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1 南海北部冷泉平端深海偏顶蛤的主要生化成分及其与热液和近岸种的对比研究 1, 郭欣雨 2 1, 2, 李超伦 (1.,, ; ) 摘要 : Bathymodiolus 属贻贝是广泛分布于全球深海冷泉和热液生态系统的优势种, 在深海化能生态系统的物质循环和能量流动中起着重要作用本文以我国南海北部冷泉的主要优势种平端深海偏顶蛤 (Bathymodiolus platifrons) 为研究对象, 采用常规生化测定方法, 研究了其主要生化成分和氨基酸组成的特征, 并就特定组织中主要生化成分的含量与生活在热液区的平端深海偏顶蛤和近岸的远东偏顶蛤 (Modiolus kurilensis) 进行了对比结果表明, 南海冷泉平端深海偏顶蛤软体部含水量 84.28%, 粗蛋白含量 7.18%, 粗脂肪含量 1.23%, 糖类含量 2.75%, 与已报道的深海贝类组成相近虽然主要生化成分含量在 3 种贻贝的鳃外套膜闭壳肌和消化腺 4 个组织中总体差异不大, 但是冷泉平端深海偏顶蛤和热液平端深海偏顶蛤的外套膜糖类含量 (25.20% 30.66%) 显著高于远东偏顶蛤 (6.97%, P<0.05), 这表明平端深海偏顶蛤的主要储存物质为外套膜中的糖类在氨基酸组成上, 冷泉平端深海偏顶蛤鳃中氨基酸总量为 44.55%( 干质量 ), 外套膜中为 34.83%( 干质量 ), 其中必需氨基酸分别占比 41.73% 和 40.52%, 总体与其他贝类相似然而, 在平端深海偏顶蛤中, 与渗透压调节相关的甘氨酸和与硫代谢相关的牛磺酸含量较高, 这与其适应深海高盐度高硫化氢浓度的环境相关综上所述, 南海冷泉平端深海偏顶蛤在常规生化组分和氨基酸组成上与近岸常见双壳类具有一定差异, 这些差异与其特殊生境的关系还需要更加深入的研究关键词 : 平端深海偏顶蛤, 生化成分, 冷泉, 南海中图分类号 : P735 文献标识码 : A 文章编号 : (2017) DOI: /hykx , [1], [2] 550m [3],,,, [4-5], 1~2 [3] Bathymodiolus [1],, [6-7] [8],,, [9-10], : ; : : (XDA ); ( ); (2017FY100803) [Foundation: Strategic Priority Research Program of the Chinese Academy of Sciences, No.XDA ; International Partnership for Innovative Team Program, No ; National Basic Resources Investigation Projects, No.2017FY100803] : (1992-),,,,, guoxinyu14@mails.ucas.ac.cn;,,, lcl@qdio.ac.cn Marine Sciences / Vol. 41, No. 6 /

2 Bathymodiolus, [11-13], Pranal 2 Bathymodiolus [14], Pruski 5 2 [15], (Bathymodiolus platifrons),, 1 材料与方法 1.1 材料 (Bathymodiolus platifrons) , (ROV),, 80 ROV SBE 25plus Sealogger CTD (Modiolus kurilensis), OSIL /, Niskin 1 表 1 样品采集地环境参数 Tab. 1 Environmental factors at collection sites /m T/ S ( E, N) ( E, 3 43 S) ( E, N) ~110 mm;, 65~95 mm; 80~100 mm, 4,,, 10,, 1.2 测定方法 ,, 50 mg 3 ml 30% KOH, 40 min, 5 ml 0.2% 15 min, 10 min, 625 nm, Kattner Fricke [16], 20 mg : :, GBT , 50 mg 5 ml 0.1 g 1 g, - 15 min, 400 nm,,, 6.25, 15 ml 7.8 ml, 100 ml mol/l,, h,, 0.02 mol/l, S-433D 17, 数据处理 SPSS 21.0, t- 3, ±, P< 结果 2.1 一般生化成分特征 84.28%, 66 / 2017 / 41 / 6

3 7.18%, 1.23%, 2.75% 1 1, 4, 12.30% 10.53% 11.94%;, 42.36% 49.96% 48.08%; ( 2),, 25.20% 30.66%,, 6.97%, (P<0.05) Fig Comparison between biochemical composition of different tissues from three mussel species a, b, c, d ; CSB HVB, MK Biochemical components of different tissues of seep mussels, vent mussels, and offshore mussels: (a) digestive gland; (b) gill; (c) mantle; (d) adductor. CSB is cold seep B. platifrons; HVB is hydrothermal vent B. platifrons; MK is M. kurilensis; black is crude lipid; densely crossed line is glycogen; sparse bias is protein. 2.2 氨基酸组成特征,,,, ( 2) 2, 17, 44.55%, 34.83%, 15.53%~15.69% 13.45%~14.51% 9.42%~ 11.07%, 3 (Thr, Val, Met, Ile, Leu, Phe, Lys, His, Arg) mg/g mg/g( ), 41.73% 40.52%, 6.47%~7.84% 6.23%~ 6.42% 6.10%~6.35% mg/g mg/g( ), 17.65mg/g 18.23mg/g, (P<0.05), Marine Sciences / Vol. 41, No. 6 /

4 3 讨论 2 Fig. 2 Glycogen content comparison between mantles of seep, vent, and off-shore mussels., (P<0.05) Value columns with same-letter superscripts indicate no significant difference; columns with different-letter superscripts indicate significant difference (P < 0.05) 表 2 南海冷泉平端深海偏顶蛤鳃和外套膜氨基酸质量比 (mg/g, 干质量, n=10) Tab. 2 Amino acid content of gill and mantle within SCS cold seep Bathymodiolus platifrons (mg/g, DW, n = 10) Asp 41.95± ±7.83 Thr 18.99± ±2.76 Ser 19.97± ±3.75 Glu 69.21± ±12.87 Gly 59.91± ±11.56 Ala 26.32± ±4.37 Cys 0.65± ±0.85 Val 21.57± ±2.80 Met 5.80± ±2.42 Ile 18.02± ±2.66 Leu 28.30± ±4.51 Tyr 19.01± ±2.31 Phe 17.10± ±2.40 His 18.70± ±1.73 Lys 28.61± ±2.78 Arg 28.84± ±6.83 Pro 22.58± ±5.62 Tau 17.65± ±3.86 EAA ± ± ± ± 基本生化成分,, ( 3), Bathymodiolus thermophiles,, B. thermophiles (Mytilus galloprovincialis) (Perna viridis) (Meretrix meretrix) (Ruditapes philippinarum),, 4, 3,, 3,, (P<0.05),,, (P<0.05) (Gafrarium tumidum) [23] (Saxidomus purpuratus) [24], (M.edulis) (~30%) [25-26], (Mytilus),, Smith Bathymodiolus childressi,, [27] 68 / 2017 / 41 / 6

5 表 3 不同双壳类间基本生化成分的比较 (%) Tab. 3 Comparison between biochemical composition of different bivalves (%) B. platifrons( ) 7.18(45.64) 1.23(7.85) 2.75(17.48) B. thermophiles [17] M. galloprovincialis [18] 10.18(57.68) 1.40(7.91) 4.90(27.74) P. viridis [19] 9.27(52.31) 2.43(13.68) 3.59(20.27) O. edulis [20] (47.50) (6.13) (25.39) M. meretrix [21] R. philippinarum [22] 9.34(55.88) 0.94(5.60) 3.92(23.43) :, 氨基酸组成, Bathymodiolus, 40% [14, 28-30], [14],, 4,, [31],,,, [32] ( ),,,,,,, 3 [33] (3.6%) [19],,,,,,, : [1] Myriam S, Karine O. Biogeography, biodiversity and fluid dependence of deep-sea cold-seep communities at active and passive margins[j]. Deep-Sea Research Part II: Topical Studies in Oceanography, 1998, 45(1-3): [2] Brooks J M, Kennicutt M C, Fay RR, et al. Thermogenic gas hydrates in the gulf of Mexico[J]. Science, 1984, 225(4660): [3] Russell C W, Powell E N. Why did ancient chemosynthetic seep and vent assemblages occur in shallower water than they do today?[j]. Int Journ Earth Sciences, 1999, 88(3): [4] James J C, Fisher C R, Brooks J M, et al. A methanotrophic marine molluscan (bivalvia, mytilidae) symbiosis: mussels fueled by gas[j]. Science, 1986, 233(4770): [5] Kochevar R E, Childress J J, Fisher C R, et al. The methane mussel: roles of symbiont and host in the metabolic utilization of methane[j]. Marine Biology, 1992, 112: [6] James M B, Kennicutt II M C, Fisher C R, et al. Deep-sea hydrocarbon seep communities: evidence for energy and nutritional carbon sources[j]. Science, 1987, 238(4830): Marine Sciences / Vol. 41, No. 6 /

6 [7] Fisher C R, Childress J J, OremLand R S, et al. The importance of methane and thiosulfate in the metabolism of the bacterial symbionts of two deep-sea mussels[j]. Marine Biology, 1987, 96: [8] Jørgensen C B. Bivalve filter feeding: hydrodynamics, bioenergetics, physiology and ecology[m]. Olsen & Olsen, [9] Kathleen M S, Chalers R F. Physiological ecology of sulfide metabolism in hydrothermal vent and cold seep vesicomyid clams and vestimentiferan tube worms[j]. American Zoologist, 1995, 35(2): [10] Von Damm K L, Buttermore L G, Oosting S E, et al. Direct observation of the evolution of a seafloor black smoker from vapor to brine[j]. Earth and Planetary Science Letters, 1997, 149: [11] Hiroaki S. Unusual novel n-4 polyunsaturated fatty acids in cold-seep mussels (Bathymodiolus japonicus and Bathymodiolus platifrons), originating from symbiotic methanotrophic bacteria[j]. Journal of Chromatography A, 2008, 1200(2): [12] Hiroaki S, Kazufumi O. Confirmation of a new food chain utilizing geothermal energy: Unusual fatty acids of a deep-sea bivalve, Calyptogena phaseoliformis[j]. Limnology and Oceanography, 2007, 52(5): [13] Hiroaki S, Masakazu M, Jun H. Lipid characteristics of a seep clam, Mesolinga soliditesta: Comparison with those of two coastal clams, Meretrix lamarckii and Ruditapes philippinarum[j]. Deep-Sea Research Part I: Oceanographic Research Papers, 2014, 94: [14] Pranal V, Fiala- Médioni A, Colomines J C. Amino acid and related compound composition in two symbiotic mytilid species from hydrothermal vents[j]. Marine Ecology Progress Series, 1995, 119(1-3): [15] Pruski A M, Fiala-Médioni A, Fisher C R, et al. Composition of free amino acids and related compounds in invertebrates with symbiotic bacteria at hydrocarbon seeps in the Gulf of Mexico[J]. Marine Biology, 2000, 136(3): [16] Gerhard K, Holger S G F. Simple gas-liquid chromatographic method for the simultaneous determination of fatty acids and alcohols in wax esters of marine organisms[j]. Journal of Chromatography, 1986, 361(1986): [17] Smith K L. Deep-sea hydrothermal vent mussels: nutritional state and distribution at the Galapagos Rift[J]. Ecology, 1985, 66(3): [18] Sedat K, Meryem Y Ç, İsmihan K, et al. Effects of stocking density on survival, growth and biochemical composition of cultured mussels (Mytilus galloprovincialis, Lamarck 1819) from an offshore submerged longline system[j]. Aquaculture Research, 2015, 46(6): [19],,. [J]., 2000, 19(1): Qing Ning, Lin Yue-guang, Jin Qi-zeng. Studies on nutritive compositions of soft part in mussel Perna viridis[j]. Tropic Oceanology, 2000, 19(1): [20] Sefa A, Aynur L, Ali K, et al. Seasonal variation in reproductive activity and biochemical composition of flat oyster (Ostrea edulis) in the Homa Lagoon, Izmir Bay, Turkey[J]. Scientia Marina, 2015, 79(4): [21],,,.., 2010, 31(23): Li Xiao-ying, Dong Zhi-guo, Yan Bin-lun, et al. Analysis and evaluation of nutritional components in Cyclina sinensis and Meretrix meretrix[j]. Food Science, 2010, 31(23): [22] MeryemY C, Saniye T Ç, Mehmet Ç, et al. Comparative study on biochemical composition of some edible marine molluscs at Canakkale coasts, Turkey[J]. Indian Journal of Geo-Marine Sciences, 2014, 43(4): [23] Babu A, Venkatesan V, Rajagopal S. Biochemical composition of different body parts of Gafrarium tumidum (Roding, 1798) from Mandapam, South East Coast of India[J]. African Journal of Biotechnology, 2012, 11(7): [24] Bi J H, Li Q, Zhang X J, et al. Seasonal variation of biochemical components in clam (Saxidomus purpuratus Sowerby 1852) in relation to its reproductive cycle and the environmental condition of Sanggou Bay, China[J]. Journal of Ocean University of China, 2016, 15(2): [25] de Zwaan A, Zandee D I. Body distribution and seasonal changes in the glycogen content of the common sea mussel Mytilus edulis[j]. Comparative Biochemistry and Physiology Part A: Physiology, 1972, 43(1): [26] Zandee D I, Kluytmans J H, Zurburg W. Seasonal variations in biochemical composition of Mytilus edulis with reference to energy metabolism and gametogenesis[j]. Netherlands Journal of Sea Research, 1980, 14(1): [27] Emily B S, Kathleen M St, Erica R N, et al. Growth and condition of seep mussels (Bathymodiolus childressi) at a gulf of Mexico Brine Pool[J]. Ecology, 2000, 81(9): [28] Tabakaeva O V, Tabakaev A V. Amino-acid profile of a mactridae bivalve mollusk from the Sea of Japan[J]. Chemistry of Natural Compounds, 2016, 52(5): [29] Tabakaeva O V, Tabakaev A V. Amino-acid composition of soft tissues of the Far-East bivalve mollusk Anadara broughtonii[j]. Chemistry of Natural Compounds, 2016, 52(3): [30] Chen D W, Su J, Liu X L, et al. Amino acid profiles of 70 / 2017 / 41 / 6

7 bvalve mollusks from Beibu Gulf, China[J]. Journal of Aquatic Food Product Technology, 2012, 21(4): [31] Page H M, Fisher C R, Childress J J. Role of filter-feeding in the nutritional biology of a deep-sea mussel with methanotrophic symbionts[j]. Marine Biology, 1990, 104: [32] Lehman L E, James M B, Kennedy T P, et al. Amino acid metabolism in euryhaline bivalves: regulation of glycine accumulation in ribbed mussel gills[j]. The Journal of Experimental Zoology, 1985, 233(3): [33] Audrey M P, Aline F M, Jean-Charles C. High amounts of sulphur-amino acids in three symbiotic mytilid bivalves from deep benthic communities[j]. Animal biology and pathology, 1997, 320(10): Biochemical components of cold seep mussel Bathymodiolus platifrons from South China Sea and comparison with hydrothermal vent and offshore mussels GUO Xin-yu 1, 2, LI Chao-lun 1, 2 (1. CAS Key Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Academy of Sciences, Qingdao , China; 2. University of Chinese Academy of Sciences, Beijing , China) Received: Dec., 24, 2016 Key words: Bathymodiolus platifrons; biochemical composition; cold seep; the South China Sea Abstract: Bathymodiolin mussels are a key species in cold seep and hydrothermal vent ecosystems where they are widely distributed and play an important role in the material cycle and energy flow of deep-sea chemotrophic ecosystems. In this paper, conventional biochemical determination methods are used to determine the fundamental biochemical components of Bathymodiolus platifrons in a South China Sea cold seep. Differences between mussels found at cold seeps, hydrothermal vents, and the offshore mussel Modiolus kurilensis are also discussed. Proportions of water, crude protein, lipids, and glycogen in B. platifrons are 84.28, 7.18, 1.23, and 2.75%, respectively, which are similar to those reported in deep-sea mussels. The glycogen content in the mantle of cold seep B. platifrons (25.20%) is significantly higher than that of M. kurilensis (6.97%) (P < 0.05), but is close to that of hydrothermal vent B. platifrons (30.66%), indicating that glycogen stored in the mantle is the energy reserve of B. platifrons. The total amino acid content in the gill and mantle of seep B. platifrons is and 34.83% of dry weight, in which the essential amino acid content accounts for and 40.52%, respectively. However, the contents of glycine and taurine are higher than in offshore bivalves. These compounds are related to osmoregulation and sulfur metabolism and may thus indicate environmental adaption. Results show certain differences between the biochemical component content of the SCS cold seep B. platifrons and offshore mussels; however, the relationship with special habitats needs further research. ( 本文编辑 : 梁德海 ) Marine Sciences / Vol. 41, No. 6 /

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i i ii 1 2 3 4 % N) 5 50.0 AMINO ACID #64 [modified by Administrator] IntAmp_1 nc 40.0 30.0 20.0 10.0 0.0-10.0 1 - ala - 10.483 2 - thr - 11.107 3 - gly - 11.872 4 - val - 13.458 5 - ser - 14.903 6 - pro