大叶藻种子萌发环境条件研究
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摘要
大叶藻海草床的退化和恢复受到国内外学者的广泛关注,而大叶藻种子萌发则是恢复实践中的一项关键技术。为了提高大叶藻种子萌发率,本研究在实验室可控条件下,采用正交实验对影响大叶藻种子萌发的环境因素(盐度、温度、N/P营养盐浓度及p H)进行了优化,并在优化条件下进一步探究了光质类型、UV-B辐射、低温层积和划破种皮对大叶藻种子萌发和平均萌发时间的影响。结果表明:盐度12、温度10°C、N/P营养盐浓度960/60μmol/L、p H 6.2是大叶藻种子相对适宜的萌发条件组合;黑暗条件下大叶藻种子萌发率最高,为92%,不同类型光质照射不促进大叶藻种子萌发率的提高,但红光照射可在短期内加速种子萌发;在一定强度范围内,种子萌发率随UV-B辐射时长的增加而升高,每日辐射8h,处理8周后种子萌发率可达93%。划破种皮和低温层积均可显著提高大叶藻种子的萌发率并缩短平均萌发时间,经划破种皮处理后,2周内种子萌发率可提高至91%。
In recent years, vegetation of seagrass species Zostera marina has been degenerated in many places around world. To solve this problem, seed germination is the key for the seagrass bed restoration. To examine the optimal combination effect of four single environmental factors(salinity, temperature, N/P concentration, and p H) on the seed germination, we conducted an orthogonal experiment under controlled indoor conditions. The results show that the optimal condition was salinity 12, 10°C, N/P concentration 960/60μmol/L, and p H 6.2. On top of that, we investigated the influences of ultraviolet B radiation, lighting(different in color, duration, and intensity), cold stratification, and seed coat scarification on the germination, and found that dark condition(the germination rate reached 92%) but lighting properties was favorable, except for short red illumination treatment which enhanced the germination somewhat. In addition, ultraviolet B radiation boosted the germination significantly(P<0.05) within a certain period of radiation. Both cold stratification and seed coat scarification could improve the germination and shorten the germination time significantly(P<0.05). The experimental results provide a theoretical basis for the indoor seeding cultivation of Z. marina, and may help the success of application to restore the ecosystem of seagrass bed.
In recent years, vegetation of seagrass species Zostera marina has been degenerated in many places around world. To solve this problem, seed germination is the key for the seagrass bed restoration. To examine the optimal combination effect of four single environmental factors(salinity, temperature, N/P concentration, and p H) on the seed germination, we conducted an orthogonal experiment under controlled indoor conditions. The results show that the optimal condition was salinity 12, 10°C, N/P concentration 960/60μmol/L, and p H 6.2. On top of that, we investigated the influences of ultraviolet B radiation, lighting(different in color, duration, and intensity), cold stratification, and seed coat scarification on the germination, and found that dark condition(the germination rate reached 92%) but lighting properties was favorable, except for short red illumination treatment which enhanced the germination somewhat. In addition, ultraviolet B radiation boosted the germination significantly(P<0.05) within a certain period of radiation. Both cold stratification and seed coat scarification could improve the germination and shorten the germination time significantly(P<0.05). The experimental results provide a theoretical basis for the indoor seeding cultivation of Z. marina, and may help the success of application to restore the ecosystem of seagrass bed.
引文
陈开宁,李文朝,潘继征,2005.不同处理对蓖齿眼子菜(Potamogeton pectinatus L.)种子萌发率的影响.湖泊科学,17(3):237-242
陈南凯,王玉兰,1995.紫外线处理农作物种子能增产.云南农业科技,(3):15-16
程丽巍,2010.三种大型海藻对海水中营养盐供应变化的生理响应研究.南京:南京农业大学硕士学位论文
方媛,于海宁,程曦等,2010.增强UV-B辐射对甘草种子萌发及幼苗形态的影响.中国农学通报,26(2):122-126
高健,熊勤,李祝等,2006.溶氧、p H及其他理化因子对蓖齿眼子菜种子萌发的影响.水生生物学报,30(5):524-529
范航清,石雅君,邱广龙,2009.中国海草植物.北京:海洋出版社
郭卫东,章小明,杨逸萍等,1998.中国近岸海域潜在性富营养化程度的评价.台湾海峡,17(1):64-70
郭彦,杨洪双,梁明,2011.紫外线照射对水稻种子萌发的影响.江苏农业科学,39(3):89-90
何海洋,胡春芹,丁强强等,2013.不同p H对光皮桦种子萌发及幼苗生长的影响.西南林业大学学报,33(5):29-33,39
胡正华,索福喜,赵晓莉等,2005.UV-B辐射增加与酸雨复合处理对菠菜种子萌发和幼苗生长的影响.中国农学通报,21(6):284-285,294
黄婷,董召荣,宋贺等,2007.模拟酸雨对紫花苜蓿种子萌发及幼苗生理特性的影响.种子,26(4):21-24
李雯琳,郁继华,杨其长,2013.不同光质对叶用莴苣种子萌发和幼苗酶活性的影响.甘肃农业大学学报,48(3):44-49
李焰焰,聂传朋,董召荣,2005.模拟酸雨对小黑麦种子萌发及幼苗生理特性的影响.生态学杂志,24(4):395-397
马红媛,梁正伟,2007.不同p H值土壤及其浸提液对羊草种子萌发和幼苗生长的影响.植物学通报,24(2):181-188
王维荣,陈松,欧阳光察等,1991.光质对黄瓜种子萌发过程中过氧化物酶活性及蛋白含量的影响.上海农业学报,7(4):17-20
王伟伟,李晓捷,潘金华等,2013.大叶藻资源动态及生态恢复面临的问题.海洋环境科学,32(2):316-320
由文辉,宋永昌,1995.淀山湖3种沉水植物的种子萌发生态.应用生态学报,6(2):196-200
叶春江,赵可夫.高等植物大叶藻研究进展及其对海洋沉水生活的适应.植物学通报,2002,19(2):184-193
郑凤英,邱广龙,范航清等,2013.中国海草的多样性、分布及保护.生物多样性,21(5):517-526
Abe M,Kurashima A,Maegawa M,2008.Temperature requirements for seed germination and seedling growth of Zostera marina from central Japan.Fisheries Science,74(3):589-593
Ailstock S,Shafer D,2006.Protocol for Large-scale Collection,Processing,and Storage of Seeds of Two Mesohaline Submerged Aquatic Plant Species.Vicksburg,MS,USA:USArmy Engineer Research and Development Center
Arts G H P,van der Heijden R A J M,1990.Germination ecology of Littorella uniflora(L.)Aschers.Aquatic Botany,37(2):139-151
Brenchley J L,Probert R J,1998.Seed germination responses to some environmental factors in the seagrass Zostera capricorni from eastern Australia.Aquatic Botany,62(3):177-188
Caye G,Bulard C,Meinesz A et al,1992.Dominant role of seawater osmotic pressure on germination in Cymodocea nodosa.Aquatic Botany,42(2):187-193
Churchill A C,1983.Field studies on seed germination and seedling development in Zostera marina L.Aquatic Botany,16(1):21-29
Conacher CA,Poiner I R,Butler J et al,1994.Germination,storage and viability testing of seeds of Zostera capricorni Aschers.from a tropical bay in Australia.Aquatic Botany,49(1):47-58
Costello C T,Kenworthy W J,2011.Twelve-year mapping and change analysis of eelgrass(Zostera marina)areal abundance in Massachusetts(USA)identifies statewide declines.Estuaries and Coasts,34(2):232-242
Delefosse M,Kristensen E,2012.Burial of Zostera marina seeds in sediment inhabited by three polychaetes:Laboratory and field studies.Journal of Sea Research,71:41-49
Dennison W C,1987.Effects of light on seagrass photosynthesis,growth and depth distribution.Aquatic Botany,27(1):15-26
Fernández-Torquemada Y,Sánchez-Lizaso J L,2013.Effects of salinity on seed germination and early seedling growth of the Mediterranean seagrass Posidonia oceanica(L.)Delile.Estuarine,Coastal and Shelf Science,119:64-70
Fong P,Fong J J,Fong C R,2004.Growth,nutrient storage,and release of dissolved organic nitrogen by Enteromorpha intestinalis in response to pulses of nitrogen and phosphorus.Aquatic Botany,78(l):83-95
Fourqurean J W,Duarte C M,Kennedy H et al,2012.Seagrass ecosystems as a globally significant carbon stock.Nature Geoscience,5(7):505-509
Harrison P G,1991.Mechanisms of seed dormancy in an annual population of Zostera marina(eelgrass)from the Netherlands.Canadian Journal of Botany,69(9):1972-1976
Hay F,Probert R,Dawson M,2008.Laboratory germination of seeds from 10 British species of Potamogeton.Aquatic Botany,88(4):353-357
Hootsmans M J M,Vermaat J E,Van Vierssen W,1987.Seed-bank development,germination and early seedling survival of two seagrass species from The Netherlands:Zostera marina L.and Zostera noltii hornem.Aquatic Botany,28(3-4):275-285
ISTA,1996.International Rules for seed testing.Seed Science and Technology.Zürich,Switzerland:ISTA
Jarvis J C,Moore K A,2008.Influence of environmental factors on Vallisneria americana seed germination.Aquatic Botany,88(4):283-294
Kahn A E,Durako M J,2005.The effect of salinity and ammonium on seed germination in Ruppia maritima from Florida Bay.Bulletin of Marine Science,77(3):453-458
Karssen C M,Hilhorst H W,Koomneef M,1990.The benefit of biosynthesis and response mutants to the study of the role of abscisic acid in plants.In:Pharis P R P,Rood P S B eds.Plant Growth Substances.Berlin Heidelberg,Germany:Springer,23-31
Ke X S,Li W,2006.Germination requirement of Vallisneria natans seeds:implications for restoration in Chinese lakes.Hydrobiologia,559(1):357-362
Kendrick G A,Waycott M,Carruthers T J B et al,2012.The central role of dispersal in the maintenance and persistence of seagrass populations.Bio Science,62(1):56-65
Kettenring K M,Galatowitsch S M,2007.Temperature requirements for dormancy break and seed germination vary greatly among 14 wetland Carex species.Aquatic Botany,87(3):209-220
Koch E W,Ailstock M S,Booth D M et al,2010.The role of currents and waves in the dispersal of submersed angiosperm seeds and seedlings.Restoration Ecology,18(4):584-595
Krause-Jensen D,Greve T M,Nielsen K,2005.Eelgrass as a bioindicator under the European Water Framework Directive.Water Resources Management,19(1):63-75
Lal C,Gopal B,1993.Production and germination of seeds in Hydrilla verticillata.Aquatic Botany,45(2-3):257-261
Lee K S,Short F T,Burdick D M,2004.Development of a nutrient pollution indicator using the seagrass,Zostera marina,along nutrient gradients in three New England estuaries.Aquatic Botany,78(3):197-216
Li W T,Kim S H,Kim J W et al,2013.An examination of photoacclimatory responses of Zostera marina transplants along a depth gradient for transplant-site selection in a disturbed estuary.Estuarine,Coastal and Shelf Science,118:72-79
Loquès F,Caye G,Meinesz A,1990.Germination in the marine phanerogam Zostera noltii Hornemann at Golfe Juan,French Mediterranean.Aquatic Botany,38(2-3):249-260
Mc Glathery K J,Pedersen M F,Borum J,1996.Changes in intracellular nitrogen pools and feedback controls on nitrogen uptake in Chaetomorpha linum(chlorophyta).Journal of Phycology,32(3):393-401
Olesen B,Sand-Jensen K,1993.Seasonal acclimatization of eelgrass Zostera marina growth to light.Marine Ecology Progress Series,94:91-99
Orth R J,Carruthers T J B,Dennison W C et al,2006.A global crisis for seagrass ecosystems.Bio Science,56(12):987-996
Orth R J,Harwell M C,Bailey E M et al,2000.A review of issues in seagrass seed dormancy and germination:implications for conservation and restoration.Marine Ecology Progress Series,200:277-288
Orth R J,Marion S R,Moore K A et al,2010.Eelgrass(Zostera marina L.)in the Chesapeake Bay region of mid-Atlantic coast of the USA:challenges in conservation and restoration.Estuaries and Coasts,33(1):139-150
Orth R J,Moore K A,1983.Seed germination and seedling growth of Zostera marina L.(eelgrass)in the chesapeake bay.Aquatic Botany,15(2):117-131
Pan J H,Jiang X,Li X J et al,2011.Influence of temperature and salinity on germination of eelgrass(Zostera marina L.)seeds.Journal of Ocean University of China,10(2):147-152
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Ranal M A,de Santana D G,2006.How and why to measure the germination process?Brazilian Journal of Botany,29(1):1-11
Reynolds L K,Waycott M,Mc Glathery K J et al,2012.Eelgrass restoration by seed maintains genetic diversity:case study from a coastal bay system.Marine Ecology Progress Series,448:223-233
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Strazisar T,Koch M S,Madden C J et al,2013.Salinity effects on Ruppia maritima L.seed germination and seedling survival at the Everglades-Florida Bay ecotone.Journal of Experimental Marine Biology and Ecology,445:129-139
Sugiura H,Hiroe Y,Suzuki T et al,2009.The carbohydrate catabolism of Zostera marina influenced by lower salinity during the pre-germination stage.Fisheries Science,75(5):1205-1217
Tanner C E,Parham T,2010.Growing Zostera marina(eelgrass)from Seeds in Land-Based Culture Systems for Use in Restoration Projects.Restoration Ecology,18(4):527-537
Taylor A R A,1957.Studies of the development of Zostera marina L.:II.Germination and seedling development.Canadian Journal of Botany,35(5):681-695
Vandelook F,Van Assche J A,2008.Temperature requirements for seed germination and seedling development determine timing of seedling emergence of three monocotyledonous temperate forest spring geophytes.Annals of Botany,102(5):865-875
陈南凯,王玉兰,1995.紫外线处理农作物种子能增产.云南农业科技,(3):15-16
程丽巍,2010.三种大型海藻对海水中营养盐供应变化的生理响应研究.南京:南京农业大学硕士学位论文
方媛,于海宁,程曦等,2010.增强UV-B辐射对甘草种子萌发及幼苗形态的影响.中国农学通报,26(2):122-126
高健,熊勤,李祝等,2006.溶氧、p H及其他理化因子对蓖齿眼子菜种子萌发的影响.水生生物学报,30(5):524-529
范航清,石雅君,邱广龙,2009.中国海草植物.北京:海洋出版社
郭卫东,章小明,杨逸萍等,1998.中国近岸海域潜在性富营养化程度的评价.台湾海峡,17(1):64-70
郭彦,杨洪双,梁明,2011.紫外线照射对水稻种子萌发的影响.江苏农业科学,39(3):89-90
何海洋,胡春芹,丁强强等,2013.不同p H对光皮桦种子萌发及幼苗生长的影响.西南林业大学学报,33(5):29-33,39
胡正华,索福喜,赵晓莉等,2005.UV-B辐射增加与酸雨复合处理对菠菜种子萌发和幼苗生长的影响.中国农学通报,21(6):284-285,294
黄婷,董召荣,宋贺等,2007.模拟酸雨对紫花苜蓿种子萌发及幼苗生理特性的影响.种子,26(4):21-24
李雯琳,郁继华,杨其长,2013.不同光质对叶用莴苣种子萌发和幼苗酶活性的影响.甘肃农业大学学报,48(3):44-49
李焰焰,聂传朋,董召荣,2005.模拟酸雨对小黑麦种子萌发及幼苗生理特性的影响.生态学杂志,24(4):395-397
马红媛,梁正伟,2007.不同p H值土壤及其浸提液对羊草种子萌发和幼苗生长的影响.植物学通报,24(2):181-188
王维荣,陈松,欧阳光察等,1991.光质对黄瓜种子萌发过程中过氧化物酶活性及蛋白含量的影响.上海农业学报,7(4):17-20
王伟伟,李晓捷,潘金华等,2013.大叶藻资源动态及生态恢复面临的问题.海洋环境科学,32(2):316-320
由文辉,宋永昌,1995.淀山湖3种沉水植物的种子萌发生态.应用生态学报,6(2):196-200
叶春江,赵可夫.高等植物大叶藻研究进展及其对海洋沉水生活的适应.植物学通报,2002,19(2):184-193
郑凤英,邱广龙,范航清等,2013.中国海草的多样性、分布及保护.生物多样性,21(5):517-526
Abe M,Kurashima A,Maegawa M,2008.Temperature requirements for seed germination and seedling growth of Zostera marina from central Japan.Fisheries Science,74(3):589-593
Ailstock S,Shafer D,2006.Protocol for Large-scale Collection,Processing,and Storage of Seeds of Two Mesohaline Submerged Aquatic Plant Species.Vicksburg,MS,USA:USArmy Engineer Research and Development Center
Arts G H P,van der Heijden R A J M,1990.Germination ecology of Littorella uniflora(L.)Aschers.Aquatic Botany,37(2):139-151
Brenchley J L,Probert R J,1998.Seed germination responses to some environmental factors in the seagrass Zostera capricorni from eastern Australia.Aquatic Botany,62(3):177-188
Caye G,Bulard C,Meinesz A et al,1992.Dominant role of seawater osmotic pressure on germination in Cymodocea nodosa.Aquatic Botany,42(2):187-193
Churchill A C,1983.Field studies on seed germination and seedling development in Zostera marina L.Aquatic Botany,16(1):21-29
Conacher CA,Poiner I R,Butler J et al,1994.Germination,storage and viability testing of seeds of Zostera capricorni Aschers.from a tropical bay in Australia.Aquatic Botany,49(1):47-58
Costello C T,Kenworthy W J,2011.Twelve-year mapping and change analysis of eelgrass(Zostera marina)areal abundance in Massachusetts(USA)identifies statewide declines.Estuaries and Coasts,34(2):232-242
Delefosse M,Kristensen E,2012.Burial of Zostera marina seeds in sediment inhabited by three polychaetes:Laboratory and field studies.Journal of Sea Research,71:41-49
Dennison W C,1987.Effects of light on seagrass photosynthesis,growth and depth distribution.Aquatic Botany,27(1):15-26
Fernández-Torquemada Y,Sánchez-Lizaso J L,2013.Effects of salinity on seed germination and early seedling growth of the Mediterranean seagrass Posidonia oceanica(L.)Delile.Estuarine,Coastal and Shelf Science,119:64-70
Fong P,Fong J J,Fong C R,2004.Growth,nutrient storage,and release of dissolved organic nitrogen by Enteromorpha intestinalis in response to pulses of nitrogen and phosphorus.Aquatic Botany,78(l):83-95
Fourqurean J W,Duarte C M,Kennedy H et al,2012.Seagrass ecosystems as a globally significant carbon stock.Nature Geoscience,5(7):505-509
Harrison P G,1991.Mechanisms of seed dormancy in an annual population of Zostera marina(eelgrass)from the Netherlands.Canadian Journal of Botany,69(9):1972-1976
Hay F,Probert R,Dawson M,2008.Laboratory germination of seeds from 10 British species of Potamogeton.Aquatic Botany,88(4):353-357
Hootsmans M J M,Vermaat J E,Van Vierssen W,1987.Seed-bank development,germination and early seedling survival of two seagrass species from The Netherlands:Zostera marina L.and Zostera noltii hornem.Aquatic Botany,28(3-4):275-285
ISTA,1996.International Rules for seed testing.Seed Science and Technology.Zürich,Switzerland:ISTA
Jarvis J C,Moore K A,2008.Influence of environmental factors on Vallisneria americana seed germination.Aquatic Botany,88(4):283-294
Kahn A E,Durako M J,2005.The effect of salinity and ammonium on seed germination in Ruppia maritima from Florida Bay.Bulletin of Marine Science,77(3):453-458
Karssen C M,Hilhorst H W,Koomneef M,1990.The benefit of biosynthesis and response mutants to the study of the role of abscisic acid in plants.In:Pharis P R P,Rood P S B eds.Plant Growth Substances.Berlin Heidelberg,Germany:Springer,23-31
Ke X S,Li W,2006.Germination requirement of Vallisneria natans seeds:implications for restoration in Chinese lakes.Hydrobiologia,559(1):357-362
Kendrick G A,Waycott M,Carruthers T J B et al,2012.The central role of dispersal in the maintenance and persistence of seagrass populations.Bio Science,62(1):56-65
Kettenring K M,Galatowitsch S M,2007.Temperature requirements for dormancy break and seed germination vary greatly among 14 wetland Carex species.Aquatic Botany,87(3):209-220
Koch E W,Ailstock M S,Booth D M et al,2010.The role of currents and waves in the dispersal of submersed angiosperm seeds and seedlings.Restoration Ecology,18(4):584-595
Krause-Jensen D,Greve T M,Nielsen K,2005.Eelgrass as a bioindicator under the European Water Framework Directive.Water Resources Management,19(1):63-75
Lal C,Gopal B,1993.Production and germination of seeds in Hydrilla verticillata.Aquatic Botany,45(2-3):257-261
Lee K S,Short F T,Burdick D M,2004.Development of a nutrient pollution indicator using the seagrass,Zostera marina,along nutrient gradients in three New England estuaries.Aquatic Botany,78(3):197-216
Li W T,Kim S H,Kim J W et al,2013.An examination of photoacclimatory responses of Zostera marina transplants along a depth gradient for transplant-site selection in a disturbed estuary.Estuarine,Coastal and Shelf Science,118:72-79
Loquès F,Caye G,Meinesz A,1990.Germination in the marine phanerogam Zostera noltii Hornemann at Golfe Juan,French Mediterranean.Aquatic Botany,38(2-3):249-260
Mc Glathery K J,Pedersen M F,Borum J,1996.Changes in intracellular nitrogen pools and feedback controls on nitrogen uptake in Chaetomorpha linum(chlorophyta).Journal of Phycology,32(3):393-401
Olesen B,Sand-Jensen K,1993.Seasonal acclimatization of eelgrass Zostera marina growth to light.Marine Ecology Progress Series,94:91-99
Orth R J,Carruthers T J B,Dennison W C et al,2006.A global crisis for seagrass ecosystems.Bio Science,56(12):987-996
Orth R J,Harwell M C,Bailey E M et al,2000.A review of issues in seagrass seed dormancy and germination:implications for conservation and restoration.Marine Ecology Progress Series,200:277-288
Orth R J,Marion S R,Moore K A et al,2010.Eelgrass(Zostera marina L.)in the Chesapeake Bay region of mid-Atlantic coast of the USA:challenges in conservation and restoration.Estuaries and Coasts,33(1):139-150
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