖和淀粉含量与蔗糖磷酸合酶、蔗糖合酶及酸性转化酶活性均呈正相关( P<0.05)。渍水逆境导致植物同化力下降和同化作用受阻, 植物蔗糖代谢改变, 而植物在渍水条件下的生存能力取决于能源物质(糖)的水平^{[ 33]}, 且植物体内的碳水化合物首先用于维持呼吸和生存能量需求^{[ 34]}, 因而推测较高的酶活性与维持植株在渍水下生存的能量需求有关^{[ 34]}, 而一旦碳水化合物含量降低甚至耗尽, 植物生长将很慢或难以恢复^{[ 35, 36, 37]}。因此棉铃对位叶酶活性增强, 用于维持生存的基本代谢消耗增加, 而减少了棉铃对位叶蔗糖外运, 随果枝部位的上升和渍水持续期延长酶活性增加的幅度增大, 是渍水条件下铃重随果枝部位上升和渍水持续期延长降低的生理原因。本文研究还发现, 渍水7 d处理下部、上部和渍水14 d处理下部果枝棉铃对位叶蔗糖磷酸合酶、蔗糖合酶活性峰值时间与可溶性糖、蔗糖和淀粉含量峰值时间一致, 表明棉铃对位叶碳水化合物含量对酶活性的响应较快; 而渍水7 d处理中部和渍水14 d处理中部、上部果枝棉铃对位叶可溶性糖和蔗糖含量峰值迟于蔗糖代谢相关酶活性和淀粉含量的峰值时间, 表明随果枝部位上升和渍水持续期的延长, 棉铃对位叶可溶性糖和蔗糖含量对酶活性调节响应变慢, 且在糖含量峰值前碳水化合物外运已受很大影响。与不同果枝部位受渍害程度、渍水起始时作用的果枝部位及渍水持续期有关。综上, 渍水下叶片蔗糖代谢激活生成的碳水化合物首先用于维持生存的能量需求, 蔗糖外运受阻。蔗糖有效利用是影响棉铃发育的重要原因。因此, 生产中渍水条件下棉花铃数和铃重的提高可从植株生长恢复和促进蔗糖的有效利用入手, 研究结果为渍水条件下制定调节措施提高棉花产量提供了理论依据。

4 结论

花铃期渍水条件下, 棉铃对位叶光合产物减少, 蔗糖代谢被激活, 碳水化合物含量大幅度增加, 维持生存需要蔗糖含量增加而外运减少, 随果枝部位的上升和渍水持续期的延长蔗糖外运受阻加剧, 最终铃重随果枝部位上升和渍水持续期延长降幅增大。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

Reference

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2004

2.474

0.0

... Range等^[1]和郭文琦等^[2,3]曾报道渍水对棉花生长影响主要集中在光合性能、干物质累积、产量及根系生长等方面 ...

2009

0.0

1.667

郭文琦, 赵新华, 陈兵林, 刘瑞显, 周治国. 氮素对花铃期短期渍水棉花根系生长的影响. 作物学报, 2009, 35: 1078-1085

Guo

W Q

, Zhao

X H

, Chen

B L

, Liu

R X

, Zhou

Z G

. Effects of nitrogen on cotton (Gossypium hirsutum L. ) root growth under short-term waterlogging during flowering and boll-forming stage. Acta Agron Sin, 2009, 35: 1078-1085 (in Chinese with English abstract)

The flowering and boll development stage is the key period for cotton quality and yield. Waterlogging during this period significantly inhibits cotton development and reduces final yield. Nitrogen fertilizer is considered to be an effective up-regulatory element for crop growth. The objective of this investigation was to evaluate effects of nitrogen on cotton resistance to soil waterlogging in terms of the changes of root dry matter accumulation, antioxidant enzyme activities and root vigor under short-term waterlogging. A pot experiment wasconducted with three N levels (0, 3.73, and 7.46 g N pot^-1 equivalent to 0, 240, and 480 kg N ha^-1, respectively), waterlogging for eight days and then recovering for fifteen days. The results showed that the root dry matter weight and the Root/Shoot ratio of plants by waterlogged decreased, and the root dry matter weight reached a peak at the 240 kg N ha^-1under waterlogging, but the root/shoot ratio decreased with the increase of N application. Under waterlogging, soluble protein content significantly decreased in comparison with well-watered cotton, and the reduced degree increased with increasing N application. Malondialdehyde (MDA) content in cotton roots was significantly (P <0.05) increased, with the lowest at the 240 kg N ha^-1 during waterlogging. Application of N increased the activities of peroxidase (POD) and catalase (CAT) of cotton roots, but decreased superoxide dismutase (SOD) activity during waterlogging. Both root vigor and CAP decreased by waterlogging, were the highest at the 240 kg N ha^-1. At the 15 d after terminating waterlogging, N application promoted root vigor and CAP, but decreased MDA content. These results suggest that appropriate N supply (240 kg N ha^-1in this investigation) may contribute to waterlogging resistance of cotton plants through adjusting the antioxidant enzyme activities of roots, decreasing lipid peroxidation and enhancing root vigor during waterlogging (waterlogging for eight days in this investigation), excessive N supply (480 kg N ha^-1) has a deleterious effect on plant waterlogging resistance, however, more N should be supplied to waterlogged cotton pants after terminating waterlogging.

Key Laboratory of Crop Physiology and Ecology in Southern China,Ministry of Agriculture/Nanjing Agricultural University,Nanjing 210095,China

于2005—2006年在江苏南京农业大学卫岗试验站进行盆栽试验，设置正常灌水(土壤含水量为田间持水量的75%左右)和棉花花铃期土壤短期渍水处理(将正常灌水的棉花增加灌水至盆内有可见明水，持续8 d，然后用导管排除表面水层，使盆内土壤含水量逐渐恢复到田间持水量的75%左右)，每个水分处理设置3个氮素水平(0、3.73、7.46 g N pot^-1，分别相当于大田0、240、480 kg N hm^-2)，研究氮素对花铃期短期渍水棉花根系生长的影响。结果表明，在渍水处理结束时，与正常灌水处理相比，根干重和根冠比(R∶S)均降低；根系可溶性蛋白含量、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性降低，过氧化物酶(POD)活性升高，丙二醛(MDA)含量升高；棉花根系活力和单株光合速率(CAP)显著降低。增加施氮可降低渍水棉花根系SOD活性，提高POD和CAT活性，以3.73 g N pot^-1(240 kg N hm^-2)施氮水平下的棉花根干物重最大，根系MDA含量最低，根系活力最强，单株光合速率(CAP)最高，相应籽棉产量最高。渍水停止15 d后，渍水棉花根系抗氧化酶活性和MDA含量与正常灌水处理的差异较小；施氮仍可提高棉花根系POD与CAT活性，降低MDA含量，增强根系活力，提高CAP。

... Range等^[1]和郭文琦等^[2,3]曾报道渍水对棉花生长影响主要集中在光合性能、干物质累积、产量及根系生长等方面 ...

2008

0.0

... Range等^[1]和郭文琦等^[2,3]曾报道渍水对棉花生长影响主要集中在光合性能、干物质累积、产量及根系生长等方面 ...

2010

0.0

... 对其他植物的研究发现, 碳水化合物在植物响应逆境胁迫的过程中具有重要意义, 尤其非结构性碳水化合物(如蔗糖和淀粉)的代谢在很大程度上影响着植株的生长以及对环境的响应^[4,5,6] ...

... 利容千等^[11]认为, 由于植物在渍水条件下缺氧代谢获取能量的效率较低, 必须消耗大量的碳水化合物, 因此, 渍水条件下叶片碳水化合物的累积可能是其耐渍水环境的重要因素^[4] ...

2010

0.0

1.989

Shi

M F

, Zeng

, Shen

J H

, Lei

S T

, Zhu

, Liu

J H

. A review of the correlation of flooding adaptability and carbohydrates in plants. Chin J Plant Ecol, 2010, 34: 855-866 (in Chinese with English abstract)

Flooding causes substantial stress for terrestrial plants, particularly if the floodwater completely submerges the shoot. The main problems during submergence are shortage of oxygen due to slow diffusion rates of gases in water and unfavorable conditions of light and carbon dioxide supply. Collectively, these factors lead to loss of biomass and eventually death of the submerged plants. The survival of plants is closely related to carbohydrates,
the substrate for respiration. There are two major aspects of the correlation of plants flooding adaptability and carbohydrates. First, there are morphological and physiological responses of terrestrial plant species that enable the positive effects of carbohydrates on underwater plant performance: plants usually elongate or reduce underwater elongation and maintain a higher level of root carbohydrates that facilitates survival. Second, plants change the expression of hormone, enzyme and gene, adjusting carbohydrate metabolism to flooding. The research advances in concentration of carbohydrates upon morphological and physiological changes and hormone, enzyme and gene expression to submergence are summarized.

<EM>Key Laboratory of Eco-environments in Three Gorges Reservoir Region (Ministry of Education), Chongqing Key Laboratory of Plant Ecology and Resources Research in Three Gorges Reservoir Region , School of Life Science, Southwest University, Chongqing 400715, China</EM>

水淹会对陆生植物存活造成本质影响, 特别是完全水淹对陆生植物的影响更为明显。水淹对陆生植物最为主要的影响是氧气不足, 这主要是由氧气在水中的扩散速率较低引起的。同时, 在水淹胁迫下植物对光和CO₂的获取都会受到限制。所有这些因素都将引起植物生物量减少, 最终导致受淹植物死亡。碳水化合物是植物的能量来源, 与植物在水淹胁迫下存活与否有着密切联系。植物水淹适应性与碳水化合物的相关性主要体现在两大方面: 在生理形态层面, 植物通过伸长生长或抑制伸长生长、地上和地下部分碳水化合物的分配比例不同来应对水淹胁迫; 在另一个层面, 植物通过改变激素、酶和基因的表达, 调整碳水化合物的代谢方式, 从而适应水淹环境。该文结合国内外研究现状, 通过对植物在水淹胁迫下生理形态、激素、酶及基因表达诸方面的变化来认识水淹耐受性与碳水化合物的关系, 并就今后的研究方向提出几点建议。

1996

0.0

Koch

K E

. Carbohydrate-modulated gene expression in plants. Annu Rev Plant Physoil Mol Biol, 1996, 47: 509-540

Plant Molecular and Cellular Biology Program, Horticultural Sciences Department, University of Florida, Gainesville, Florida 32611.

Plant gene responses to changing carbohydrate status can vary markedly. Some genes are induced, some are repressed, and others are minimally affected. As in microorganisms, sugar-sensitive plant genes are part of an ancient system of cellular adjustment to critical nutrient availability. However, in multicellular plants, sugar-regulated expression also provides a mechanism for control of resource distribution among tissues and organs. Carbohydrate depletion upregulates genes for photosynthesis, remobilization, and export, while decreasing mRNAs for storage and utilization. Abundant sugar levels exert opposite effects through a combination of gene repression and induction. Long-term changes in metabolic activity, resource partitioning, and plant form result. Sensitivity of carbohydrate-responsive gene expression to environmental and developmental signals further enhances its potential to aid acclimation. The review addresses the above from molecular to whole-plant levels and considers emerging models for sensing and transducing carbohydrate signals to responsive genes.

PMID: 15012299

2000

6.736

0.0

... Malik等^[7]、Chen等^[8]和Liao等^[9]研究表明, 淹水胁迫下大多数不耐淹植物光合产物运输到根和竞争性库器官的量减少, 导致叶片中可溶性糖和淀粉积累, 也有研究认为植物在渍水条件下由于缺氧代谢获取的能量效率较低, 必须消耗大量的碳水化合物, 叶片非结构性糖含量降低甚至糖分迅速耗尽^[10,11] ...

... 本文研究发现, 花铃期渍水条件下棉铃对位叶可溶性糖、蔗糖和淀粉含量均大幅度增加, 与小麦^[7]、宽叶独行菜(Lepidium latifolium)^[8]等大多数不耐渍水物种具有相同的特征 ...

2002

6.736

0.0

2005

1.593

0.0

2001

0.0

Liao

C T

, Lin

C H

. Physiological adaptation of crop plants to flooding stress. Proc Natl Sci Counc Roc (B), 2001, 25: 148-157

Taichung District Agricultural Improvement Station, Changhua, Taiwan, ROC.

When crop plants are subjected to soil waterlogging, or an anaerobic condition, their root and shoot systems respond differently. A variety of morphological and anatomical alterations develop in the root system. Reduction of the root respiration rate has been reported in both flooding-tolerant and intolerant species. Besides alcoholic fermentation, several diverse fermentative bypasses take place, which ameliorate the poisoning through excessive accumulation of specific metabolic intermediates. Root systems starved of oxygen are also poor providers of mineral nutrients for both themselves and the shoot systems. Stomatal closure and non-stomatal metabolic alterations are responsible for the reduction of leaf CO2 incorporation. Plant hormones are much involved in regulation of these physiological adaptations.

PMID: 11480770

2003

2.32

0.0

Singla

N K

, Jain

, Sawhney

S K

. Activities of glycolytic enzymes in leaves and roots of contrasting cultivars of sorghum during flooding. Biol Plant, 2003, 47: 555-560

Activities of phosphofructokinase (PFK), fructose-1,6-bisphosphate aldolase (FBP aldolase) and pyruvate kinase (PK) increased progressively in the roots of flood-tolerant SSG-59-3 cultivar during flooding. In contrast, only a slight change in activities of PFK and FBP aldolase was discerned in the roots of flood-sensitive S-308 cultivar during initial stages of flooding followed by a decline in the activities of these enzymes. Although the activity of hexokinase (HK) was transiently elevated in roots of both the cultivars during flooding, the magnitude of increase was much more in SSG-59-3 than in the S-308. In leaves of SSG-59-3, HK activity increased during 12 h of flooding whereas only a minor change occurred in the case of S-308. Flooding resulted in depressed activities of PFK and PK in leaves of S-308 but that in SSG-59-3 rose following imposition of waterlogged conditions. Activity of FBP aldolase in leaves of tolerant cultivar also showed a steady enhancement during flooding. The total and reducing sugars content decreased in leaves and roots of the S-308 during flooding but in SSG-59-3 the amount was more or less comparable to that in corresponding non-flooded plants.

1.CCS Haryana Agricultural University<br/>

2002

0.0

... 一般认为, 植物叶片中蔗糖磷酸合酶是蔗糖合成的关键限速酶^[12], 其活性与蔗糖含量密切相关^[13,14] ...

2005

3.347

0.0

Chen

, Hajirezaei

, Peisker

, Tschiersch

, Sonnewald

, Börnke

. Decreased sucrose-6-phosphate phosphatase level in transgenic tobacco inhibits photosynthesis, alters carbohydrate partitioning, and reduces growth. Planta, 2005, 221: 479-492

The aim of this work was to examine the role of sucrose-6-phosphate phosphatase (SPP; EC 3.1.3.24) in photosynthetic carbon partitioning. SPP catalyzes the final step in the pathway of sucrose synthesis; however, until now the importance of this enzyme in plants has not been studied by reversed-genetics approaches. With the intention of conducting such a study, transgenic tobacco plants with reduced SPP levels were produced using an RNA interference (RNAi) strategy. Transformants with less than 10% of wild-type SPP activity displayed a range of phenotypes, including those that showed inhibition of photosynthesis, chlorosis, and reduced growth rates. These plants had strongly reduced levels of sucrose and hexoses but contained 3–5 times more starch than the control specimens. The leaves were unable to export transient starch during extended periods of darkness and as consequence showed a starch- and maltose-excess phenotype. This indicates that no alternative mechanism for carbon export was activated. Inhibition of SPP resulted in an approximately 1,000-fold higher accumulation of sucrose-6-phosphate (Suc6P) compared to wild-type leaves, whereas the content of hexose-phosphates was reduced. Although the massive accumulation of Suc6P in the cytosol of transgenic leaves was assumed to impair phosphate-recycling into the chloroplast, no obvious signs of phosphate-limitation of photosynthesis became apparent. 3-Phosphoglycerate (3-PGA) levels dropped slightly and the ATP/ADP ratio was not reduced in the transgenic lines under investigation. It is proposed that in SPP-deficient plants, long-term compensatory responses give rise to the observed acceleration of starch synthesis, increase in total cellular Pi content, decrease in protein content, and related reduction in photosynthetic activity.

1.Institut für Pflanzengenetik und Kulturpflanzenforschung (IPK) Corrensstrasse 3 06466 Gatersleben Germany<br/>

... 一般认为, 植物叶片中蔗糖磷酸合酶是蔗糖合成的关键限速酶^[12], 其活性与蔗糖含量密切相关^[13,14] ...

1998

0.0

... 一般认为, 植物叶片中蔗糖磷酸合酶是蔗糖合成的关键限速酶^[12], 其活性与蔗糖含量密切相关^[13,14] ...

2001

0.0

1.667

李永庚, 于振文, 姜东, 余松烈. 冬小麦旗叶蔗糖和籽粒淀粉合成动态及与其有关的酶活性的研究. 作物学报, 2001, 27: 658-664

Y G

, Yu

Z W

, Jiang

, Yu

S L

. Studies on the dynamic changes of the synthesis of sucrose in the flag leaf and starch in the grain and related enzymes of high-yielding wheat. Acta Agron Sin, 2001, 27: 658-664 (in Chinese with English abstract).

鲁麦22和鲁麦14相比,旗叶的光合能力强,蔗糖含量高,催化蔗糖合成的磷酸蔗糖合成酶(SPS)活性高;籽粒中蔗糖供应充足,降解蔗糖的蔗糖合成酶(SS)活性高;与淀粉合成有关的酶:腺苷二磷酸葡萄糖焦磷酸化酶(ADPG-PPase)、游离态(或可溶性)淀粉合成酶(S-STS)和束缚态淀粉合成酶(B-STS)均有较高的活性,这是其粒重高的原因.通过对两个品

... 蔗糖合酶既可催化蔗糖合成又可催化蔗糖分解, 植物光合组织中的蔗糖合酶催化蔗糖合成的活性较高^[15,16] ...

1993

3.347

0.0

Geigenberger

, Stitt

. Sucrose synthase catalyses a reversible reaction in vivo in developing potato tubers and other plant tissues. Planta, 1993, 189: 329-339

Lehrstuhl für Pflanzenphysiologie, Universität Bayreuth, Universitätstrasse 30, W-8580, Bayreuth, FRG.

Experiments were carried out to investigate whether sucrose synthase (Susy) catalyses a readily reversible reaction in vivo in potato (Solanum tuberosum L.) tubers, Ricinus communis L. cotyledons, and heterotrophic Chenopodium rubrum L. cell-suspension cultures. (i) The contents of sucrose, fructose, UDP and UDP-glucose were measured and the mass-action ratio compared with the theoretical equilibrium constant. In all three tissues the values were similar. (ii) Evidence for rapid turnover of label in the sucrose pool was obtained in pulse-chase experiments with potato discs and with intact tubers attached to the plant. The unidirectional rates of sucrose synthesis and degradation were considerably higher than the net flux through the sucrose pool in the tubers. (iii) Labelling of the glucosyl and fructosyl moieties of sucrose from [(14)C]glucose in the presence of unlabelled fructose provided evidence that Susy contributes to the movement of label into sucrose. Methods for estimating the contribution of sucrose-phosphate synthase and Susy are presented and it is shown that their relative contribution varies. For example, the contribution of Susy is high in developing tubers and is negligible in harvested tubers which contain low Susy activity. (iv) The absolute values of the forward (v(+1)) and backward (v(-1)) reaction direction of Susy are calculated from the kinetic labelling data. The estimated values of v(+1) and v(-1) are comparable, and much higher than the net flux through the sucrose pool. (v) The estimated concentrations of the substrates and products of Susy in tubers are comparable to the published K m values for potato-tuber Susy. (vi) It is concluded that Susy catalyses a readily reversible reaction in vivo and the relevance of this conclusion is discussed with respect to the regulation of sucrose breakdown and the role of Susy in phloem unloading.

PMID: 24178489

... 蔗糖合酶既可催化蔗糖合成又可催化蔗糖分解, 植物光合组织中的蔗糖合酶催化蔗糖合成的活性较高^[15,16] ...

1997

5.135

0.0

... 叶片中酸性转化酶是蔗糖降解主要酶^[17] ...

2006

2.578

0.0

... 棉铃对位叶作为棉铃养分的主要供应者, 其生理特性直接影响棉铃发育所需养分的合成和转运^[18] ...

1988

6.555

0.0

Wells

. Response of leaf ontogeny and photosynthetic activity to reproductive growth in cotton. Plant Physiol, 1988, 87: 274-279

United States Department of Agriculture-Agricultural Research Service, Cotton Physiology and Genetics Research, Stoneville, Mississippi 38776.

This study was conducted to determine if reproductive growth in cotton (Gossypium hirsutum L.) affects concurrent leaf development. Apparent photosynthesis (AP), stomatal conductance (Cs), soluble protein (SP), ribulose bisphosphate carboxylase (RuBisCO), and chlorophyll (Chl) were monitored in four main-stem cotton leaves which emerged at approximately 2 week intervals. The leaf which emerged during vegetative growth (48 days after planting) had higher AP, SP, and RuBisCO levels than that present in any leaves which emerged during fruit development. The last leaf studied (89 days after planting) was still present after boll maturation was completed and exhibited a rejuvenation in AP, SP, RuBisCO, and Chl starting at 30 days after leaf emergence. At 96 days after planting, the P700 Chl a-protein complex (PSI) was virtually absent from the leaves that emerged at 48 and 62 days after planting. The light harvesting Chl a/b complex was still present in these leaves, indicating greater degradation of PSI. The data emphasize the influence of developing fruit on concurrently developing leaves, an effect which was alleviated after boll maturation was completed. The declining AP per unit leaf area and smaller leaf size at the top of the plant results in a reduced photosynthetic potential of successively later emerging leaves. This reduction in leaf AP is consistent with earlier reported seasonal canopy photosynthesis patterns.

PMID: 16666118

... 张文静等^[19]研究表明棉铃对位叶的生理特性存在基因型差异且与铃重的形成关系密切, 棉铃对位叶C/N适宜、内源保护酶活性高有利于铃重提高^[20], 最近研究发现, 外源6-BA显著提高棉铃对位叶蔗糖含量和转化率及淀粉含量, 促进光合产物向棉铃的运输, 从而提高铃重^[21] ...

2007

0.0

1.462

2009

0.0

2013

0.0

1.667

刘敬然, 孟亚利, 王友华, 陈兵林, 张国伟, 周治国. 外源6-BA和ABA对不同播种期棉花产量和品质及其棉铃对位叶光合产物的影响. 作物学报, 2013, 39: 1078-1088

Liu

J R

, Liu

J J

, Meng

Y L

, Wang

Y H

, Chen

B L

, Zhang

G W

, Zhou

Z G

. Effect of 6-BA and ABA applications on yield, quality and photosynthate contents in the subtending leaf of cotton with different planting dates. Acta Agron Sin, 2013, 39: 1078-1088

To study the effects of 6-BA and ABA applications on carbohydrate contents and endogenous hormones in subtending leaf, and their relationship to cotton yield and qualities under different planting dates, we conducted a field experiment with two cotton cultivars Kemian 1 and NuCOTN 33B in the Lower Reaches of Yangtze River (Nanjing) and in the Yellow River Valley (Anyang), China. The cotton seeds were planted on Apr. 25th and May 25th, 2006, respectively, which could result in different growth temperatures for the bolls and their subtending leaves at the same site, and the growth regulators were sprayed at flowering stage. The results showed that the application of 6-BA significantly increased contents of sucrose and starch, and sucrose transformation rate in subtending leaf, while the application of ABA only regulated the balance of endogenous hormones. The application of 6-BA increased boll weight and enhanced fiber qualities in cotton plants with planting dates of Apr. 25th and May 25th, whereas the decrease ranges of cotton yield and qualities in Anyang were less for the application of ABA in cotton plants with planting date of May 25th, compared to those with planting date of Apr. 25th. However, the qualities of cottonseeds were not affected by the applications of 6-BA or ABA significantly. The application of 6-BA or ABA improved relative cotton qualities of late planted cotton, whereas the action mechanisms were different. The application of 6-BA improved cotton yield and quality via enhancing the photosynthate contents and sucrose transformation rate. However, the application of ABA improved cotton yield and qualities via increasing the stress resistance of cotton plants.

<div style="line-height: 150%">Key Laboratory of Crop Physiology & Ecology in Southern China, Ministry of Agriculture / Nanjing Agricultural University, Nanjing 210095, China</div>

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MDT_min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。

... 用蒽酮比色法^[22]测定可溶性总糖和淀粉含量 ...

... 用间苯二酚比色法^[22]测定蔗糖含量 ...

... 用DNS显色法^[22]测定酸性转化酶活性 ...

1999

0.0

... 用果糖和UDPG比色法^[23]测定蔗糖磷酸合酶和蔗糖合酶(合成方向)活性 ...

1999

0.0

... 叶片中蔗糖的水平取决于光合速率、蔗糖代谢和蔗糖外运的协调作用^[24] ...

1989

6.555

0.0

Huber

S C

. Biochemical mechanism for regulation of sucrose accumulation in leaves during photosynthesis. Plant Physiol, 1989, 91: 656-662

U.S. Department of Agriculture, Agricultural Research Service, North Carolina State University, Raleigh, North Carolina 27695-7631.

It is not known why some species accumulate high concentrations of sucrose in leaves during photosynthesis while others do not. To determine the possible basis, we have studied 10 species, known to differ in the accumulation of sucrose, in terms of activities of sucrose hydrolyzing enzymes. In general, acid invertase activity decreased as leaves expanded; however, activities remaining in mature, fully expanded leaves ranged from low (<10 micromoles per gram fresh weight per hour) to very high (>100 micromoles per gram fresh weight per hour). In contrast, sucrose synthase activities were low and relatively similar among the species (4-10 micromoles per gram fresh weight per hour). Importantly, leaf sucrose concentration, measured at midafternoon, was negatively correlated with acid invertase activity. We propose that sucrose accumulation in vacuoles of species such as soybean and tobacco is prevented by acid invertase-mediated hydrolysis. Initial attempts were made to characterize the relatively high activity of acid invertase from mature soybean leaves. Two apparent forms of the enzyme were resolved by Mono Q chromatography. The two forms had similar affinity for substrate (apparent K(m) [sucrose] = 3 millimolar) and did not interconvert upon rechromatography. It appeared that the loss of whole leaf invertase activity during expansion was largely the result of changes in one of the enzyme forms. Overall, the results provide a mechanism to explain why some species do not accumulate sucrose in their leaves. Some futile cycling between sucrose and hexose sugars is postulated to occur in these species, and thus, the energy cost of sucrose production may be higher than is generally thought.

PMID: 16667083

... 郭文琦等^[25]盆栽试验结果表明, 渍水条件下棉株光合速率下降, 光合产物减少, 致使铃数减少、铃重降低 ...

2010

0.0

1.883

郭文琦, 刘瑞显, 周治国, 陈兵林. 施氮量对花铃期短期渍水棉花叶片气体交换参数和叶绿素荧光参数的影响. 植物营养与肥料学报, 2010, 16: 362-369

Guo

W Q

, Liu

R X

, Zhou

Z G

, Chen

B L

. Effects of nitrogen fertilization on gas exchange and chlorophyll fluorescence parameters of leaf during the flowering and boll-forming stage of cotton under short-term waterlogging. Plant Nutr Fert Sci, 2010, 16: 362-369 (in Chinese with English abstract)

试验于2005～2006在南京农业大学卫岗试验站进行,采用盆栽方法,没置正常灌水(土壤相对含水最为75%±5%)和棉花花铃期短期渍水处理(将正常灌水的棉花增加灌水至盆内有可见明水,持续8 d,然后用导管排出表面水层,使盆内土壤相对含水量逐渐恢复到75%±5%),每个水分处理设置3个施氮水平(N 0、3.73、7.46 g/pot,分别相当于大田N 0、240、480 ks/hm~2),研究施氮量对棉花叶片气体交换参数和叶绿素荧光参数的影响.结果表明,与正常灌水处理相比,渍水棉花的叶绿素(Chl)含量、类胡萝卜素(Car)含量和Chl/Car比值显著降低.渍水降低了棉花净光合速率(P_n)和气孔导度(G_s),降低幅度随施氮水平的提高而增大;渍水结束时,渍水棉花的P_n和G_s均随施氮水平的提高而降低.渍水提高了棉花叶片叶绿素仞始荧光(F_o)和非光化学猝灭系数(qN),降低了最大光化学效率(F_v/F_m)、光系统Ⅱ(PS Ⅱ)量子产量(Φ_(PSⅡ))和光化学猝火系数(qP),F_o和qN的升高幅度与F_v/F_m、Φ_(PSⅡ)和qP的降低幅度均随施氮水平的提高而增大.花铃期短期渍水显著降低了棉花生物量和籽棉产量,适量施氮(N 240ks/hm~2)可提高渍水棉化的生物量和籽棉产量.

Abstract：

The flowering and boll development stage is the key period F_or cotton yielding.Waterlogging during this stage inhibits cotton development and reduces final yield significantly.Nitrogen is considered to be an effective up-regulatory nutrient F_or crop growth.The objective of this investigation was to evaluate the effects of nitrogen fertilization on the gas exchange and chlorophyll fluorescence parameters under a short-term waterlogging for about eight days.Pot experiments were conducted with three N levels,namely N 0,3.73 and 7.46 g/pot(equivalent to N 0,240 and 480 kg/ha).The results show that chlorophyll and carotenoid contents and the Chl/Car ratio are significantly decreased under the waterlogging,and net photosynthetic rate(P_n) and stomatal conductance(G_s) are reduced and the reduced degree of P_n and G_s are increased with the increase of N application.The minimum fluorescence(F_o) and non-photochemical quenching coefficient (qN) are increased under the waterlogging,while the maximum photochemical efficiency of open photosystem(F_v/F_m),the quantum yield of electron transport(Φ_(PSⅡ)) and the photochemical quenching(qP)are decreased.The increased degree of F_o and qN,and the reduced degree of F_v/F_m,Φ_(PSⅡ) and qP are enhanced with the increase of N application.At the time of terminating the waterlogging,the P_n,G_s,F_v/F_m,Φ_(PSⅡ) and qP of the waterlogged cottonare decreased with the increase of N application,while the F_o and q N behave contrarily.Both biomass and seed cotton yield,which are affected by the waterlogging,are the highest under the N 240 kg/ha fertilization.

... Albrecht等^[26]和Islam等^[27]研究还表明, 渍水条件下叶片碳水化合累积是碳水化合物向库器官和根系运输受阻的结果, 因此, 棉铃对位叶碳水化合物累积不利于库器官-棉铃的发育 ...

2004

6.555

0.0

Albrecht

, Mustroph

, Fox

T C

. Sugar and fructan accumulation during metabolic adjustment between respiration and fermentation under low oxygen conditions in wheat roots. Physiol Plant, 2004, 120: 93-105

Humboldt-University Berlin, Institute of Biology, Department of Botany, Spaethstr. 80/81, D-12437 Berlin, Germany University of West Florida, Department of Biology, Pensacola, FL 32514-5751, USA.

In terms of gene expression and carbohydrate metabolism, the response of wheat seedlings to hypoxia is dramatically different from the anoxic response. Total carbohydrate content of roots increased 4-fold during 6 days of hypoxia, with a 17-fold increase in fructans. In contrast, anoxically treated roots depleted all soluble carbohydrates and died within 72 h. Gas exchange measurements (CO(2) release vs. O(2) uptake) demonstrate that hypoxia establishes a new balance between fermentation and aerobic respiration in the roots without altering the flux of carbon through glycolysis. Furthermore, the respiratory component of this new balance is 55% higher in roots that have been hypoxically pretreated compared to non-hypoxically pretreated roots. The establishment of this new homeostasis under hypoxia involves the induction of glycolytic (aldolase and enolase) and fermentative enzymes (pyruvate decarboxylase, alcohol dehydrogenase, and lactate dehydrogenase). Enzyme induction is generally complete within 24 h with mRNA induction occurring primarily during Period I (0-6 h of hypoxia), and maximal enzymes activities attained during Period II (6-24 h of hypoxia). Accumulation rates of Suc, hexoses, and fructans also change during Periods I and II. By the start of Period III (24-144 h of hypoxia), the metabolic adjustments are complete and fructans are the major carbohydrate accumulated. In anoxia, the pattern of enzyme induction was dramatically different: aldolase was not induced and declined throughout the treatment. Alcohol dehydrogenase, pyruvate decarboxylase, and lactate dehydrogenase were induced as in hypoxia, but rapidly declined within 72 h of anoxia. Only enolase exhibited a similar expression pattern in both anoxia and hypoxia.

PMID: 15032881

2004

1.925

0.0

Islam

M A

, Macdonald

S E

. Ecophysiological adaptations of black spruce (Picea mariana) and tamarack (Larix laricina) seedlings to flooding. Trees, 2004, 18: 35-42

Black spruce [ Picea mariana (Mill.) B.S.P.] and tamarack [ Larix laricina (Du Roi) K. Koch] are the predominant tree species in boreal peatlands. The effects of 34 days of flooding on morphological and physiological responses were investigated in the greenhouse for black spruce and tamarack seedlings in their second growing season (18 months old). Flooding resulted in reduced root hydraulic conductance, net assimilation rate and stomatal conductance and increased needle electrolyte leakage in both species. Flooded tamarack seedlings maintained a higher net assimilation rate and stomatal conductance compared to flooded black spruce. Flooded tamarack seedlings were also able to maintain higher root hydraulic conductance compared to flooded black spruce seedlings at a comparable time period of flooding. Root respiration declined in both species under flooding. Sugar concentration increased in shoots while decreasing in roots in both species under flooding. Needles of flooded black spruce appeared necrotic and electrolyte leakage increased over time with flooding and remained significantly higher than in flooded tamarack seedlings. No visible damage symptoms were observed in flooded tamarack seedlings. Flooded tamarack seedlings developed adventitious roots beginning 16 days after the start of flooding treatment. Adventitious roots exhibited significantly higher root hydraulic conductivity than similarly sized flooded tamarack roots. Flooded black spruce lacked any such morphological adaptation. These results suggest that tamarack is better able to adjust both morphologically and physiologically to prolonged soil flooding than black spruce seedlings.

1.Department of Renewable Resources University of Alberta 751 General Services Building Edmonton Alberta T6G 2H1 Canada<br/>

... Yang等^[28]和王维等^[29]研究表明, 轻度、适度的土壤水分胁迫可诱导水稻茎部蔗糖磷酸合酶活性升高而酸性转化酶下降, 利于贮藏碳水化合物的输出, Hirano等^[30]研究发现淹水下水稻上层叶片的蔗糖磷酸合酶活性被高度激活, Harada等^[31]发现厌氧条件下水池草芽蔗糖磷酸合酶、蔗糖合酶和酸性转化酶活性均增加 ...

2001

5.242

0.0

Yang

, Zhang

, Wang

, Zhu

. Activities of starch hydrolytic enzymes and sucrose-phosphate synthase in the stems of rice subjected to water stress during grain filling. J Exp Bot, 2001, 52: 2169-2179

College of Agriculture, Yangzhou University, Yangzhou, Jiangsu, China.

To understand the effect of water stress on the remobilization of prestored carbon reserves, the changes in the activities of starch hydrolytic enzymes and sucrose-phosphate synthase (SPS) in the stems of rice (Oryza sativa L.) during grain filling were investigated. Two rice cultivars, showing high lodging-resistance and slow remobilization, were grown in the field and subjected to well-watered (WW, psi(soil)=0) and water-stressed (WS, psi(soil)=-0.05 MPa) treatments 9 d after anthesis (DAA) till maturity. Leaf water potentials of both cultivars markedly decreased during the day as a result of WS treatment, but completely recovered by early morning. WS treatment accelerated the reduction of starch in the stems, promoted the reallocation of prefixed (14)C from the stems to grains, shortened the grain filling period, and increased the grain filling rate. More soluble sugars including sucrose were accumulated in the stems under WS than under WW treatments. Both alpha- and beta-amylase activities were enhanced by the WS, with the former enhanced more than the latter, and were significantly correlated with the concentrations of soluble sugars in the stems. The other two possible starch-breaking enzymes, alpha-glucosidase and starch phosphorylase, showed no significant differences in the activities between the WW and WS treatments. Water stress also increased the SPS activity that is responsible for sucrose production. Both V(limit) and V(max), the activities of the enzyme at limiting and saturating substrate concentrations, were enhanced and the activation state (V(limit)/V(max)) was also increased as a result of the more significant enhancement of V(limit). The enhanced SPS activity was closely correlated with an increase of sucrose accumulation in the stems. The results suggest that the fast hydrolysis of starch and increased carbon remobilization were attributed to the enhanced alpha-amylase activity and the high activation state of SPS when the rice was subjected to water stress.

PMID: 11604456

2004

0.0

1.667

王维, 张建华, 杨建昌, 朱庆森. 水分胁迫对贪青迟熟水稻茎贮藏碳水化合物代谢及产量的影响. 作物学报, 2004, 30: 196-204

Wang

, Zhang

J H

, Yang

J C

, Zhu

Q S

. Effect of water stress on metabolism of stored carbohydrate of stem and yield in rice grown under unfavorable- delayed senescence. Acta Agron Sin, 2004, 30: 196-204 (in Chinese with English abstract)

The effect of water stress in pot conditions on stem-stored carbohydrate metabolism associated with key enzymes and grain yield in rice grown under unfavorable-delayed Senescence were investigated. The rice grown under unfavorable-delayed Senescence by applied with heavy nitrogen resulted in marked

Key Lab of Crop Cultivation & Physiology, Ministry of Agriculture, Yangzhou University, Yangzhou 225009, Jiangsu

研究了水分胁迫对贪青迟熟水稻的产量及其茎节间贮藏碳水化合代谢和相关酶活性的影响。结果表明：水稻贪青迟熟显著地降低结实率和粒重而导致产量显著降低，适度土壤水分胁迫抑制了高氮的不良效应，有效地促进了茎贮藏碳水化合物的输出，增加对籽粒产量的贡献，降低了产量损失。花后的土壤干旱明显诱导α-淀粉酶、蔗糖磷酸合

1997

0.0

2003

2.638

0.0

Harada

, Ishizawa

. Starch degradation and sucrose metabolism during anaerobic growth of pondweed (Potamogeton distinctus A. Benn. ) turions. Plant Soil, 2003, 253: 125-135

Stem growth in pondweed (Potamogeton distinctus A. Benn.) turions was stimulated in anaerobic conditions. Decreases in starch contents in turion cells, the number of amyloplasts in a cell and sizes of starch granules in amyloplasts clearly showed enhancement of starch consumption in turions during anaerobic growth of stems of pondweed. Although the total activities of both agr

-amylase and starch phosphorylase were not stimulated by anaerobic conditions, the activities of one amylase and two isozymes of starch phosphorylase, separated by native polyacrylamide gel electrophoresis, were enhanced in anaerobic conditions. The sucrose content in turion cells decreased rapidly in the early stage of anaerobic growth but then remained constant after the enhanced growth had started, suggesting active sucrose metabolism. The activities of sucrose-phosphate synthase, sucrose synthase and acid invertase, measured in vitro, were enhanced in extracts from turions in anaerobic conditions. The induction of the activity of sucrose synthase was most significant, suggesting that sucrose synthase plays an important role in sucrose metabolism in pondweed turions growing in anaerobic conditions.

1.Department of Developmental Biology and Neuroscience, Graduate School of Life Sciences Tohoku University 980-8578 Sendai Japan<br/>

... 表明渍水条件下棉铃对位叶蔗糖代谢较活跃, 蔗糖合成和降解代谢同时进行^[32] ...

2009

0.0

Rosa

, Prado

, Podazza

, Interdonato

, González

J A

, Hilal

, Prado

F E

. Soluble sugars-Metabolism, sensing and abiotic stress. Plant Signal Behav, 2009, 4: 388-393

Facultad de Ciencias Naturales e IML, Tucumán, Argentina.

Plants are autotrophic and photosynthetic organisms that both produce and consume sugars. Soluble sugars are highly sensitive to environmental stresses, which act on the supply of carbohydrates from source organs to sink ones. Sucrose and hexoses both play dual functions in gene regulation as exemplified by the upregulation of growth-related genes and downregulation of stress-related genes. Although coordinately regulated by sugars, these growth- and stress-related genes are upregulated or downregulated through HXK-dependent and/or HXK-independent pathways. Sucrose-non-fermenting-1- (SNF1-) related protein pathway, analogue to the protein kinase (SNF-) yeast-signalling pathway, seems also involved in sugar sensing and transduction in plants. However, even if plants share with yeast some elements involved in sugar sensing, several aspects of sugar perception are likely to be peculiar to higher plants. In this paper, we have reviewed recent evidences how plants sense and respond to environmental factors through sugar-sensing mechanisms. However, we think that forward and reverse genetic analysis in combination with expression profiling must be continued to uncover many signalling components, and a full biochemical characterization of the signalling complexes will be required to determine specificity and cross-talk in abiotic stress signalling pathways.

PMID: 19816104

... 相关分析发现, 渍水处理棉铃对位叶可溶性糖、蔗糖和淀粉含量与蔗糖磷酸合酶、蔗糖合酶及酸性转化酶活性均呈正相关(P[33], 且植物体内的碳水化合物首先用于维持呼吸和生存能量需求^[34], 因而推测较高的酶活性与维持植株在渍水下生存的能量需求有关^[34], 而一旦碳水化合物含量降低甚至耗尽, 植物生长将很慢或难以恢复^[35,36,37] ...

2001

0.0

1973

0.0

2001

0.0

1996

5.242

0.0

2001

5.242

0.0

Schlüter

, Crawford

R M M

. Long term anoxia tolerance in leaves of Acorus calamus L. and Iris pseudacorus L. J Exp Bot, 2001, 52: 2213-2225

Sir Harold Mitchell Building, University of St Andrews, St Andrews KY16 9AJ, Fife, UK. schluter@mpimp-golm.mpg.de

Mature green leaves of Acorus calamus and Iris pseudacorus have been shown to survive at least 28 d of total anoxia in the dark during the growing season, increasing up to 75 d and 60 d in overwintering leaves in A. calamus and I. pseudacorus, respectively. During the period of anaerobic incubation the glycolytic rate is reduced, carbohydrate reserves are conserved and ethanol levels in the tissues reached an equilibrium. Prolonged anoxia significantly suppressed leaf capacity for respiration and photosynthesis. After 28 d of anoxia, respiratory capacity was reduced in A. calamus and I. pseudacorus by 80% and 90%, respectively. The photosynthetic capacity of leaves decreased by 83% in A. calamus and by 97% in I. pseudacorus after 28 d of anoxia. This reduction in photosynthetic capacity was accompanied by a modification of the chlorophyll fluorescence pattern indicating damage to the PSII reaction centre and subsequent electron transport. Chlorophyll content was only slightly reduced after 28 d under anoxia and darkness in A. calamus, whereas there was a 50% reduction in I. pseudacorus. On return to air A. calamus leaves that endured 28 d of anoxia recovered full photosynthetic activity within 7 d while those of I. pseudacorus had a lag phase of 3-10 d. This well-developed ability to endure prolonged periods of oxygen deprivation in both these species is associated with a down-regulation in metabolic activity in response to the imposition of anaerobiosis. It is suggested that when leaf damage eventually does take place in these species after protracted oxygen deprivation, it is anoxic rather than post-anoxic stress that is responsible.

PMID: 11604461