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淹水胁迫对植物生长的抑制作用除了低氧环境引起的根系活力下降、呼吸抑制以及矿质元素吸收受阻外,长时间淹水胁迫引起的叶绿素合成受阻与降解加速导致了叶光合色素含量下降、光能利用与转化活性改变,进而引起光合能力的大幅度下降[1]。光合作用是植物生存和繁衍的物质基础,在这个复杂的生理生化过程中,受到伤害的最原初部位是与光系统Ⅱ(PSⅡ)紧密联系的[2-5]。植物淹水后会导致PSⅡ光化学活性和电子传递速率降低[6],PSⅡ捕光色素蛋白复合物(LHCⅡa,LHCⅡb,LHCⅡc)各组分的变化,从而引起光合二氧化碳同化效率的降低[7]。另一方面,植物也可以以热的形式耗散过剩光能[8],PSⅡ反应中心的失活和周转[9]及Mehler反应[10]等减轻光抑制过程,从而保护光合机构免受破坏。叶绿素荧光参数最大光化学效率(Fv/Fm),PSⅡ实际光化学效率(Fv′/Fm′),光化学荧光猝灭系数(qP)和非光化学猝灭系数(qN),PSⅡ的实际光化学量子产量(Yyield),表观光合电子传递速率(RET)等的变化可反映逆境胁迫对PSⅡ的损伤程度[11-12],已经广泛应用于光抑制、水分、高温、低温等逆境生理研究[13-14]。竹子是集经济、生态和社会效益于一体的优良林种,是区域农村经济社会发展的重要资源和生态环境保护的重要屏障。水分、温度、光照等环境条件的变化直接影响着竹子的生长发育和分布。随着全球气候的变化,水资源不均匀分布造成近年来极端干旱和洪涝灾害频发,水分胁迫已经成为影响竹子生长发育的主要逆境因子之一,研究竹子对水分胁迫的适应能力越来越受到关注[15]。目前,国内外相关研究主要集中在短期干旱或水淹对竹子生长和生理生态的影响[16-21],而对于长期处于浸渍环境中的竹子生理生态响应及其机制研究甚少[22]。河竹Phyllostachys rivalis隶属禾本科Gramineae倭竹族Shibataeeae刚竹属Phyllostachys,主要分布于浙江、福建等地,生于溪涧边、山沟旁,性喜水湿,鞭根系统极为发达,竹鞭韧皮部密生一圈肉眼可见的气孔,具有耐淹植物的特征。我们前期的研究表明,河竹鞭根系统可以通过抗氧化系统平衡调节、生物量合理分配和异速生长调节等来适应长期淹水环境,维持生长和更新[23-24],而长期水淹胁迫下河竹叶绿素荧光变化特征、能量耗散过程及其与河竹耐受水淹的关系尚不清楚。为此,本研究以2年生河竹盆栽苗为试材,设置不同的水淹深度处理,测定分析不同水淹时间下叶片荧光参数和能量耗散的变化规律,探讨持续淹水对河竹光能的吸收和转化、能量的传递与分配、反应中心的活性、过剩能量的耗散以及光合作用的光抑制和光破坏等的影响,并从光合系统“内在性”揭示河竹对持续淹水的响应与适应机制。
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由图 1可知:随淹水时间的延长,淹水处理的河竹叶片Fo总体呈升高趋势,而Fm总体呈下降趋势。短期淹水处理(30 d)对河竹叶片Fo和Fm并无明显影响,但淹水时间进一步延长,处理间差异增大,水深效应也日趋明显,至淹水90 d和180 d时,处理Ⅱ的河竹叶片Fo显著高于处理Ⅰ和ck(P<0.05),而Fm显著低于处理Ⅰ和ck(P<0.05),且后两者Fo和Fm均无显著差异(P>0.05),其后至淹水处理结束,淹水处理的河竹叶片Fo持续升高,Fm总体上持续下降,且水位效应更加明显,处理间差异均达显著水平(P<0.05)。
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随着淹水时间的延长和淹水深度的增大,河竹叶片Fv/Fm呈下降趋势,Fv′/Fm′呈先升高后降低的趋势(图 2)。相对ck,短期淹水(30 d),河竹叶片Fv/Fm降低,但处理间差异并不显著(P>0.05),而Fv′/Fm′则升高,且处理Ⅱ显著高于ck(P<0.05);至淹水90 d和180 d时,淹水处理河竹叶片Fv/Fm明显下降,处理Ⅱ显著低于处理Ⅰ和ck(P<0.05),而后两者间无显著差异(P>0.05);其后至淹水处理结束,淹水处理河竹叶片Fv/Fm持续下降,处理间差异达显著水平(P<0.05),水位效应明显,但处理Ⅰ和处理Ⅱ仍有ck的89.4%和55.4%。河竹叶片Fv′/Fm′较Fv/Fm对淹水胁迫更敏感,短期淹水即会引起Fv′/Fm′的明显升高,淹水处理30 d时,处理Ⅱ就显著高于ck(P<0.05),处理90 d时,处理Ⅰ和处理Ⅱ均显著高于ck(P<0.05),但至处理180 d时各处理的Fv′/Fm′均明显下降,且处理间并无显著差异(P>0.05),其后淹水处理的Fv′/Fm′显著下降,水位效应较为明显。
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由图 3可知:随着淹水时间的延长,不同淹水处理下的河竹叶片qP总体呈下降趋势,qN呈升高趋势,不同处理间变化幅度不同。相对ck,淹水30 d时,处理Ⅱ的qP显著升高(P<0.05);淹水90 d时,河竹叶片qP开始降低,但处理Ⅰ和处理Ⅱ与ck差异不显著(P>0.05);淹水180 d时至处理结束,河竹叶片qP为ck>处理Ⅰ>处理Ⅱ,各处理间均有显著差异(P<0.05),水位效应明显。整个淹水过程中,河竹叶片qN基本上为ck<处理Ⅰ<处理Ⅱ,总体上淹水处理显著高于ck(P<0.05),水位效应也较为明显。
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由图 4可知:不同淹水处理的河竹叶片Yyield和RET的变化不同。随着淹水时间的延长,处理Ⅰ的Yyield先升高后降低,RET先升高后降低再升高,在淹水90 d时均达最高值。在处理180 d后,各处理间河竹叶片Yyield和RET总体上差异显著(P<0.05);处理Ⅱ的Yyield和RET均随着淹水时间的延长而逐渐降低,各处理时间点上均显著低于ck(P<0.05)。至淹水180 d后,河竹叶片的Yyield和RET均随着淹水深度的增大而显著降低(P<0.05),存在明显的水位效应。
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由表 1可知:随着淹水时间的延长,处理Ⅰ和处理Ⅱ的河竹叶片光化学反应能量(P)均呈先升高后下降的变化趋势。淹水30 d和90 d时,处理Ⅰ和处理Ⅱ的P较ck升高,且处理Ⅱ与ck差异显著(P<0.05);淹水180,270和360 d时,处理Ⅰ和处理Ⅱ的P均显著低于ck(P<0.05),且处理Ⅰ和处理Ⅱ间差异显著(P<0.05),水位效应明显。天线色素耗散能量(D)随着淹水时间的延长呈先降低后升高的变化趋势。淹水30 d和90 d时,处理Ⅰ和处理Ⅱ的D均显著低于ck(P<0.05);淹水180 d时,处理Ⅰ和处理Ⅱ的D仍低于ck,但未达显著差异水平(P>0.05);淹水270 d和360 d时,处理Ⅰ和处理Ⅱ的D均显著高于ck(P<0.05)。整个淹水处理过程中,河竹叶片天线色素耗散能量(D)的水位效应总体上并不明显。在淹水30,90和180 d时,处理Ⅰ和处理Ⅱ的非光化学反应耗散能量(E)较ck显著升高(P<0.05),但处理Ⅰ和处理Ⅱ之间差异并不显著(P>0.05);淹水270 d时,处理Ⅰ的E较ck显著降低(P<0.05);淹水360 d时,处理Ⅰ和处理Ⅱ的E较对照降低(P>0.05),河竹叶片非光化学反应耗散能量(E)的水位效应总体上也并不明显。
表 1 持续淹水对河竹叶片吸收光能分配的影响
Table 1. Effects of long-term flooding on characteristics fractions of absorbed light utilized in leaves of Phyllostachys rivalis
参数 处理 不同淹水时间河竹叶叶片吸收光能/% 30 90 180 270 360 d 光化学反应能量(P) ck 22.8 ± 0.7 b 24.6 ± 0.6 b 23.5 ± 1.1 a 26.3 ± 0.3 a 28.1 ± 3.4 a Ⅰ 23.8 ± 0.7 b 26.1 ± 0.2 a 20.0 ± 1.2 b 19.5 ± 0.8 b 15.1 ± 0.7 b Ⅱ 25.2 ± 0.6 a 26.4 ± 0.2 a 12.6 ± 1.0 c 7.6 ± 0.3 c 8.5 ± 0.7 c 天线色素耗散能量(D) ck 45.1 ± 1.3 a 43.7 ± 3.0 a 51.3 ± 2.1 a 36.5 ± 1.1 c 41.2 ± 5.7 b Ⅰ 42.6 ± 1.6 b 38.4 ± 0.9 b 46.8 ± 3.5 a 47.3 ± 2.0 b 57.3 ± 2.5 a Ⅱ 42.2 ± 0.5 b 37.0 ± 0.5 b 51.5 ± 3.3 a 55.1 ± 2.4 a 61.7 ± 0.9 a 非光化学反应耗散能量(E) ck 32.1 ± 0.7 b 31.8 ± 2.6 b 25.2 ± 1.0 b 37.2 ± 0.8 a 30.7 ± 2.3 a Ⅰ 33.7 ± 0.9 a 35.6 ± 0.7 a 33.3 ± 2.3 a 33.3 ± 1.2 b 27.5 ± 2.8 a Ⅱ 32.5 ± 0.2 ab 36.6 ± 0.3 a 35.9 ± 2.4 a 37.3 ± 2.1 a 29.8 ± 0.6 a 说明:同列不同字母表示在0.05水平存在显著性差异。
Chlorophyll fluorescence and excitation energy dissipation of pot-grown Phyllostachys rivalis leaves after long-term flooding
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摘要: 为揭示河竹Phyllostachys rivalis对持续淹水的生理生态响应与适应机制,为河竹在水陆交错带的应用提供理论依据,以2年生河竹盆栽苗为试材,设置不同深度的淹水处理[水位高出栽培基质5 cm(处理Ⅰ),10 cm(处理Ⅱ)和正常供水(ck)],测定持续淹水30,90,180,270和360 d时河竹叶片叶绿素荧光参数,分析叶片光能的吸收和转化、能量的传递与分配、反应中心的活性、过剩能量的耗散对持续淹水胁迫的响应。结果表明:① 持续淹水胁迫下,河竹叶片能通过维持相对较高的表观光合电子传递速率(RET),光化学荧光猝灭(qP)和光化学反应能量(P),增强非光化学猝灭(qN)来调节自身能量代谢,以热耗散形式散失过多的光能,有效地避免或减轻光抑制和光氧化,河竹吸收光强主要以天线色素耗散(D)为主要光能分配途径,淹水后期PSⅡ反应中心的非光化学反应耗散(E)的恢复起着重要作用,持续淹水一定程度上会损害河竹叶片光系统Ⅱ(PSⅡ),但对于PSⅡ的功能反应中心影响较小。② 不同淹水处理叶绿素荧光参数存在差异,持续淹水30,90 d时,处理Ⅰ的初始荧光(Fo),最大荧光(Fm),最大光化学效率(Fv/Fm),qP和RET等与ck差异不显著(P > 0.05),而处理Ⅱ在淹水90 d时与ck差异显著(P < 0.05),至淹水270,360 d时,各处理间差异均达显著水平(P < 0.05),持续淹水胁迫对叶绿素荧光参数及能量耗散的影响存在明显的水位效应。Abstract: To determine the physiological and biochemical responses and adaptive mechanisms of Phyllostachys rivalis to long-term soil flooding, two-year-old potted seedlings of Ph. rivalis were subjected to different flooding depths. The flooding treatment was set with water levels of 5 cm (TreatmentⅠ) and 10 cm (TreatmentⅡ) higher than the soil surface and normal water supply (ck). Then the chlorophyll fluorescence parameters in leaves were determined after continuous flooding for 30, 90, 180, 270, and 360 d. Responses to the continuous flooding stress for light energy absorption and transformation, energy transfer and distribution, reaction center activity, and excitation energy dissipation in leaves of three leaves per seedlings and three seedlings were measured and analyzed. A one-way analysis of variance with duncan's test was conducted at a significant level of 0.05. Results showed that the chlorophyll fluorescence parameters varied with different flooding levels. When flooding for 30 d and 90 d, minimal fluorescence (Fo), maximum fluorescence (Fm), photochemical maximum efficiency of PS Ⅱ (Fv/Fm), photochemical quenching coefficient (qP), and electron transport rate(RET) of Treatment Ⅰ were not significantly different (P > 0.05) compared with ck; however, Treatment Ⅱ when flooding for 90 d compared with ck was significantly increased for Fo while significantly decreased for Fm, Fv/Fm and RET (P < 0.05). With flooding treatments of 270 d and 360 d, Fm, Fv/Fm, qP and RET of Treatment Ⅰ and Ⅱ were both significant decreased while Fo were significant increased than that of ck (P < 0.05). The effect of flooding on chlorophyll fluorescence parameters and energy dissipation was also dependent on the water level. In leaves of Ph. rivalis, Fm, Fv/Fm, and qP were significantly decreased (P < 0.05) with continuous flooding stress; whereas, Fo and non-photochemical quenching coefficient (qN) increased gradually but not reached significant level (P > 0.05). The amount of absorbed light in photochemistry (P) and excess energy (E) increased first and then decreased (P < 0.05), but the energy of dissipation of the antenna heat dissipation (D) observed were opposite. Taken together, Ph. rivalis could maintain a relatively high RET, qP, and P in the early stages of flooding, and enhance qN to regulate their energy metabolism, dissipate excess light energy via heat dissipation, thereby alleviating the light photoinhibition and photooxidation; however, continuous long-term soil flooding could damage photosystem Ⅱ (PS Ⅱ) even though there was minimal adverse effect on the functional center of PS Ⅱ (P < 0.05). It can be inferred that Ph. rivalis can tolerate a short term flooding which facilitate its possible application in plantation restoration of riparion zone.
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表 1 持续淹水对河竹叶片吸收光能分配的影响
Table 1. Effects of long-term flooding on characteristics fractions of absorbed light utilized in leaves of Phyllostachys rivalis
参数 处理 不同淹水时间河竹叶叶片吸收光能/% 30 90 180 270 360 d 光化学反应能量(P) ck 22.8 ± 0.7 b 24.6 ± 0.6 b 23.5 ± 1.1 a 26.3 ± 0.3 a 28.1 ± 3.4 a Ⅰ 23.8 ± 0.7 b 26.1 ± 0.2 a 20.0 ± 1.2 b 19.5 ± 0.8 b 15.1 ± 0.7 b Ⅱ 25.2 ± 0.6 a 26.4 ± 0.2 a 12.6 ± 1.0 c 7.6 ± 0.3 c 8.5 ± 0.7 c 天线色素耗散能量(D) ck 45.1 ± 1.3 a 43.7 ± 3.0 a 51.3 ± 2.1 a 36.5 ± 1.1 c 41.2 ± 5.7 b Ⅰ 42.6 ± 1.6 b 38.4 ± 0.9 b 46.8 ± 3.5 a 47.3 ± 2.0 b 57.3 ± 2.5 a Ⅱ 42.2 ± 0.5 b 37.0 ± 0.5 b 51.5 ± 3.3 a 55.1 ± 2.4 a 61.7 ± 0.9 a 非光化学反应耗散能量(E) ck 32.1 ± 0.7 b 31.8 ± 2.6 b 25.2 ± 1.0 b 37.2 ± 0.8 a 30.7 ± 2.3 a Ⅰ 33.7 ± 0.9 a 35.6 ± 0.7 a 33.3 ± 2.3 a 33.3 ± 1.2 b 27.5 ± 2.8 a Ⅱ 32.5 ± 0.2 ab 36.6 ± 0.3 a 35.9 ± 2.4 a 37.3 ± 2.1 a 29.8 ± 0.6 a 说明:同列不同字母表示在0.05水平存在显著性差异。 -
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