烟气对泥炭藓孢子萌发影响的模拟研究
|
摘要
适量烟气能促进种子萌发,但对苔藓植物孢子的作用尚不清楚.选取采自长白山区泥炭地的粗叶泥炭藓和中位泥炭藓的孢蒴为试验材料,通过燃烧泥炭地植物产生烟气,制备烟溶液,分别与不同大小(大:直径为2.10～2.50 mm;小:直径为1.50～1.90 mm)以及不同保存时长(旧:4.3和6.3年;新:0.3年)的孢蒴进行两组双因素试验,经不同时长的烟溶液浸泡和萌发试验,模拟研究烟气、孢蒴大小和保存时长对苔藓植物孢子萌发的影响.结果表明:烟溶液浸泡影响孢子萌发,培养10 d时,不同时长的烟溶液浸泡均可使孢子的萌发率提高5倍以上,小孢蒴孢子的萌发率高;培养21 d时,仅适度浸泡(3 d)表现出促进萌发的作用,孢蒴大小对孢子萌发率无影响;烟溶液浸泡对长时间保存(4.3和6.3年)的孢蒴孢子萌发无促进作用.研究表明,适量烟气可加速新泥炭藓孢子以及小孢蒴孢子的萌发.在存在不定期火烧干扰的泥炭地中,与对种子植物的作用类似,烟气可能在苔藓植物种群的有性更新和种群维持中发挥重要作用.
Moderate smoke could facilitate seed germination,but its effects on bryophyte spore germination is still unknown. Here,we analyzed the effects of smoke,capsule size and storage time on the spore germination of Sphagnum squarrosum and S. magellanicum,with the capsules of which being collected from two peatlands of the Changbai Mountains. The smoke solution prepared by burning peatland plants was combined with the capsules with different sizes( large,2.10-2.50 mm in diameter; small,1.50-1.90 mm in diameter) and storage time( old,being stored for 4.3 or 6.3a; new,being stored for 0. 3 a) to conduct a factorial experiment. The spores were soaked with smoke solution for different durations and then cultured for germination. The results showed that smoke solution affected spore germination. After 10 d cultivation,germination rate of spores soaking with smoke solution for all duration was increased by more than 5-fold,with the small spores having higher germination rate. After 21 d cultivation,the facilitative effect was only observed in moderate soaking( 3 d),and spore size showed no effect on germination. Smoke solution could not increase the germination of spores from the capsules with long storage time( 4.3 and 6.3 a). Our results indicated that moderate smoke solution soaking might accelerate germination of Sphagnum spores including small pores. In the ecosystems with casual fire disturbance such as peatlands,similar with its effects on the seed plants,smoke might play a key role in the regeneration and persistence of bryophyte population.
Moderate smoke could facilitate seed germination,but its effects on bryophyte spore germination is still unknown. Here,we analyzed the effects of smoke,capsule size and storage time on the spore germination of Sphagnum squarrosum and S. magellanicum,with the capsules of which being collected from two peatlands of the Changbai Mountains. The smoke solution prepared by burning peatland plants was combined with the capsules with different sizes( large,2.10-2.50 mm in diameter; small,1.50-1.90 mm in diameter) and storage time( old,being stored for 4.3 or 6.3a; new,being stored for 0. 3 a) to conduct a factorial experiment. The spores were soaked with smoke solution for different durations and then cultured for germination. The results showed that smoke solution affected spore germination. After 10 d cultivation,germination rate of spores soaking with smoke solution for all duration was increased by more than 5-fold,with the small spores having higher germination rate. After 21 d cultivation,the facilitative effect was only observed in moderate soaking( 3 d),and spore size showed no effect on germination. Smoke solution could not increase the germination of spores from the capsules with long storage time( 4.3 and 6.3 a). Our results indicated that moderate smoke solution soaking might accelerate germination of Sphagnum spores including small pores. In the ecosystems with casual fire disturbance such as peatlands,similar with its effects on the seed plants,smoke might play a key role in the regeneration and persistence of bryophyte population.
引文
[1] De Lange JH, Boucher C. Autecological studies on Audouinia capitata(Bruniaceae). I. Plant-derived smoke as a seed germination cue. South African Journal of Botany,1990,56:700-703
[2] Brown NAC. Seed germination in the fynbos fire ephemeral,Syncarpha vestita(L)B-Nord is promoted by smoke,aqueous extracts of smoke and charred wood derived from burning the ericoid-leaved shrub,Passerinavulgaris Thoday. International Journal of Wildland Fire,1993,3:203-206
[3] Baxter BJM,Staden JV. Plant-derived smoke:An effective seed pre-treatment. Plant Growth Regulation,1994,14:279-282
[4] Flematti GR,Ghisalberti EL,Dixon KW,et al. A compound from smoke that promotes seed germination.Science,2004,305:977
[5] Van Staden J,Brown NAC,Jger AK,et al. Smoke as germination cue. Plant Species Biology,2000,15:167-178
[6] Dixon KW,Merritt DJ,Flematti GR,et al. Karrikinolide:A phytoreactive compound derived from smoke with applications in horticulture,ecological restoration and agriculture. Acta Horticulturae,2009,813:155-170
[7] Nelson DC,Flematti GR,Ghisalberti EL,et al. Regulation of seed germination and seedling growth by chemical signals from burning vegetation. Annual Review of Plant Biology,2012,63:107-130
[8]atav SS,Kü9ükakyüz K,Akba? K,et al. Smokeenhanced seed germination in Mediterranean Lamiaceae.Seed Science Research,2014,24:257-264
[9] Zuloaga-Aguilar S,Briones O,Orozco-Segovia A. Seed germination of montane forest species in response to ash,smoke and heat shock in Mexico. Acta Oecologica,2011,37:256-262
[10] Dayamba SD,Sawadogo L,Tigabu M,et al. Effects of aqueous smoke solutions and heat on seed germination of herbaceous species of the Sudanian savanna-woodland in Burkina Faso. Flora:Morphology,Distribution,Functional Ecology of Plants,2010,205:319-325
[11] Dixon KW,Roche S. The role of combustion products(smoke)in stimulating ex situ and in situ germination of Western Australian plants. Proceedings of the International Plant Propagation Society,1995,45:53-56
[12] Chou YF,Cox RD,Wester DB. Smoke water and heat shock influence germination of shortgrass prairie species.Rangeland Ecology&Management,2012,65:260-267
[13] Ferraz IDK,Arruda YMBC,Van Staden J. Smoke-water effect on the germination of Amazonian tree species.South African Journal of Botany,2013,87:122-128
[14] Gresta F,Avola G,Anastasi U,et al. Effect of maturation stage,storage time and temperature on seed germination of Medicago species. Seed Science and Technology,2007,35:698-708
[15] Delgado JA,Serrano JM,López F,et al. Seed size and seed germination in the Mediterranean fire-prone shrub Cistus ladanifer. Plant Ecology,2008,197:269-276
[16] Smith CC,Fretwell SD. The optimal balance between size and number of offspring. American Naturalist,1974,108:499-506
[17] Joosten H,Clarke D. Wise use of mires and peatlands.International Mire Conservation Group and International Peat Society,Helsinki,2002:304
[18] Sundberg S,Rydin H. Experimental evidence for a persistent spore bank in Sphagnum. New Phytologist,2000,148:105-116
[19] Feng L(冯璐),Tang Y(汤袁),Bu Z-J(卜兆君),et al. A direct experimental proof for long-term persistent spore bank in Hani Peatland of the Changbai Mountains. Bulletin of Botanical Research(植物研究),2013,33(2):248-251(in Chinese)
[20] Sundberg S,Rydin H. Spore number in Sphagnum and its dependence on spore and capsule size. Journal of Bryology,1998,20:1-16
[21] Feng Y-M(冯亚敏),Bu Z-J(卜兆君),Feng L(冯璐),et al. Effects of nutrient concentration and light intensity on the sexual regeneration of Sphagnum palustre.Ecological Science(生态科学),2016(5):31-37(in Chinese)
[22] Minayeva T,Sirin AA,Stracher GB. The peat fires of Russia//Stracher GB,Prakak A,Sokol EV,eds. Coal and Peat fires:A Global Perspective vol. 2. Amsterdam,the Netherlands:Elsevier,2012:375-394
[23] Zhao K-Y(赵魁义). The Record of Mires in China.Beijing:Science Press,1999(in Chinese)
[24] Bu ZJ,Li Z,Liu LJ,et al. Bryophyte spore germinability is inhibited by peatland substrates. Acta Oecologica,2017,78:34-40
[25] Feng L,Bu ZJ,Mallik A,et al. Continuous waterlogging may not facilitate germinability maintenance of Sphagnum spores. Wetlands,2017,37:1015-1022
[26] Van Staden J,Jger AK,Light ME,et al. Isolation of the major germination cue from plant-derived smoke.South African Journal of Botany,2004,70:654-659
[27] Adkins SW,Peters NCB. Smoke derived from burnt vegetation stimulates germination of arable weeds. Seed Science Research,2001,11:213-222
[28] Daws MI,Davies J,Pritchard HW,et al. Butenolide from plant-derived smoke enhances germination and seedling growth of arable weed species. Plant Growth Regulation,2007,51:73-82
[29] Light ME,Daws MI,Van Staden J. Smoke-derived butenolide:Towards understanding its biological effects.South African Journal of Botany,2009,75:1-7
[30] Fenner M,Thompson K. The Ecology of Seeds. Cambridge:Cambridge University Press,2005
[31] Berger A. Seed dimorphism and germination behaviour in Salicornia patula. Vegetatio,1985,61:137-143
[32] Guzzon F,Orsenigo S,Gianella M,et al. Seed heteromorphy influences seed longevity in Aegilops. Seed Science Research, 2018, doi. org/10. 1017/S096025851800034X
[33] Wu G-L(武高林),Du G-Z(杜国祯),Shang Z-H(尚占环). Contribution of seed size and its fate to vegetation renewal:A review. Chinese Journal of Applied Ecology(应用生态学报),2006,17(10):1969-1972(in Chinese)
[34] Wu G-L(武高林),Du G-Z(杜国祯). Relationships between seed size and seedling growth strategy of herbaceous plant:A review. Chinese Journal of Applied Ecology(应用生态学报),2008,19(1):191-197(in Chinese)
[35] Mogensen GS. The biological significance of morphological characters in bryophytes:The spore. The Bryologist,1981,84:187-207
[36] Demir I,Ozden E,Yildirim KC,et al. Priming with smoke-derived karrikinolide enhances germination and transplant quality of immature and mature pepper seed lots. South African Journal of Botany,2018,115:264-268
[37] Bu ZJ,Sundberg S,Feng L,et al. The Methuselah of plant diaspores:Sphagnum spores can survive in nature for centuries. New Phytologist,2017,214:1398-1402
[38] Crosti R,Ladd PG,Dixon KW,et al. Post-fire germination:The effect of smoke on seeds of selected species from the central Mediterranean basin. Forest Ecology and Management,2006,221:306-312
[39] Marcisz K,Gaka M,Pietrala P,et al. Fire activity and hydrological dynamics in the past 5700 years reconstructed from Sphagnum peatlands along the oceanic-continental climatic gradient in northern Poland. Quaternary Science Reviews,2017,177:145-157
[40] Turetsky MR,Benscoter B,Page S,et al. Global vulnerability of peatlands to fire and carbon loss. Nature Geoscience,2015,8:11-14
[2] Brown NAC. Seed germination in the fynbos fire ephemeral,Syncarpha vestita(L)B-Nord is promoted by smoke,aqueous extracts of smoke and charred wood derived from burning the ericoid-leaved shrub,Passerinavulgaris Thoday. International Journal of Wildland Fire,1993,3:203-206
[3] Baxter BJM,Staden JV. Plant-derived smoke:An effective seed pre-treatment. Plant Growth Regulation,1994,14:279-282
[4] Flematti GR,Ghisalberti EL,Dixon KW,et al. A compound from smoke that promotes seed germination.Science,2004,305:977
[5] Van Staden J,Brown NAC,Jger AK,et al. Smoke as germination cue. Plant Species Biology,2000,15:167-178
[6] Dixon KW,Merritt DJ,Flematti GR,et al. Karrikinolide:A phytoreactive compound derived from smoke with applications in horticulture,ecological restoration and agriculture. Acta Horticulturae,2009,813:155-170
[7] Nelson DC,Flematti GR,Ghisalberti EL,et al. Regulation of seed germination and seedling growth by chemical signals from burning vegetation. Annual Review of Plant Biology,2012,63:107-130
[8]atav SS,Kü9ükakyüz K,Akba? K,et al. Smokeenhanced seed germination in Mediterranean Lamiaceae.Seed Science Research,2014,24:257-264
[9] Zuloaga-Aguilar S,Briones O,Orozco-Segovia A. Seed germination of montane forest species in response to ash,smoke and heat shock in Mexico. Acta Oecologica,2011,37:256-262
[10] Dayamba SD,Sawadogo L,Tigabu M,et al. Effects of aqueous smoke solutions and heat on seed germination of herbaceous species of the Sudanian savanna-woodland in Burkina Faso. Flora:Morphology,Distribution,Functional Ecology of Plants,2010,205:319-325
[11] Dixon KW,Roche S. The role of combustion products(smoke)in stimulating ex situ and in situ germination of Western Australian plants. Proceedings of the International Plant Propagation Society,1995,45:53-56
[12] Chou YF,Cox RD,Wester DB. Smoke water and heat shock influence germination of shortgrass prairie species.Rangeland Ecology&Management,2012,65:260-267
[13] Ferraz IDK,Arruda YMBC,Van Staden J. Smoke-water effect on the germination of Amazonian tree species.South African Journal of Botany,2013,87:122-128
[14] Gresta F,Avola G,Anastasi U,et al. Effect of maturation stage,storage time and temperature on seed germination of Medicago species. Seed Science and Technology,2007,35:698-708
[15] Delgado JA,Serrano JM,López F,et al. Seed size and seed germination in the Mediterranean fire-prone shrub Cistus ladanifer. Plant Ecology,2008,197:269-276
[16] Smith CC,Fretwell SD. The optimal balance between size and number of offspring. American Naturalist,1974,108:499-506
[17] Joosten H,Clarke D. Wise use of mires and peatlands.International Mire Conservation Group and International Peat Society,Helsinki,2002:304
[18] Sundberg S,Rydin H. Experimental evidence for a persistent spore bank in Sphagnum. New Phytologist,2000,148:105-116
[19] Feng L(冯璐),Tang Y(汤袁),Bu Z-J(卜兆君),et al. A direct experimental proof for long-term persistent spore bank in Hani Peatland of the Changbai Mountains. Bulletin of Botanical Research(植物研究),2013,33(2):248-251(in Chinese)
[20] Sundberg S,Rydin H. Spore number in Sphagnum and its dependence on spore and capsule size. Journal of Bryology,1998,20:1-16
[21] Feng Y-M(冯亚敏),Bu Z-J(卜兆君),Feng L(冯璐),et al. Effects of nutrient concentration and light intensity on the sexual regeneration of Sphagnum palustre.Ecological Science(生态科学),2016(5):31-37(in Chinese)
[22] Minayeva T,Sirin AA,Stracher GB. The peat fires of Russia//Stracher GB,Prakak A,Sokol EV,eds. Coal and Peat fires:A Global Perspective vol. 2. Amsterdam,the Netherlands:Elsevier,2012:375-394
[23] Zhao K-Y(赵魁义). The Record of Mires in China.Beijing:Science Press,1999(in Chinese)
[24] Bu ZJ,Li Z,Liu LJ,et al. Bryophyte spore germinability is inhibited by peatland substrates. Acta Oecologica,2017,78:34-40
[25] Feng L,Bu ZJ,Mallik A,et al. Continuous waterlogging may not facilitate germinability maintenance of Sphagnum spores. Wetlands,2017,37:1015-1022
[26] Van Staden J,Jger AK,Light ME,et al. Isolation of the major germination cue from plant-derived smoke.South African Journal of Botany,2004,70:654-659
[27] Adkins SW,Peters NCB. Smoke derived from burnt vegetation stimulates germination of arable weeds. Seed Science Research,2001,11:213-222
[28] Daws MI,Davies J,Pritchard HW,et al. Butenolide from plant-derived smoke enhances germination and seedling growth of arable weed species. Plant Growth Regulation,2007,51:73-82
[29] Light ME,Daws MI,Van Staden J. Smoke-derived butenolide:Towards understanding its biological effects.South African Journal of Botany,2009,75:1-7
[30] Fenner M,Thompson K. The Ecology of Seeds. Cambridge:Cambridge University Press,2005
[31] Berger A. Seed dimorphism and germination behaviour in Salicornia patula. Vegetatio,1985,61:137-143
[32] Guzzon F,Orsenigo S,Gianella M,et al. Seed heteromorphy influences seed longevity in Aegilops. Seed Science Research, 2018, doi. org/10. 1017/S096025851800034X
[33] Wu G-L(武高林),Du G-Z(杜国祯),Shang Z-H(尚占环). Contribution of seed size and its fate to vegetation renewal:A review. Chinese Journal of Applied Ecology(应用生态学报),2006,17(10):1969-1972(in Chinese)
[34] Wu G-L(武高林),Du G-Z(杜国祯). Relationships between seed size and seedling growth strategy of herbaceous plant:A review. Chinese Journal of Applied Ecology(应用生态学报),2008,19(1):191-197(in Chinese)
[35] Mogensen GS. The biological significance of morphological characters in bryophytes:The spore. The Bryologist,1981,84:187-207
[36] Demir I,Ozden E,Yildirim KC,et al. Priming with smoke-derived karrikinolide enhances germination and transplant quality of immature and mature pepper seed lots. South African Journal of Botany,2018,115:264-268
[37] Bu ZJ,Sundberg S,Feng L,et al. The Methuselah of plant diaspores:Sphagnum spores can survive in nature for centuries. New Phytologist,2017,214:1398-1402
[38] Crosti R,Ladd PG,Dixon KW,et al. Post-fire germination:The effect of smoke on seeds of selected species from the central Mediterranean basin. Forest Ecology and Management,2006,221:306-312
[39] Marcisz K,Gaka M,Pietrala P,et al. Fire activity and hydrological dynamics in the past 5700 years reconstructed from Sphagnum peatlands along the oceanic-continental climatic gradient in northern Poland. Quaternary Science Reviews,2017,177:145-157
[40] Turetsky MR,Benscoter B,Page S,et al. Global vulnerability of peatlands to fire and carbon loss. Nature Geoscience,2015,8:11-14