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1 45 3 Vol.45, No OCEANOLOGIA ET LIMNOLOGIA SINICA May, 2014 * 1, 2 11 ( ; ), Phyto-PAM,,, :, 0.69, ,, 1000μmol/(s m 2 ), ; 1500μmol/(s m 2 ),,,,,,,, ; ; ; ; Q178.1 doi: /hyhz ,, 45%(Field et al, 1998),, (Lalli et al, 1993),, : (HNLC),,, (Kolber et al, 1994; Boyd et al, 2000); (McMinn et al, 2004); (, 2012), ;,, I(PSI) b 6 f, ( ), (Strzepek et al, 2004) * (973), 2011CB ;, XDA ;, , [email protected] :,,, [email protected] : , :

2 3 : 469 : ; ; (Papageorgiou et al, 2007),,, (PS ) (Schreiber et al, 1995) (Bolhar- Nordenkampf et al, 1989),, F v /F m (Häder, 1998), F v /F m, 0.65(Kolber et al, 1988),, F v /F m (Babin et al, 1996),,,, (Strzepek et al, 2004; Jakob et al, 2005; Napoléon et al, 2012),,,, (, 1992), (, 2005),, 12 1,, 2 (, 2002;, 2005;, 2006;, 2007),,,,,,, , 10 ( 1), ULM-500 (Walz, Germany) Niskin, 50mL ; 1000mL Logul s ; 500mL GF/F, ( 20 C) a CTD(SBE25-02) 1 Fig.1 Sampling stations in the Jiaozhou Bay 1.2 Phyto-PAM(Walz, Effeltrich, Germany), Phytowin2.13(Walz) (Skeletonema costatum),, 0.2μm,, 2mL, 10min, (Measuring

3 Radiation) F 0, (Saturation Pulse) F m, PSII F v /F m =(F m F 0 )/F m, PAR 1μmol/(s m 2 ), 20s (Actinic Light) 100μmol/(s m 2 ) 2000μmol/ (s m 2 ), RLC Platt (1980): α PAR/ Pm ( ) β PAR/ Pm P = Pm 1 e e P, (retr); P m (retr max ); α P-I, ; β ; P m α I k Φ PS (qp) (NPQ)(Bilger et al, 1990): Φ PS =(F m F t )/F m qp=(f m F t )/(F m F 0 ) NPQ=(F m F m )/F m F m, F t mL (Hydro- Bios, Germany) 24h Olympus IX51, 90% 12 24h, Turner-Designs- Model 10 a (QuAAtro, Germany) : (NO 3 -N) (NO 2 -N)(NH + 4 -N) (PO 3 4 -P) (SiO 2 3 -Si) a(chl a),, 1 Tab.1 ( C) 表 1 胶 州 湾 调 查 站 位 温 盐 光 照 营 养 盐 叶 绿 素 a Measurements of water temperature, salinity, irradiance, nutrients, and chlorophyll a in the bay stations (μmol/(s m 2 )) (μmol/l) (μmol/l) (μmol/l) (μmol/l) (μmol/l) A A B C C C D D D D a (mg/m 3 ) 2.2,, 12,,,, ( 2), A2 B1 C1 C3, cell/m 3, C1, cell/m 3,, (Skeletonema costatum) A2 B1 C1 C3 D2 D3, 50% ; (Asteronella kariana) (Nitzschia paradoxa Gmelin), C2 D1 D4,,, A1 ( 2) 12 a, PSII F v /F m PSII F v /F m, A2 0.8, C1 0.59, 0.69 ( 5)

4 3 : 471 表 2 12 月 胶 州 湾 表 层 浮 游 植 物 群 落 结 构 组 成 Tab.2 Species composition of phytoplankton in the Jiaozhou Bay Skeletonema costatum (Grev.) Cleve Schroederella delicatula Paralia sulcata (Ehr.) Cleve Rhizosolenia delicatula Cleve Rhizosolenia setigera Brightwell Biddulpha sinensis Greville Ditylum brightwelli (West) Grunow Thalasiosira rotula Meunier Thalasiosira sp. Melosira moniliformis Chaetoceros densus Cleve C. sp. Nitzschia paradoxa Gmelin N. lorenziana Grunow N. closterium W. Smith N. sp. Pseudonitzschia pungens Coscinodiscus granii Coscinodiscus sp. Cyclotella sp. Asteronella kariana Grunow Synedra sp. Diploneis bombus Ehrenberg Amphiprora alata (Ehr.) Kuetzing Pinnularia sp. Pleurosigma sp. Ceratium fusus (Ehrenberg) Dujardin Prorocentrum micans Ehrenb Euglena sp. Dictyocha fibula Ehrenberg 3 Fig.3 Horizontal distribution of phytoplankton 10 4 cell/m 3 4 a Fig.4 Horizontal distribution of concentration of chlorophyll a mg/m 3 2 Fig.2 Phytoplankton community structure, distribution, and abundance RLC 6,,,, retr, 1000μmol/(s m 2 ), retr (retr max ), 1500μmol/(s m 2 ), 3, (α) , A2, 0.33; retr max

5 , C3, 211.4; (I k ) μmol/(s m 2 ), A2, C Φ PS 8, Φ PS, 0μmol/(s m 2 ) 2000μmol/(s m 2 ), Φ PS Fig.5 5 The maximum quantum yield of phytoplankton F v /F m 0, PS PS, NPQ 9, Fig.6 6 The rapid light curves of phytoplankton

6 3 : 473 表 3 浮 游 植 物 快 速 光 曲 线 系 数 Tab.3 The photosynthetic parameters obtained from the rapid light curves (α) retr max (μmol/(s m 2 )) I k (μmol/(s m 2 )) A A B C C C D D D D NPQ,, qp qp, qp, PSII, (Kooten et al, 1990) 0 qp<1, C3 D4 1 ( ), qp, NPQ, Fig.7 7 The actual photochemical efficiency of PSII in the light

7 Fig.8 Non-photochemical quenching of phytoplankton, qp,,, PS,, ( 10) 3,, F v /F m,, F v /F m (Cermeno et al, 2005),

8 3 : Fig.9 The photochemical quenching of phytoplankton, (Rosenbergl, 1993; Tamigneaux et al, 1999), F v /F m,, /, (package effect)(kiørboe, 1993; Raven, 1998; Sosik, 2002) F v /F m F v /F m 10 20m, PS (Neale et al, 1991; Bergmann et al, 2002; Bouchard et al, 2005),, PSII, (Bracher et al, 2000; Kaiblinger et al, 2007), 20 40m F v /F m, 0.65, F v /F m, F v /F m 20% 30%, 40m F v /F m, 400m 0.1(McMinn et al, 2004) Metz, F v /F m (Vaillancourt et al, 2003), DIN/DIP DSi/DIP F v /F m (Goto et al, 2008), ATP, PSII (Lippemeier et al, 2001), F v /F m

9 476 45, (F v /F m 0.65) (McMinn et al, 2004),, F v /F m 0.61, (Suzuki et al, 2002) PS F v /F m, , 0.69, DIN 37.4μmol/L( ), DIP 0.97μmol/L, DSi 9.71μmol/L, DIN/DIP 38:1, DSi/DIP 10:1, DSi/DIN 0.26, Redfield, 2000 DIN, DIP DSi, (, 2011), (, 2002b),, ( 700μmol/(s m 2 )), 12 RLC,, μmol/(s m 2 ), retr (retr max ), 1500μmol/(s m 2 ),, PSII (White et al, 1999) 12, (α) , retr max ; (I k ) μmol/(s m 2 ) α,, (Dubinsky et al, 2009), retr CO 2 (Figueroa et al, 2003), I k,, α 0.2, 1500μmol/(s m 2 ), (, 2012), F v /F m 0.125, α 0.022, 385.1μmol/(s m 2 ),, (McMinn et al, 2004), (α) ,, I k retr max, PS (, 1999),, NPQ (Schreiber, 2004) 20m,,, NPQ ( 8), 0μmol/(s m 2 ) 2000μmol/(s m 2 ), NPQ,,, (Olaizola et al, 1994; Kashino et al, 2002), (Dugdale et al, 1998),, /, (Logan et al, 1989),,,,, ;,,,

10 3 : 477,, 12,, (, 2002a;, 2006;, 2007) (2005), 12 1,,,, F v /F m, RLC, α,,, 2 (, 2002a;, 2006;, 2007),,,,, 2 4, :, PSII F v /F m, , 0.69 (α) ,, 1000μmol/(s m 2 ), ; 1500μmol/(s m 2 ),,,,,, I k retr max, /,,,, 2 致 谢 本 研 究 中 的 海 上 调 查 和 样 品 采 集 工 作 得 到 了 本 课 题 组 赵 增 霞 王 世 伟 刘 群 万 艾 勇 老 师 和 宁 娟 杨 洋 同 学 以 及 创 新 号 考 察 船 全 体 工 作 人 员 的 大 力 支 持, 谨 致 谢 忱,,, , 43(3): ,, 2002a. Ⅱ. :, 32(3): ,,, 2002b.., (2): 39 44,,, , 36(6): ,,, , 42(5): ,,, , 23(3): ,,, , 36(6): , , 16(4): , :, 10 17,, :.. :, Babin M, Morel A, Claustre H et al, Nitrogen-and irradiance-dependent variations of the maximum quantum yield of carbon fixation in eutrophic, mesotrophic and oligotrophic marine systems. Deep Sea Research Part I: Oceanographic Research Papers, 43(8): Bergmann T, Richardson T, Paerl H et al, Synergy of light and nutrients on the photosynthetic efficiency of phytoplankton populations from the Neuse River estuary, North Carolina. Journal of Plankton Research, 24(9): Bilger W, Björkman O, Role of the xanthophyll cycle in photoprotection elucidated by measurements of lightinduced absorbance changes, fluorescence and photosynthesis in leaves of Hedera canariensis. Photosynthesis Research, 25(3): Bolhar-Nordenkampf H, Long S, Baker N et al, Chlorophyll fluorescence as a probe of the photosynthetic competence of leaves in the field: a review of current

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Aquatic Microbial Ecology, 56(2 3): Dugdale R C, Wilkerson F P, Silicate regulation of new production in the equatorial Pacific upwelling. Nature, 391(6664): Field C B, Behrenfeld M J, Randerson J T et al, Primary production of the biosphere: integrating terrestrial and oceanic components. Science, 281(5374): Figueroa F L, Conde A R, Gómez I, Relations between electron transport rates determined by pulse amplitude modulated chlorophyll fluorescence and oxygen evolution in macroalgae under different light conditions. Photosynthesis Research, 75(3): Goto N, Kihira M, Ishida N, Seasonal distribution of photosynthetically active phytoplankton using pulse amplitude modulated fluorometry in the large monomictic Lake Biwa, Japan. Journal of Plankton Research, 30(10): Häder D P, Porst M, Santas R, Photoinhibition by solar radiation in the Mediterranean alga Peyssonnelia squamata measured on site. 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12 3 : 479 (Western Gulf of St Lawrence, Canada). Estuarine, Coastal and Shelf Science, 48(2): Vaillancourt R D, Sambrotto R N, Green S et al, Phytoplankton biomass and photosynthetic competency in the summertime Mertz Glacier Region of East Antarctica. Deep Sea Research Part II: Topical Studies in Oceanography, 50(8): White A J, Critchley C, Rapid light curves: a new fluorescence method to assess the state of the photosynthetic apparatus. Photosynthesis Research, 59(1): PHOTOSYNTHETIC CHARACTERISTICS OF PHYTOPLANKTON IN WINTER IN THE JIAOZHOU BAY LI Jun-Lei 1, 2, SUN Xiao-Xia 1 (1. Jiaozhou Bay Marine Ecosystem Research Station, Institute of Oceanology, Chinese Academy of Sciences, Qingdao , China; 2. University of Chinese Academy of Sciences, Beijing , China) Abstract The photosynthetic characteristics of phytoplankton in Chinese offshore was less known because the studies were few. In this paper, Phyto-PAM (Pulse Amplitude Modulation) was applied to measure F v /F m (maximum quantum yield of PSII), RLCs (rapid light curves), qp, and NPQ (photochemical and non-photochemical quenching of phytoplankton). Phytoplankton community structure, distribution, and abundance, and the environmental parameters were also investigated to study the in situ photochemical vitality of phytoplankton in December (wintertime) in the Jiaozhou Bay. Results show that the photochemical vitality and potential of phytoplankton were active during this period; F v /F m maintained at about 0.69; most of the light utilization efficiencies (α) were between 0.2 to 0.3. Typical RLCs suggest that saturated electron transport appeared at PAR of 1000μmol/(s m 2 ), when PAR rose to 1500μmol/(s m 2 ), retr (relative electron transport rate) began to decline, Φ PS (the actual photochemical efficiency of PS ) reduced to the minimum and NPQ increased. Excess light energy was dissipated in heat, protecting marine phytoplankton photosynthesis apparatus from the damage of high intensity irradiation. Though phytoplankton abundance was extremely low in December. Strong photochemical activity combined with relatively stable hydrological, biological environment, and rich nutrients would lead to the peak of phytoplankton abundance in February in the Jiaozhou Bay. Key words Jiaozhou Bay; phytoplankton; community structure; photosynthetic characteristics; nutrient