根灌乙酸及葡萄酒对海水胁迫下葡萄光抑制的影响

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王辉^,, 高玉录, 于梦, 杜远鹏, 孙永江^,, 翟衡^,山东农业大学园艺科学与工程学院/作物生物学国家重点实验室,山东泰安 271018

Effect of Root Irrigation of Acetic Acid and Wine on Photoinhibition of Grape Under Seawater Stress

WANG Hui^,, GAO YuLu, YU Meng, DU YuanPeng, SUN YongJiang^,, ZHAI Heng^,College of Horticultural Science and Engineering, Shandong Agricultural University/State Key Laboratory of Crop Biology, Taian 271018, Shandong

通讯作者: 王辉,E-mail： 1924412287@qq.com。孙永江,Tel：0538-8241335;E-mail： sunhyongjiang12@163.com。翟衡,Tel：0538-8241335;E-mail： zhaih@sdau.edu.cn

责任编辑: 岳梅
收稿日期:2018-05-7接受日期:2018-08-1网络出版日期:2018-11-01

基金资助:

国家现代农业产业技术体系建设专项.CARS-29-zp-2
山东省重大科技创新工程.2018CXG0306
山东省“双一流”建设奖补资金.SYL2017YSTD10

Received:2018-05-7Accepted:2018-08-1Online:2018-11-01

摘要
【目的】为了扩大盐渍化土壤的高效利用方式,提高葡萄在盐渍化土壤上的生长发育质量,探讨乙酸与葡萄酒对15%海水胁迫下葡萄叶片光抑制的影响,为盐渍化土壤葡萄栽培技术改良提供理论依据和技术参考。【方法】以一年生‘摩尔多瓦’葡萄(‘Moldova’)盆栽苗为试材,定植于直径为20 cm、高度为18 cm的塑料盆内,每盆土和基质的比例为1﹕1,置于日光温室内。培养条件为日均温在25℃左右,温室内透光率不低于自然光照强度的50%,其中最高光强在800 μmol·m ^-2·s ^-1,相对湿度在40%—60%,正常肥水管理至8—10片完全展开叶时,进行海水和外源乙酸处理。进行15%海水浇灌的同时,分别浇灌30 mmol·L ^-1乙酸、2%葡萄酒,以浇灌清水为对照,每个处理重复5盆,各处理均浇灌3次,每隔2 d浇灌一次,浇灌量为基质持水量的2倍,约1/3的溶液流出, 每次浇灌后第2天测定叶绿素荧光指标,处理到第10天后,解析植株取样分析二者对海水胁迫下丙二醛(MDA)含量、叶绿素含量以及根系活力的影响。【结果】15%海水灌溉显著增加了葡萄各器官的MDA含量,显著降低了葡萄叶片叶绿素含量和根系活力,其中,与清水对照相比,根、茎、叶中MDA含量分别提高了1.10、0.27、0.41倍,叶绿素含量降低了18.5%,根系活力降低了41.9%,而浇灌30 mmol·L ^-1乙酸及2%葡萄酒处理显著降低了葡萄各器官中的MDA含量,其中,浇灌乙酸处理的根、茎、叶中MDA含量比海水胁迫分别降低了29.3%、20.6%、15.8%;浇灌葡萄酒处理的根、茎、叶中MDA含量分别比海水胁迫的降低了29.4%、20.2%、25.2%。浇灌乙酸及葡萄酒处理显著提高了葡萄叶片的叶绿素含量和根系活力,其中,根系活力分别比海水胁迫处理的显著提高了68.4%和56.9%,叶绿素含量则分别提高了18.8%和20.3%。叶绿素荧光分析表明,15%海水灌溉导致叶片光系统II（PSII）最大光化学效率(F_v/F_m)及最大光氧化P700 (P_m)明显下降,其中第3、6、9天的P_m分别比对照下降了5.0%、9.6%、13.0%,光化学淬灭系数(qP)和PSII实际光化学效率Y(II)逐渐下降,而PSII处可调节性能量耗散的量子产额Y(NPQ)及非调节性能量耗散的量子产额Y(NO)均逐渐升高,PSI供体端侧抑制程度Y(ND)未发生明显变化,但是PSI受体侧抑制程度Y(NA)逐渐升高,导致PSI的实际光化学效率Y(I)降低。根灌乙酸及葡萄酒可以显著缓解15%海水胁迫,PSII及PSI实际光化学效率显著升高,而用于热耗散的能量分配明显降低,从而提高了葡萄叶片的光合性能,其中以2%葡萄酒处理缓解效果较明显。【结论】根灌乙酸及葡萄酒显著提高15%海水胁迫下葡萄根系活力及叶绿素含量,缓解叶片光抑制程度,从而提高了葡萄对海水胁迫环境下的适应性,研究结果为残次果加工果酒、果渣及醋等的综合利用改良盐渍化土壤提供了理论依据。
关键词： 葡萄;海水;光抑制;乙酸;葡萄酒;叶绿素荧光

Abstract

【Objective】In order to expand the effective utilization of salinized soil and improve the quality of grape growth and development in salinized soil, the effects of acetic acid and wine on leaf photoinhibition under 15% seawater stress were studied. The result will provide a theoretical basis and technical reference for the improvement of grape cultivation techniques in salinized soil. 【Method】One-year old potted ‘Moldova’ seedlings were grown in plastic pots (with a diameter of 20 cm, and a height of 18 cm). The ratio of soil to substrate in each pot was 1﹕1 and the pots were placed in greenhouse. The culture conditions of the greenhouse were as follows: the average daily temperature was about 25℃, the light transmittance was more than 50% of natural light intensity, the maximum light intensity was 800 μmol·m ^-2·s ^-1, the relative humidity was 40%-60%, and the normal fertilizer and water was managed until 8-10 pieces of fully expanded leaves, then the seawater and exogenous acetic acid were used. The plants were irrigated with 15% seawater, 30 mmol·L ^-1 exogenous acetic acid and 2% wine at the same time, each treatment repeated 5 times, the controls were treated with clear water. Each treatment was irrigated every 2 days and 3 times, the irrigation amount was 2 times of water holding capacity, and about 1/3 of the solution leaked out, chlorophyll fluorescence indicators were determined on the 2nd day of irrigation, after 10 days treatment, the effects of the two methods on malondialdehyde (MDA) content, chlorophyll content, and root activity under seawater stress were analyzed.【Result】15% seawater treatment significantly decreased the chlorophyll content of leaf and root activity, but increased MDA content of root, stem and leaf. Compared with clear water control, the chlorophyll content decreased by 18.5% and root activity decreased by 41.9%. MDA content of root, stem and leaf increased by 1.10, 0.27 and 0.41 times, respectively. However, 30 mmol·L ^-1 acetic acid and 2% wine treatments significantly decreased the MDA content in grape. The MDA content in root, stem and leaf of grape treated with acetic acid decreased by 29.3%, 20.6% and 15.8%, respectively, compared with that of seawater stress, and the MDA content in root, stem and leaf of grape treated with wine decreased by 29.4%, 20.2% and 25.2%, respectively, compared with that of seawater stress. Irrigated with acetic acid and wine significantly increased the leaf chlorophyll content and root activity. Compared with seawater stress treatment, the root activity increased by 68.4% and 56.9%, and the chlorophyll content increased by 18.8% and 20.3%, respectively. The analysis of chlorophyll fluorescence showed that 15% seawater treatment decreased the levels of photosystem II (PSII) maximum photochemical efficiency (F_v/F_m) and the maximum photo-oxidizable P700 (P_m). The Pm on the 3rd, 6th, and 9th day decreased by 5.0%, 9.6% and 13.0% compared with the control, the levels of photochemical quenching coefficient (qP) and PSII actual photochemical efficiency (YII) decreased gradually, but the quantum yield of regulated energy dissipation Y (NPQ) and the quantum yield of non-regulated energy dissipation Y (NO) increased gradually. The nonphotochemical quantum yield due to PSI donor side limitation Y (ND) changed little, but the nonphotochemical quantum yield due to PSI acceptor side limitation Y (NA) increased gradually, resulting in the decrease of PSI actual photochemical efficiency Y (I). The irrigation of 30 mmol·L ^-1 acetic acid and 2% wine significantly relieved the stress of 15% seawater. The actual photochemical efficiency of PSI and PSII increased significantly, meanwhile the energy dissipated through heat decreased, then enhanced the efficiency of light energy utilization of grape leaves, in which 2% wine treatment showed a more obvious alleviated effect.【Conclusion】Irrigation of acetic acid and wine significantly improved root activity and chlorophyll content, which alleviated the photoinhibition level and improved the adaptability of grape to saline environment. The results provided a theoretical basis for the use of extract of wine lees to improvement of coastal saline soil.

Keywords：grape;seawater;photoinhibition;acetic acid;wine;chlorophyll fluorescence

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本文引用格式
王辉, 高玉录, 于梦, 杜远鹏, 孙永江, 翟衡. 根灌乙酸及葡萄酒对海水胁迫下葡萄光抑制的影响[J]. 中国农业科学, 2018, 51(21): 4210-4218 doi:10.3864/j.issn.0578-1752.2018.21.019
WANG Hui, GAO YuLu, YU Meng, DU YuanPeng, SUN YongJiang, ZHAI Heng. Effect of Root Irrigation of Acetic Acid and Wine on Photoinhibition of Grape Under Seawater Stress[J]. Scientia Acricultura Sinica, 2018, 51(21): 4210-4218 doi:10.3864/j.issn.0578-1752.2018.21.019

0 引言

【研究意义】我国耕地中盐渍化面积已达到920.9万公顷^[1]。土壤盐渍化是影响农业生产和生态环境的全球性问题^[2],盐渍化耕地中积累的盐主要来自海水、盐碱湖或灌溉水中的氯化钠^[3]。随着国家把黄河三角洲开发上升为国家战略,滨海盐渍化土壤改良与高效经济作物的适应性栽培驯化又重新提上研究日程,采用工程技术进行排盐和洗盐地,降低地下水位,避免海水倒灌是滨海盐渍化土地改良及利用的基础,选择耐盐的经济作物,进行耐盐育种及采用一定的技术措施等成为改良土壤盐渍化的发展方向^[4]。研究乙酸及葡萄酒缓解海水胁迫的响应机制对改良盐碱地具有重要意义,并可为充分利用各种残次水果及酒糟浸提液来改良盐碱地提供理论依据。【前人研究进展】盐胁迫中产生的离子毒害会降解叶绿素^[5,6],造成植物光合电子传递和光系统活力受到抑制^[7],研究也发现盐胁迫可以下调参与光反应和卡尔文循环的酶和蛋白活性^[8],导致光合受阻,严重的甚至导致植株死亡^[9]。对于盐渍化土地的改良,研究者也采取了一定的措施,如袁隆平培育成的‘海水稻’;利用栽培技术措施进一步提高了经济作物在盐渍化土壤上的生长发育,从而提高商品生产水平,如设施栽培、局部改良土壤的根域限制等。生长季节施加外源物质是缓解盐胁迫的一种有效栽培技术方式,较为常用的外源物质有渗透调节物质、激素类等^[10,11]。KIM等^[12]最新研究发现,通过浇灌乙酸可以显著缓解逆境对拟南芥、水稻、小麦、玉米和油菜生长发育的抑制,其结果为研究者提供了一个新思路,是否乙酸以及含有较高乙酸含量的葡萄酒也能够缓解盐胁迫?【本研究切入点】外源乙酸能够缓解植物受到的非生物胁迫,但乙酸及葡萄酒对海水胁迫下葡萄叶片光合作用的影响未见报道。【拟解决的关键问题】探讨乙酸以及含有较高乙酸含量的葡萄酒缓解葡萄盐胁迫的机制,以期为盐渍化土壤葡萄园栽培管理提供新思路。

1 材料与方法

试验于2017年在山东农业大学葡萄园内进行。

1.1 试材及处理

以一年生‘摩尔多瓦’葡萄（‘Moldova’,Guzali kala×SV.12-375）自根苗为试材,定植于直径为20 cm、高度为18 cm的塑料盆内,每盆土和基质的比例为1﹕1,置于日光温室内。培养条件：日均温在25℃左右,温室内透光率不低于自然光照强度的50%,其中最高光强在800 μmol·m^-2·s^-1,相对湿度在40%—60%,正常肥水管理至8—10片完全展开叶,此时叶片光合作用的饱和光强在1 100 μmol·m^-2·s^-1左右,进行海水和外源乙酸处理。选取无病虫害、长势一致的苗子分成4组,处理1为浇灌海水（取自渤海湾,用清水稀释到15% v/v,各种离子以mg·L^-1为单位,稀释后分别是Na⁺ 1 584,Cl^- 2 847,SO₄^2- 384,HCO₃^- 21.3,Ca²⁺ 45,Mg²⁺ 253.5,pH 7.88,电导率为12.32 ms·cm^-1,盐离子浓度为8.1 mg·L^-1）;处理2为15%海水+30 mmol·L^-1乙酸（购于天津市凯通化学试剂有限公司,预试验确定适宜浓度）;处理3为15%海水+2% v/v葡萄酒（葡萄酒产自笔者实验室葡萄园,其中乙酸的含量为0.43 g·L^-1,酸度为7.3 g·L^-1）,对照为清水。每个处理重复5盆,各处理均浇灌3次,每隔2 d浇灌一次,浇灌量为基质持水量的2倍,约1/3的溶液流出,每次浇灌后第2天测定叶绿素荧光指标（从上数第5片展开叶）,即第0、3、6、9天测定;处理到第10天,解析植株取样测定丙二醛（malondialdehyde,MDA）含量、根系活力和叶绿素含量,同时取土测定土壤pH等指标。

1.2 测定指标及方法

1.2.1 土壤pH的测定用土壤水浸液法,称取通过2 mm孔径筛的风干试样20 g（精确至0.1 g）于50 mL高型烧杯中,加入超纯水20 mL,以搅拌器搅拌1 min,使土粒充分分散,放置30 min用PB-10型pH计（北京赛多利斯科学仪器有限公司）进行测定。

1.2.2 土壤容重的测定用环刀法测定,100 cm³取样烘干后,按公式容重=干土重（g）/体积（cm³）计算。

1.2.3 土壤含水量的测定采用HBN-BXSF型土壤水分速测仪进行测定。

1.2.4 丙二醛含量、根系活力和叶绿素含量的测定参照赵世杰等^[13]的硫代巴比妥酸比色法测定MDA含量;利用氯化三苯基四氮唑法测定根系活力;按照赵世杰等^[13]方法计算叶绿素含量。

1.2.5 叶绿素荧光参数的测定使用Dual-PAM100叶绿素荧光仪（Walz,Germany）进行叶绿素荧光的活体测定,测定前对叶片进行15 min的暗适应。先测定初始荧光产量（F_o）与最大荧光产量（F_m）,之后远红光照射10 s后,测定P700最大量子产量（P_m）,对光下完全光启动的叶片打光强度为600 μmol·m^-2·s^-1的光化光,待荧光信号达到稳态后（大约4—5 min）打开饱和脉冲光（8 000 μmol·m^-2·s^-1）,测定任意时间的实际荧光产量（F_t）、光下最大量子产量（P_m′）和光适应下的最大荧光产量（F_m′）。根据以下公式进行各参数的计算：PSII最大光化学效率F_v/F_m=(F_m- F_o)/F_m,PSII光化学量子产量Y(II)=(F_m′-F_s)/F_m′,光化学淬灭系数qP=(F_m′-F_s)/(F_m'-F_o'),非调节性能量耗散Y(NO)=F_s/F_m,调节性能量耗散Y(NPQ)=F_s/F_m′-F_s/F_m,PSI光化学量子产量Y(I)=(P_m′-P)/P_m,PSI供体测抑制程度Y(ND)=P/P_m,受体测抑制程度Y(NA)=(P_m- P_m')/P_m^[14]。

1.3 数据统计与分析

数据测定取3次生物学重复,分别用Microsoft Excel和Sigma Plot处理数据和作图,用DPS软件对数据进行单因素方差分析。数据以平均值±标准误表示。

2 结果

2.1 浇灌乙酸及葡萄酒对海水胁迫下土壤理化性状的影响

土壤pH是衡量土壤酸碱性的重要指标,由表1可以看出,浇灌15%海水的土壤pH与清水对照相比升高了0.67%,但没有显著性影响,浇灌乙酸和葡萄酒的处理土壤pH仅分别比海水处理降低了1.5%和1.2%,也未出现显著性降低。对土壤容重以及土壤含水量的测定表明,浇灌15%海水后,与对照相比并未出现显著性变化,浇灌乙酸和葡萄酒后,也未出现显著性变化。

Table 1
表1
表1浇灌乙酸及葡萄酒对海水胁迫下土壤理化性状的影响
Table 1Effects of irrigating acetic acid and wine on soil physical and chemical properties under seawater stress

处理 Treatment	土壤pH Soil pH	土壤容重 Soil bulk density (g·cm^-2)	土壤含水量 Soil moisture content (%)
清水对照Clear water control	7.48±0.06a	0.89±0.04a	21.6±1.6a
海水Seawater	7.53±0.08a	0.94±0.03a	24.6±3.4a
乙酸+海水Acetic acid + seawater	7.42±0.06a	0.86±0.09a	23.0±1.1a
葡萄酒+海水Wine + seawater	7.44±0.06a	0.83±0.04a	22.5±3.4a

Different lowercases in the same column indicate significant difference among treatments (P<0.05). The same as below
同列数据后不同小写字母表示处理间差异显著（P<0.05）。下同

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2.2 浇灌乙酸及葡萄酒对海水胁迫下葡萄根、茎、叶中丙二醛含量的影响

丙二醛（MDA）是膜脂过氧化作用的产物之一,可以作为鉴定植物遭受盐胁迫程度的生理指标。由表2可见,浇灌3次15%海水后对‘摩尔多瓦’葡萄造成了明显胁迫,显著增加了各器官MDA含量,根、茎、叶中MDA含量分别是清水对照的2.10、1.27、1.41倍,而根灌乙酸及葡萄酒均显著缓解了海水胁迫下葡萄根、茎、叶中MDA的含量,其中,浇灌乙酸比海水胁迫分别降低了29.3%、20.6%、15.8%;浇灌葡萄酒比海水胁迫分别降低了29.4%、20.2%、25.2%。

Table 2
表2
表2浇灌乙酸及葡萄酒对海水胁迫下葡萄根、茎、叶中丙二醛含量的影响
Table 2Effects of irrigating acetic acid and wine on malondialdehyde content in grape root, stem and leaf under seawater stress

处理 Treatment	丙二醛含量Malondialdehyde content (μmol·g^-1FW )
处理 Treatment	根Root	茎Stem	叶Leaf
清水对照Clear water control	6.76±0.06c	8.34±0.04b	12.44±0.04d
海水Seawater	14.20±0.03a	10.58±0.09a	17.52±0.09a
乙酸+海水Acetic acid + seawater	10.04±0.03b	8.40±0.03b	14.76±0.07b
葡萄酒+海水Wine + seawater	10.02±0.03b	8.44±0.06b	13.10±0.09c

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2.3 浇灌乙酸及葡萄酒对海水胁迫下葡萄根系活力的影响

由图1可以看出,与清水对照相比,浇灌3次15%海水显著降低了葡萄的根系活力,比对照降低了41.9%。根灌30 mmol·L^-1乙酸及2%葡萄酒浇灌均显著升高了根系活力,与15%海水处理相比,分别提高了68.4%、56.9%。

图1

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图1浇灌乙酸及葡萄酒处理对海水胁迫下葡萄根系活力的影响

C：清水对照Clear water control;S：海水Seawater;A+S：乙酸+海水Acetic acid + seawater;W+S：葡萄酒+海水Wine + seawater。下同The same as below
Fig. 1Effects of irrigating acetic acid and grape wine treatment on grape root activity under seawater stress

2.4 浇灌乙酸及葡萄酒对海水胁迫下葡萄叶片叶绿素含量的影响

叶绿素含量是反映植物光合能力的重要指标^[15]。图2表明,15%海水胁迫显著降低了葡萄叶片中叶绿素含量,与对照相比降低了18.5%,而根灌乙酸和葡萄酒后叶绿素含量与海水处理相比分别提高了18.8%、20.3%,与清水对照相比没有显著差异。

图2

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图2浇灌乙酸及葡萄酒处理对海水胁迫下葡萄叶片叶绿素含量的影响

Fig. 2Effects of irrigating acetic acid and grape wine treatment on chlorophyll content in grape leaves under seawater stress

2.5 浇灌乙酸及葡萄酒对海水胁迫下葡萄叶片叶绿素荧光的影响

叶绿素荧光是反映植物光合性能的有效探针,最大光化学效率（F_v/F_m）反映了光系统Ⅱ（photosystem II, PSII）反应中心受损情况,P700最大量子产额（P_m）可以反映PSI活性^[16]。如图3所示,15%海水处理下P_m、F_v/F_m呈现下降趋势。其中第3、6、9天的P_m分别比对照下降了5.0%、9.6%、13.0%;F_v/F_m在海水胁迫后第3天出现明显下降,根灌乙酸和葡萄酒处理均缓解了F_v/F_m和P_m的降低幅度,表明根灌乙酸和葡萄酒可以有效缓解15%海水胁迫下葡萄叶片的光抑制程度,其中以葡萄酒处理效果较明显。

图3

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图3浇灌乙酸及葡萄酒处理对海水胁迫下葡萄叶片F_v/F_m、P_m的影响

Fig. 3Effects of irrigating acetic acid and grape wine treatment on F_v/F_m and P_m in grape leaves under seawater stress

2.6 浇灌乙酸及葡萄酒处理对海水胁迫下葡萄叶片能量分配的影响

如图4所示,15%海水胁迫明显降低了葡萄叶片光下最大光化学效率（F_v′/F_m′）（图4-A）,并且光化学淬灭系数（qP）（图4-B）逐渐降低,表明PSII反应中心在海水胁迫环境中处于关闭状态,从而导致用于光化学途径的能量减少,光合作用受到抑制。根灌乙酸及葡萄酒的F_v′/F_m′、qP与海水胁迫相比均明显提高,说明乙酸及葡萄酒可以提高PSII的光化学效率。15%海水灌溉也改变了葡萄叶片PSII和PSI的能量分配,与对照相比,15%海水灌溉下PSII实际光化学效率[Y(II)]（图4-C）明显下降,但可调节性能量耗散[Y(NPQ)]（图4-G）及非调节性耗散[Y(NO)]（图4-E）均明显升高;PSI供体端侧抑制程度[Y(ND)]（图4-H）发生明显变化,但PSI受体侧抑制程度[Y(NA)]（图4-F）逐渐升高,导致PSI的实际光化学效率[Y(I)]（图4-D）降低。根灌乙酸和葡萄酒明显提高了15%海水灌溉下葡萄叶片的Y(II)及Y(I),降低了Y(NPQ)及Y(NO)水平。进一步表明乙酸及葡萄酒可以缓解15%海水胁迫下葡萄叶片光抑制程度,从而改善能量的分配比例。

图4

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图4浇灌乙酸及葡萄酒处理对海水胁迫下葡萄叶片能量分配的影响

Fig. 4Effects of irrigating acetic acid and grape wine treatment on energy distribution in grape leaves under seawater stress

3 讨论

众所周知,乙酸是一种简单的基本生物化合物,由于含有的羧基具有较强的络合作用,对土壤理化性质、矿物溶解和成土作用、根际营养、重金属毒害等方面产生着深刻的影响,同时也在植物化感作用方面起着重要的作用^[17]。有研究发现,土壤中也存在乙酸^[18],且在一定浓度范围内土壤中乙酸浓度越高植物体内累积的钠、钾离子浓度越低^[19],而作为协调植物生存能力的初始因子,将植物的基本代谢、表观遗传调控和激素信号传导联系起来,最终赋予植物抗逆性,KIM等^[12]在拟南芥及水稻上的研究表明,外源乙酸对逆境胁迫的缓解作用与茉莉酸（JA）信号途径有关,通过促进JA合成和组蛋白H4乙酰化的富集,从而影响JA信号通路对植物耐逆境性能的启动。这种新颖的醋酸功能在进化上是保守的,是植物作为抵抗环境变化的生存策略。

由于乙酸同时具有的酸性特征,最直接的考虑是其在盐渍化土壤上的应用,前期调查发现,黄河三角洲盐渍化土壤上栽培的葡萄春季发芽晚,生长慢,叶色浅淡。前人研究表明,土壤盐胁迫抑制根系的生长发育,会引起细胞膜质过氧化^[20],增大膜通透性,破坏膜结构的完整性,降低根系活力。本研究发现海水胁迫导致葡萄根茎叶中的丙二醛含量均显著升高,根系活力下降,而浇灌乙酸和葡萄酒显著降低了各器官的膜脂过氧化,维持了较高的根系活力,与15%海水处理相比,分别提高了68.4%、56.9%。

盐胁迫对植物光系统活性产生抑制,严重的会发生光破坏,导致植物整株死亡^[21]。叶绿素是光合作用的物质基础,其含量的高低在一定程度上反应了光合作用的强弱^[22],盐胁迫对叶绿素含量的影响因植物种类、盐类型、处理浓度等因素的不同而异,其分别表现为下降^[23]、升高^[24]或先升高后降低的变化趋势^[25,26]。本研究表明,15%海水胁迫显著降低了叶绿素的含量,可能因为海水胁迫下植物细胞色素系统遭到破坏,叶绿素酶活性提高,从而加速了叶绿素的降解^[27,28]。当植物吸收的光能超过所能利用的光能时,过剩光能便会导致光合结构的光抑制,甚至光破坏^[29]。本研究中,15%海水处理导致了叶片最大光化学效率（F_v/F_m）及P700量子产额（P_m）的显著降低,而浇灌乙酸及葡萄酒均明显提高了海水胁迫下叶片的荧光参数,缓解了PSII及PSI的光抑制程度^[30]。与清水对照相比,15%海水胁迫明显改变了葡萄光系统的光能分配,叶片可调节的能量耗散[Y(NPQ)]显著升高,一方面表明其接受的光强过剩,另一方面说明叶片启动了与高能态淬灭有关的叶黄素循环过程^[31,32,33],但是Y(NPQ)在海水胁迫条件下自身调节能力有限,并不能耗散掉所有的光能,随着盐胁迫时间的延长,过剩光能的比例越来越高,非调节性能量耗散[Y(NO)]逐渐升高,叶片发生严重光抑制^[34]。而浇灌乙酸及葡萄酒后,葡萄叶片Y(NPQ)与Y(NO)均维持在降低水平,从而提高了PSII的实际光化学效率[Y(II)]。

不同浓度海水浇灌会引起土壤中及植物体内离子的失衡^[35],合理的海水灌溉浓度不会引起土壤理化性状的显著改变^[36]。本研究中浇灌15%海水胁迫及用30 mmol·L^-1乙酸和2%葡萄酒缓解均未导致土壤pH、土壤容重和土壤含水量的显著变化,表明15%海水处理可能通过离子毒害进行伤害。

本研究中,浇灌乙酸及葡萄酒均显著降低了15%海水胁迫下葡萄各器官的膜脂过氧化程度,缓解了葡萄叶片的光抑制,促进了叶片吸收的光能用于光化学反应的比例,对以后用盐渍化土壤果园残次果酿造的葡萄酒、果酒或者果醋等的综合利用提供了理论支撑。

4 结论

15%海水灌溉导致葡萄根系活力及叶绿素含量下降,加剧膜脂过氧化程度,光系统活性受到了抑制。根灌乙酸及葡萄酒可以显著提高15%海水胁迫下根系活力及叶绿素含量,改善光系统能量分配并提高光能利用效率,从而缓解光抑制程度,提高了葡萄对盐环境的适应性。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

参考文献原文顺序
文献年度倒序
文中引用次数倒序
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Plant Science, 2016,244:1-7.

DOI:10.1016/j.plantsci.2015.11.009 URLPMID:26810448 [本文引用: 1]

The functions of jasmonic acid (JA) in various stress responses have been uncovered in details, but its role in salt tolerance remains unclear. Here, we characterize the function ofArabidopsislipoxygenase3 (LOX3), an enzyme involved in JA synthesis, in salt stress response. The transcriptional analysis indicated thatLOX3was dramatically induced under salt treatment. Compared with wild type, thelox3mutant exhibited hypersensitivity to salt stress in germination and different developmental stages. Interestingly, methyl jasmonate (MeJA) rescued the salt sensitivity phenotypes of thelox3mutant, suggesting the impairment of salinity response in the mutant may be mediated by JA. Furthermore, the lateral root number of thelox3mutant was similar with that in wild type under normal condition, but less than that in wild type during salt treatment, and this lateral root sensitivity phenotype was also complemented by exogenous MeJA. In addition, the measurement of oxylipins in thelox3mutant and the analysis on germination of the JA receptorcoi1mutant under salt stress supported that JA may regulate the early response to salinity. In conclusion, we characterized the novel function of LOX3 in salinity stress response, and found that the salt hypersensitivity of thelox3mutant can be complemented by MeJA, providing new evidence for the association between JA and salt tolerance.

[11]

JIANG C

, CUI Q

, FENG

, XU D

, LI C

, ZHENG Q

. Melatonin improves antioxidant capacity and ion homeostasis and enhances salt tolerance in maize seedlings
Acta Physiologiae Plantarum, 2016,38:82.

DOI:10.1007/s11738-016-2101-2 URL [本文引用: 1]

Melatonin (N-acetyl-5-methoxytryptamine) is a biological hormone involved in many important physiological processes in plants. To better understand the protective roles of melatonin in plants against salt stress, we determined the effects of exogenous melatonin (10208M) on plant growth, photosynthesis, antioxidant enzyme activity and ion homeostasis in maize plants under 10002mM NaCl. Results showed that salt stress decreased plant dry matter accumulation, net photosynthetic rate (P 63) and chlorophyll contents. However, 10208M melatonin application significantly alleviated this growth inhibition and enhanced P 63 by 1902%. Melatonin application also enhanced the activities of antioxidative enzymes of salt-stressed maize leaves, and decreased their electrolyte leakage and MDA content by 25 and 2202%, respectively. In addition, melatonin application significantly increased K62 contents and K62/Na62 ratios in salt-stressed maize shoots by 18 and 5202%, respectively. However, the Na62 content was decreased significantly in melatonin-treated leaves under salinity. The results suggested that the melatonin enhanced maize salt tolerance in maize were most likely due to the improvement of photosynthetic capacity, antioxidative capacity and ion homeostasis in leaves. This study provides, for the first time, the evidence that support the protective roles of exogenous melatonin in maize against salinity.

[12]

KIM J

, TO T

, MATSUI

, TANOI

, KOBAYASHI N

, MATSUDA

, HABU

, OGAWA

, SAKAMOTO

, MATSUNAGA

, BASHIR

, RASHEED

, ANDO

, TAKEDA

, KAWAURA

, KUSANO

, FUKUSHIMA

, ENDO T

, KUROMORI

, ISHIDA

, MOROSAWA

, TANAKA

, TORII

, TAKEBAYASHI

, SAKAKIBARA

, OGIHARA

, SAITO

, SHINOZAKI

, DEVOTO

, SEKI

. Acetate-mediated novel survival strategy against drought in plants
Nature Plants, 2017, 3: Article number 17097.

DOI:10.1038/nplants.2017.97 URLPMID:28650429 [本文引用: 2]

Water deficit caused by global climate changes seriously endangers the survival of organisms and crop productivity, and increases environmental deterioration(1,2). Plants' resistance to drought involves global reprogramming of transcription, cellular metabolism, hormone signalling and chromatin modification(3-8). However, how these regulatory responses are coordinated via the various pathways, and the underlying mechanisms, are largely unknown. Herein, we report an essential drought-responsive network in which plants trigger a dynamic metabolic flux conversion from glycolysis into acetate synthesis to stimulate the jasmonate (JA) signalling pathway to confer drought tolerance. In Arabidopsis, the ON/OFF switching of this whole network is directly dependent on histone deacetylase HDA6. In addition, exogenous acetic acid promotes de novo JA synthesis and enrichment of histone H4 acetylation, which influences the priming of the JA signalling pathway for plant drought tolerance. This novel acetate function is evolutionarily conserved as a survival strategy against environmental changes in plants. Furthermore, the external application of acetic acid successfully enhanced the drought tolerance in Arabidopsis, rapeseed, maize, rice and wheat plants. Our findings highlight a radically new survival strategy that exploits an epigenetic switch of metabolic flux conversion and hormone signalling by which plants adapt to drought.

[13]

赵世杰, 史国安, 董新纯 . 植物生理学实验指导. 北京: 中国农业科学技术出版社, 2015: 55-57, 142-143.

[本文引用: 2]

ZHAO S

, SHI G

, DONG X

. Experimental Instruct of Plant Physiology. Beijing: China Agricultural Science and Technology Press, 2015: 55-57, 142-143. (in Chinese)

[本文引用: 2]

[14]

PFüNDEL

, KLUGHAMMER

, SCHREIBER

. Monitoring the effects of reduced PS II antenna size on quantum yields of photosystems I and II using the Dual-PAM-100 measuring system
PAM Application Notes, 2008,1:21-24.

[本文引用: 1]

[15]

蒋金豹, 陈云浩, 黄文江 . 用高光谱微分指数估测条锈病胁迫下小麦冠层叶绿素密度
光谱学与光谱分析, 2010,30(8):2243-2247.

URL [本文引用: 1]

通过人工田间诱发不同等级条锈病, 在不同生育期测定感染不同严重程度条锈病的冬小麦冠层光谱与冠层叶绿素密度(canopy chlorophyll density, CCD)。把CCD与高光谱指数进行相关性分析, 选取相关系数大于0.7的指数构建反演模型, 并对模型进行检验, 结果表明微分指数(D750-D550)/(D750+D550)反演精度以及稳定性最好, 其次是微分指数(D725-D702)/(D725+D702)。对上述两个微分指数分别进行饱和度分析, 发现当CCD大于12 μg·cm-2时微分指数(D750-D550)/(D750+D550)易达到饱和, 因此当CCD小于12 μg·cm-2时, 微分指数(D750-D550)/(D750+D550)反演CCD结果较好；但当CCD大于12μg·cm-2时, 利用微分指数(D725-D702)/(D725+D702)反演CCD较好, 该指数不易达到饱和状态。由于CCD与小麦病情指数(disease index, DI)之间存在极显著负相关性, 利用高光谱遥感精确估测小麦冠层CCD, 不仅可以帮助判断作物的长势, 而且可为识别小麦病害提供辅助信息。因此, 该研究对于农业防灾减灾也具有重要现实意义。

JIANG J

, CHEN Y

, HUANG W

. Using hyperspectral remote sensing to estimate canopy chlorophyll density of wheat under yellow rust stress
Spectroscopy and Spectral Analysis, 2010,30(8):2243-2247. (in Chinese)

URL [本文引用: 1]

HUANG

, FU P

, JIANG Y

, ZHANG J

, ZHANG S

, HU

, CAO K

. Differences in the responses of photosystem I and photosystem II of three tree species Cleistanthus sumatranus, Celtis philippensis and Pistacia weinmannifolia exposed to a prolonged drought in a tropical limestone forest.
Tree Physiology, 2013,33(2):211-220.

URL [本文引用: 1]

[17]

赵鹏志, 陈祥伟, 杨小燕, 齐思明, 王恩姮 . 低分子有机酸对东北黑土酶活性与养分关系的影响
南京林业大学学报(自然科学版), 2018,42(1):105-112.

URL [本文引用: 1]

ZHAO P

, CHEN X

, YANG X

, QI S

, WANG E

. Relationship between enzyme activities and nutrients of black soil subjected to low molecular organic acid
Journal of Nanjing Forestry University (Natural Sciences Edition), 2018,42(1):105-112. (in Chinese)

URL [本文引用: 1]

[18]

丁永祯, 李志安, 邹碧 . 土壤低分子量有机酸及其生态功能
土壤, 2005,37(3):243-250.

DOI:10.3321/j.issn:0253-9829.2005.03.004 URL [本文引用: 1]

本文综述了土壤低分子量有机酸的研究进展.它主要来源于动植物残体的分解、植物根系的分泌和微生物的合成,其组成复杂,主要包括脂肪族和芳香族两大类,含量一般为微摩尔至毫摩尔级.土壤低分子量有机酸具有重要的生态功能,如促进矿物溶解和成土作用;降低土壤根际环境pH;影响土壤微生物和酶的活性;促进根际环境养分转化,增加植物对P、Fe等营养元素的吸收;缓解Al离子对植物的毒害;影响重金属在土体中的迁移行为和在植物中的吸收量,以及具有重要的植物化感效应.文章最后提出了今后值得加强研究的方面.

DING Y

, LI Z

, ZOU

. Low-molecular-weight organic acids and their ecological roles in soil
Soils, 2005,37(3):243-250. (in Chinese)

DOI:10.3321/j.issn:0253-9829.2005.03.004 URL [本文引用: 1]

吴雪, 杨晓婷, 王冰, 王林, 疏伟慧, 张丽丽, 韩玉林 . 外源乙酸和EDTA对铜尾矿矿砂中芦苇幼苗生长及部分金属元素积累的影响
植物资源与环境学报, 2011,20(4):29-34.

[本文引用: 1]

, YANG X

, WANG

, SHU W

, ZHANG L

, HAN Y

. Effects of exogenous acetic acid and EDTA on growth and accumulation of some metal elements of Phragmites australis seedling in copper tailing ore.
Journal of Plant Resources and Environment, 2011,20(4):29-34. (in Chinese)

[本文引用: 1]

[20]

刘建新, 王鑫, 王瑞娟, 李东波 . 碱胁迫对黑麦草幼苗根系活性氧代谢和渗透溶质积累的影响
植物研究, 2011,31(6):674-679.

Magsci [本文引用: 1]

为了探讨牧草对碱胁迫的耐受程度，采用营养液砂培方法，研究了不同浓度NaHCO3（0、50、100、150和200 mmol·L-1）胁迫对黑麦草幼苗根系生长、活性氧代谢和渗透溶质积累的影响。结果表明：NaHCO3胁迫显著抑制黑麦草幼苗根系的生长，其抑制程度随胁迫浓度提高而增强，黑麦草可耐受的最高NaHCO3浓度约为150 mmol·L-1。随着NaHCO3胁迫浓度的增加，黑麦草根中超氧阴离子(O-·2)、过氧化氢(H2O2)和丙二醛(MDA)含量明显上升，超氧化物歧化酶(SOD)活性和谷胱甘肽(GSH)含量显著下降，过氧化氢酶(CAT)、过氧化物酶(POD)和抗坏血酸过氧化物酶(APX)活性及抗坏血酸(ASA)含量先升后降。黑麦草根中Na+含量随NaHCO3浓度增大而增加，K+含量和K+/Na+比降低，可溶性糖含量先升后降，脯氨酸含量则先降后升，游离氨基酸含量呈先升后降再升高变化。表明碱胁迫导致的活性氧代谢失调和Na+、K+失衡及积累有机溶质进行渗透调节时更多能量的消耗可能是黑麦草根系生长受抑的重要因素。

LIU J

, WANG

, WANG R

, LI D

. Effects of alkaline stress on the metabolism of reactive oxygen species and osmotic accumulation in ryegrass seedling roots
Bulletin of Botanical Research, 2011,31(6):674-679. (in Chinese)

Magsci [本文引用: 1]

王振兴, 吕海燕, 秦红艳, 赵滢, 刘迎雪, 艾军, 曹建冉, 杨义明, 沈育杰 . 盐碱胁迫对山葡萄光合特性及生长发育的影响
西北植物学报, 2017,37(2):339-345.

DOI:10.7606/j.issn.1000-4025.2017.02.0339 URL [本文引用: 1]

以棚内盆栽山葡萄品种‘双优’(抗逆性弱)和‘左山一’(抗逆性强)扦插苗为材料,利用气体交换分析和叶绿素荧光诱导动力学分析手段,研究了轻度、中度和重度盐碱胁迫对不同抗逆性山葡萄品种的光合特性及PSⅡ活性以及生长发育的影响,为山葡萄耐盐碱品种的评价提供理论依据。结果显示:(1)‘双优’各胁迫程度下的净光合速率(P_n)、气孔导度(Gs)和重度胁迫下的蒸腾速率(T_r)显著低于对照,而‘左山一’重度胁迫下的Pn和T_r、中重度胁迫下的Gs、各胁迫程度下的胞间二氧化碳浓度(Ci)显著低于对照,净光合速率下降主要归因于气孔限制和非气孔限制。(2)轻度、中度盐碱胁迫‘双优’叶片单位反应中心吸收的光能(ABS/RS)、单位面积内有活性反应中心数目(RS/CS)和吸收光能为基础的性能指数(PIABS),以及重度盐碱胁迫下‘左山一’的ABS/RS、RS/CS、用于电子传递的光能(ETo/CS)均显著低于相应对照,其余均无显著变化。(3)2品种山葡萄植株茎流率及株高、叶片数和根系生物量等生长指标在不同盐碱胁迫水平下均比对照显著降低,并以‘双优’品种降低幅度更大。研究表明,盐碱胁对山葡萄‘左山一’的叶片净光合速率、PSⅡ光能吸收和传递效率以及生长的抑制作用均小于‘双优’,‘左山一’的耐盐碱性更强。

WANG Z

, Lü H

, QIN H

, ZHAO

, LIU Y

, AI

, CAO J

, YANG Y

, SHEN Y

. Photosynthetic characteristics and growth development of Amur grape (Vitis amurensis Rupr.) under saline- alkali stress.
Acta Botanica Boreali-Occidentalia Sinica, 2017,37(2):339-345. (in Chinese)

DOI:10.7606/j.issn.1000-4025.2017.02.0339 URL [本文引用: 1]

王伟华, 张希明, 闫海龙, 梁少民, 杨小林 . 盐处理对多枝怪柳光合作用和渗调物质的影响
干旱区研究, 2009,26(4):561-568.

URL Magsci [本文引用: 1]

通过对人工控制条件下,不同浓度盐水(0,4,12,20,28 g/L)处理55 d后,多枝柽柳(Tamarix ramosissima Ledeb.)的气体交换参数、光合色素、脯氨酸和可溶性糖含量变化情况的分析.结果表明:①随盐水浓度的增加,多枝柽柳净光合速率(Pn)、气孔导度(Gs)的日变化曲线均由"双峰"型变为"单峰"型;Pn,Gs、胞间CO2浓度(Ci)日均值呈先上升后下降趋势,而气孔限制值(Ls)则相反;4 g/L浓度盐水处理促进多枝柽柳光合作用,12～28 g/L浓度盐水处理则使其因气孔因素限制而光合作用下降.②4 g/L浓度盐水处理显著促进多枝柽柳叶绿素a(chla)、叶绿素b(chlb)、总叶绿素(chl)、类胡萝卜素(car)的合成;随盐水浓度的增加,光合色素分解加剧,chla/b比值下降,而其car/chl值上升,且在20 g/L和28 g/L高盐水浓度处理下均达到极显著水平.③盐水处理使多枝柽柳可溶性糖含量增加,脯氨酸含量下降,但差异均不显著;随盐水浓度的增加,两物质含量均上升,并在高浓度盐水处理下达到极显著水平.较低盐水浓度(4 g/L)有利于多枝柽柳的生长,而随盐水浓度的升高均对其产生盐分胁迫.

WANG W

, ZHANG X

, YAN H

, LIANG S

, YANG X

. Effects of salt stress on photosynthesis and osmoregulation substance of Tamarix ramisissma Ledeb.
Arid Zone Research, 2009,26(4):561-568. (in Chinese)

URL Magsci [本文引用: 1]

王斌 . NaCl和Na₂SO₄胁迫下沼泽小叶桦的生理响应
南京林业大学学报 (自然科学版), 2013,37(1):132-136.

DOI:10.3969/j.issn.1000-2006.2013.01.024 URL [本文引用: 1]

以沼泽小叶桦为对象,分析了不同浓度NaCl和Na2SO4处理下沼泽小叶桦叶绿素和脯氨酸、可溶性糖、蛋白质等渗透调节物质含量的变化。结果表明：沼泽小叶桦在NaCl作用下其叶绿素、可溶性糖和脯氨酸含量的变化与Na2SO4胁迫下相似,随着盐浓度的增加,叶绿素含量显著下降,可溶性糖和脯氨酸含量均显著增加;叶片蛋白质含量在较低Na＋浓度下表现为先降低然后升高的趋势,在高浓度NaCl胁迫下蛋白质含量随着胁迫时间的增加逐渐下降;在相同浓度Na＋处理时,NaCl胁迫下叶片生理指标变幅比Na2SO4胁迫时大,这可能与沼泽小叶桦因长期生长在以硫酸盐为主的环境下已适应一定浓度的硫酸盐有关。

WANG

. Effects of NaCl and Na₂SO₄ stress on physiological characteristics in Betula microphylla.
Journal of Nanjing Forestry University (Natural Science Edition), 2013,37(1):132-136. (in Chinese)

DOI:10.3969/j.issn.1000-2006.2013.01.024 URL [本文引用: 1]

范希峰, 侯新村, 朱毅, 武菊英 . 盐胁迫对柳枝稷苗期生长和生理特性的影响
应用生态学报, 2012,23(6):1476-1480.

Magsci [本文引用: 1]

2010年,在人工气候室中设置了0、50、100、150和200 mmol·L-1 5种NaCl浓度处理,分析盐胁迫对柳枝稷苗期生长的影响.结果表明: 随着NaCl浓度的增加,柳枝稷的生长明显受到抑制,株高降低、叶片变小、光合叶面积减少、净光合速率下降,干物质积累量显著降低,表现出甜土植物的特点.柳枝稷的耐盐能力较强,在200 mmol·L-1NaCl溶液中处理30 d后仍能存活,单株绿叶面积为491.9 cm2,净光合速率为0.93 μmol CO2·m-2·s-1.本试验条件下,以生长量下降50%为标准求得柳枝稷的耐盐阈值为178.6 mmol·L-1.

FAN X

, HOU X

, ZHU

, WU J

. Impacts of salt stress on the growth and physiological characteristics of Panicum virgatum seedlings.
Chinese Journal of Applied Ecology, 2012,23(6):1476-1480. (in Chinese)

Magsci [本文引用: 1]

李宏, 邓江宇, 张振春, 杨森, 曹林 . 盐胁迫对盐桦幼树光合特性的影响
新疆农业科学, 2010,47(2):213-217.

URL [本文引用: 1]

【目的】盐胁迫对盐桦幼树光合特性的影响。【方法】利用土壤含盐量为1．0％、1．2％、1．4％、1．6％、1．8％、2．0％和2．2％七个盐浓度梯度，处理1年生盐桦实生苗，0％为对照CK，在生长的不同阶段分别测定叶片光合速率及其它生理指标。【结果】盐处理后，叶片净光合速率和气孔导度（Cond）均随NaCl浓度提高而显著降低；胞间C02浓度（Ci）、蒸腾速率Tr，随NaCl浓度提高呈先降低后升高的趋势；试验中1．6％NaCl胁迫下，盐桦幼树叶绿素含量提高，而1．6％。2．2％NaCl胁迫使叶绿素含量开始降低。【结论】NaCl胁迫下叶绿素含量提高是由于Na^＋的吸收促进了生长，也有利于叶绿素的合成；1．4％-1．6％NaCl胁迫下叶绿素含量提高是由于盐胁迫下叶片生长缓慢，导致叶绿体收缩，产生“浓缩”效应；1．4％NaCl胁迫下叶绿素含量仍然提高则可能是NaCl胁迫提高了叶绿素酶的活性，使光合色素合成减少、分解增加，从而使光合色素易于提取而导致了测得的叶绿素含量提高。 ...全文

, DENG J

, ZHANG Z

, YANG

, CAO

. Influence of salt stress on physiological adaptability of young Betula halophila trees.
Xinjiang Agricultural Sciences, 2010,47(2):213-217. (in Chinese)

URL [本文引用: 1]

徐静, 董宽虎, 高文俊, 谢开云 . NaCl和Na₂SO₄胁迫下冰草幼苗的生长及生理响应
中国草地学报, 2011,33(1):36-41.

URL [本文引用: 1]

在温室条件下用不同浓度的NaCl和Na2SO4（60 mmol/L、120 mmol/L、180 mmol/L、240 mmol/L、300 mmol/L）对冰草进行胁迫,结果表明：在两种盐处理下随着胁迫浓度递增,冰草的MDA含量增加,生物量、根冠比、叶绿素含量、脯氨酸含量和POD活性都呈先增后降的变化趋势;胁迫浓度为300 mmol/L时,冰草幼苗的生物量、根冠比、叶绿素含量、POD活性和脯氨酸含量显著下降,MDA含量达到最大值。120 mmol/L以下的低浓度盐胁迫对冰草生长有一定的促进作用,NaCl对冰草的胁迫作用大于Na2SO4。

, DONG K

, GAO W

, XIE K

. Growth and physiological responses of Agropyron cristatum seedlings under NaCl and Na₂SO₄ stress.
Chinese Journal of Grassland, 2011,33(1):36-41. (in Chinese)

URL [本文引用: 1]

SANTOS C

. Regulation of chlorophyll biosynthesis and degradation by salt stress in sunflower leaves
Scientia Horticulturae, 2004,103(1):93-99.

DOI:10.1016/j.scienta.2004.04.009 URL [本文引用: 1]

Helianthus annuus L. plants when exposed to increasing concentrations of NaCl showed a reduction of chlorophyll content and fluorescence (mainly due to a reduction of Fm). Chlorophyllase activity was higher in 15-day-old unstressed leaves. It increased during the first days in 25 mM NaCl stressed leaves decreasing with higher concentrations. NaCl effect on chlorophyll synthesis was then studied: only 50 and 100 mM NaCl inhibited synthesis of 5-aminolaevulinic acid, a precursor of chlorophyll. Our results show that: (1) chlorophyllase is stimulated during the first days in leaves under moderate stress but it is affected by high salt concentrations; (2) salt stress affects more drastically chlorophyll synthesis (decreasing ALA synthesis) than chlorophyllase-mediated degradation.

[28]

王斌, 巨波, 赵慧娟, 张群, 朱义, 崔心红 . 不同盐梯度处理下沼泽小叶桦的生理特征及叶片结构研究
林业科学, 2011,47(10):29-36.

DOI:10.11707/j.1001-7488.20111005 URL Magsci [本文引用: 1]

设计盐梯度(0,2&permil;,4&permil;,6&permil;,8&permil;盐溶液的质量比为NaCl ∶Na2SO4 ∶NaHCO3=75% ∶15% ∶10%)试验分析沼泽小叶桦生理表现和叶片解剖结构。结果显示: 在低于4&permil;盐度时沼泽小叶桦能正常生长,6&permil;盐度时沼泽小叶桦生长发育不良,当盐度8&permil;时沼泽小叶桦只能存活30天左右; 在低盐度处理下沼泽小叶桦光合生理表现和叶片解剖结构正常; 随着盐度增加,其叶片净光合速率、气孔导度、胞间CO2浓度、蒸腾速率、叶绿素含量等指标均下降,而叶片中可溶性糖含量、脯氨酸含量、蛋白质含量升高; 当盐度≥0.4%时,沼泽小叶桦叶片栅栏细胞逐渐纵向伸长且排列更紧密,叶绿体从椭圆形变成球形,叶绿体中出现粗大的淀粉粒和脂质球,表明其对滨海盐土具有一定的适应能力。

WANG

, JU

, ZHAO H

, ZHANG

, ZHU

, CUI X

. Photosynthetic performance and variation in leaf anatomic structure of Betula microphylla var. paludosa under different saline conditions.
Scientia Silvae Sinicae, 2011,47(10):29-36. (in Chinese)

DOI:10.11707/j.1001-7488.20111005 URL Magsci [本文引用: 1]

REINBOTHE

, REINBOTHE

. The regulating of enzymes involved in chlorophyll biosynthesis
European Journal of Biochemistry, 1996,237(2):323-343.

DOI:10.1111/j.1432-1033.1996.00323.x URLPMID:8647070 [本文引用: 1]

Abstract All living organisms contain tetrapyrroles. In plants, chlorophyll (chlorophyll a plus chlorophyll b ) is the most abundant and probably most important tetrapyrrole. It is involved in light absorption and energy transduction during photosynthesis. Chlorophyll is synthesized from the intact carbon skeleton of glutamate via the C 5 pathway. This pathway takes place in the chloroplast. It is the aim of this review to summarize the current knowledge on the biochemistry and molecular biology of the C 5 -pathway enzymes, their regulated expression in response to light, and the impact of chlorophyll biosynthesis on chloroplast development. Particular emphasis will be placed on the key regulatory steps of chlorophyll biosynthesis in higher plants, such as 5-aminolevulinic acid formation, the production of Mg 2+ -protoporphyrin IX, and light-dependent protochlorophyllide reduction.

[30]

HUANG

, YANG Y

, ZHANG J

, HU

, ZHANG S

. Superoxide generated in the chloroplast stroma causes photoinhibition of photosystem I in the shade-establishing tree species Psychotria henryi.
Photosynthesis Research, 2017,132(3):293-303.

DOI:10.1007/s11120-017-0389-4 URLPMID:28432538 [本文引用: 1]

Abstract Our previous studies indicated that high light induced significant photoinhibition of photosystem I (PSI) in the shade-establishing tree species Psychotria henryi. However, the underlying mechanism has not been fully clarified. In the present study, in order to investigate the mechanism of PSI photoinhibition in P. henryi, we treated detached leaves with constant high light in the presence of methyl viologen (MV) or a soluble -tocopherol analog, 2,2,5,7,8-pentamethyl-6-chromanol (PMC). We found that MV significantly depressed photochemical quantum yields in PSI and PSII when compared to PMC. On condition that no PSI photoinhibition happened, although cyclic electron flow (CEF) was abolished in the MV-treated samples, P700 oxidation ratio was maintain at higher levels than the PMC-treated samples. In the presence of PMC, PSI photoinhibition little changed but PSII photoinhibition was significantly alleviated. Importantly, PSI photoinhibition was largely accelerated in the presence of MV, which stimulates the production of superoxide and subsequently other reactive oxygen species at the chloroplast stroma by accepting electrons from PSI. Furthermore, MV largely aggravated PSII photoinhibition when compared to control. These results suggest that high P700 oxidation ratio cannot prevent PSI photoinhibition in P. henryi. Furthermore, the superoxide produced in the chloroplast stroma is critical for PSI photoinhibition in the higher plant P. henryi, which is opposite to the mechanism underlying PSI photoinhibition in Arabidopsis thaliana and spinach. These findings highlight a new mechanism of PSI photoinhibition in higher plants.

[31]

GIIMORE A

. Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves
Physiologia Plantarum, 1997,99:197-209.

DOI:10.1111/j.1399-3054.1997.tb03449.x URL [本文引用: 1]

Higher plants must dissipate absorbed light energy that exceeds the photosynthetic capacity to avoid molecular damage to the pigments and proteins that comprise the photosynthetic apparatus. Described in this minireview is a current view of the biochemical, biophysical and bioenergetic aspects of the primary photoprotective mechanism responsible for dissipating excess excitation energy as heat from photosystem II (PSII). The photoprotective heat dissipation is measured as nonphotochemical quenching (NPQ) of the PSII chlorophyll a (Chl a ) fluorescence. The NPQ mechanism is controlled by the trans-thylakoid membrane pH gradient ( pH) and the special xanthophyll cycle pigments. In the NPQ mechanism, the de-epoxidized endgroup moieties and the trans-thylakoid membrane orientations of antheraxanthin (A) and zeaxanthin ( Z ) strongly affect their interactions with protonated chlorophyll binding proteins (CPs) of the PSII inner antenna. The CP protonation sites and steps are influenced by proton domains sequestered within the proteo-lipid core of the thylakoid membrane. Xanthophyll cycle enrichment around the CPs may explain why changes in the peripheral PSII antenna size do not necessarily affect either the concentration of the xanthophyll cycle pigments on a per PSII unit basis or the NPQ mechanism. Recent time-resolved PSII Chi a fluorescence studies suggest the NPQ mechanism switches PSII units to an increased rate constant of heat dissipation in a series of steps that include xanthophyll de-epoxidation, CP-protonation and binding of the xanthophylls to the protonated CPs; the concerted process can be described with a simple two-step, pH-activation model. The xanthophyll cycle-dependent NPQ mechanism is profoundly influenced by temperatures suboptimal for photosynthesis via their effects on the trans-thylakoid membrane energy coupling system. Further, low temperature effects can be grouped into either short term (minutes to hours) or long term (days to seasonal) series of changes in the content and composition of the PSII pigment-proteins. This minireview concludes by briefly highlighting primary areas of future research interest regarding the NPQ mechanism.

[32]

NIYOGI K

, GROSSMAN A

, BJöRKMAN

. Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion
The Plant Cell, 1998,10(7):1121-1134.

[本文引用: 1]

[33]

YAMAMOTO H

, BUGOS R

, HIEBER A

. Biochemistry and molecular biology of the xanthophyll cycle//The Photochemistry of Carotenoids. Springer
Dordrecht, 1999: 293-303.

DOI:10.1007/0-306-48209-6_16 URL [本文引用: 1]

The xanthophyll cycle is the cyclical interconversion of violaxanthin, antheraxanthin and zeaxanthin in plants and green algae. The existence of the cycle has been known for many years but has attract

[34]

HORTON

, HAGUE

. Studies on the induction of chlorophyll fluorescence in isolated barley protoplasts. IV. Resolution of non-photochemical quenching
Biochimica et Biophysica Acta (BBA)- Bioenergetics, 1988,932:107-115.

DOI:10.1016/0005-2728(88)90144-2 URL [本文引用: 1]

Using DCMU addition and light-saturation pulses, chlorophyll fluorescence quenching by isolated barley protoplasts has been examined upon illumination with different light intensities. Total quenching was constant at all intensities, photochemical quenching predominating in low light and non-photochemical quenching becoming increasingly important at higher intensity. The components of non-photochemical quenching were resolved by examination of kinetics of relaxation of quenching upon DCMU addition or darkening. It was found that: (a) energy-dependent quenching saturated with characteristics similar to photosynthesis; (b) a slowly relaxing, NaF-sensitive component attributed to protein phosphorylation saturated in low light and decreased at high light; (c) an irreversible component ascribed to photoinhibition was a major quencher as light levels were increased above saturation; (d) the decreased photochemical quenching at high light was not strictly correlated with alteration in rate of O 2 evolution. Analysis of these data showed that the quantum yield of Photosystem II declined as the light intensity was increased from zero to that saturating for photosynthesis and was associated with the presence of energy-dependent quenching. The regulation of excitation dissipation by the thylakoid membrane is discussed.

[35]

寇伟锋, 刘兆普, 郑宏伟 . 海水胁迫对向日葵苗期生长及矿质营养吸收特性的影响
生态学杂志, 2006,25(5):521-525.

URL Magsci [本文引用: 1]

采用砂培法，研究了海水胁迫对向日葵幼苗生长及矿质营养吸收特性的影响。结果表明，海水胁迫下，向日葵幼苗株高、茎粗、干物重明显降低。幼苗根茎叶中Cl-，茎和叶中Mg2+、叶中Na+和Ca2+含量随海水浓度的增加而增加，根茎叶中K+、全氮和全磷含量随海水浓度升高而降低，但在10%和20%海水胁迫下，向日葵体内Na+、Cl-主要集中于根和茎中，叶中较少。海水胁迫下，向日葵幼苗各部位K+ / Na+始终是叶部最高，根部最低，且根茎叶中SK,Na值均大于1。因此，低浓度海水胁迫下向日葵幼苗对Na+ 和Cl-的截流作用、海水胁迫下幼苗根部对K+强的选择性吸收以及K+向地上部的选择性运输是向日葵具有一定耐盐性的主要原因。

KOU W

, LIU Z

, ZHENG H

. Effects of sea water stress on Helianthus annuus L. seedlings growth and mineral nutrition.
Chinese Journal of Ecology, 2006,25(5):521-525. (in Chinese)

URL Magsci [本文引用: 1]

王杨 . 长期海水灌溉对赤霞珠果实品质及土壤性质的影响
[D]. 泰安: 山东农业大学, 2016.

[本文引用: 1]

WANG

. The effect of long term seawater irrigation on the quality of Cabernet Sauvignon fruit and soil properties
[D]. Taian: Shandong Agricultural University, 2016. ( in Chinese)

[本文引用: 1]