作物杂志, 2018, 34(6): 76-82 doi: 10.16035/j.issn.1001-7283.2018.06.012

生理生化·植物营养·栽培耕作

脱落酸对低温胁迫下玉米幼苗生长和光合特性的影响

田礼欣1, 杨晔1, 左师宇1, 刘旋1, 魏湜1, 孙磊2, 李晶1

1 东北农业大学农学院,150030,黑龙江哈尔滨

2 黑龙江省农业科学院土壤肥料与环境资源研究所,150086,黑龙江哈尔滨

Effects of Abscisic Acid on Growth and Photosynthetic Characteristics of Maize Seedlings under Low Temperature Stress

Tian Lixin1, Yang Ye1, Zuo Shiyu1, Liu Xuan1, Wei Shi1, Sun Lei2, Li Jing1

1 College of Agronomy, Northeast Agricultural University, Harbin 150030, Heilongjiang, China

2 Institute of Soil Fertilizer and Environment Resources, Heilongjiang Academy of Agricultural Sciences, Harbin 150086, Heilongjiang, China

通讯作者: 李晶,副教授,主要从事作物逆境生理研究

收稿日期: 2018-06-4   修回日期: 2018-09-5   网络出版日期: 2018-12-15

基金资助: 国家重点研发计划.  2017YFD0300400

Received: 2018-06-4   Revised: 2018-09-5   Online: 2018-12-15

作者简介 About authors

田礼欣,硕士研究生,主要从事玉米栽培生理研究 。

摘要

脱落酸(ABA)是作物生产中广泛应用的一种植物激素。探究ABA对低温胁迫下玉米幼苗光合特性的影响,为阐明低温胁迫下玉米幼苗叶片光合结构的防御保护机制提供理论依据。以玉米品种“久龙5号”为试验材料,采用盆栽试验,探讨了低温胁迫0、2、4、6、8d下不同浓度脱落酸对玉米幼苗生长及光合特性的影响。结果表明,低温胁迫显著抑制玉米幼苗的正常生长,导致SPAD值下降,叶片净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、PSⅡ光化学最大效率(Fv/Fm)和潜在光化学效率(Fv/Fo)水平下降,初始荧光(Fo)显著升高。外源喷施适当浓度的ABA能够显著缓解由低温胁迫导致的玉米叶片Pn、Tr和Gs下降幅度,提高Fv/Fm和Fv/Fo,降低Fo水平。由此可见,外源施加适当浓度的ABA能提高幼苗叶片的光合能力,提高PSⅡ反应中心活性,促进物质积累,提高玉米幼苗的耐冷性。同时,外源施加ABA存在一定的浓度效应,即“低促高抑”。本研究所设计浓度范围中15mg/L ABA浓度对光合抑制缓解效果最显著。

关键词: 玉米 ; 脱落酸 ; 低温胁迫 ; 光合特性

Abstract

Abscisic acid (ABA) is a plant hormone widely used in the physiological and biochemical processes of plants. The effects of ABA on photosynthetic characteristics of maize seedlings under low temperature stress were explored, which provided a theoretical basis for elucidating the defense and protection mechanism of photosynthetic structure of maize seedling leaves under low temperature stress. A pot experiment of maize seedlings (variety Jiulong 5) was conducted to investigate the changes of growth and photosynthetic characteristics of maize seedlings under different concentrations of abscisic acid at 0, 2, 4, 6 and 8 days after low temperature stress. The results showed that, the normal growth of maize seedlings was inhibited significantly under chilling stress. All the decreases in SPAD value, Pn, Tr, Gs, Fv/Fm, Fv/Fo and the increases in Fo reflected the inhibitory effects of chilling stress. Exogenous appropriate concentration of ABA led to a marked increase in Pn, Tr, Gs, Fv/Fm, Fv/Fo and the reduction in Fo of maize seedlings. Therefore, exogenous appropriate concentration of ABA could promote the photosynthesis, PSⅡ reactivity and substance accumulation, as well as enhance the chilling resistance of maize seedlings. Meanwhile, exogenous ABA has a certain concentration effect, that is, “low promotion and high inhibition”. The concentration of 15mg/L ABA in the designed concentration range was the most effective in alleviating the photosynthesis inhibition of chilling stress.

Keywords: Maize ; Abscisic acid ; Chilling stress ; Photosynthetic characteristics

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本文引用格式

田礼欣, 杨晔, 左师宇, 刘旋, 魏湜, 孙磊, 李晶. 脱落酸对低温胁迫下玉米幼苗生长和光合特性的影响[J]. 作物杂志, 2018, 34(6): 76-82 doi:10.16035/j.issn.1001-7283.2018.06.012

Tian Lixin, Yang Ye, Zuo Shiyu, Liu Xuan, Wei Shi, Sun Lei, Li Jing. Effects of Abscisic Acid on Growth and Photosynthetic Characteristics of Maize Seedlings under Low Temperature Stress[J]. Crops, 2018, 34(6): 76-82 doi:10.16035/j.issn.1001-7283.2018.06.012

玉米是中国第一大粮食作物,2016年种植面积达0.37×108hm2[1]。玉米是典型的喜温作物,对低温胁迫比较敏感。黑龙江省地处高纬度,热量资源有限,早春低温冷害是玉米生产上主要的气象灾害之一[2,3]。低温是限制作物生长及其产量形成的重要环境因素[4,5]。研究表明,植物干重的90%来自光合作用,而光合系统对低温最敏感[6],低温导致叶片叶绿素吸收的光能减少,叶片光系统Ⅱ(PSⅡ)活性中心受损,从而抑制玉米进行光合作用[7,8]。脱落酸(ABA)作为一种植物激素,在植物的生理生化过程中起着重要作用,也是低温胁迫的一个重要信号因子,对细胞结构有保护作用[9,10]。因此,研究外源ABA对低温胁迫下玉米幼苗光合特性的影响具有重要意义。已有研究表明,外源ABA可以缓解胁迫环境对植物造成的伤害,目前已在绿豆[11]、象草[12]、辣椒[13]、水稻[14]和小麦[15]等植物中得到验证。任旭琴等[16]研究发现,外源施加ABA可以提高辣椒在低温胁迫下的SPAD值,缓解低温对幼苗叶绿素合成的限制,保护叶绿素结构的完整性。Karimi等[17]研究表明,低温胁迫下施加ABA提高了葡萄幼苗的净光合速率(Pn)、蒸腾速率(Tr)和气孔导度(Gs),缓解了低温逆境对光合机构的破坏。黄宇等[18]研究表明,低温条件下外源应用ABA后雷公藤幼苗叶片的初始荧光(Fo)和实际光化学量子效率(ΦPSⅡ)降低,最大荧光(Fm)和PSⅡ最大光化学量子产量(Fv/Fm)升高,说明低温下外源ABA可以提升叶片的光合能力,从而提高幼苗的耐冷性。前人已经从氧化系统、渗透调节和激素含量等方面对ABA调控低温胁迫下玉米幼苗的生理特性进行了详细研究[19],但从光合结构和光合分子机制方向深入探究外源ABA对低温胁迫下光合作用的调节机制报道较少。本研究以耐低温玉米品种“久龙5号”为试验材料,探究外源ABA对低温条件下玉米幼苗生长及光合特性的影响,为阐明低温条件下玉米幼苗叶片光合结构的防御保护机制提供理论依据。

1 材料与方法

1.1 试验材料与设计

基于本研究团队前期筛选鉴定,选用耐低温玉米品种“久龙5号”为试验材料,该品种由黑龙江久龙种业有限公司提供。试验土壤取自东北农业大学香坊实验实习基地常规玉米田(黑土)0~20cm耕层。土样含全氮1.78g/kg、速效钾183.48mg/kg、速效磷66.13mg/kg、有机质26.31g/kg、碱解氮122.34mg/kg,pH 6.88。试验所用ABA购自哈尔滨伟业生物工程有限公司,纯度为95%~98%。

试验于2014年4月25日在东北农业大学玉米研究所生理生态实验室进行室外盆栽种植。盆直径20cm、高23cm。于室外正常培养至三叶一心,选择无病、健壮、生长势一致的植株进行试验。试验设置6个处理:自来水+常温,记为CK;自来水+低温,记为C0;ABA+低温,ABA浓度梯度为5、15、25、35mg/L,分别记为C5、C15、C25、C35,每株叶片均匀喷施5mL,喷施时加入吐温-20作为展着剂(含CK和C0)。每个处理20株,取3株长势一致的幼苗计算各指标平均值。喷施诱导24h后,将植株放入HPG-280HX型人工智能培养箱中进行低温胁迫[昼温/夜温=(14±0.5)℃/(5±0.5)℃]处理,光照时间为12h,光强为140μmol/(m2·s),于低温胁迫0、2、4、6、8d时取样。

1.2 测定项目与方法

1.2.1 干重 将叶片鲜样放入烘箱内105℃杀青30min,80℃烘干至恒重并称重。

1.2.2 SPAD值 应用CCM-200 PLUS型(Opti-Sciences,Int.,U.S.A)便携式叶绿素测定仪测定SPAD值。每个处理随机选取3株,测定完全展开倒2叶中上部的同一位置。

1.2.3 光合参数 应用LI-6400便携式光合仪测定叶片Pn、Gs、Tr,测定部位同1.2.2。测定条件为:大气CO2浓度为550μmol/(m2·s),温度为(27±1)℃,光照强度为250μmol/(m2·s)。

1.2.4 叶绿素荧光参数 应用PAM-2500型(heinz walz GmbH,Germany)便携式调制叶绿素荧光仪测定各项荧光参数(叶片暗适应20min后测量),测定部位同1.2.2。首先用弱光测定Fo,随后通过一个强闪光[6 000μmol/(m2·s),脉冲时间0.7s]测得Fm,计算Fv/Fm、Fv/Fo,其中Fv=Fm-Fo

1.3 数据分析

采用SPSS 17.0和Microsoft Excel 2010软件分析试验数据,各个处理间采用LSD法进行数据方差分析和显著性检验。

2 结果与分析

2.1 低温胁迫下外源施加ABA对玉米幼苗叶片SPAD值的影响

研究表明,叶绿素含量与SPAD值呈极显著正相关,SPAD值越大,表明叶绿素含量越高。由图1可知,常温下培养的幼苗(CK)的SPAD值随培养天数增加呈逐渐升高的趋势,表明非逆境胁迫下幼苗叶片SPAD值随生长会提高。低温胁迫条件下,5个不同浓度ABA处理的玉米幼苗叶片SPAD值随着处理时间的延长逐渐降低。其中C15处理的SPAD值下降幅度最小,分别较CK下降了6.28%、12.89%、25.37%和41.98%;C35处理下降最明显,分别较CK下降了21.47%、27.32%、47.80%和57.08%。说明C35处理的玉米幼苗自身保护系统受到严重抑制,细胞中叶绿素被降解。

图1

图1   外源ABA对低温胁迫下玉米幼苗叶片SPAD的影响

CK:自来水+常温;C0:自来水+低温;C5:低温ABA浓度5mg/L;C15:低温ABA浓度15mg/L;C25:低温ABA浓度25mg/L;C35:低温ABA浓度35mg/L。柱形图上不同字母表示处理间存在显著性差异(P<0.05)。下同

Fig.1   Effects of exogenous ABA on the SPAD in the leaves of maize seedlings under chilling stress

CK: treatment of clear water and room temperature; C0: treatment of clear water and low temperature; C5: treatment of low temperature and 5mg/L ABA; C15: treatment of low temperature and 15mg/L ABA; C25: treatment of low temperature and 25mg/L ABA; C35: treatment of low temperature and 35mg/L ABA. Different letters above the bar mean significant difference between them (P<0.05). The same below


2.2 低温胁迫下外源施加ABA对玉米幼苗叶片光合参数的影响

图2可知,随着低温胁迫时间的延长,外源施加不同浓度ABA处理的玉米幼苗叶片Pn含量均呈下降趋势。与对照相比,低温胁迫降低了玉米幼苗叶片的Pn。但是不同浓度ABA处理降低的幅度不同,其中C0处理和C35处理在第2天和第8天有一个迅速下降的过程,C15处理的下降最为缓慢,C5处理和C25处理下降幅度均低于C0处理和C35处理。低温胁迫的第8天,与CK处理相比,C0、C5、C15、C25、C35处理Pn分别下降了36.08%、35.53%、25.00%、28.46%、40.96%。

图2

图2   外源ABA对低温胁迫下玉米幼苗叶片Pn的影响

Fig.2   Effects of exogenous ABA on the Pn in the leaves of maize seedlings under chilling stress


图3可知,低温胁迫条件下,外源施加不同浓度ABA处理的玉米幼苗叶片Tr的变化呈逐渐下降趋势。在低温胁迫下,C15处理和C25处理在各时间段较其他浓度处理下降幅度最小,说明C15处理和C25处理减轻了低温胁迫对玉米幼苗的伤害,提高了玉米幼苗的耐低温能力。C35处理加剧了低温胁迫下玉米幼苗Tr的下降速度,说明C35处理可能抑制了玉米幼苗自身的耐低温能力,降低了其抗冷性。低温胁迫的第8天,与CK处理相比,C0、C5、C15、C25、C35处理Tr分别下降了39.20%、38.00%、25.69%、29.96%、42.77%。

图3

图3   外源ABA对低温胁迫下玉米幼苗叶片Tr的影响

Fig.3   Effects of exogenous ABA on the Tr in the leaves of maize seedlings under chilling stress


图4可知,与CK相比,外源施加不同浓度ABA处理的玉米幼苗叶片Gs均随低温胁迫时间的增加呈下降趋势,且与Pn和Tr的变化规律一致。低温胁迫初期各处理叶片Gs均呈线性下降,低温胁迫时间延长至8d时,Gs均显著下降。各处理相比,尽管变化规律相似,但C15处理和C25处理在各时间点的下降幅度均小于其他3个处理,C35处理下降的幅度最为显著。这说明C15和C25不同程度缓解了低温下玉米幼苗叶片的Gs降低,而C35处理加快了其下降幅度。低温胁迫的第8天,与CK处理相比,C0、C5、C15、C25、C35处理Gs分别下降了26.39%、26.11%、15.28%、31.11%、16.39%。

图4

图4   外源ABA对低温胁迫下玉米幼苗叶片Gs的影响

Fig.4   Effects of exogenous ABA on the Gs in the leaves of maize seedlings under chilling stress


2.3 低温胁迫下外源施加ABA对玉米幼苗叶片叶绿素荧光参数的影响

Fo为初始荧光,代表在叶片暗反应过程中光合体系的PSⅡ反应中心完全行使功能时的荧光水平,用于评估植物在逆境条件下叶片PSⅡ永久性伤害程度[20]。由图5可知,随着低温胁迫时间的延长,外源施加不同浓度ABA处理的玉米幼苗叶片Fo均呈上升趋势,表现为C15处理和C25处理呈线性缓慢上升;C0、C5和C35处理在0~2d先迅速上升,在2~6d时上升速度减缓,6d后迅速上升;CK变化不显著。各低温处理时间段,C35处理玉米幼苗叶片的Fo明显高于其他浓度处理,同时除CK外,C15处理和C25处理的Fo均显著低于其他浓度处理。说明C15和C25处理有效缓解了低温胁迫引起的原初光能转化效率的降低,C35处理则加速光抑制过程。

图5

图5   外源ABA对低温胁迫下玉米幼苗叶片Fo的影响

Fig.5   Effects of exogenous ABA on the Fo in the leaves of maize seedlings under chilling stress


Fv/Fm指PSⅡ最大光化学量子产量,反映PSⅡ最大光能转换效率和光合潜能。由图6可知,随着低温胁迫时间的延长,外源不同浓度ABA处理的玉米幼苗叶片Fv/Fm也随之下降。其中Fv/Fm在0~2d时下降迅速,2~4d时下降速度减缓,4d后迅速下降,对照无明显变化。Fv/Fm下降,说明玉米幼苗PSⅡ原初光能转化率降低,叶片光合系统受到抑制,2~4d时Fv/Fm变化表明玉米幼苗在低温胁迫下启动了自我保护机制。常温下,玉米幼苗叶片的Fv/Fm随处理时间的延长变化幅度不明显。不同低温胁迫时间,C35处理玉米幼苗叶片的Fv/Fm下降幅度最大,C15处理和C25处理的Fv/Fm下降最为缓慢。C35处理在没有经过低温处理时相较其他处理Fv/Fm就已经出现下降趋势,说明高浓度ABA处理在常温下引起了玉米幼苗叶片的光抑制。

图6

图6   外源ABA对低温胁迫下玉米幼苗叶片Fv/Fm的影响

Fig.6   Effects of exogenous ABA on the Fv/Fm in the leaves of maize seedlings under chilling stress


Fv/Fo反映光合电子传递链中最敏感成分PSⅡ的水溶性复合体的效率。Fv/Fo和Fv/Fm都是表明光化学反应状况的重要参数,且二者变化趋势类似。由图7可知,对照在常温下正常培养,Fv/Fo保持稳定。随着低温胁迫时间的延长,各处理玉米叶片的Fv/Fo均呈下降趋势,且外源不同浓度ABA处理的变化存在差异。表现为C35处理下降最明显,C15处理下降幅度最小。除低温4d时,C35处理玉米幼苗叶片的Fv/Fo在不同低温胁迫时间的下降幅度都最大。C15处理和C25处理的Fv/Fo下降最为缓慢,说明C15处理和C25处理保护了低温胁迫下玉米幼苗PSⅡ的潜在活性。但在低温胁迫8d时,C15处理和C25处理玉米幼苗叶片的Fv/Fo较CK下降迅速。

图7

图7   外源ABA对低温胁迫下玉米幼苗叶片Fv/Fo的影响

Fig.7   Effects of exogenous ABA on the Fv/Fo in the leaves of maize seedlings under chilling stress


2.4 低温胁迫下外源施加ABA对玉米幼苗干重的影响

表1可知,随着低温胁迫时间的延长,玉米幼苗的干重呈逐渐增加趋势。低温胁迫初期CK与C0差异不显著,随着胁迫程度的加重而达显著水平,CK处理与C5、C15和C25处理的幼苗干重差异不显著;C0处理与C5、C25和C35处理的幼苗干重均差异不显著。在低温胁迫条件下,C15处理幼苗干重均显著高于C0处理(P<0.05),说明外源施加适宜浓度的ABA可以缓解低温对幼苗生长的抑制作用,也有利于玉米幼苗干物质的积累。

Table 1   Effects of exogenous ABA on the leaf dry weight of maize seedling under chilling stress g/株 g/plant

处理
Treatment
低温胁迫天数Days of chilling stress (d)
02468
CK0.35±0.03ab0.45±0.02ab0.50±0.04a0.58±0.03a0.83±0.03a
C00.32±0.03b0.36±0.09b0.42±0.01b0.49±0.02b0.72±0.02b
C50.34±0.03ab0.41±0.02ab0.45±0.04ab0.56±0.02a0.76±0.05ab
C150.38±0.02a0.45±0.03a0.48±0.03a0.57±0.02a0.82±0.03a
C250.34±0.05ab0.44±0.03a0.47±0.03ab0.51±0.03ab0.77±0.07ab
C350.31±0.03b0.39±0.02ab0.43±0.02b0.55±0.02a0.72±0.02b

Note: Different letters indicate significant difference (P<0.05)

注:不同字母表示处理间存在显著差异(P<0.05)

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2.5 不同性状之间的相关性分析

表2可知,SPAD与其他7个性状之间相关性均达到显著水平(P<0.05);Fo与SPAD、Pn、Tr、Gs、Fv/Fm和Fv/Fo之间达到显著或极显著负相关水平,与干重达到显著正相关水平;干重与SPAD、Pn、Fo和Fv/Fm相关性达到显著水平;Fv/Fm与Tr和Fo相关性达到极显著水平(P<0.01);Fv/Fo与Pn和Fv/Fm相关性也达到极显著水平(P<0.01),表明气体交换参数和叶绿素荧光参数之间存在差异,幼苗的干重与气体交换参数和叶绿素荧光参数之间也存在差异。

表2   不同性状之间的相关性分析

Table 2  The correlation analysis between different characters

相关系数Correlation coefficientSPADPnTrGsFoFv/FmFv/Fo叶片干重Leaf dry weight
SPAD1
Pn0.52*1
Tr0.64*0.501
Gs0.60*0.51*0.63*1
Fo-0.65*-0.65*-0.66**-0.62*1
Fv/Fm0.76*0.63*0.79**0.69*-0.78**1
Fv/Fo0.49*0.62**0.57*0.53*-0.65*0.70**1
叶片干重Leaf dry weight-0.37*-0.48*-0.44-0.430.49*-0.46*-0.421

Note: "*" significant at the 0.05 probability level; "**" significant at the 0.01 probability level

注:“*”代表P<0.05水平,“**”代表P<0.01水平

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3 讨论

叶绿素是植物进行光合作用的重要色素,也是评价光合活性的重要指标[21]。低温胁迫降低光合色素的合成速率,进而加速叶绿素的降解,引起膜脂过氧化,进一步导致光能的吸收与传递受到影响,最后导致光合速率降低[22,23]。Pagter等[24]和Kalisz等[25]研究表明,低温胁迫显著抑制了绣球(Hydrangea)和罗勒(Ocimum basilicum L.)叶片叶绿素的积累。然而,Zeng等[26]研究表明,低温胁迫显著促进了水稻幼苗叶片中叶绿素的积累,可能是由于低温造成了水稻幼苗极度脱水,进而保持了更好的绿色形态和光合能力。本研究表明,低温胁迫破坏了叶绿体质膜结构,抑制了叶片进行光合作用,导致玉米幼苗叶片叶绿素含量持续下降,延缓玉米生长。前人研究表明,外源施加激素类物质可以延缓植株衰老,保护叶绿素免受低温胁迫的影响[27]。Karimi等[17]研究表明,外源施加ABA可以促进低温胁迫下葡萄幼苗叶片中叶绿素的合成及积累。本研究表明,低温条件下,外源施加适当浓度的ABA处理在一定程度上可以缓解幼苗叶片叶绿素的下降幅度,这可能是外源施加ABA能够缓解叶绿体的质膜过氧化,促进叶片中原叶绿素酸酯氧化还原酶的活性,保持叶片中光合色素含量,利于叶绿素的合成。

光合作用是植物生长发育的基础,也是受低温影响最敏感的性状之一[28]。低温胁迫会影响Gs、Tr、Pn和PSⅡ的光化学效率等[29]。低温胁迫条件下,植株叶片的光合参数(Pn、Tr、Gs)均小于常温处理[25,30]。本研究结果表明,低温胁迫导致叶片Pn降低,可能是由于低温胁迫降低了Gs,限制了CO2向叶绿体的运输,导致幼苗叶片CO2浓度降低,进而限制了运输光合原料和产物的能力所致[31]。低温胁迫下,Karimi等[17]用200µmol/L ABA处理葡萄叶片后显著提高了Pn、Tr和Gs。Souza等[32]研究发现,外源施加ABA可以提高干旱胁迫下玉米叶片的Tr和Pn。本研究表明,低温胁迫下施加适当浓度的ABA可以提高玉米叶片Pn、Tr和Gs,这与前人的研究结果一致。这可能是由于ABA可以缓解由低温引起的气孔限制来减少低温对玉米叶片的不利影响。

PSⅡ反应中心是对低温敏感的光合机构的主要组成部分,因此,叶绿素荧光参数常被用作细胞应激的指标,并也用于评价各种环境胁迫下的光合活性[21]。低温胁迫降低植物叶片对光能的利用率,Fo值增大,Fv/Fm和Fv/Fo值显著下降[21,24-25]。本研究结果表明,低温胁迫下,玉米幼苗叶片Fo升高,Fv/Fm和Fv/Fo下降,原因可能是低温胁迫造成原初光能转化效率降低,植株受到光抑制,PSⅡ反应中心受到伤害。外源施加适当浓度的ABA后,玉米幼苗遭受的低温胁迫抑制均有所缓解。Zhou等[30]和Souza等[32]研究表明,施加一定浓度的ABA均能够缓解逆境对植物的光抑制现象。说明ABA能够通过保护PSⅡ反应中心来缓解低温诱导的光抑制作用,提高光能转化率,保持较高的叶绿素荧光产额,保证玉米幼苗能在低温胁迫下进行光合作用。

植株的生长受到抑制是植物对逆境胁迫的综合表现。其中,低温是限制植物生长和发育的主要因素之一[33]。本研究结果表明,低温胁迫显著降低了玉米幼苗的干重,外源施加适当浓度的ABA可以缓解低温胁迫对玉米幼苗生长的抑制作用,当ABA浓度过低时,对玉米幼苗的生长达不到缓解作用,浓度过高对玉米幼苗的生长起到抑制作用。Zeng等[26]和Zhou等[30]研究发现,外源施加亚精胺和ABA均可以缓解低温胁迫对幼苗的抑制作用,本研究与前人的结果一致。可能是由于外源施加ABA后促进了叶绿素的合成,提高了幼苗的光合作用,从而缓解了低温胁迫对玉米幼苗的抑制作用。

4 结论

外源施用适当浓度的ABA可以促进低温胁迫下玉米幼苗的生长及物质积累,促进叶绿素的合成,提高幼苗叶片Pn、Tr、Gs,提高叶片PSⅡ的Fv/Fm和Fv/Fo,降低Fo,保护了PSⅡ反应中心活性,进而缓解低温胁迫对玉米幼苗的抑制作用,有效地提高了幼苗的耐冷性。同时,外源施加ABA存在一定的浓度效应,即“低促高抑”,本文所设计浓度范围中15mg/L ABA浓度对光合抑制缓解效果最显著。

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

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中国农业科学, 2014,47(11):2109-2125.

DOI:10.3864/j.issn.0578-1752.2014.11.005      URL     Magsci     [本文引用: 1]

【目的】探讨花后高温和外源脱落酸(ABA)对不同持绿型小麦籽粒胚乳细胞增殖、籽粒灌浆和内源激素的影响,为高温逆境下采用激素调控措施提高粒重提供理论依据。【方法】选用持绿型汶农6号和非持绿型济麦20,花后1&mdash;5 d,用透明聚乙烯塑料膜搭设增温棚进行高温处理,同时花后1&mdash;3 d喷施10 mg?L-1的ABA于穗部,用量100 mL?m-2,3次重复。定期取籽粒样,用高效液相色谱法测定4种内源激素,用简易胚乳细胞计数法测定胚乳细胞数目,Richard方程对籽粒增重及胚乳细胞增殖动态模拟并计算相关参数。【结果】高温处理显著降低了两品种强弱势籽粒的胚乳细胞数目,降低胚乳细胞增殖速率,但延长了籽粒胚乳细胞活跃分裂期和实际分裂终期;显著降低两品种弱势籽粒的灌浆速率,缩短了两品种弱势粒的生长活跃期及实际灌浆终期。高温处理显著降低两品种千粒重和穗粒数,其中汶农6号强、弱势粒分别减少3.7和8.2 粒/穗,济麦20强、弱势粒分别减少1.3和4.3 粒/穗;显著降低两品种产量,汶农6号和济麦20产量分别降低19.65%和26.22%。常温及高温下喷施ABA均显著提高了两品种灌浆速率,提高了籽粒胚乳细胞增殖速率,扩大胚乳细胞数目。高温处理降低了强弱势籽粒ZR含量,显著提高了济麦20强、弱势粒花后3&mdash;27 d的GA3含量,显著提高汶农6号花后12&mdash;27 d的GA3含量;但降低了弱势粒花后15&mdash;27 d的IAA含量。高温处理下喷施ABA,降低了济麦20强势粒花后3&mdash;9 d ZR含量,但显著提高济麦20强势粒花后3&mdash;28 d内源ABA含量,显著提高汶农6号强势粒花后3&mdash;18 d ABA含量。常温下喷施ABA显著降低了济麦20和汶农6号强、弱势粒的GA3含量;高温下喷施ABA,显著降低了汶农6号强弱势粒的GA3含量,降低济麦20强势粒花后3&mdash;12 d的GA3含量,显著降低弱势粒花后6&mdash;15 d的GA3含量。常温下喷施ABA显著提高济麦20强势粒花后12&mdash;18 d的IAA含量;提高了汶农6号强势粒花后6&mdash;18 d IAA含量,显著提高两品种弱势粒花后6&mdash;27 d IAA含量。持绿型汶农6号的千粒重和产量均显著大于非持绿型济麦20。【结论】高温胁迫对非持绿型品种的产量和两品种弱势粒粒数影响较大,高温降低了两品种籽粒胚乳细胞数目,降低籽粒灌浆速率,最终导致粒重及产量降低。喷施外源ABA通过调节内源激素水平,促进胚乳细胞分裂,扩大了常温及高温下籽粒库容量,提高了籽粒灌浆速率,从而提高了籽粒产量。

任旭琴, 缪旻珉 .

低温下喷施ABA对辣椒光合特性和N、P、K的影响

湖北农业科学, 2008,47(12):1461-1463.

[本文引用: 1]

Karimi R, Ershadi A, Nejad A R , et al.

Abscisic acid alleviates the deleterious effects of cold stress on ‘Sultana’ grapevine (Vitis vinifera L.) plants by improving the anti-oxidant activity and photosynthetic capacity of leaves

Journal of Horticultural Science & Biotechnology, 2016,91(2):1-10.

DOI:10.1080/14620316.2016.1162027      URL     [本文引用: 3]

The effects of exogenous application of abscisic acid (ABA) on anti-oxidant enzyme activities and photosynthetic capacity in 090004Sultana090005 grapevine (Vitis vinifera L.) were investigated under cold stress. When vines had an average of 15 leaves, 0 (control), 50, 100, or 200 0008M ABA was sprayed to run-off on all leaves of each plant. Twenty-four hours after foliar spraying with ABA, half (n = 5) of the water-only control vines and half (n = 5) of each group of ABA-treated plants were subjected to 400°C for 12 h, followed by a recovery period of 3 d under greenhouse conditions (2500°/1800°C day/night). The remaining plants in each treatment group were kept at 2400°C. Cold stress increased H2O2 and malondialdehyde (MDA) concentrations in vine leaves, whereas all foliar ABA treatments significantly reduced their levels. Chilled plants showed marked increases in their total soluble protein contents in response to each ABA treatment. ABA significantly increased the activities of superoxide dismutase, peroxidase, catalase, and ascorbate peroxidase in cold-stressed grapevine leaves. In contrast, cold stress markedly decreased the rates of leaf photosynthesis (A) and evaporation (E), stomatal conductance (gs), and chlorophyll concentrations in leaves, but increased intercellular CO2 concentrations (Ci) in leaves. Treatment with all concentrations of ABA resulted in lower leaf A, E, and gs values, but higher Ci values at 2400°C. However, following cold stress, ABA-treated vines showed higher leaf A, E, and gs values, but lower Ci values compared to control vines without ABA treatment. The application of 50090009200 0008M ABA allowed chilled vines to recover more quickly when re-exposed to normal temperatures, enabling the vines to resume their photosynthetic capacity more efficiently following cold stress. These results showed that, by stimulating anti-oxidant enzyme systems and alleviating cold-induced stomatal limitations, ABA reduced the inhibitory effect of cold stress on the rate of CO2 fixation in 090004Sultana090005 grapevine plants.

黄宇, 林智勇, 荣俊冬 , .

脱落酸对低温下雷公藤幼苗光合作用及叶绿素荧光的影响

应用生态学报, 2011,22(12):3150-3156.

Magsci     [本文引用: 1]

以1年生雷公藤扦插苗为试材,研究低温胁迫下不同浓度外源脱落酸(ABA,0、5、10、15、20、25 mg&middot;L<sup>-1</sup>)叶面喷施处理对雷公藤叶片光合作用及叶绿素荧光参数的影响.结果表明:喷施20 mg&middot;L<sup>-1</sup>的ABA能显著提高雷公藤幼苗的抗冷性,减缓低温下雷公藤叶片净光合速率(<em>P</em><sub>n</sub>)、蒸腾速率(<em>T</em><sub>r</sub>)、气孔导度(<em>g</em><sub>s</sub>)、胞间CO<sub>2</sub>浓度(<em>C</em><sub>i</sub>)的下降幅度,提高幼苗叶片的光合能力.低温处理6 d后,随着ABA浓度上升,雷公藤叶片的初始荧光(<em>F</em><sub>o</sub>)下降,最大荧光(<em>F</em><sub>m</sub>)和PSII最大光化学效率(<em>F</em><sub>v</sub>/<em>F</em><sub>m</sub>)上升,PSII实际光化学量子产量(<em>&Phi;</em><sub>PS</sub><sub>Ⅱ</sub>)、光化学猝灭系数(<em>q</em><sub>P</sub>)先下降后上升,而非光化学猝灭系数(<em>q</em><sub>N</sub>)呈下降-上升-下降趋势.<em>P</em><sub>n</sub>、<em>g</em><sub>s</sub>、<em>q</em><sub>P</sub>、<em>F</em><sub>m</sub>和<em>F</em><sub>v</sub>/<em>F</em><sub>m</sub>均在20 mg&middot;L<sup>-1</sup>ABA处理时达到峰值.不同浓度ABA的相对电子传递速率(rETR)随着光化光强度增加呈先上升后下降的趋势,当光化光强度(PAR)达到395 &mu;mol&middot;m<sup>-2</sup>s<sup>-1</sup>时,各处理的rETR达到最高值,其中25 mg&middot;L<sup>-1</sup>和20 mg&middot;L<sup>-1</sup>ABA处理分别比对照高17.1%和5.2%.雷公藤叶片<em>&Phi;</em><sub>PSⅡ</sub>的光响应曲线均随光化光强度升高而下降,<em>q</em><sub>N</sub>的光响应曲线则呈相反趋势.

李馨园, 杨晔, 张丽芳 , .

外源ABA对低温胁迫下玉米幼苗内源激素含量及Asr1基因表达的调节

作物学报, 2017,43(1):141-148.

DOI:10.3724/SP.J.1006.2017.00141      URL     [本文引用: 1]

脱落酸(ABA)是低温逆境下的重要信号因子,为了探讨外源 ABA 对低温胁迫下玉米幼苗的生长调节作用,以耐低温玉米品种久龙5号为试验材料,采用不同浓度(5、15、25、35 mg L–1) ABA于玉米三叶一心时喷雾于叶片,并进行低温梯次处理。分析处理后玉米叶片相对电导率、抗氧化酶活性及内源激素ABA、IAA的含量变化,并采用Real-time PCR明确Asr1基因表达水平变化。结果表明,低温胁迫下不同浓度外源ABA处理的玉米叶片相对电导率整体呈上升趋势, SOD和POD活性加强,15 mg L–1和25 mg L–1ABA处理的SOD活性均显著高于未应用ABA处理,玉米內源ABA和IAA合成水平上升,应用ABA后Asr1基因相对表达水平上调,其中5、15和25 mg L–1浓度处理基因表达上调显著。相关分析表明, ABA含量与Asrl基因相对表达量、SOD活性均表现为极显著正相关,与POD活性显著正相关。说明Asr1基因表达受ABA的介导调控, Asr1基因表达量的提升,也促进了内源ABA的合成,抗氧化酶活性加强,提升了应用ABA后玉米的抗低温能力。但外源ABA的介导调控具有一定浓度效应,表现为低促高抑。

杨猛, 魏玲, 庄文锋 , .

低温胁迫对玉米幼苗电导率和叶绿素荧光参数的影响

玉米科学, 2012,20(1):90-94.

DOI:10.3969/j.issn.1005-0906.2012.01.017      URL     [本文引用: 1]

以3个玉米品种为试验材料,通过盆栽试验,研究低温胁迫下电导率和叶绿素荧光参数的变化。结果表明,3个品种的抗低温性为金玉5〉兴垦3〉吉单198,随着低温处理时间的延长,3个品种叶片电导率和初始荧光(F0)上升,Fv/Fm、Fv/F0、NPQ整体上均呈下降趋势。说明低温胁迫使叶绿素所吸收的光能被用于光合作用的部分减少,玉米叶光系统Ⅱ(PSⅡ)活性中心受损。

Mishra A, Mishra K B, Höermiller I I , et al.

Chlorophyll fluorescence emission as a reporter on cold tolerance in Arabidopsis thaliana accessions

Plant Signaling & Behavior, 2011,6(2):301-310.

DOI:10.4161/psb.6.2.15278      URL     PMID:3121992      [本文引用: 3]

Non-invasive, high-throughput screening methods are valuable tools in breeding for abiotic stress tolerance in plants. Optical signals such as chlorophyll fluorescence emission can be instrumental in developing new screening techniques. In order to examine the potential of chlorophyll fluorescence to reveal plant tolerance to low temperatures, we used a collection of nine Arabidopsis thaliana accessions and compared their fluorescence features with cold tolerance quantified by the well established electrolyte leakage method on detached leaves. We found that, during progressive cooling, the minimal chlorophyll fluorescence emission rose strongly and that this rise was highly dependent on the cold tolerance of the accessions. Maximum quantum yield of PSII photochemistry and steady state fluorescence normalized to minimal fluorescence were also highly correlated to the cold tolerance measured by the electrolyte leakage method. In order to further increase the capacity of the fluorescence detection to reveal the low temperature tolerance, we applied combinatorial imaging that employs plant classification based on multiple fluorescence features. We found that this method, by including the resolving power of several fluorescence features, can be well employed to detect cold tolerance already at mild sub-zero temperatures. Therefore, there is no need to freeze the screened plants to the largely damaging temperatures of around -15 C. This, together with the method easy applicability, represents a major advantage of the fluorescence technique over the conventional electrolyte leakage method.

Mohanty S, Grimm B, Tripathy B C .

Light and dark modulation of chlorophyll biosynthetic genes in response to temperature

Planta, 2006,224(3):692-699.

DOI:10.1007/s00425-006-0248-6      URL     [本文引用: 1]

Yadegari L Z, Heidari R, Carapetian J .

The influence of cold acclimation on proline,malondialdehyde (MDA),total protein and pigments contents in soybean (Glycine max) seedlings

Journal of Biological Sciences, 2008,3(1):74-79.

[本文引用: 1]

Pagter M, Liu F, Jensen C R , et al.

Effects of chilling temperatures and short photoperiod on PSⅡ function,sugar concentrations and xylem sap ABA concentrations in two hydrangea species

Plant Science, 2008,175(4):547-555.

DOI:10.1016/j.plantsci.2008.06.006      URL     [本文引用: 2]

Cold injuries are frequently seen in Hydrangea macrophylla but not in Hydrangea paniculata. This may be ascribed to different levels of hardiness in the non-acclimated and the acclimated state, and to differences in responses to short day (SD) and low temperature (LT) and hence in the ability to cold acclimate. In this study H. macrophylla ssp. macrophylla (Thunb.) Ser. ‘Blaumeise’ and H. paniculata Sieb. ‘Kyushu’ were exposed to short photoperiod (10-h) and 4 °C in controlled conditions for 25 days, with measurements and samplings carried out at regular intervals. Chlorophyll fluorescence measurements revealed significant alterations in O–J–I–P fluorescence kinetics and decreases in the photochemical efficiency of photosystem II in stressed plants, followed by less chlorophyll contents. Perturbations of the photosynthetic apparatus were relatively greater in H. macrophylla than in H. paniculata. Likewise, induction of a transient increase in xylem sap abscisic acid concentrations ([ABA] xylem) and accumulation of soluble sugars in leaves and stems were different in the two species. Stem cold hardiness in the non-acclimated state did not differ between H. macrophylla and H. paniculata, indicating equal sensitivity to sudden temperature drops in the growing season. Despite adaptive responses induced by the treatment neither species developed increased stem cold hardiness, suggesting that cold acclimation in Hydrangea may require exposure to temperatures below ca. 4 °C.

Kalisz A, Jezdinský A, Pokluda R , et al.

Impacts of chilling on photosynthesis and chlorophyll pigment content in juvenile basil cultivars

Horticulture Environment & Biotechnology, 2016,57(4):330-339.

DOI:10.1007/s13580-016-0095-8      URL     [本文引用: 3]

The objective of this study was to examine several cultivars of Ocimum basilicum L. (green, red, cinnamon, lettuce leaf, lemon, and Thai basils) for photosynthetic performance, chlorophyll a fluorescence, and chlorophyll content under chilling stress conditions of 6°C in comparison to non-stressed controls (18°C). The basil plants were grown in a peat substrate for 8 weeks and then exposed to chilling for 8 or 16 days, under a 300 μmol61m -2 61s -1 photosynthetic photon flux. After chilling, significant reductions in both the transpiration ( E ) and net photosynthetic ( P N ) rates were observed in basil plants, while the intercellular CO 2 concentration ( C i ) was higher in the plants treated with 6°C in comparison to the controls. The decrease in P N and E was associated with decreased water use efficiency (WUE) and stomatal conductance ( g s ). The greatest impairment of photosynthesis for Thai basil leaves was observed after 8 days of chilling, and for green basil after the 16-day low temperature treatment. The photosystem II (PSII) activity (F v /F m ) and variable-to-initial chlorophyll fluorescence (F v /F 0 ) were decreased after chilling. PSII activity was most affected in lettuce leaf basil after 8 days, and in Thai and red basil plants after the prolonged temperature treatment. Low temperatures did not significantly alter the chlorophyll concentration but did increase the Chl a/b ratio in leaves of basil. The results indicated that the decrease in photosynthesis was not attributable mainly to damage to PSII, but rather to chilling-induced photoinhibition of PSI. The knowledge gained in this study on the genotypic variation in basil response should be helpful for future selection of plants with low chilling sensitivity.

Zeng Y H, Zhang Y P, Xiang J , et al.

Effects of chilling tolerance induced by spermidine pretreatment on antioxidative activity,endogenous hormones and ultrastructure of indica-japonica hybrid rice seedlings

Journal of Integrative Agriculture, 2015,15(2):295-308.

DOI:10.1016/S2095-3119(15)61051-6      URL     [本文引用: 2]

Spermidine (Spd) is known to be involved in the regulation of plant responses to chilling stress and counteract the adverse effect of stress conditions. Antioxidant activities, endogenous hormones and ultrastructure change under chilling stress were investigated inindica-japonicahybrid rice seedlings. 12-d-old seedlings were subjected to exogenous Spd (1 mmol L 1) and then a chilling stress (6 C, 4 d) was induced, followed by a subsequent recovery (25 C, 4 d). Results showed that malondialdehyde (MDA) and proline content were enhanced significantly, whereas shoot fresh and dry weights decreased during chilling stress and after recovery; chlorophyll content of chilling-stressed seedlings increased slightly but declined after recovery; additionally, total soluble sugar, sucrose, fructose and starch contents increased significantly during chilling stress, and only soluble sugar and fructose contents were observed in increase after recovery; chilling stress-induced increases in superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT) activities, but declined after recovery, and the level of ascorbate peroxidase was lower during chilling stress and after recovery; however, endogenous indole-3-acetic acid (IAA), zeatin riboside (ZR), gibberellic acid (GA3), and abscisic acid (ABA) levels were induced decreased compared with Spd pretreatment. The microscopic analysis revealed that chilling stress-induced destruction of the chloroplast envelope during chilling stress and increased the number of plastoglobuli along with aberrations in thylakoid membranes after recovery. In contrast, exogenous Spd protected rice seedlings from chilling-induced injuries in terms of lower malondialdehyde, proline and carbohydrates accumulation coupled with increased endogenous hormones metabolism. After recovery, Spd pretreatment chilling-exposed seedlings showed higher activities of antioxidant enzymes and normal physiological function of chloroplasts. These results suggest that Spd could promote effectively chilling tolerance which might be largely attributable to the integrity of cell structure and normal metabolism of endogenous hormones inindica-japonicahybrid rice seedlings.

Mohabbati F, Paknejad F, Vazan S , et al.

Protective effect of exogenous PGRs on chlorophyll fluorescence and membrane integrity of rice seedlings under chilling stress

Research Journal of Applied Sciences Engineering & Technology, 2013,5(1):146-153.

[本文引用: 1]

Hendrickson L, Ball M C, Wood J T , et al.

Low temperature effects on photosynthesis and growth of grapevine.

Plant, Cell & Environment, 2004,27(7):795-809.

DOI:10.1111/j.1365-3040.2004.01184.x      URL     [本文引用: 1]

Growth and photosynthesis of grapevine ( Vitis vinifera L.) planted on two sloping cool climate vineyards were measured during the early growth season. At both vineyards, a small difference in mean minimum air temperature (1–3 °C) between two microsites accumulated over time, producing differences in shoot growth rate. The growth rates of the warmer (upper) microsite were 34–63% higher than the cooler (lower) site. Photosynthesis measurements of both east and west canopy sides revealed that the difference in carbon gain between the warmer and cooler microsites was due to low temperatures restricting the photosynthetic contribution of east-facing leaves. East-facing leaves at the warmer microsite experienced less time at suboptimal temperature while being exposed to high irradiance, contributing to an average 10% greater net carbon gain compared to the east-facing leaves at the cooler microsite. This chilling-induced reduction in photosynthesis was not due to net photo-inhibition. Further analysis revealed that CO 2 - and light-saturated photosynthesis of grapevines was restricted by stomatal closure from 15 to 25 °C and by a limitation of RuBP regeneration and/or end-product limitation from 5 to 15 °C. Changes in photosynthetic carboxylation efficiency implied that Rubisco activity may also play a regulatory role at all temperatures. This restriction of total photosynthetic carbon gain is proposed to be a major contributor to the temperature dependence of growth rate at both vineyards during the early season growth period.

Allen D J, Ort D R .

Impacts of chilling temperatures on photosynthesis in warm-climate plants

Trends in Plant Science, 2001,6(1):36-42.

[本文引用: 1]

Zhou B Y, Guo Z F, Lin L .

Effects of abscisic acid application on photosynthesis and photochemistry of Stylosanthes guianensis under chilling stress

Plant Growth Regulation, 2006,48(3):195-199.

DOI:10.1007/s10725-006-0005-7      URL     [本文引用: 3]

In our previous study, it was found that abscisic acid (ABA) improved the chilling resistance of Stylosanthes guianensis . In order to determine the effects of ABA on photosynthesis and photochemistry of S. guianensis , an experiment was conducted under controlled condition to determine the effects of exogenous ABA on stomatal conductance ( g s ), transpiration ( E ), photosynthetic rate ( A ) and chlorophyll a fluorescence of this pasture legume. The results showed that ABA treatment reduced A , g s , and E under both chilling (802°C) and control temperature (2802°C). A of the ABA treated plants returned to a high rate, while that of the water-treated plants remained low when plants were rewarmed after chilling treatment. ABA-treated plants had higher maximum photochemical efficiency ( F v / F m ), non-photochemical quenching (NPQ), quantum efficiency of PS II photochemistry (Φ ps02ii ) than water-treated ones during chilling. Although the biomass of S. guianensis was reduced by ABA under control temperature, ABA-treated plants had higher biomass than water-treated ones after 702days of recovery.

Bertamini M, Zulini L, Muthuchelian K , et al.

Low night temperature effects on photosynthetic performance on two grapevine genotypes

Biologia Plantarum, 2007,51(2):381-385.

DOI:10.1007/s10535-007-0080-2      URL     [本文引用: 1]

The functional activities of the photosynthetic apparatus of two grapevine genotypes ( Vitis vinifera L. cvs. Müller-Thurgau and Lagrein) were investigated after low night temperature (LNT) treatment for 7 d. LNT caused important reductions of the net photosynthetic rate (P N ) of Lagrein plants due to non-stomatal components. These non-stomatal effects were not evident in Müller-Thurgau. At LNT treatment, the contents of photosynthetic pigments decreased significantly in Lagrein, but in Müller-Thurgau the contents of chlorophyll (Chl) remained unchanged whereas the contents of carotenoids (Car) increased. An increase and decrease of Chl a/b was shown in Mü ller-Thurgau and Lagrein stressed plants, respectively. RuBPC activity and content of soluble proteins were also significantly reduced in Lagrein. Under LNT treatment, photosystem (PS) 2 was markedly more inhibited in Lagrein than in Müller-Thurgau. The decrease in PS 2 activity in Lagrein was mostly due to the marked loss of D1, 47, 43, 33, 28-25, 23 and 17 kDa proteins determined by immunological and SDS-PAGE studies.

Souza T C D, Magalhães P C, Castro E M D , et al.

The influence of ABA on water relation,photosynthesis parameters,and chlorophyll fluorescence under drought conditions in two maize hybrids with contrasting drought resistance

Acta Physiologiae Plantarum, 2013,35(2):515-527.

DOI:10.1007/s11738-012-1093-9      URL     [本文引用: 2]

AbstractDrought is a major limitation of maize cultivation in Brazil. Agronomic and physiological practices have been considered to overcome this stress and consequently, increase grain production. The present study investigated the role of abscisic acid (ABA) application in some physiological parameters, in two hybrids with contrasting drought resistance (DKB 390 and BRS 1030 resistant and sensitive, respectively). Contrasting resistance to drought in these genotypes was determined in previous studies. Water deficit was imposed for 10 days at flowering stage, in association with the application of 100 M abscisic acid on plant canopy. Evaluations of gas exchange, chlorophyll fluorescence, relative water content (RWC), and endogenous ABA content were performed during stress period and also at water recovery (recovery irrigation). A significant functional relationship was observed between RWC and the parameters of gas exchange and fluorescence. During water recovery, no differences were observed among the treatments. DKB 390 presented higher photosynthesis rate () and electron transport rate (ETR) under water stress, while BRS 1030 presented higher intercellular CO concentration () and lower photochemical quenching (qP), non-photochemical quenching (NPQ), and lower / ratio. DBK 390 was more responsive to ABA application than BRS 1030, presenting higher endogenous ABA content in the first day of stress. DBK 390 with ABA application reduced the effect of water stress through maintenance of water status, an increase of photosynthetic parameters, and a decrease of decline in the functions of photosystem II during stress.

Baker N R .

Chlorophyll fluorescence:a probe of photosynthesis in vivo

Annual Review of Plant Biology, 2008,59(1):89-113.

DOI:10.1146/annurev.arplant.59.032607.092759      URL     [本文引用: 1]

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