外源NO对NaHCO3胁迫下黑麦草幼苗光合生理响应的调节

采用营养液砂培方法,研究了外源一氧化氮(NO)对100 mmol/L NaHCO3胁迫下黑麦草幼苗叶片叶绿素含量、光合气体交换和叶绿素荧光参数、光能分配及叶黄素循环的影响。结果表明:(1)外施60μmol/L NO供体硝普钠(SNP)显著缓解了NaHCO3胁迫下叶绿素含量、净光合速率(Pn)、气孔导度(Gs)和气孔限制值(Ls)的下降及胞间CO2浓度(Ci)的升高,提高了光系统Ⅱ(PSⅡ)的潜在活性(Fv/Fo)、最大光化学效率(Fv/Fm)、实际光化学效率(ΦPSⅡ)和光化学猝灭(qP),降低了初始荧光(Fo)和非光化学猝灭(NPQ)。(2)NaHCO3胁迫下,外施SNP显著抑制了天线转换效率(Fv’/Fm’)的下降,降低了光系统间激发能分配的不平衡性(β/α-1)和天线热耗散的比例(D),提高了吸收光能中用于光化学反应的比例(P),而对PSⅡ反应中心的过剩光能(Ex)无明显影响。(3)外施SNP显著降低了NaHCO3胁迫下叶黄素循环库(V+A+Z)下降和叶黄素循环脱环氧化状态(A+Z)/(V+A+Z)上升的幅度。但SNP对NaHCO3胁迫的缓解效应可被NO清除剂血红蛋白(Hb)部分或完全地逆转,SNP的分解产物NaNO2处理对NaHCO3胁迫无明显改善。表明外源NO可能通过提高光化学效率,缓解了碱胁迫引起的光抑制对光合机构的破坏,从而提高黑麦草的光合效率。     

濒危植物长序榆(Ulmus elongata)幼苗叶绿素荧光特性的初步研究

用Li-6400XT便携式光合作用仪对濒危植物长序榆幼苗的各叶绿素荧光参数的日变化和快速光响应曲线进行了测定。结果发现,光系统Ⅱ(PSⅡ)的实际光化学效率(ΦPSⅡ)、电子传递速率(ETR)在整个白天阶段较稳定,下午18:00显著下降。光化学淬灭(qP)先增大后减小。非光化学淬灭(NPQ)呈现出与光化学淬灭(qP)相反的变化趋势,中午最低,说明长序榆幼苗光能利用率较高。快速光曲线表明实际光化学效率(ΦPSⅡ)和光化学淬灭(qP)随着光合有效辐射(PAR)的增大而减小,电子传递速率(ETR)和非光化学淬灭(NPQ)随着光合有效辐射(PAR)的增大而增大。使用幂函数能够很好的拟合实际光化学效率(ΦPSⅡ)和电子传递速率(ETR)随光强的变化,而对数函数能较好的拟合实际光化学淬灭(qP)和非光化学淬灭(NPQ)随光强的变化。     

Effects of Exogenous Putrescine on Chlorophyll Fluorescence Imaging and Heat Dissipation Capacity in Cucumber (Cucumis sativus L.) Under Salt Stress

The objective of this study was to identify the effects of exogenous putrescine on photosynthetic performance and heat dissipation capacity in cucumber seedlings under salt stress. The stress of 75 mM NaCl for 7 days caused a significant decrease in net photosynthetic rate (P _N ). The experiment employed a chlorophyll fluorescence imaging technique and demonstrated that the maximal quantum yield of photosystem II photochemistry (Fv/Fm) and the actual photochemical efficiency of photosystem II (ΦPSII) were reduced by salt stress. Moreover, salt stress markedly reduced the photochemical quenching coefficient (qP) and non-photochemical quenching coefficient (qN), and significantly increased non-regulated heat dissipation (ΦNO). However, stressed plants supplied with exogenous putrescine exhibited higher P _N and ΦPSII, which indicated that putrescine can alleviate the detrimental effects on photosynthesis induced by salt stress. Putrescine sprayed on stressed plants significantly enhanced the regulated energy dissipation (ΦNPQ) and decreased ΦNO. Application of exogenous putrescine also changed the levels of xanthophyll cycle components and further enhanced the de-epoxidation state of xanthophyll cycle pigments under salt stress. Under control conditions, putrescine exerted little influence on the photosynthetic parameters in cucumber leaves. In conclusion, the application of exogenous putrescine may improve the heat dissipation capacity by promoting the xanthophyll cycle to reduce the damage caused by excess excitation energy, thus enhancing the salt tolerance of cucumber seedlings.     

外源ATP对NaCl胁迫下菜豆叶片叶绿素荧光特性的调节

盐胁迫是影响植物生长的主要逆境因子之一,外源ATP被发现可作为信号分子参与植物对逆境胁迫生理反应的调节。为了探明外源ATP在植物盐胁迫响应中的作用,以增强植物对土壤盐渍化的耐性,更好地应用于土壤盐渍化修复。该研究以菜豆( Phaseolus vulgaris)为材料,通过叶绿素荧光技术探讨了外源ATP 对菜豆叶片在NaCl胁迫下叶绿素荧光特性的变化规律。结果表明：在NaCl胁迫下,叶片光系统Ⅱ( PSⅡ)潜在最大光化学量子效率( Fv/Fm)、光适应下最大光化学效率( Fv′/Fm′)、PSⅡ光适应下实际光化学效率[ Y (Ⅱ)]、光化学荧光猝灭( qP)、电子传递速率( ETR)与对照组相比均有显著性下降,而非光化学猝灭( NPQ)和( qN)较对照组有显著性增加,这表明NaCl胁迫导致菜豆叶片光系统Ⅱ光化学效率的下降和光能耗散的增加。而外源ATP(eATP)的处理能有效缓解NaCl胁迫所造成的Fv/Fm、Fv′/Fm′、Y(Ⅱ)、qP、ETR下降和NPQ、qN的上升。该研究结果表明在NaCl胁迫下外源ATP可以有效地提高菜豆幼苗光系统Ⅱ( PSⅡ)的光化学反应效率。     

硝基苯酚胁迫下外源褪黑素对水稻幼苗生长及生理特性的影响

采用液体培养实验方法,研究硝基苯酚胁迫对水稻(Oryza sativa L.)幼苗生长、抗氧化特性、光系统Ⅱ(PSⅡ)光合特性的影响,以及添加外源褪黑素对缓解硝基苯酚胁迫的作用。结果显示,随着硝基苯酚胁迫浓度的升高,水稻幼苗株高、根长、地下部干重、地上部干重、全株干重和叶片PSⅡ实际光化学效率[Y(Ⅱ)]、光化学淬灭系数(q P)、PSⅡ电子传递速率(ETR)、叶绿素含量均有所下降,而叶片非光化学淬灭系数(qN、NPQ)上升;同时,根系活性氧[过氧化氢(H_2O_2)和超氧阴离子(O·-2)]积累量、抗氧化酶[超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)]活性,以及渗透调节物质(可溶性蛋白和可溶性糖)含量呈先升高后降低的趋势。在非硝基苯酚胁迫下,与对照组相比,添加外源褪黑素显著提高了幼苗地下部干重、根系可溶性糖含量和SOD活性、叶片PSⅡ光化学效率和叶绿素含量。与单独添加硝基苯酚处理相比,硝基苯酚+褪黑素复合处理显著缓解了硝基苯酚胁迫对幼苗生长、叶片PSⅡ光化学效率和叶绿素合成的抑制作用;降低了根系活性氧水平、抗氧化酶活性和渗透调节物质含量。研究结果表明添加外源褪黑素能够显著缓解硝基苯酚胁迫对水稻幼苗生长、根系活性氧水平、抗氧化酶活性、叶片PSⅡ光化学效率及叶绿素合成的不良影响,提高水稻幼苗对硝基苯酚胁迫的适应性。     

不同干旱胁迫下欧李光合及叶绿素荧光参数的响应

采用人为控制土壤含水量的方法对欧李进行轻度和重度干旱的处理,测定叶片的气体交换和叶绿素荧光参数的日变化。结果表明,干旱胁迫下欧李叶片净光合速率、蒸腾速率、水分利用效率、气孔导度、PSII最大光化学效率、光化学量子效率显著下降,但胞间CO2浓度、非光化学猝灭系数以及叶黄素循脱环氧化状态（Z＋0.5A）/（V＋A＋Z）和Z含量升高。两干旱处理植株的影响程度存在差异。这表明在长时间干旱条件下,欧李叶片光合作用的降低受到气孔与非气孔因素的双重影响,叶黄素循环的启动增加了胁迫条件下的热耗散能力以保护光合机构免受干旱胁迫的进一步伤害。     

木棉叶片叶绿素荧光参数和SPAD值对干旱胁迫的响应

采用盆栽控水干旱法模拟干旱逆境,对1年生木棉(Bombax ceiba)盆栽苗进行不同程度的胁迫处理,测定干旱胁迫时其叶片叶绿素SPAD值和荧光参数动态变化。结果表明,干旱胁迫下,木棉叶片叶绿素相对含量逐渐降低;初始荧光(Fo)随着干旱程度的加剧而呈上升趋势,而最大荧光(Fm)、潜在光化学效率(Fv /Fo)、最大光化学效率(Fv /Fm)、光化学淬灭系数(qP)和表观光合电子传递速率(ETR)下降,在中度和重度干旱胁迫时差异显著(P＜0.05);非光化学淬灭系数(NPQ)变化不显著。干旱胁迫导致木棉植株出现光抑制,植株可通过降低光化学淬灭、增加热耗散的形式增强对干旱的适应能力。     

Spermidine application enhances tomato seedling tolerance to salinity-alkalinity stress by modifying chloroplast antioxidant systems

The purpose of this study was to elucidate whether exogenous spermidine (Spd) protection of tomato (Solanum lycopersicum L.) seedlings under salinity-alkalinity stress is associated with antioxidant enzymes in the chloroplast. The effects of exogenous Spd on antioxidant enzyme activity and antioxidant content in the chloroplast were evaluated in seedlings of salt-sensitive ecotype (Zhongza 9) grown in a 75 mM salinity-alkalinity solution, with or without 0.25 mM Spd foliar spraying. Results showed that salinity-alkalinity stress increased MDA content, superoxide anion O₂^?- generation rate, superoxide dismutase (SOD), ascorbate peroxidase (APX), monodehydroascorbate reductase (MDHAR) activities and ratio of AsA/DHA and reduced contents of ascorbate (AsA), dehydroascorbate (DHA), AsA+DHA, glutathione (GSH), oxidized glutathione (GSSG), GSH+GSSG, dehydroascorbate reductase (DHAR) activity and ratio of GSH/GSSG in chloroplasts. The exogenous Spd application combined with salinity-alkalinity stress decreased the O₂^?- generation rate and MDA content compared to salinity-alkalinity stress alone. The exogenous Spd also increased AsA-GSH cycle components and increased all antioxidant enzyme activities in most cases. Therefore, exogenous Spd alleviates salinity-alkalinity stress damage using antioxidant enzymes and non-enzymatic systems in chloroplasts.     

麻疯树幼苗对干旱胁迫的响应

以不同浓度(5%～25%)的聚乙二醇(PEG-6000)模拟干旱胁迫处理麻疯树三叶期幼苗,研究了不同程度干旱胁迫下麻疯树叶片光合特性及其对干旱的耐受能力.结果表明:在较低浓度PEG(≤15%)处理下,随PEG浓度的增加,麻疯树叶片净光合速率(P_n)、气孔导度(G_s)、胞间CO₂浓度(C_i)、PSⅡ实际光化学量子产量(Φ_PSⅡ)、光化学猝灭(q_P)和表观光合电子传递速率(ETR)下降,PSⅡ原初光能转化效率(F_v/F_m)轻微下降,水分利用效率(WUE)则逐渐升高,非光化学猝灭(NPQ)明显上升,初始荧光(F_o)无显著变化(P＞0.05);在高浓度PEG(＞15%)处理下,C_i随PEG浓度的增加而显著上升,P_n、G_s和WUE持续下降,F_v/F_m、Φ_PSⅡ、q_P和ETR下降幅度明显增大,F_o显著上升,而NPQ下降.低浓度PEG处理导致麻疯树叶片P_n下降主要是由气孔因素造成的;在高浓度PEG处理下,P_n的下降则是由非气孔和气孔因素的共同限制作用造成的.当PEG浓度<20%时,虽然出现P_n下降,但光合机构未受损伤.经15 d高浓度PEG处理的植株叶片,在胁迫解除后光合活性能够迅速恢复,且植株可以存活.说明麻疯树对干旱胁迫有较强的耐受能力.     

重金属对盐生草光合生理生长特性的影响

以盐生草幼苗为试验材料,分别设置0(CK)、50、100、200、400μg?g-1的Ni2+、Cu2+处理,研究重金属Ni2+和Cu2+对盐生草光合生理特性的影响.结果表明:盐生草叶片光合色素含量、净光合速率(Pn)、气孔导度Gs、蒸腾速率Tr、PSⅡ最大光化学效率Fv/Fm、非光化学猝灭系数qN及生长指标(株高、地上部干重和鲜重)在50μg?g-1的Ni2+处理时均达到最大值,后随Ni2+浓度继续增加,其叶片叶绿素a、叶绿素b、Pn、Gs、Tr、Fv/Fm、PSⅡ电子传递量子产率ΦPSⅡ、光化学猝灭系数qP、qN及各项生长指标逐步下降并低于对照水平,而细胞间隙CO2浓度(Ci)较对照呈增加趋势.在50μg?g-1的Cu2+处理时,盐生草叶片光合色素含量、Pn、Gs、Tr、Ci、Fv/Fm、ΦPSⅡ、qP、qN及各项生长指标均达峰值;在100μg?g-1Cu2+处理时,光合色素含量、Pn、Gs、Tr、Fv/Fm、ΦPSⅡ、qN及各项生长指标较对照仍有增加,而后随Cu2+浓度继续增加,其叶绿素a、叶绿素b、各光合参数、叶绿素荧光参数及生长指标均逐步降低并低于对照.可见,盐生草Pn在Ni2+胁迫下的下降主要是由非气孔限制所致,而Cu2+胁迫下的下降主要是由气孔限制所致;低浓度Ni2+和Cu2+对盐生草生长具有一定促进作用,过高浓度Ni2+和Cu2+则会通过抑制盐生草叶片叶绿素合成,影响其光合作用,从而抑制植株生长.     

外源亚精胺对Ca(NO3)2胁迫下番茄幼苗光合特性和抗氧化酶活性的影响

以耐盐性较弱的番茄品种上海‘合作903’幼苗为试验材料,采用营养液栽培方法,研究了叶面喷施1mmol.L-1亚精胺(Spd)对75mmol.L-1 Ca(NO32)胁迫下番茄幼苗生长、叶绿素含量、光合及荧光参数、叶片抗氧化酶活性的影响,以探讨外源亚精胺缓解Ca(NO3)2胁迫伤害的机制。结果显示:Ca(NO3)2胁迫能够显著抑制番茄幼苗的生长;与Ca(NO32)胁迫处理相比,叶面喷施外源Spd 9d后,受胁迫番茄幼苗的株高、茎粗、干重、鲜重分别显著增加70.9%、15.8%、43.4%、41.4%;叶绿素a、b的含量分别提高17.7%、13.8%;净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、光合电子传递效率(rETR)和光化学猝灭系数(qP)分别升高6.6%、18.0%、31.0%、4.9%、5.0%,而胞间二氧化碳浓度(Ci)、非光化学猝灭系数(qN)分别降低21.5%、8.1%;超氧化物歧化酶(SOD)、过氧化氢酶(CAT)活性分别增加10.7%和37.5%,而丙二醛(MDA)含量和超氧阴离子(O2.-)产生速率分别显著下降34.5%、17.1%。研究表明,外源Spd通过提高番茄幼苗抗氧化酶活性来有效清除体内活性氧,维持光合机构的稳定性,提高其光合速率,从而缓解Ca(NO3)2胁迫对番茄幼苗的伤害。     

高温胁迫下外源NO对高灌蓝莓PSⅡ光化学活性和抗氧化系统的影响

以高灌蓝莓品种‘都克’试管苗移栽到田间6个月的植株为试材,研究了高温胁迫条件下,外源一氧化氮(NO)对高灌蓝莓幼苗生长、PSⅡ光化学活性和抗氧化系统的影响.结果表明:0.2、0.5和1.0 mmol·L^-1外源NO供体硝普钠(SNP)处理显著减缓了高温胁迫对高灌蓝莓光合系统Ⅱ(PSⅡ)的抑制,高灌蓝莓PSⅡ最大光化学效率(F_v/F_m)、实际光化学效率(Φ_PSⅡ)、光化学猝灭系数(q_P)和非光化学猝灭系数(NPQ)下降缓慢,使光合机构免受高温胁迫的伤害;叶片质膜透性和丙二醛含量降低,而超氧化物歧化酶和过氧化氢酶活性升高,从而促进了脯氨酸的积累.适当浓度的SNP能显著缓解高温胁迫对高灌蓝莓植株的伤害,其中0.5 mmol·L^-1 SNP的缓解效果最好.     

Effects of gamma-aminobutyric acid on the photosynthesis and chlorophyll fluorescence parameters of muskmelon seedlings under hypoxia stress

By the method of hydroponic culture, this paper studied the effects of exogenous gamma-aminobutyric acid (GABA) on the photosynthetic pigment contents, photosynthesis, and chlorophyll fluorescence parameters of muskmelon seedlings under hypoxia stress. Hypoxia stress induced a significant decrease of photosynthetic pigment contents, resulting in the decrease of photosynthesis. Applying GABA could significantly increase the photosynthetic pigment contents, net photosynthetic rate (P(n)), stomatal conductance (G(s)), intercellular CO2 concentration (C(i)), carboxylation efficiency (CE), maximal photochemical efficiency of PS II (F(v)/F(m)), photochemical quenching (q(P)), apparent photosynthetic electron transfer rate (ETR), and quantum yield of PS II electron transport (phi(PS II)), and decrease the stomatal limitation value (L(s)), minimal fluorescence (F(o)), and non-photochemical quenching (NPQ) under both hypoxic and normal conditions. The alleviation effect of GABA on photosynthetic characteristics was more obvious under hypoxia stress. However, simultaneously applying GABA and VGB could significantly decrease the alleviation effect of GABA under hypoxia stress.     

不同浓度NaCl和Na2CO3处理对菊芋幼苗光合及叶绿素荧光的影响

 以砂培菊芋(Helianthus tuberosus)幼苗作为试验材料,分别进行不同浓度NaCl (50、 100、150、200、250 mmol&;#8226;L^－1)和Na₂CO₃ (25、50、 75、100、125 mmol&;#8226;L^－1)胁迫处理,以1/2全营养液作为对照,处理7 d后研究NaCl和Na₂CO₃胁迫处理对菊芋幼苗叶片光合作用及叶绿素动力学 参数的影响。结果表明：1)在NaCl处理下,当浓度小于150 mmol&;#8226;L^－1时,增加了菊芋的叶绿素含量、净光合速率(Net photosynthetic rate, P_n)和气孔导度(Stomatal conductivity, G_s),对荧光参数PSⅡ的电子传递情况( F_m/F_o)、PSⅡ原初光能转换效率(F_v/F_m)、PSⅡ量子效率 (Actual quantum yield of PSⅡ under actinic irradiation,φPSⅡ)和光化学猝灭系数(Photochemical quenching coefficient, qP)和非 光化学猝灭系 数(Non-photochemical quenching coefficient, NPQ)没有显著影响,随着浓度的增加,各项生理指标与对照相比除了NPQ显著 增加,其余均显著降低;2)在Na₂CO₃胁迫处理下,随着Na₂CO₃浓度的增加,与对照相比菊芋幼苗叶绿素含量、P_n、G_s以及叶绿素a荧光诱导动力 学参数F_m/F_o、F_v/F_m、φPSⅡ和qP均显著降低,NPQ显著增加;3)就NaCl和Na₂CO₃相比而言,在相同Na⁺浓度情况下,处于Na₂CO₃胁迫下的菊芋 幼苗的叶绿素含量、P_n、G_s以及叶绿素a荧光诱导动力学参数F_m/F_o、F_v/F_m、φPSⅡ和qP下降幅度和NPQ的增加幅度均显著大于NaCl,这说明 NaCl和Na₂CO₃胁迫均对菊芋幼苗造成不同程度的伤害,但在相同Na⁺浓度情况下,Na₂CO₃的伤害程度大于NaCl。由此说明菊芋对盐的忍耐程度高 于碱。     

外源GSH对NaCl胁迫下番茄幼苗光合特性及碳同化关键酶基因表达的影响

对NaCl胁迫下番茄幼苗叶片分别喷施还原型谷胱甘肽(GSH)、氧化型谷胱甘肽(GSSG)和谷胱甘肽合成抑制剂(BSO)构建不同氧化还原水平的番茄植株,研究外源GSH介导的氧化还原状态对NaCl胁迫下番茄幼苗光合作用的影响.结果表明: 外源喷施GSH诱导NaCl胁迫下番茄幼苗叶片的还原力水平提高,叶片净光合速率(P_n)、气孔导度(g_s)、蒸腾速率(T_r)及最大光化学效率(F_v/F_m)、实际光化学效率(Ф_PSⅡ)、光化学猝灭系数(q_P)、非光化学猝灭系数(NPQ)值均提高,核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)活性及大亚基(RbcL)、小亚基(RbcS)和Rubisco活化酶(RCA)的基因表达水平上调,从而有效保护了光合系统,促进了(光系统Ⅱ)PSⅡ光化学反应活性、降低了NaCl胁迫对光合暗反应的抑制,缓解了NaCl胁迫对番茄植株的危害.喷施GSSG显著降低了NaCl胁迫下番茄幼苗叶片还原力水平,造成叶片光损伤和光抑制加剧,但RbcS和RbcL的基因表达水平上调可能是导致NaCl+GSSG处理下叶片P_n未下降的原因.喷施BSO对NaCl胁迫下番茄幼苗叶片氧化还原状态、CO₂传导能力和PSⅡ反应中心无显著影响,但BSO上调碳同化关键酶Rubisco初始活性、总活性及RCA和RbcS表达水平是导致P_n提高的原因.     

GABA Enhances Muskmelon Chloroplast Antioxidants to Defense Salinity-Alkalinity Stress

Salinity-alkalinity stress is a pivotal factor influencing plant growth, development, and yield. γ-Aminobutyric acid (GABA) protects plants against a variety of environmental stresses. However, it is remains largely unknown whether exogenous GABA increases the tolerance of Cucumis melon L. seedlings via effects on the chloroplast antioxidant system. In this study, the role of exogenous GABA application on the malondialdehyde content and antioxidant enzyme activities and the ascorbate-glutathione (AsA-GSH) cycle in seedlings of muskmelon was investigated. Plants were treated with foliar spraying of GABA (50 mM) under control or salinity-alkalinity stress conditions. Salinity-alkalinity stress induced cellular membrane damage. Treatment with GABA protected muskmelon seedlings from salinity-alkalinity stress by enhancing antioxidant enzyme activity and reducing malondialdehyde content. These effects of GABA resulted in maintenance of the membrane integrity of the muskmelon seedling. In addition, the status of both GSH and AsA redox played key roles in the regulation of the oxidative stress response in muskmelon seedlings under salinity-alkalinity stress.     

Application of trehalose ameliorates heat stress and promotes recovery of winter wheat seedlings

Trehalose was supplied to wheat (Triticum aestivum L.) seedlings just before a high temperature (40 °C) treatment and some physiological parameters were measured during the heat stress and recovery. The application of trehalose decreased the net photosynthetic rate (P_N) of wheat seedlings under the heat stress, but to a small extent increased the dry mass (DM) and leaf water content (LWC) after recovery from the heat stress. The trehalose-induced decrease in PN under the heat stress was not associated with a stomatal response. The heat stress slightly decreased the maximal efficiency of photosystem II (PS II) photochemistry (the variable to maximum chlorophyll a fluorescence ratio, F_v/F_m) similarly in the trehalose treated or non-treated plants. Under the heat stress, the actual efficiency of PS II photochemistry (Φ_PSII) and the efficiency of excitation energy capture by open reaction centers (F_v′/F_m′) were lower in the trehalose-pretreated seedlings, whereas they were higher after the recovery. The patterns of changes in nonphotochemical quenching (NPQ) were contrary to those of ?_{PS II} and F_v′/F_m′. The chlorophyll content was lower, whereas the β-carotene content and the degree of de-epoxidation (DEPS) of xanthophyll cycle pigments were higher in the trehalose-pretreated wheat seedlings under the heat stress. These results suggest that exogenous trehalose partially promotes recovery of wheat by the increase of NPQ, β-carotene content, and DEPS.