保存光强对耐干藓生理特性与营养繁殖的影响
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摘要
【目的】研究不同保存光强对耐干藓生理特性与营养繁殖的影响,为耐干藓营养繁殖及人工培养提供理论依据。【方法】以黄土丘陵区常见3种耐干藓优势种(扭口藓、土生对齿藓、短叶对齿藓)为材料,在0 lx(遮光,L0)、20 lx(避光,L1)、4 000 lx(温室,L2)下以干燥状态保存89 d,测定保存前后配子体生理指标(叶绿素、可溶性糖、可溶性蛋白质、丙二醛含量)与萌发参数(配子体萌发率、配子体增加量、配子体活力指数),研究不同保存光强对耐干藓生理特性与营养繁殖的影响。【结果】除了土生对齿藓和短叶对齿藓的叶绿素含量以及短叶对齿藓L0、L2处理的可溶性糖含量外,不同光照处理的3个藓种的不同生理指标均高于保存前。随着光强的增加,土生对齿藓和短叶对齿藓的叶绿素含量总体均呈现下降的趋势,扭口藓叶绿素含量则先增加后下降;扭口藓可溶性糖含量下降,土生对齿藓可溶性糖含量无显著变化,短叶对齿藓可溶性糖含量先增加后下降;扭口藓和土生对齿藓的可溶性蛋白质含量增大,短叶对齿藓的可溶性蛋白质含量则先增加后下降;扭口藓和短叶对齿藓的丙二醛含量先增后减,土生对齿藓的丙二醛含量呈现先下降后增加的趋势。随着光强的增加,扭口藓萌发参数均呈增加的趋势,土生对齿藓的萌发参数变化均不显著,短叶对齿藓的萌发参数均呈降低趋势。叶绿素含量与配子体萌发率存在显著正相关(P<0.05),可溶性蛋白质含量则与配子体萌发率存在极显著正相关(P<0.01),可溶性糖含量与配子体增加量有极显著负相关关系(P<0.01)。可知叶绿素、可溶性糖、可溶性蛋白质含量变化对藓类复水后的营养繁殖结果影响较大。【结论】长期干燥期间,光强影响藓类细胞膜渗透性,使其生理特性与营养繁殖能力发生明显变化。不同耐干藓适宜的保存光强存在差异。
【Objective】 This study investigated the effects of different light intensities on physiological characteristics and vegetative propagation of desiccation-tolerant mosses after storage to provide theoretical foundation for vegetative propagation and artificial cultivation of desiccation-tolerant mosses.【Method】 Three dominant species of common desiccation-tolerant mosses in the Loess hilly region of China,including Barbula unguiculata,Didymodon vinealis and Didymodon tectorum,were collected and stored at three light intensity gradients in a dry environment for 89 days.The three light intensity gradients were 0 lx(shading,L0),20 lx(away from light,L0) and 4 000 lx(in greenhouse,L2).The physiological indices(contents of chlorophyll, soluble sugar,soluble protein and malondialdehyde) and germination parameters(gametophyte germination percentage,gametophyte increment and gametophyte vigor index) of moss gametophytes were investigated before and after storage to determine the effects of different light intensities on the physiological characteristics and vegetative propagation of desiccation-tolerant mosses after storage.【Result】 Except for chlorophyll contents of D.vinealis and D.tectorum and soluble sugar contents of D.tectorum in L0 and L2 treatments,all physiological indices of the three mosses at each light intensity after storage were higher than before storage.When light intensity increased, chlorophyll contents of D. vinealis and D. tectorum generally decreased,while contents of B. unguiculata increased at first and decreased thereafter. The soluble sugar content of B.unguiculata decreased with increasing light intensity, while content of D. tectorum increased at first and then decreased.The soluble sugar content of D. vinealis showed no significant difference among light intensities.The soluble protein content of B.unguiculata and D.tectorum continuously increased when the light intensity increased,while that of D.tectorum increased at first followed by decrease.The malondialdehyde contents of B.unguiculata and D.tectorum increased firstly and then decreased with increasing light intensity, whereas that of D.vinealis had an opposite trend.When light intensity increased,the germination parameters of B.unguiculata increased,while that of D.tectorum decreased.There were no significant differences in germination parameters among the light intensities in D.vinealis.Gametophyte germination percentage was significantly positively correlated with chlorophyll content(P<0.05) and soluble protein content(P<0.01),while gametophyte increment was significantly negatively correlated with soluble sugar content(P<0.01).These correlations indicate changes in contents of chlorophyll,soluble sugar and soluble protein had significant effect on vegetative propagation of mosses after rehydration.【Conclusion】 Light intensity had significant effect on membrane permeability of mosses and caused significant changes in physiological characteristics and vegetative propagation capacity of mosses during long-term dry.The adaptive storage light intensity of desiccation-tolerant mosses varied with species.
【Objective】 This study investigated the effects of different light intensities on physiological characteristics and vegetative propagation of desiccation-tolerant mosses after storage to provide theoretical foundation for vegetative propagation and artificial cultivation of desiccation-tolerant mosses.【Method】 Three dominant species of common desiccation-tolerant mosses in the Loess hilly region of China,including Barbula unguiculata,Didymodon vinealis and Didymodon tectorum,were collected and stored at three light intensity gradients in a dry environment for 89 days.The three light intensity gradients were 0 lx(shading,L0),20 lx(away from light,L0) and 4 000 lx(in greenhouse,L2).The physiological indices(contents of chlorophyll, soluble sugar,soluble protein and malondialdehyde) and germination parameters(gametophyte germination percentage,gametophyte increment and gametophyte vigor index) of moss gametophytes were investigated before and after storage to determine the effects of different light intensities on the physiological characteristics and vegetative propagation of desiccation-tolerant mosses after storage.【Result】 Except for chlorophyll contents of D.vinealis and D.tectorum and soluble sugar contents of D.tectorum in L0 and L2 treatments,all physiological indices of the three mosses at each light intensity after storage were higher than before storage.When light intensity increased, chlorophyll contents of D. vinealis and D. tectorum generally decreased,while contents of B. unguiculata increased at first and decreased thereafter. The soluble sugar content of B.unguiculata decreased with increasing light intensity, while content of D. tectorum increased at first and then decreased.The soluble sugar content of D. vinealis showed no significant difference among light intensities.The soluble protein content of B.unguiculata and D.tectorum continuously increased when the light intensity increased,while that of D.tectorum increased at first followed by decrease.The malondialdehyde contents of B.unguiculata and D.tectorum increased firstly and then decreased with increasing light intensity, whereas that of D.vinealis had an opposite trend.When light intensity increased,the germination parameters of B.unguiculata increased,while that of D.tectorum decreased.There were no significant differences in germination parameters among the light intensities in D.vinealis.Gametophyte germination percentage was significantly positively correlated with chlorophyll content(P<0.05) and soluble protein content(P<0.01),while gametophyte increment was significantly negatively correlated with soluble sugar content(P<0.01).These correlations indicate changes in contents of chlorophyll,soluble sugar and soluble protein had significant effect on vegetative propagation of mosses after rehydration.【Conclusion】 Light intensity had significant effect on membrane permeability of mosses and caused significant changes in physiological characteristics and vegetative propagation capacity of mosses during long-term dry.The adaptive storage light intensity of desiccation-tolerant mosses varied with species.
引文
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[2] 徐杰,白学良,田桂泉,等.腾格里沙漠固定沙丘结皮层藓类植物的生态功能及与土壤环境因子的关系 [J].中国沙漠,2005,25(2):234-242. Xu J,Bai X L,Tian G Q,et al.Ecological function of mosses in biotic crusts on fixed dunes on the Tengger Desert and its relation with soil factors [J].Journal of Desert Research,2005,25(2):234-242.
[3] 王雪芹,张元明,张伟民,等.古尔班通古特沙漠生物结皮对地表风蚀作用影响的风洞实验 [J].冰川冻土,2004,26(5):632-638. Wang X Q,Zhang Y M,Zhang W M,et al.Wind tunnel experiment of biological crust effect on wind erodibility of sand surface in Gurbantnnggnt Desert,Xinjiang [J].Journal of Glaciology and Geocryology,2004,26(5):632-638.
[4] Gao L Q,Bowker M A,Xu M X,et al.Biological soil crusts decrease erodibility by modifying inherent soil properties on the Loess Plateau,China [J].Soil Biology & Biochemistry,2017,105:49-58.
[5] 王爱国,赵允格,许明祥,等.黄土丘陵区不同演替阶段生物结皮对土壤CO2通量的影响 [J].应用生态学报,2013,24(3):659-666. Wang A G,Zhao Y G,Xu M X,et al.Effects of biological soil crust at different succession stages in hilly region of Loess Plateau on soil CO2 flux [J].Chinese Journal of Applied Ecology,2013,24(3):659-666.
[6] Reed S C,Coe K K,Sparks J P.Changes to dryland rainfall result in rapid moss mortality and altered soil fertility [J].Nature,2012,10(2):752-755.
[7] Xiao B,Zhao Y G,Wang Q H,et al.Development of artificial moss-dominated biological soil crusts and their effects on runoff and soil water content in a semi-arid environment [J].Journal of Arid Environments,2015,117:75-83.
[8] Zhao Y G,Bowker M A,Zhang Y M,et al.Biological soil crusts an organizing principle in drylands [M].Berlin:Springer,2016:499-523.
[9] 陈彦芹,赵允格,冉茂勇.黄土丘陵区藓结皮人工培养方法试验研究 [J].西北植物学报,2009,29(3):586-592. Chen Y Q,Zhao Y G,Ran M Y.Experimental research on artificial culture of mosses crusts in hilly Loess Plateau region [J].Acta Botanica Boreali-Occidentalia Sinica,2009,29(3):586-592.
[10] 杨永胜,冯伟,袁方,等.快速培育黄土高原苔藓结皮的关键影响因子 [J].水土保持学报,2015,29(4):289-294. Yang Y S,Feng W,Yuan F,et al.Key influential factors of rapid cultivation of moss crust on Loess Plateau region [J].Journal of Soil and Water Conservation,2015,29(4):289-294.
[11] 杨延哲,张侃侃,杨永胜,等.毛乌素沙地苔藓结皮的野外人工培育技术 [J].水土保持通报,2016,36(2):165-170. Yang Y Z,Zhang K K,Yang Y S,et al.Field artificial cultivation technology of moss dominated crust in Mu Us Sandland [J].Bulletin of Soil and Water Conservation,2016,36(2):165-170.
[12] Antoninka A J,Bowker M A,Reed S C,et al.Production of greenhouse-grown biocrust mosses and associated cyanobacteria to rehabilitate dryland soil function [J].Restoration Ecology,2016,24(3):324-335.
[13] Stark L R,Greenwood J L,Brinda J C.Desiccated Syntrichia ruralis shoots regenerate after 20 years in the herbarium [J].Journal of Bryology,2016,39(1):85-93.
[14] Stark L R,Nichols L,Mcletchie D N,et al.Age and sex-specific rates of leaf regeneration in the Mojave Desert moss Syntrichia caninervis [J].American Journal of Botany,2004,91(1):1-9.
[15] Burch J.Some mosses survive cryopreservation without prior pretreatment [J].Bryologist,2003,106(2):270-277.
[16] Guo Y W,Zhao Y G.Effects of storage temperature on the physiological characteristics and vegetative propagation of desiccation-tolerant mosses [J].Biogeosciences,2018,15:797-808.
[17] Seel W,Hendry G,Atherton N,et al.Radical formation and accumulation in vivo,in desiccation tolerant and intolerant mosses [J].Free Radic Res Commun,1991,15(3):133-141.
[18] Zhang G H,Liu G B,Wang G L,et al.Effects of vegetation cover and rainfall intensity on sediment-bound nutrient loss,size composition and volume fractal dimension of sediment particles [J].Pedosphere,2011,21(5):676-684.
[19] 赵允格,许明祥,王全九,等.黄土丘陵区退耕地生物结皮理化性状初报 [J].应用生态学报,2006,17(8):1429-1434. Zhao Y G,Xu M X,Wang Q J,et al.Physical and chemical propertis of soil bio-crust on rehabilitated grassland in hilly Loess Plateau of China [J].Chinese Journal of Applied Ecology,2006,17(8):1429-1434.
[20] 杨雪伟,赵允格,许明祥.黄土丘陵区藓结皮优势种形态结构差异 [J].生态学杂志,2016,35(2):370-377. Yang X W,Zhao Y G,Xu M X.Variation of morphological structure of dominant species in moss crusts in hilly Loess Plateau region [J].Chinese Journal of Ecology,2016,35(2):370-377.
[21] 高谦.扭口藓属 [M]//高谦. 中国苔藓志:第2卷.北京:科学出版社,1996:171-193. Gao Q.Barbula Hedw [M]//Gao Q.Bryophytes in China:Volume 2.Beijing:Science Press,1996:171-193.
[22] 郭玥微.不同季节及保存条件对耐干藓生理特性及营养繁殖的影响 [D].陕西杨凌:西北农林科技大学,2018. Guo Y W.Effects of seasons and storage conditions on the physiological characteristics and vegetative propagation of desiccation-tolerant mosses [D].Yangling,Shaanxi:Northwest A&F University,2018.
[23] 黄学林,陈润政.种子生理实验手册 [M].北京:农业出版社,1990:83-85. Huang X L,Chen R Z.Experimental handbook of seed physiology [M].Beijing:China Agriculture Press,1990:83-85.
[24] Proctor M C F,Oliver M J,Wood A J,et al.Desiccation-tolerance in bryophytes:a review [J].Bryologist,2007,110:595-621.
[25] Keever C.Establishment of Grimmia laevigata on bare granite [J].Ecology,1957,38(3):422-429.
[26] Deltoro V I,Calatayud A,Gimeno C,et al.Water relations,chlorophyll fluorescence, and membrane permeability during desiccation in bryophytes from xeric,mesic,and hydric environments [J].Canadian Journal of Botany,1998,76(11):1923-1929.
[27] Csintalan Z,Proctor M C F,Tuba Z.Chlorophyll fluorescence during drying and rehydration in the mosses Rhytidiadelphus loreus (Hedw.) Warnst.,Anomodon viticulosus (Hedw.) Hook.and Tayl.and Grimmia pulvinata (Hedw.) Sm [J].Annals of Botany,1999,84(2):235-244.
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