盐旱耦合胁迫对费尔干猪毛菜种子萌发及其幼苗生长的影响
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摘要
本研究采用双因素组合实验,探讨盐(NaCl)、干旱(PEG 6000)对费尔干猪毛菜(Salsola ferganica)种子萌发和幼苗生长的影响。结果显示:①费尔干猪毛菜种子萌发能耐受较高浓度NaCl和PEG胁迫。低浓度NaCl或PEG胁迫下,费尔干猪毛菜去翅种子和具翅种子萌发率与对照无显著差异,高浓度NaCl或PEG处理下,去翅种子萌发率35%以上,显示较强的耐盐、耐旱性;盐旱耦合处理下,不同低浓度PEG能够显著缓解中浓度NaCl对种子萌发的抑制作用。②随NaCl浓度升高,具翅种子恢复萌发率逐渐上升,而去翅种子则维持较低水平;随着PEG浓度的升高,2种种子恢复萌发能力显著降低;盐旱耦合处理下2种种子的恢复萌发率与NaCl胁迫具有相同趋势。③NaCl或PEG对去翅种子幼苗的影响比具翅种子强,特别是PEG处理;随NaCl或PEG浓度升高,2种种子幼苗的株高和根长均降低;盐旱耦合处理下低浓度NaCl对去翅种子幼苗的影响比具翅种子小(除了100 mmol·L~(-1)NaCl+25 g·L~(-1)PEG处理),中浓度NaCl对去翅种子幼苗的影响比具翅种子强,高浓度NaCl处理下随PEG浓度升高,2种种子幼苗的株高和根长均降低。以上结果显示,费尔干猪毛菜在萌发期和幼苗期具有较强的耐盐耐旱性,盐旱耦合在一定程度上能够缓解盐离子毒害效应,这对其度过早期恶劣环境具有重要的生态学意义;果翅对种子在胁迫下进入次级休眠,从而补充土壤种子库起到重要作用。
In this study,the effects of different stress conditions,e. g.,salt (NaCl),drought (PEG 6000) or salt coupled with drought,on seed germination (SG) and seedling growth (SGr) of Salsola ferganica were researched. Results showed that:① SG of S. ferganica could tolerate relatively high NaCl and PEG concentrations. There was no effect of low NaCl and PEG concentrations on SG; at higher concentration,however,SG was significantly decreased but its percentage still exceeded 35%,which suggested that S. ferganica was salt-or drought-tolerant. Under salt stress coupled with drought stress,the inhibition of seed germination was significantly relieved by increasing the PEG concentration under the low and medium concentration of NaCl.② Under NaCl stress,the recovery rate of winged perianth (WWP) seeds was increased with increasing NaCl concentration,while that of dewing perianth (NWP) seeds was remained at low level. With increasing the PEG concentration,however,the recovery rates of both WWP and NWP seeds were significantly decreased. The recovery of germination of WWP and NWP seeds under salt coupled with drought stress was similar as that under NaCl stress.③ The effect of NaCl or PEG treatment on the growth of the seedlings from NWP seeds of S. ferganica was stronger than that from WWP seeds. With increasing the NaCl or PEG concentration,the growth of plant height and root length of seedlings from WWP and NWP were inhibited. The growth of S. ferganica seedlings Under salt stress coupled with drought stress,the effect of low NaCl concentration on the growth of the WWP seedlings was less than that of NWP seeds (except for 100 mmol ·L~(-1) NaCl + 25 g·L~(-1) PEG). The effect of NaCl concentration on the seedlings of NWP seeds was stronger than that of WWP seeds. With increasing the PEG concentration at high NaCl level,the growth of plant height and root length of seedlings from WWP and NWP were inhibited. In conclusion,S. ferganica was much more tolerant to salt and drought in seed germination and seedling growth. Salt stress coupled with drought stress could relieve salt ion toxicity to some extent,which has important ecological significance for S. ferganica to survive the early harsh environment. Moreover,the fruit wing plays an important role in puting the seed to dormancy under stress and then replenishing the soil seed bank.
In this study,the effects of different stress conditions,e. g.,salt (NaCl),drought (PEG 6000) or salt coupled with drought,on seed germination (SG) and seedling growth (SGr) of Salsola ferganica were researched. Results showed that:① SG of S. ferganica could tolerate relatively high NaCl and PEG concentrations. There was no effect of low NaCl and PEG concentrations on SG; at higher concentration,however,SG was significantly decreased but its percentage still exceeded 35%,which suggested that S. ferganica was salt-or drought-tolerant. Under salt stress coupled with drought stress,the inhibition of seed germination was significantly relieved by increasing the PEG concentration under the low and medium concentration of NaCl.② Under NaCl stress,the recovery rate of winged perianth (WWP) seeds was increased with increasing NaCl concentration,while that of dewing perianth (NWP) seeds was remained at low level. With increasing the PEG concentration,however,the recovery rates of both WWP and NWP seeds were significantly decreased. The recovery of germination of WWP and NWP seeds under salt coupled with drought stress was similar as that under NaCl stress.③ The effect of NaCl or PEG treatment on the growth of the seedlings from NWP seeds of S. ferganica was stronger than that from WWP seeds. With increasing the NaCl or PEG concentration,the growth of plant height and root length of seedlings from WWP and NWP were inhibited. The growth of S. ferganica seedlings Under salt stress coupled with drought stress,the effect of low NaCl concentration on the growth of the WWP seedlings was less than that of NWP seeds (except for 100 mmol ·L~(-1) NaCl + 25 g·L~(-1) PEG). The effect of NaCl concentration on the seedlings of NWP seeds was stronger than that of WWP seeds. With increasing the PEG concentration at high NaCl level,the growth of plant height and root length of seedlings from WWP and NWP were inhibited. In conclusion,S. ferganica was much more tolerant to salt and drought in seed germination and seedling growth. Salt stress coupled with drought stress could relieve salt ion toxicity to some extent,which has important ecological significance for S. ferganica to survive the early harsh environment. Moreover,the fruit wing plays an important role in puting the seed to dormancy under stress and then replenishing the soil seed bank.
引文
[1]Ma Y,Zhang J,Li X,et al.Effects of environmental stress on seed germination and seedling growth of Salsola ferganica(Chenopodiaceae)[J].Acta Ecologica Sinica,2016,36(6):456-463.
[2]Volis S.Seed heteromorphism in Triticum dicoccoides:Association between seed positions within a dispersal unit and dormancy[J].Oecologia,2016,181(2):401-412.
[3]Jaganathan G K.Influence of maternal environment in developing different levels of physical dormancy and its ecological significance[J].Plant Ecology,2016,217(1):71-79.
[4]Gul B,Ansari R,Flowers T J,et al.Germination strategies of halophyte seeds under salinity[J].Environmental&Experimental Botany,2013,92(4):4-18.
[5]渠晓霞,黄振英.盐生植物种子萌发对环境的适应对策[J].生态学报,2005,25(9):2 389-2 398.[Qu Xiaoxia,Huang Zhenying.The adaptive strategies of halophyte seed germination[J].Acta Ecologica Sinica,2005,25(9):2 389-2 398.]
[6]Li R,Shi F,Fukuda K.Interactive effects of salt and alkali stresses on seed germination,germination recovery,and seedling growth of a halophyte Spartina alterniflora(Poaceae)[J].South African Journal of Botany,2010,76(2):380-387.
[7]Katembe W J,Ungar I A,Mitchell J P.Effect of salinity on germination and seedling growth of two Atriplex species(Chenopodiaceae)[J].Annals of Botany,1998,82(2):167-175.
[8]Khan M A,Ungar I A.Influence of salinity and temperature on the germination of Haloxylon recurvum Bunge ex.Boiss[J].Annals of Botany,1996,78(5):547-551.
[9]李宽亮,刘彤,李勇冠,等.褐翅猪毛菜种子异型性及其萌发特性研究[J].石河子大学学报(自然科学版),2012,30(3):276-281.[Li Kuanliang,Liu Tong,Li Yongguan,et al.Seed polymorphism and germination of Salsola korshinskyi Drob[J].Journal of Shihezi University(Natural Science Edition),2012,30(3):276-281.]
[10]Baskin J M,Lu J J,Baskin C C,et al.Diaspore dispersal ability and degree of dormancy in heteromorphic species of cold deserts of northwest China:A review[J].Perspectives in Plant Ecology Evolution&Systematics,2014,16(2):93-99.
[11]Mamut J,Tan D Y,Baskin C C,et al.Role of trichomes and pericarp in the seed biology of the desert annual Lachnoloma lehmannii(Brassicaceae)[J].Ecological Research,2014,29(1):33-44.
[12]Baskin J M,Baskin C C.A classification system for seed dormancy[J].Seed Science Research,2004,14(1):1-16.
[13]何美香,杜晓峰,陈玲,等.盐分、变温和激素处理对盐生植物异子蓬异型性种子萌发及成苗的影响[J].生态学杂志,2013,32(1):45-51.[He Meixiang,Du Xiaofeng,Chen Ling,et al.Effects of salt,alternating temperature and hormone treatments on seed germination and seedling establishment of Suaeda aralocaspica(Chenopodiaceae)dimorphic seeds[J].Chinese Journal of Ecology,2013,32(1):45-51.]
[14]Li J,Yin L Y,Jongsma M A,et al.Effects of light,hydropriming and abiotic stress on seed germination,and shoot and root growth of pyrethrum(Tanacetum cinerariifolium)[J].Industrial Crops&Products,2011,34(3):1 543-1 549.
[15]Wang Y,Jiang G Q,Han Y N,et al.Effects of salt,alkali and saltalkali mixed stresses on seed germination of the halophyte Salsola ferganica(Chenopodiaceae)[J].Acta Ecologica Sinica,2013,33(6):354-360.
[16]黄俊华.中国猪毛菜属(Salsola L.)植物的地理分布特点[J].干旱区地理,2005,28(3):325-329.[Huang Junhua.Geographical distribution of Salsola L.in China[J].Arid Land Geography,2005,28(3):325-329.]
[17]Ma Y L,Wang J,Zhang J H,et al.Seed heteromorphism and effects of light and abiotic stress on germination of a typical annual halophyte Salsola ferganica in cold desert[EB/OL].https://www.frontiersin.org/articles/10.3389/fpls.2017.02257/full,2018/2018-8-29.
[18]Yao S,Lan H,Zhang F.Variation of seed heteromorphism in Chenopodium album and the effect of salinity stress on the descendants[J].Annals of Botany,2010,105(6):1 015-1 025.
[19]Coons J M,Kuehl R O,Simons N R.Tolerance of ten lettuce cultivars to high temperature combined with Na Cl during germination[J].Journal of the American Society for Horticultural Science,1990,115(6):1 004-1 007.
[20]Kaufmann M R,Eckard A N.Evaluation of water stress control with polyethylene glycols by analysis of guttation[J].Plant Physiology,1971,47(4):453-456.
[21]Dodd G L,Donovan L A.Water potential and ionic effects on germination and seedling growth of two cold desert shrubs[J].American Journal of Botany,1999,86(8):1 146-1 153.
[22]Naylor R E L.An evaluation of various germination indices for predicting differences in seed vigour in Italian ryegrass[J].Seed Science&Technology,1981,9:593-600.
[23]Khan M A,Gul B,Weber D J.Seed germination in the Great Basin halophyte Salsola iberica[J].Revue Canadienne De Botanique,2002,80(6):650-655.
[24]Wei Y,Dong M,Huang Z,et al.Factors influencing seed germination of Salsola affinis(Chenopodiaceae),a dominant annual halophyte inhabiting the deserts of Xinjiang,China[J].Flora,2008,203(2):134-140.
[25]Ahmed M Z,Khan M A.Tolerance and recovery responses of playa halophytes to light,salinity and temperature stresses during seed germination[J].Flora,2010,205(11):764-771.
[26]Yildiz M,Cenkci S,Kargioglu M.Effects of salinity,temperature,and light on seed germination in two Turkish endemic halophytes,Limonium iconicum and L.lilacinum(Plumbaginaceae)[J].Seed Science&Technology,2008,36(3):646-656.
[27]Bhaskar G,Huang B R.Mechanism of salinity tolerance in plants:Physiological,biochemical,and molecular characterization[EB/OL].https://www.hindawi.com/journals/ijg/2014/701596/,2014/2018-8-29.
[28]Yao X,Chu J G.Effects of selenium on wheat seedlings under drought stress[J].Biological Trace Element Research,2009,130(3):283-290.
[29]Friedman J,Stein Z,Rushkin E.Drought tolerance of germinating seeds and young seedlings of Anastatica hierochuntica L.[J].Oecologia,1981,51(3):400-403.
[30]杨景宁,王彦荣.PEG模拟干旱胁迫对四种荒漠植物种子萌发的影响[J].草业学报,2012,21(6):23-29.[Yang Jingning,Wang Yanrong.Effects of drought stress simulated by PEG on seed germination of four desert plant species[J].Acta Prataculturae Sinica,2012,21(6):23-29.
[31]魏振艳,黄海霞,欧阳尔乾,等.水盐胁迫对裸果子种子萌发的影响[J].干旱区研究,2018,35(6):1 444-1 449.[Wei Zhenyan,Huang Haixia,Ouyang Erqian,et al.Effects of salt and water stress on seed germination of Gymnocarpos przewalskii[J].Arid Zone Research,2018,35(6):1 444-1 449.]
[32]Pigliucci M.Evolution of phenotypic plasticity:Where are we going now?[J].Trends in Ecology&Evolution,2005,20(9):481-486.
[33]Almansouri M,Kinet J M,Lutts S.Effect of salt and osmotic stresses on germination in durum wheat(Triticum durum Desf.)[J].Plant&Soil,2001,231(2):243-254.
[34]Bajji M,Kinet J M,Lutts S.Salt stress effects on roots and leaves of Atriplex halimus L.and their corresponding callus cultures[J].Plant Science,1998,137(2):131-142.
[35]Bajji M,Kinet J M,Lutts S.Osmotic and ionic effects of Na Cl on germination,early seedling growth,and ion content of Atriplex halimus(Chenopodiaceae)[J].Canadian Journal of Botany,2002,80(3):297-304.
[36]Lai F M,Lecouteux C G,Mckersie B D.Germination of alfalfa(Medicago sativa L.)seeds and desiccated somatic embryos:I.Mobilization of storage reserves[J].Journal of Plant Physiology,1995,145(4):507-513.
[37]王习勇,魏岩,严成.温周期及果翅对梭梭种子萌发行为的调控[J].干旱区研究,2006,23(4):558-561.[Wang Xiyong,Wei Yan,Yan Cheng.Study on the effects of thermoperiods and fruiting wings on the germination of Haloxylon ammodendron seeds[J].Arid Zone Research,2006,23(4):558-561.]
[2]Volis S.Seed heteromorphism in Triticum dicoccoides:Association between seed positions within a dispersal unit and dormancy[J].Oecologia,2016,181(2):401-412.
[3]Jaganathan G K.Influence of maternal environment in developing different levels of physical dormancy and its ecological significance[J].Plant Ecology,2016,217(1):71-79.
[4]Gul B,Ansari R,Flowers T J,et al.Germination strategies of halophyte seeds under salinity[J].Environmental&Experimental Botany,2013,92(4):4-18.
[5]渠晓霞,黄振英.盐生植物种子萌发对环境的适应对策[J].生态学报,2005,25(9):2 389-2 398.[Qu Xiaoxia,Huang Zhenying.The adaptive strategies of halophyte seed germination[J].Acta Ecologica Sinica,2005,25(9):2 389-2 398.]
[6]Li R,Shi F,Fukuda K.Interactive effects of salt and alkali stresses on seed germination,germination recovery,and seedling growth of a halophyte Spartina alterniflora(Poaceae)[J].South African Journal of Botany,2010,76(2):380-387.
[7]Katembe W J,Ungar I A,Mitchell J P.Effect of salinity on germination and seedling growth of two Atriplex species(Chenopodiaceae)[J].Annals of Botany,1998,82(2):167-175.
[8]Khan M A,Ungar I A.Influence of salinity and temperature on the germination of Haloxylon recurvum Bunge ex.Boiss[J].Annals of Botany,1996,78(5):547-551.
[9]李宽亮,刘彤,李勇冠,等.褐翅猪毛菜种子异型性及其萌发特性研究[J].石河子大学学报(自然科学版),2012,30(3):276-281.[Li Kuanliang,Liu Tong,Li Yongguan,et al.Seed polymorphism and germination of Salsola korshinskyi Drob[J].Journal of Shihezi University(Natural Science Edition),2012,30(3):276-281.]
[10]Baskin J M,Lu J J,Baskin C C,et al.Diaspore dispersal ability and degree of dormancy in heteromorphic species of cold deserts of northwest China:A review[J].Perspectives in Plant Ecology Evolution&Systematics,2014,16(2):93-99.
[11]Mamut J,Tan D Y,Baskin C C,et al.Role of trichomes and pericarp in the seed biology of the desert annual Lachnoloma lehmannii(Brassicaceae)[J].Ecological Research,2014,29(1):33-44.
[12]Baskin J M,Baskin C C.A classification system for seed dormancy[J].Seed Science Research,2004,14(1):1-16.
[13]何美香,杜晓峰,陈玲,等.盐分、变温和激素处理对盐生植物异子蓬异型性种子萌发及成苗的影响[J].生态学杂志,2013,32(1):45-51.[He Meixiang,Du Xiaofeng,Chen Ling,et al.Effects of salt,alternating temperature and hormone treatments on seed germination and seedling establishment of Suaeda aralocaspica(Chenopodiaceae)dimorphic seeds[J].Chinese Journal of Ecology,2013,32(1):45-51.]
[14]Li J,Yin L Y,Jongsma M A,et al.Effects of light,hydropriming and abiotic stress on seed germination,and shoot and root growth of pyrethrum(Tanacetum cinerariifolium)[J].Industrial Crops&Products,2011,34(3):1 543-1 549.
[15]Wang Y,Jiang G Q,Han Y N,et al.Effects of salt,alkali and saltalkali mixed stresses on seed germination of the halophyte Salsola ferganica(Chenopodiaceae)[J].Acta Ecologica Sinica,2013,33(6):354-360.
[16]黄俊华.中国猪毛菜属(Salsola L.)植物的地理分布特点[J].干旱区地理,2005,28(3):325-329.[Huang Junhua.Geographical distribution of Salsola L.in China[J].Arid Land Geography,2005,28(3):325-329.]
[17]Ma Y L,Wang J,Zhang J H,et al.Seed heteromorphism and effects of light and abiotic stress on germination of a typical annual halophyte Salsola ferganica in cold desert[EB/OL].https://www.frontiersin.org/articles/10.3389/fpls.2017.02257/full,2018/2018-8-29.
[18]Yao S,Lan H,Zhang F.Variation of seed heteromorphism in Chenopodium album and the effect of salinity stress on the descendants[J].Annals of Botany,2010,105(6):1 015-1 025.
[19]Coons J M,Kuehl R O,Simons N R.Tolerance of ten lettuce cultivars to high temperature combined with Na Cl during germination[J].Journal of the American Society for Horticultural Science,1990,115(6):1 004-1 007.
[20]Kaufmann M R,Eckard A N.Evaluation of water stress control with polyethylene glycols by analysis of guttation[J].Plant Physiology,1971,47(4):453-456.
[21]Dodd G L,Donovan L A.Water potential and ionic effects on germination and seedling growth of two cold desert shrubs[J].American Journal of Botany,1999,86(8):1 146-1 153.
[22]Naylor R E L.An evaluation of various germination indices for predicting differences in seed vigour in Italian ryegrass[J].Seed Science&Technology,1981,9:593-600.
[23]Khan M A,Gul B,Weber D J.Seed germination in the Great Basin halophyte Salsola iberica[J].Revue Canadienne De Botanique,2002,80(6):650-655.
[24]Wei Y,Dong M,Huang Z,et al.Factors influencing seed germination of Salsola affinis(Chenopodiaceae),a dominant annual halophyte inhabiting the deserts of Xinjiang,China[J].Flora,2008,203(2):134-140.
[25]Ahmed M Z,Khan M A.Tolerance and recovery responses of playa halophytes to light,salinity and temperature stresses during seed germination[J].Flora,2010,205(11):764-771.
[26]Yildiz M,Cenkci S,Kargioglu M.Effects of salinity,temperature,and light on seed germination in two Turkish endemic halophytes,Limonium iconicum and L.lilacinum(Plumbaginaceae)[J].Seed Science&Technology,2008,36(3):646-656.
[27]Bhaskar G,Huang B R.Mechanism of salinity tolerance in plants:Physiological,biochemical,and molecular characterization[EB/OL].https://www.hindawi.com/journals/ijg/2014/701596/,2014/2018-8-29.
[28]Yao X,Chu J G.Effects of selenium on wheat seedlings under drought stress[J].Biological Trace Element Research,2009,130(3):283-290.
[29]Friedman J,Stein Z,Rushkin E.Drought tolerance of germinating seeds and young seedlings of Anastatica hierochuntica L.[J].Oecologia,1981,51(3):400-403.
[30]杨景宁,王彦荣.PEG模拟干旱胁迫对四种荒漠植物种子萌发的影响[J].草业学报,2012,21(6):23-29.[Yang Jingning,Wang Yanrong.Effects of drought stress simulated by PEG on seed germination of four desert plant species[J].Acta Prataculturae Sinica,2012,21(6):23-29.
[31]魏振艳,黄海霞,欧阳尔乾,等.水盐胁迫对裸果子种子萌发的影响[J].干旱区研究,2018,35(6):1 444-1 449.[Wei Zhenyan,Huang Haixia,Ouyang Erqian,et al.Effects of salt and water stress on seed germination of Gymnocarpos przewalskii[J].Arid Zone Research,2018,35(6):1 444-1 449.]
[32]Pigliucci M.Evolution of phenotypic plasticity:Where are we going now?[J].Trends in Ecology&Evolution,2005,20(9):481-486.
[33]Almansouri M,Kinet J M,Lutts S.Effect of salt and osmotic stresses on germination in durum wheat(Triticum durum Desf.)[J].Plant&Soil,2001,231(2):243-254.
[34]Bajji M,Kinet J M,Lutts S.Salt stress effects on roots and leaves of Atriplex halimus L.and their corresponding callus cultures[J].Plant Science,1998,137(2):131-142.
[35]Bajji M,Kinet J M,Lutts S.Osmotic and ionic effects of Na Cl on germination,early seedling growth,and ion content of Atriplex halimus(Chenopodiaceae)[J].Canadian Journal of Botany,2002,80(3):297-304.
[36]Lai F M,Lecouteux C G,Mckersie B D.Germination of alfalfa(Medicago sativa L.)seeds and desiccated somatic embryos:I.Mobilization of storage reserves[J].Journal of Plant Physiology,1995,145(4):507-513.
[37]王习勇,魏岩,严成.温周期及果翅对梭梭种子萌发行为的调控[J].干旱区研究,2006,23(4):558-561.[Wang Xiyong,Wei Yan,Yan Cheng.Study on the effects of thermoperiods and fruiting wings on the germination of Haloxylon ammodendron seeds[J].Arid Zone Research,2006,23(4):558-561.]