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    沙棘形态和光合系统对干旱、盐碱及其复合胁迫的响应

    Morphology and photosynthetic system responses to single and combined drought-saline-alkali stress in Hippophae rhamnoides

    • 摘要:
      目的 国家审定良种‘深秋红’沙棘兼具重要的生态和经济价值,探究其在干旱、盐碱和干旱−盐碱复合3个胁迫维度下形态和光合能力的变化情况,旨在揭示沙棘逆境胁迫下的响应机制,为干旱盐碱区植被恢复与沙棘良种推广提供理论依据。
      方法 以‘深秋红’沙棘一年生无性系幼苗为研究对象,利用自然控水法和逐渐浇灌盐碱法,设置不同土壤体积含水量、不同盐碱浓度以及复合胁迫处理,观察不同处理下其形态变化,测定其光合系统相关指标,并进行聚类和相关性分析。
      结果 (1)干旱、盐碱及其复合胁迫对沙棘幼苗形态和光合系统一定产生抑制作用,但复合胁迫效应显著强于单一胁迫。(2)单一胁迫初期影响有限,但随胁迫时间和程度增加,叶绿素含量(SPAD)、电子传递效率(ETR)、光能利用效率(α)、PSII最大光化学效率(Fv/Fm)下降;复合胁迫下,SPAD、ETR、Fv/Fm显著降低,PSII反应中心密度(RC/CSo)同步减小,电子传递受阻,能量吸收与捕获受限。(3)单一胁迫下,沙棘主要通过增强调节性能量耗散(Y(NPQ))维持光系统稳定;而在重度复合胁迫后期,Y(NPQ)明显下降,非调节性能量耗散(Y(NO))显著升高,最终导致光合系统功能失稳。(4)复合胁迫显著改变了类囊体膜能量流动结构,随着复合胁迫强度增加,吸收能量通量(ABS/CSm)和捕获能量通量(TRo/CSm)及电子传递通量(ETo/CSm)均明显降低,且下降幅度显著高于单一胁迫处理。(5)高强度复合胁迫组在 Per3 ~ Per4 阶段聚为单独一组,显著区别于单一胁迫。且PSII稳定性指标(Fv/Fm和RC/CSo)与光化学及电子传递参数呈正相关,而与K点相对可变荧光(Vk)和J点相对可变荧光(Vj)及 Y(NO) 显著负相关,表明PSII活性下降伴随着供体侧与受体侧电子传递受阻以及非调控能量耗散的增强。
      结论 干旱、盐碱及复合胁迫均对沙棘幼苗的形态及光合系统造成不同程度的损伤,且复合胁迫影响最为严重。受到干旱或盐碱胁迫时,沙棘可以通过增加热耗散、启动光保护机制以抵御胁迫损害,但重度复合胁迫会造成不可逆的损伤。研究结果对木本植物逆境适应机制进行了补充,对干旱盐碱区生态修复树种选育具有参考价值。

       

      Abstract:
      Objective This study investigated the morphological and photosynthetic responses of the nationally certified cultivar ‘Shenqiu Hong’ of Hippophae rhamnoides under drought, saline-alkali, and combined drought-saline-alkali stresses.
      Methods One-year-old clonal seedlings were used as the experimental material. Natural water deficit control and gradient saline-alkali irrigation were applied to establish treatments with different soil volumetric water contents, different saline-alkali concentrations, and combined stresses. Morphological changes were observed and photosynthetic parameters were measured.
      Results (1) Drought stress, saline-alkali stress, and their combined stress suppressed morphological performance and photosynthetic function in H. rhamnoides seedlings, with the combined stress exerting markedly stronger effects than either single stress. (2) The effects of single stress were minimal at early stages but intensified with increasing stress duration and severity, leading to declines in chlorophyll content (SPAD), electron transport rate (ETR), light-use efficiency (α), and the maximum photochemical efficiency of PSII (Fv/Fm). Under combined stress, SPAD, ETR, and Fv/Fm were markedly reduced, accompanied by a concurrent decrease in PSII reaction center density (RC/CSo), resulting in impaired electron transport and constrained light energy absorption and trapping. (3) Under drought stress or saline-alkali stress, seedlings predominantly maintained photosystem stability by enhancing regulated non-photochemical energy dissipation Y(NPQ). In contrast, during the late stage of severe combined stress, Y(NPQ) declined sharply while non-regulated energy dissipation Y(NO) increased substantially, ultimately leading to functional destabilization of the photosynthetic apparatus. (4) Combined drought–salinity stress markedly altered the energy flux organization of the thylakoid membrane. With increasing stress intensity, the absorbed energy flux (ABS/CSm), trapped energy flux (TRo/CSm), and electron transport flux (ETo/CSm) all declined substantially, with reductions significantly greater than those observed under single-stress treatments. (5) High-intensity combined stress treatments clustered independently at the Per3–Per4 stages, clearly separated from single-stress conditions. PSII stability indices (Fv/Fm, RC/CSo) showed strong positive correlations with photochemical performance and electron transport metrics, and significant negative correlations with Vk, Vj, and Y(NO), indicating that the decline in PSII activity is accompanied by impaired electron transport at both the donor and acceptor sides and by enhanced non-regulated energy dissipation.
      Conclusion Morphological and photosynthetic damage in H. rhamnoides indeced by combined stress are the most severe compared with individual drought or saline-alkali stresses. H. rhamnoides activates photoprotective mechanisms through enhanced thermal dissipation to mitigate stress damage under drought or saline-alkali conditions. However, these adaptions were compromised under combined stress, resulting in irreversible photodamage. The results enhance our understanding of stress adaptation in woody plants and serve as a reference for selecting and breeding tree species for ecological restoration in drought-and saline-alkali regions.

       

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