光强和硝态氮水平对铜藻(Sargassum horneri)生长和光合生理特性的影响
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摘要
以金潮种铜藻(Sargassum horneri)为对象,探讨了不同硝态氮水平下光照强度对其生长和光合生理特性的影响,以期为金潮暴发机制的研究提供一定的参考。实验设置了高(100μmol·L-1)、低(自然海水,50μmol·L-1) 2个硝态氮浓度和高(150μmol photons·m-2·s-1)、中(60μmol photons·m-2·s-1)、低(10μmol photons·m-2·s-1)三个光照水平,检测了藻体在不同硝态氮浓度和光照条件下适应培养7 d后生长、色素含量、可溶性糖含量、可溶性蛋白含量、氮吸收速率及最大光合放氧速率(Pmax)等生理指标的变化。结果表明:氮浓度相同时,铜藻的生长速率、可溶性糖含量、可溶性蛋白含量、氮吸收速率、Pmax均随光照强度的增加而增加,而色素含量随着光照强度的增加而降低;低氮条件下,除类胡萝卜素外,以上指标在高光处理下均显著高于低光处理(P<0.05);高氮条件下,除可溶性糖外,以上指标在高光条件下也均显著高于低光处理(P<0.05);同一光照强度下,色素及可溶性蛋白含量、氮吸收速率及Pmax随氮浓度的增加而增加;可溶性糖含量随着硝态氮浓度的增加而减少;且在3个光照条件下,高氮处理下藻体的Pmax均显著高于低氮处理。总之,不同光照和氮浓度处理7 d后,高光、高氮适应后的藻体表现出较高的生长和光合作用潜能。铜藻断裂后漂浮于海面极易获得较高光强条件,而富营养海水中的高氮条件会刺激铜藻的光合潜能而使其加速生长,进而加剧铜藻金潮的暴发。
The effects of light intensity on the growth and photosynthetic characteristics of Sargassum horneri,a golden tide causing species,were investigated under different nitrate( NO3--N)levels to reveal the physiological mechanism underlying the outbreak of golden tide. There were two NO3--N levels and three light intensities: i.e. high-N( 100 μmol·L-1),low-N( 50 μmol·L-1,the normal NO3-concentration in natural seawater),high-light( 150 μmol photons·m-2·s-1),medium-light( 60 μmol photons·m-2·s-1) and low-light( 10 μmol photons·m-2·s-1). The growth,pigment concentration,soluble sugar concentration,soluble protein concentration,nitrogen absorption rate and the maximum photosynthetic oxygen evolution rate( Pmax) of S.horneri thalli were measured after they had acclimated to different NO3--N and light levels for seven days. The results showed that the growth rate,soluble sugar concentration,soluble protein concentration,nitrogen absorption rate,and Pmaxof S. horneri thalli all increased with increased light intensities,while the pigment concentration decreased with the increasing light intensities.For the low-N acclimated S. horneri thalli,except for carotenoids,all the above mentioned indicators under the high light treatment were significantly higher than those under the low light treatment( P<0.05). For the high-N acclimated ones,except for the soluble sugar,all the indicators under high light treatment were significantly higher than those under the low light treatment( P<0.05). For the S. horneri thalli acclimated under same light intensities,pigment concentration,soluble protein concentration,nitrogen absorption rate and Pmaxincreased with the increasing NO3--N level,while the soluble sugar concentration decreased with the increasing NO3--N level.For the S. horneri thalli acclimated to the three light conditions,the Pmaxunder the high-N treatment was significantly higher than that under the low-N treatment. In conclusion,after seven days acclimation to different light intensities and NO3--N levels,S. horneri thalli showed higher potentials in growth and photosynthesis under the condition of high light intensity and high nitrogen level. Therefore,eutrophication in seawater would aggravate the occurrence of golden tide in that S. horneri thalli can obtain higher light intensities when they break from the stems and drift on the sea surface,which would stimulate the potentials of growth and photosynthesis.
The effects of light intensity on the growth and photosynthetic characteristics of Sargassum horneri,a golden tide causing species,were investigated under different nitrate( NO3--N)levels to reveal the physiological mechanism underlying the outbreak of golden tide. There were two NO3--N levels and three light intensities: i.e. high-N( 100 μmol·L-1),low-N( 50 μmol·L-1,the normal NO3-concentration in natural seawater),high-light( 150 μmol photons·m-2·s-1),medium-light( 60 μmol photons·m-2·s-1) and low-light( 10 μmol photons·m-2·s-1). The growth,pigment concentration,soluble sugar concentration,soluble protein concentration,nitrogen absorption rate and the maximum photosynthetic oxygen evolution rate( Pmax) of S.horneri thalli were measured after they had acclimated to different NO3--N and light levels for seven days. The results showed that the growth rate,soluble sugar concentration,soluble protein concentration,nitrogen absorption rate,and Pmaxof S. horneri thalli all increased with increased light intensities,while the pigment concentration decreased with the increasing light intensities.For the low-N acclimated S. horneri thalli,except for carotenoids,all the above mentioned indicators under the high light treatment were significantly higher than those under the low light treatment( P<0.05). For the high-N acclimated ones,except for the soluble sugar,all the indicators under high light treatment were significantly higher than those under the low light treatment( P<0.05). For the S. horneri thalli acclimated under same light intensities,pigment concentration,soluble protein concentration,nitrogen absorption rate and Pmaxincreased with the increasing NO3--N level,while the soluble sugar concentration decreased with the increasing NO3--N level.For the S. horneri thalli acclimated to the three light conditions,the Pmaxunder the high-N treatment was significantly higher than that under the low-N treatment. In conclusion,after seven days acclimation to different light intensities and NO3--N levels,S. horneri thalli showed higher potentials in growth and photosynthesis under the condition of high light intensity and high nitrogen level. Therefore,eutrophication in seawater would aggravate the occurrence of golden tide in that S. horneri thalli can obtain higher light intensities when they break from the stems and drift on the sea surface,which would stimulate the potentials of growth and photosynthesis.
引文
包楠欧,史定刚,关万春,等.2014.CO2及光强对南麂列岛铜藻生长的影响.浙江农业学报,26(3):649-655.
曹春晖,刘文岭,施定基,等.2010.不同氮磷浓度对米氏凯伦藻生长的影响.天津科技大学学报,25(2):22-25.
陈自强,孙庆海,陈全震,等.2013.酸胁迫对铜藻早期生长阶段的影响.生态毒理学报,8(6):864-870.
方婷轩,马增岭.2016.温度变化对铜绿微囊藻(Microcystis aeruginosa)生长和光合作用的影响.生态科学,35(6):14-23.
李慧,丁刚,辛美丽,等.2017.不同氮、磷浓度及配比对铜藻(Sargassum horneri)幼苗生长的影响.海洋与湖沼,48(2):368-372.
李科,潘耀茹,吴嘉平,等.2017.不同光照周期下LED光源对铜藻生长的影响.浙江农业学报,29(4):631-636.
李磊,关万春,陈少波,等.2016.水体N/P对塔玛亚历山大藻响应紫外辐射的影响.生态学杂志,35(2):395-400.
李信书,伏光辉,陈百尧,等.2012.氮、磷加富对条斑紫菜生长及生化组成的影响.水产科学,31(9):544-548.
司冉冉,关万春,蔡景波,等.2017.氮源对塔玛亚历山大藻生长和毒性的影响.生态学杂志,36(10):2880-2885.
孙建璋,庄定根,孙庆海,等.2009.铜藻人工栽培的初步研究.南方水产科学,5(6):41-46.
王丽梅,李世国,柴雨,等.2011.鼠尾藻幼苗的室内培养及有性生殖同步化.水产学报,35(3):395-404.
王巧晗.2008.环境因子节律性变动对潮间带大型海藻孢子萌发、早期发育和生长的影响及其生理生态学机制(硕士学位论文).青岛:中国海洋大学.
徐智广,吴海一,孙福新,等.2016.不同氮源加富对海带生长、光合固碳和氮吸收特性的影响.水产学报,40(4):577-584.
徐智广,邹定辉,高坤山,等.2011.不同温度、光照强度和硝氮浓度下龙须菜对无机磷吸收的影响.水产学报,35(7):1023-1029.
徐智广,邹定辉,张鑫,等.2008.CO2和硝氮加富对龙须菜(Gracilaria lemaneiformis)生长、生化组分和营养盐吸收的影响.生态学报,28(8):3752-3759.
杨安强,史定刚,陈少波,等.2017.高CO2条件下温度和光照对铜藻吸收N、P的短期影响.浙江农业科学,58(2):303-307.
张鹏,王铁杆,谢起浪,等.2014.培养条件对铜藻幼苗生长发育的影响.上海海洋大学学报,23(2):200-207.
张玉荣,刘峰,单体锋,等.2009.利用叶绿素荧光技术揭示人工培育的铜藻幼苗对胁迫温度、光照和盐度的反应.南方水产科学,5(2):1-9.
周百成,郑舜琴,曾呈奎.1974.几种绿藻、褐藻和红藻的吸收光谱的比较研究.植物学报,16(2):146-155.
Davison JR,Jordan TL,Fegley JC,et al.2007.Response of Laminaria saccharina(Phaeophyta)growth and photosynthesis to simultaneous ultraviolet radiation and nitrogen limitation.Journal of Phycology,43:636-646.
Falkowski PG,Raven JA.2007.Aquatic Photosynthesis.Princeton:Princeton University Press.
Geun CC,Geun KH,Hyun SC.2003.Transplantation of young fronds of Sargassum horneri for construction of seaweed beds.Journal of Geotechnical&Geoenvironmental Engineering,36:469-473.
Giordano M,Beardall J,Raven JA.2005.CO2concentrating mechanisms in algae:Mechanisms,environmental modulation,and evolution.Annual Review of Plant Biology,56:99-131.
Hanelt D,Figueroa FL.2012.Physiological and photomorphogenic effects of light on marine macrophytes//Wiencke C,Bischof K,eds.Seaweed Biology:Novel Insights into Ecophysiology,Ecology and Utilization.Berlin:Springer:3-23.
Hangil C,Hyunil KH,Kang PJ,et al.2008.Physiological differences in the growth of Sargassum horneri between the germling and adult stages.Journal of Applied Phycology,20:729-735.
Hellebust JA,Craigie JS.1978.Handbook of Phycological Methods:Physiological and Biochemical Methods.Cambridge:Cambridge University Press.
Henley WJ.1993.Measurement and interpretation of photosynthetic light-response curves in algae in the context of photoinhibition and diel changes.Journal of Phycology,29:729-739.
Hwang RL,Tsai CC,Lee TM.2010.Assessment of temperature and nutrient limitation on seasonal dynamics among species of Sargassum from a coral reef in southern Taiwan.Journal of Phycology,40:463-473.
Lobban CS,Chapman DJ,Kremer BP.1988.Experimental Phycology:A Laboratory Manual.Cambridge:Cambridge University Press.
Luo MB,Liu F,Xu ZL.2012.Growth and nutrient uptake capacity of two co-occurring species,Ulva prolifera and Ulva linza.Aquatic Botany,100:18-24.
Provasoli L.1968.Media and prospects for the cultivation of marine algae//Watanabe A,Hattori A,eds.Cultures and Collections of Algae.Hakone:Japanese Society of Plant Physiology Press:63-75.
Qi L,Hu C,Wang M,et al.2017.Floating algae blooms in the East China Sea.Geophysical Research Letters,44:501-509.
Rivera EC,Díaz MF,Hawkins A.2015.A fish kill coincident with dense Sargassum accumulation in a tropical bay.Bulletin of Marine Science,91:455-456.
Schell JM,Goodwin DS,Siuda ANS.2015.Recent Sargassum inundation events in the Caribbean:Shipboard observations reveal dominance of a previously rare form.Oceanography,28:8-10.
Sfriso A,Facca C.2013.Annual growth and environmental relationships of the invasive species Sargassum muticum,and Undaria pinnatifida,in the lagoon of Venice.Estuarine,Coastal&Shelf Science,129:162-172.
Smetacek V,Zingone A.2013.Green and golden seaweed tides on the rise.Nature,504:84-88.
Uchida T.2010.The life circle of Sargassum horneri(Phaeophyta)in laboratory culture.Journal of Phycology,29:231-235.
Umezaki I.1984.Ecological studies of Sargassum horneri(Turner)C.Agardh in Obama Bay,Japan Sea.Bulletin of the Japanese Society of Scientific Fisheries,50:1193-1200.
Wirtz KW,Pahlow M.2010.Dynamic chlorophyll and nitrogen:Carbon regulation in algae optimizes instantaneous growth rate.Marine Ecology Progress Series,402:81-96.
Xu Z,Gao G,Xu J,et al.2017.Physiological response of a golden tide alga(Sargassum muticum)to interaction of ocean acidification and phosphorus enrichment.Biogeosciences,14:1-36.
Xu Z,Gao K.2012.NH4+enrichment and UV radiation interact to affect the photosynthesis and nitrogen uptake of Gracilaria lemaneiformis(Rhodophyta).Marine Pollution Bulletin,64:99-105.
Xu Z,Wu H,Zhan D,et al.2014.Combined effects of light intensity and NH4+enrichment on growth,pigmentation,and photosynthetic performance of Ulva prolifera(Chlorophyta).Chinese Journal of Oceanology and Limnology,32:1016-1023.
Xu Z,Zou D,Gao K,et al.2010.Effects of elevated CO2and phosphorus supply on growth,photosynthesis and nutrient uptake in the marine macroalga Gracilaria lemaneiformis(Rhodophyta).Botanica Marina,53:123-129.
Yoshida G,Murase N,Arai S,et al.2004.Ecotypic differentiation in maturation seasonality among Sargassum horneri(Fucales,Phaeophyta)populations in Hiroshima Bay,Seto Inland Sea,Japan.Phycologia,43:703-710.
Zou D,Gao K.2009.Effects of elevated CO2on the red seaweed Gracilaria leman.Phycologia,48:510-517.
曹春晖,刘文岭,施定基,等.2010.不同氮磷浓度对米氏凯伦藻生长的影响.天津科技大学学报,25(2):22-25.
陈自强,孙庆海,陈全震,等.2013.酸胁迫对铜藻早期生长阶段的影响.生态毒理学报,8(6):864-870.
方婷轩,马增岭.2016.温度变化对铜绿微囊藻(Microcystis aeruginosa)生长和光合作用的影响.生态科学,35(6):14-23.
李慧,丁刚,辛美丽,等.2017.不同氮、磷浓度及配比对铜藻(Sargassum horneri)幼苗生长的影响.海洋与湖沼,48(2):368-372.
李科,潘耀茹,吴嘉平,等.2017.不同光照周期下LED光源对铜藻生长的影响.浙江农业学报,29(4):631-636.
李磊,关万春,陈少波,等.2016.水体N/P对塔玛亚历山大藻响应紫外辐射的影响.生态学杂志,35(2):395-400.
李信书,伏光辉,陈百尧,等.2012.氮、磷加富对条斑紫菜生长及生化组成的影响.水产科学,31(9):544-548.
司冉冉,关万春,蔡景波,等.2017.氮源对塔玛亚历山大藻生长和毒性的影响.生态学杂志,36(10):2880-2885.
孙建璋,庄定根,孙庆海,等.2009.铜藻人工栽培的初步研究.南方水产科学,5(6):41-46.
王丽梅,李世国,柴雨,等.2011.鼠尾藻幼苗的室内培养及有性生殖同步化.水产学报,35(3):395-404.
王巧晗.2008.环境因子节律性变动对潮间带大型海藻孢子萌发、早期发育和生长的影响及其生理生态学机制(硕士学位论文).青岛:中国海洋大学.
徐智广,吴海一,孙福新,等.2016.不同氮源加富对海带生长、光合固碳和氮吸收特性的影响.水产学报,40(4):577-584.
徐智广,邹定辉,高坤山,等.2011.不同温度、光照强度和硝氮浓度下龙须菜对无机磷吸收的影响.水产学报,35(7):1023-1029.
徐智广,邹定辉,张鑫,等.2008.CO2和硝氮加富对龙须菜(Gracilaria lemaneiformis)生长、生化组分和营养盐吸收的影响.生态学报,28(8):3752-3759.
杨安强,史定刚,陈少波,等.2017.高CO2条件下温度和光照对铜藻吸收N、P的短期影响.浙江农业科学,58(2):303-307.
张鹏,王铁杆,谢起浪,等.2014.培养条件对铜藻幼苗生长发育的影响.上海海洋大学学报,23(2):200-207.
张玉荣,刘峰,单体锋,等.2009.利用叶绿素荧光技术揭示人工培育的铜藻幼苗对胁迫温度、光照和盐度的反应.南方水产科学,5(2):1-9.
周百成,郑舜琴,曾呈奎.1974.几种绿藻、褐藻和红藻的吸收光谱的比较研究.植物学报,16(2):146-155.
Davison JR,Jordan TL,Fegley JC,et al.2007.Response of Laminaria saccharina(Phaeophyta)growth and photosynthesis to simultaneous ultraviolet radiation and nitrogen limitation.Journal of Phycology,43:636-646.
Falkowski PG,Raven JA.2007.Aquatic Photosynthesis.Princeton:Princeton University Press.
Geun CC,Geun KH,Hyun SC.2003.Transplantation of young fronds of Sargassum horneri for construction of seaweed beds.Journal of Geotechnical&Geoenvironmental Engineering,36:469-473.
Giordano M,Beardall J,Raven JA.2005.CO2concentrating mechanisms in algae:Mechanisms,environmental modulation,and evolution.Annual Review of Plant Biology,56:99-131.
Hanelt D,Figueroa FL.2012.Physiological and photomorphogenic effects of light on marine macrophytes//Wiencke C,Bischof K,eds.Seaweed Biology:Novel Insights into Ecophysiology,Ecology and Utilization.Berlin:Springer:3-23.
Hangil C,Hyunil KH,Kang PJ,et al.2008.Physiological differences in the growth of Sargassum horneri between the germling and adult stages.Journal of Applied Phycology,20:729-735.
Hellebust JA,Craigie JS.1978.Handbook of Phycological Methods:Physiological and Biochemical Methods.Cambridge:Cambridge University Press.
Henley WJ.1993.Measurement and interpretation of photosynthetic light-response curves in algae in the context of photoinhibition and diel changes.Journal of Phycology,29:729-739.
Hwang RL,Tsai CC,Lee TM.2010.Assessment of temperature and nutrient limitation on seasonal dynamics among species of Sargassum from a coral reef in southern Taiwan.Journal of Phycology,40:463-473.
Lobban CS,Chapman DJ,Kremer BP.1988.Experimental Phycology:A Laboratory Manual.Cambridge:Cambridge University Press.
Luo MB,Liu F,Xu ZL.2012.Growth and nutrient uptake capacity of two co-occurring species,Ulva prolifera and Ulva linza.Aquatic Botany,100:18-24.
Provasoli L.1968.Media and prospects for the cultivation of marine algae//Watanabe A,Hattori A,eds.Cultures and Collections of Algae.Hakone:Japanese Society of Plant Physiology Press:63-75.
Qi L,Hu C,Wang M,et al.2017.Floating algae blooms in the East China Sea.Geophysical Research Letters,44:501-509.
Rivera EC,Díaz MF,Hawkins A.2015.A fish kill coincident with dense Sargassum accumulation in a tropical bay.Bulletin of Marine Science,91:455-456.
Schell JM,Goodwin DS,Siuda ANS.2015.Recent Sargassum inundation events in the Caribbean:Shipboard observations reveal dominance of a previously rare form.Oceanography,28:8-10.
Sfriso A,Facca C.2013.Annual growth and environmental relationships of the invasive species Sargassum muticum,and Undaria pinnatifida,in the lagoon of Venice.Estuarine,Coastal&Shelf Science,129:162-172.
Smetacek V,Zingone A.2013.Green and golden seaweed tides on the rise.Nature,504:84-88.
Uchida T.2010.The life circle of Sargassum horneri(Phaeophyta)in laboratory culture.Journal of Phycology,29:231-235.
Umezaki I.1984.Ecological studies of Sargassum horneri(Turner)C.Agardh in Obama Bay,Japan Sea.Bulletin of the Japanese Society of Scientific Fisheries,50:1193-1200.
Wirtz KW,Pahlow M.2010.Dynamic chlorophyll and nitrogen:Carbon regulation in algae optimizes instantaneous growth rate.Marine Ecology Progress Series,402:81-96.
Xu Z,Gao G,Xu J,et al.2017.Physiological response of a golden tide alga(Sargassum muticum)to interaction of ocean acidification and phosphorus enrichment.Biogeosciences,14:1-36.
Xu Z,Gao K.2012.NH4+enrichment and UV radiation interact to affect the photosynthesis and nitrogen uptake of Gracilaria lemaneiformis(Rhodophyta).Marine Pollution Bulletin,64:99-105.
Xu Z,Wu H,Zhan D,et al.2014.Combined effects of light intensity and NH4+enrichment on growth,pigmentation,and photosynthetic performance of Ulva prolifera(Chlorophyta).Chinese Journal of Oceanology and Limnology,32:1016-1023.
Xu Z,Zou D,Gao K,et al.2010.Effects of elevated CO2and phosphorus supply on growth,photosynthesis and nutrient uptake in the marine macroalga Gracilaria lemaneiformis(Rhodophyta).Botanica Marina,53:123-129.
Yoshida G,Murase N,Arai S,et al.2004.Ecotypic differentiation in maturation seasonality among Sargassum horneri(Fucales,Phaeophyta)populations in Hiroshima Bay,Seto Inland Sea,Japan.Phycologia,43:703-710.
Zou D,Gao K.2009.Effects of elevated CO2on the red seaweed Gracilaria leman.Phycologia,48:510-517.
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