Subsurface Chlorophyll Maximum in Ocean：Its Characteristics and Influencing Factors
Received date: 2011-09-21
Revised date: 2012-02-17
Online published: 2012-05-10
The vertical distribution of marine phytoplankton is complicated. However, a bell-shape vertical profile of phytoplankton in mass concentration is frequently observed in coastal seas and open oceans. The corresponding Subsurface Chlorophyll Maximum Layers (SCML) have caused a very high scientific interest and have been studied well. SCM can be characterized by three parameters, i.e., depth, thickness and intensity of the SCML. However, challenges still exist on how to best characterize the three parameters. They are determined by marine hydrology, nutrient concentrations and phytoplankton species, etc. A few studies have been conducted to explore variability of three parameters among different oceans. In general, compared with coastal seas，seasonal variation of the three parameters in open oceans is insignificant. It is also indicated that the three parameters are deeper，thicker and smaller in open oceans than coastal seas. In oligotrophic ocean, the depth of SCML was mainly a function of light and turbulent diffusion. Its intensity was determined by light, turbulent diffusion and biogenic processes associated with zooplankton grazing. However, the quantitative relationship between these biogenic processes and the three parameters are not well established. For example, how the sinking rate of phytoplankton, physiological adaptation capabilities, motility, trophic interactions affect them are poorly understood, and further investigations are needed. This review also finds that the SCML study in coastal seas is rare, which could be due to the lack of a long-term and high-timeresolution observation in those areas. By use of satallite data, in situ data and physical-biological models, it might be useful for further understanding SCM furtherly.
Gong Xiang，Shi Jie，Gao Huiwang . Subsurface Chlorophyll Maximum in Ocean：Its Characteristics and Influencing Factors[J]. Advances in Earth Science, 2012 , 27(5) : 539 -548 . DOI: 10.11867/j.issn.1001-8166.2012.05.0539
[1]Varela R A, Cruzado A, Tintore J, et al. Modelling the deep-chlorophyll maximum: A coupled physical-biological approach[J]. Journal of Marine Research,1992, 50(3): 441-463.
[2]Cullen J J. The deep chlorophyll maximum: Comparing vertical profiles of chlorophyll a[J]. Canadian Journal of Fisheries Aquatic Sciences,1982, 39(5): 791-803.[3]Geek S, Legovi T. Nutrients and grazing in modeling the deep chlorophyll maximum[J]. Ecological Modelling,2001, 138(1/3): 143-152.
[4]Fennel K, Boss E. Subsurface maxima of phytoplankton and chlorophyll: Steady-state solutions from a simple model[J]. Limnology and Oceanogoraphy,2003, 48(4): 1 521-1 534.
[5]Steele J H. Environmental control of photosynthesis in the sea[J]. Limnology and Oceanography,1962, 7(2): 137-150.
[6]Pak H, Kiefer D A, Kitchen J C. Meridional variations in the concentration of chlorophyll and microparticles in the North Pacific Ocean[J]. Deep Sea Research Part A:Oceanography Research Papers,1988, 35(7): 1 151-1 171.
[7]Kitchen J C, Zaneveld J. On the noncorrelation of the vertical structure of light scattering and chlorophyll α in case I waters[J]. Journal of Geophysical Research,1990, 95(C11): 20 237-20 246.
[8]Hodges B A, Rudnick D L. Simple models of steady deep maxima in chlorophyll and biomass[J]. Deep Sea Research Part I:Oceanographic Research Papers,2004, 51(8): 999-1 015.
[9]Hanson C E, Pesant S, Waite A M, et al. Assessing the magnitude and significance of deep chlorophyll maxima of the coastal eastern Indian Ocean[J]. Deep Sea Research Part II:Topical Studies in Oceanography,2007, 54(8/10): 884-901.
[10]Siswanto E, Ishizaka J, Yokouchi K. Estimating chlorophyll-a vertical profiles from satellite data and the implication for primary production in the kuroshio front of the East China Sea[J]. Journal of Oceanography,2005, 61(3): 575-589.
[11]Takahashi M, Hori T. Abundance of picophytoplankton in the subsurface chlorophyll maximum layer in subtropical and tropical waters[J]. Marine Biology,1984, 79(2): 177-186.
[12]Hanson C E, Waite A M, Thompson P A, et al. Phytoplankton community structure and nitrogen nutrition in Leeuwin Current and coastal waters off the Gascoyne region of Western Australia[J]. Deep Sea Research Part II: Topical Studies in Oceanography,2007, 54(8/10): 902-924.
[13]Hopkinson B M, Barbeau K A. Interactive influences of iron and light limitation on phytoplankton at subsurface chlorophyll maxima in the eastern North Pacific[J]. Limnology and Oceanography,2008, 53(4): 1 303-1 318.
[14]Huisman J, Thi N, Karl D M, et al. Reduced mixing generates oscillations and chaos in the oceanic deep chlorophyll maximum[J]. Naure,2006, 439(7 074): 322-325.
[15]Mellard J P, Yoshiyama K, Litchman E, et al. The vertical distribution of phytoplankton in stratified water columns[J]. Journal of Theoretical Biology,2010,269(1):16-30.
[16]Ni Xiaobo, Huang Daji. Subsurface chlorophyll maximum：Its temporal-spatial distribution and formation mechanism in the ocean[J]. Marine Science,2006, 30(5): 58-64.[倪晓波，黄大吉. 海洋次表层叶绿素最大值的分布和形成机制研究[J]. 海洋科学，2006, 30(5): 58-64.]
[17]Beckmann A, Hense I. Beneath the surface: Characteristics of oceanic ecosystems under weak mixing conditions—A theoretical investigation[J]. Progress in Oceanography,2007, 75(4): 771-796.
[18]Jamart B M, Winter D F, Banse K. Sensitivity analysis of a mathematical model of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern US coast[J]. Journal of Plankton Research,1979, 1(3): 267-290.
[19]Holm-Hansen O, Hewes C D. Deep chlorophyll-a maxima (DCM) in Antarctic waters[J]. Polar Biology,2004, 27(11): 699-710.
[20]Letelier R M, Karl D M, Abbott M R, et al. Light driven seasonal patterns of chlorophyll and nitrate in the lower euphotic zone of the North Pacific Subtropical Gyre[J]. Limnology and Oceanography,2004, 49(2): 508-519.
[21]Pérez V, Fernández E, Maraén E, et al. Vertical distribution of phytoplankton biomass, production and growth in the Atlantic subtropical gyres[J]. Deep Sea Research Part I: Oceanographic Research Papers,2006, 53(10): 1 616-1 634.
[22]Lewis M R, Cullen J J, Platt T. Phytoplankton and thermal structure in the upper ocean: Consequences of nonuniformity in chlorophyll profile[J]. Journal of Geophysical Research,1983, 88(C4): 2 565-2 570.
[23]Kononen K, Huttunen M, Hllfors S, et al. Development of a deep chlorophyll maximum of Heterocapsa triquetra Ehrenb. at the entrance to the Gulf of Finland[J]. Limnology and Oceanography,2003,48(2): 594-607.
[24]Pedrós-Alió C, Calderón-Paz J I, Guixa-Boixereu N, et al. Bacterioplankton and phytoplankton biomass and production during summer stratification in the northwestern Mediterranean Sea[J]. Deep Sea Research Part I: Oceanographic Research Papers,1999, 46(6): 985-1 019.
[25]Varela R A, Cruzado A, Tintoré J. A simulation analysis of various biological and physical factors influencing the deep-chlorophyll maximum structure in oligotrophic areas[J]. Journal of Marine Systems,1994, 5(2): 143-157.
[26]Klausmeier C A, Litchman E. Algal games: The vertical distribution of phytoplankton in poorly mixed water columns[J]. Limnology and Oceanography,2001,46(8): 1 998-2 007.
[27]Murty V, Gupta G, Sarma V V, et al. Effect of vertical stability and circulation on the depth of the chlorophyll maximum in the Bay of Bengal during May-June, 1996[J]. Deep Sea Research Part I: Oceanographic Research Papers,2000, 47(5): 859-873.
[28]Gong G C, Shiah F K, Liu K K, et al. Spatial and temporal variation of chlorophyll a, primary productivity and chemical hydrography in the southern East China Sea[J]. Continental Shelf Research,2000, 20(4/5): 411-436.
[29]Fu Mingzhu, Wang Zongling, Sun Ping, et al. Spatial distribution characteristics and the environmental regulation mechanisms of phytoplankton chlorophyll a in Southern Yellow Sea during summer 2006[J]. Acta Ecologica Sinica,2009, 29(10): 5 366-5 375.[傅明珠，王宗灵，孙萍，等. 2006 年夏季南黄海浮游植物叶绿素 a 分布特征及其环境调控机制[J]. 生态学报,2009, 29(10): 5 366-5 375.]
[30]Martin J, Tremblay J, Gagnon J, et al. Prevalence, structure and properties of subsurface chlorophyll maxima in Canadian Arctic waters[J]. Marine Ecology Progress Series,2010, 412: 69-84.
[31]Li Baohua. Variation of chlorophyll a contens at the wharf of the Great Wall Station, Antactica[J]. Chinese Journal of Polar Research,2004, 16(4):332-337.[李宝华. 南极长城站码头及临近海域夏季叶绿素 a 含量及变化[J]. 极地研究,2004, 16(4): 332-337.]
[32]Hense I, Beckmann A. Revisiting subsurface chlorophyll and phytoplankton distributions[J]. Deep Sea Research Part I: Oceanographic Research Papers,2008, 55(9): 1 193-1 199.
[33]Radenac M H, Rodier M. Nitrate and chlorophyll distributions in relation to thermohaline and current structures in the western tropical Pacific during 1985-1989[J]. Deep Sea Research Part II: Topical Studies in Oceanography,1996, 43(4/6): 725-752.
[34]Teira E, Mouri O B, Maran E, et al. Variability of chlorophyll and primary production in the eastern North Atlantic Subtropical Gyre: Potential factors affecting phytoplankton activity[J]. Deep-Sea Research Part I:Oceanographic Research Papers,2005, 52(4): 569-588.
[35]Lu Saiying. Chlorophyll a maximum distribution and variabilityes in the northern East China Sea[J]. Acta Oceanologica Sinica,1998, 20(3): 64-75.[陆赛英. 东海北部叶绿素a极大值的分布规律[J]. 海洋学报,1998, 20(3): 64-75.]
[36]Ruiz J, García C M, Rodriguez J. Vertical patterns of phytoplankton size distribution in the Cantabric and Balearic Seas[J]. Journal of Marine Systems,1996, 9(3/4): 269-282.
[37]Mantyla A W, Bograd S J, Venrick E L. Patterns and controls of chlorophyll-a and primary productivity cycles in the Southern California Bight[J]. Journal of Marine Systems,2008, 73(1/2): 48-60.
[38]Chen Xingqun, Chen Qihuan, Zhuang Liangzhong. Relationship between distribution of chlorophyll a, photosynthesis and main environmental factors in the central South China Sea[J]. Acta Oceanologica Sinica,1989, 11(3): 349-355.[陈兴群，陈其焕，庄亮钟. 南海中部叶绿素分布和光合作用及其与环境因子的关系[J]. 海洋学报,1989, 11(3): 349-355.]
[39]Anderson G C. Subsurface chlorophyll maximum in the northeast Pacific Ocean[J]. Limnology and Oceanography,1969, 14(3): 386-391.
[40]Sharples J, Moore C M, Rippeth T P, et al. Phytoplankton distribution and survival in the thermocline[J]. Limnology and Oceanography,2001, 46(3): 486-496.
[41]Gabric A J, Parslow J. Effect of physical factors on the vertical distribution of phytoplankton in eutrophic coastal waters[J]. Australian Journal of Marine & Fishwater Research,1989, 40(5): 559-569.
[42]Eppley R W. Temperature and phytoplankton growth in the sea[J]. Fishery Bulletin,1972, 4(70): 1 063-1 085.
[43]Primavera K, San Diego-Mcglone M L, Villanoy C. Development of a simple biological model of vertical phytoplankton distribution[J]. Science Diliman,2007, 18(1): 43-52.
[44]El Venrick J, Mantyla A W. Deep maxima of photosynthetic chlorophyll in the Pacific Ocean[J]. Fishery Bulletin,1973,71:41-52.
[45]Lu Z, Gan J, Dai M, et al. The influence of coastal upwelling and a river plume on the subsurface chlorophyll maximum over the shelf of the northeastern South China Sea[J]. Journal of Marine Systems,2010,82(1/2): 35-46.
[46]Kirk J. Light and Photosynthesis in Aquatic Ecosystems[M]. Cambridge:Cambridge University Press, 1994.
[47]Lund-Hansen L C. Diffuse attenuation coefficients Kd (PAR) at the estuarine North Sea-Baltic Sea transition: Time-series, partitioning, absorption, and scattering[J]. Estuarine, Coastal and Shelf Science,2004, 61(2): 251-259.
[48]Xia Jie, Gao Huiwang. Simulation on seasonal cycle vertical structure of plankton ecosystem in eastern area of South Yellow Sea[J]. Journal of Safety and Environment,2006, 6(4): 59-65.[夏洁，高会旺. 南黄海东部海域浮游生态系统要素垂直分布的模拟研究[J]. 安全与环境学报,2006, 6(4): 59-65.]
[49]Zhang Shuwen. Effect mechanisms on chlorophyll a verticaldistribution in the summer within Yellow Sea Cold Water Mass[J]. Oceanologia et Limnologa Sinica,2003, 34(2): 179-186.[张书文. 黄海冷水团夏季叶绿素垂向分布结构的影响机制[J]. 海洋与湖沼,2003, 34(2): 179-186.]
[50]Ryabov A B, Rudolf L, Blasius B. Vertical distribution and composition of phytoplankton under the influence of an upper mixed layer[J]. Journal of Theoretical Biology, 2010, 263(1): 120-133.
[51]Carr M E, Friedrichs M A M, Schmeltz M, et al. A comparison of global estimates of marine primary production from ocean color[J]. Deep Sea Research Part II: Topical Studies in Oceanography,2006, 53(5/7): 741-770.
[52]Margalef R. Life-forms of phytoplankton as survival alternatives in unstable environments[J]. Month,1978,1(4):493-509.
[53]Kaiser W, Postel L. Importance of the vertical nutrient flux for biological production in the equatorial undercurrent region at 30°W[J]. Marine Biology,1979, 55(1): 23-27.
[54]Sarmiento J L, Hughes T, Stouffer R J, et al. Simulated response of the ocean carbon cycle to anthropogenic climate warming[J]. Nature,1998, 393(6 682): 245-249.
[55]Rosenzweig C, Parry M L. Potential impact of climate change on world food supply[J]. Nature,1994, 367(6 459): 133-138.
[56]Huisman J, Arrayás M, Ebert U, et al. How do sinking phytoplankton species manage to persist?[J]. The American Naturalist,2002, 159(3): 245-254.
[57]Bienfang P K. Phytoplankton Dynamics in Oceanic Waters off Ke-ahole Point, Hawaii[M].Oakland: University of California,1980.
[58]Cullen J J, Horrigan S G. Effects of nitrate on the diurnal vertical migration, carbon to nitrogen ratio, and the photosynthetic capacity of the dinoflagellate Gymnodinium splendens[J]. Marine Biology,1981, 62(2): 81-89.
[59]Yoshiyama K, Mellard J P, Litchman E, et al. Phytoplankton competition for nutrients and light in a stratified water column[J]. American Naturalist,2009, 174(2): 190-203.
[60]Bienfang P K. Phytoplankton sinking rates in oligotrophic waters off Hawaii, USA[J]. Marine Biology,1980, 61(1): 69-77.
[61]Jamart B M, Winter D F, Banse K, et al. A theoretical study of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern US coast[J]. Deep Sea Research,1977, 24(8): 753-773.
[62]Taylor A H. Characteristic properties of models for the vertical distribution of phytoplankton under stratification[J]. Ecological Modelling,1988, 40(3/4): 175-199.
[63]Venrick E L. Phytoplankton seasonality in the central North Pacific: The endless summer reconsidered[J]. Limnology and Oceanography,1993, 38(6):1 135-1 149.
[64]Sharples J, Tett P. Modelling the effect of physical variability on the midwater chlorophyll maximum[J]. Journal of Marine Research,1994, 52(2): 219-238.
[65]Sharples J. Investigating the seasonal vertical structure of phytoplankton in shelf seas[J]. Marine Models,1999, 1(1/4): 3-38.
[66]Zhang Shuwen, Xia Changshui, Yuan Yeli. Effect of the upwelling current on the vertical chlorophyll distribution in the Yellow Sea cold water mass[J].Advances in Marine Science,2002, 20(3): 9-14.[张书文，夏长水，袁业力. 黄海冷水团上升流对叶绿素垂向分布的影响[J]. 海洋科学进展,2002, 20(3): 9-14.]
[67]Lindholm T. Ecological role of depth maxima of phytoplankton[J]. Archiv fu Hydrobiologie Beiheft Ergebnisseder Limnologie,1992, 35: 33-45.
[68]Richardson A J, Silulwane N F, Mitchell-Innes B A, et al. A dynamic quantitative approach for predicting the shape of phytoplankton profiles in the ocean[J]. Progress in Oceanography,2003, 59(2/3): 301-319.
[69]Fei Zunle, Zhu Mingyuan. The study on Subsurface Chlorophyll-a Maximum (SCM) and its relationship with light and nutrients[C]∥Collection of Papers on Kuroshio Study and Research (I).Beijing:Ocean Press,1990:67-74.[费尊乐，朱明远. 次表层叶绿素a极大值（SCM）分布与其光—营养盐环境相关关系的研究[C]∥黑湖调查研究论文选.北京: 海洋出版社, 1990:67-74.]
[70]Wang Xiaoyu, Zhu Biying. Discussion of the subsurface chlorophyll maximum forming in the continental and Kuroshio area of the East China Sea in the summer[C]∥Collection of Papers on Kuroshio Survey and Research. Beijing: Ocean Press. Selected Works on the Kuroshio Study(III),1991:297-304.[王小羽，朱碧英. 东海陆架海及黑潮区夏季次表层叶绿素最大值成因探讨[C]∥黑湖调查研究论文选.北京: 海洋出版社, 1991:297-304.]
[71]Zhang Fan, Ruan Wuqi, Huang Bangqin. A study of chlorophyll a maximum in the southern part of the Taiwan Strait[J]. Journal of Xiamen University(Natural Science),2000, 39(5): 664-668.[张钒，阮五崎，黄邦钦. 台湾海峡南部叶绿素a最大值的研究[J]. 厦门大学学报: 自然科学版,2000, 39(5): 664-668.]
[72]Wu Rongjun, Lü Ruihua, Zhu Mingyuan, et al. Impacts of sea water mixing and stratification on the vertical profile of chlorophyll a[J].Ecology and Environment,2004, 13(4): 515-519.[吴荣军，吕瑞华，朱明远，等. 海水混合和层化对叶绿素 a 垂直分布的影响[J]. 生态环境,2004, 13(4): 515-519.]
[73]Hu Yi, Chen Jian. Vertical distribution of chlorophyll a fluorescence and its response to temperature and salinity near Taiwan Bank in summer[J]. Journal of Tropical Oceanography,2008, 27(2): 18-24.[胡毅，陈坚. 夏季台湾浅滩周边海域叶绿素a 荧光垂直分布对温盐的响应分析[J]. 热带海洋学报,2008, 27(2): 18-24.]
[74]Chen D, Horrigan S G, Wang D P. The late summer vertical nutrient mixing in Long Island Sound[J].Journal of Marine Research, 1988, 46(4): 753-770.
[75]Holloway G, Denman K. Influence of internal waves on primary production[J].Journal of Plankton Research, 1989, 11(2): 409.
[76]Shigesada N, Okubo A. Analysis of the self-shading effect on algal vertical distribution in natural waters[J]. Journal of Mathematical Biology,1981, 12(3): 311-326.
/
| 〈 |
|
〉 |
甘公网安备62010202000687
