俞宙菲, 類彥立, 李鐵剛, 3, 4

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青島灣潮間帶活體底棲有孔蟲Ammonia aomoriensis(Asano, 1951)殼體18O值的季節(jié)變化

俞宙菲1, 2, 類彥立1, 李鐵剛1, 3, 4

(1. 中國(guó)科學(xué)院海洋研究所, 山東青島266071; 2. 中國(guó)科學(xué)院大學(xué), 北京100049; 3. 國(guó)家海洋局第一海洋研究所, 海洋沉積與環(huán)境地質(zhì)國(guó)家海洋局重點(diǎn)實(shí)驗(yàn)室, 山東青島266061; 4. 青島海洋科學(xué)與技術(shù)國(guó)家實(shí)驗(yàn)室海洋地質(zhì)過程與環(huán)境功能實(shí)驗(yàn)室, 山東青島266061)

近岸淺水底棲有孔蟲是陸架海區(qū)古環(huán)境重建的重要手段, 但是至今對(duì)其活體的研究非常缺乏。2014年6月至2015年5月對(duì)青島灣潮間帶活體底棲有孔蟲進(jìn)行了連續(xù)12個(gè)月的采樣, 用虎紅染色以確認(rèn)活體。對(duì)殼徑范圍在200 ~ 550 μm的活體殼體, 以每增加50 μm殼徑為一組, 進(jìn)行18O測(cè)試。結(jié)果顯示,殼體18O的月平均值的季節(jié)變化趨勢(shì)與溫度和鹽度一致, 與溫度反相關(guān), 與鹽度正相關(guān), 即青島灣殼體18O值受溫度和海水18O的影響, 其殼體18O值表現(xiàn)出了很好的季節(jié)性波動(dòng)。但是殼體18O的變化滯后于所測(cè)溫度和鹽度約2個(gè)月, 其記錄的是蟲體在生長(zhǎng)周期內(nèi)在真實(shí)鈣化溫度下分餾所得的18O值。

潮間帶;; 殼體氧同位素; 季節(jié)變化

溫度控制無機(jī)碳酸鈣沉降時(shí)的氧同位素分餾[1-4]。McCrea通過在4~80℃環(huán)境下文石和文石-碳酸鈣混合物的沉降實(shí)驗(yàn), 證實(shí)了溫度對(duì)無機(jī)碳酸鈣沉降同位素組成的控制作用[5]。隨后, Epstein等[6]用生長(zhǎng)在7~30℃環(huán)境的生物碳酸鈣進(jìn)行校正, 顯示出溫度與生物殼體18O之間負(fù)相關(guān)的關(guān)系。20世紀(jì)80年代, 這種控制作用進(jìn)一步通過野外和培養(yǎng)實(shí)驗(yàn)在浮游有孔蟲得到了證實(shí)[7-9]。通過對(duì)深海鉆孔頂部樣品中底棲有孔蟲殼體18O的測(cè)試, 建立了諸多底棲有孔蟲殼體18O與水溫的關(guān)系式, 包括[10-11],和[11-12]。利用深海底棲有孔蟲殼體18O來反映深海底層水溫度和全球冰量的變化, 在古海洋環(huán)境重建中已得到廣泛的應(yīng)用[13-17]。

但是, 由于對(duì)淺水底棲有孔蟲的野外調(diào)查研究甚少, 底棲有孔蟲活體18O對(duì)時(shí)間序列(季節(jié))響應(yīng)的記錄十分缺乏。Fontanier等對(duì)采自法國(guó)比斯開灣的活體底棲有孔蟲和等進(jìn)行了兩年的調(diào)查, 沒有觀察到殼體18O的季節(jié)變化[18]。Filipsson等對(duì)Havstens灣20 m水深和Gullmar灣119 m水深的有孔蟲進(jìn)行了16個(gè)月的調(diào)查, 也沒有發(fā)現(xiàn)溫度和鹽度的季節(jié)變化與殼體18O的關(guān)聯(lián)[19]。但Wefer 和Berger[20]的研究發(fā)現(xiàn)熱帶淺水大型底棲有孔蟲和的殼體18O表現(xiàn)出明顯的季節(jié)變化。

主要分布在北海和日本沿海的潮間帶、濕地和咸水湖環(huán)境[21-22]; 是波羅的海泥質(zhì)沉積物里的常見種[23]; 在中國(guó)黃海廣泛分布, 是青島灣的優(yōu)勢(shì)種之一[24]。分子和生態(tài)調(diào)查研究顯示, 在中國(guó)陸架海域?yàn)榭陀^存在的種類, 其形態(tài)變化與溫度無相關(guān)性, 但微球型初房比率與海水鹽度顯著正相關(guān)[25]。在世界上關(guān)于的研究較少, 很可能是因?yàn)檫@個(gè)種常被誤鑒定為[26]或[22, 24, 27]。

青島灣潮間帶環(huán)境的季節(jié)性變化很明顯, 鑒于淺水底棲有孔蟲野外調(diào)查數(shù)據(jù)的匱乏, 本文對(duì)青島灣活體有孔蟲的優(yōu)勢(shì)種進(jìn)行了連續(xù)12個(gè)月的殼體18O采樣測(cè)試, 探討了其殼體18O值對(duì)潮間帶環(huán)境季節(jié)變化的響應(yīng)。

1 材料與方法

1.1 樣品的采集和處理

沉積物樣品采自青島灣泥砂質(zhì)潮間帶(圖1a, b), 從2014年6月至2015年5月每月中旬采樣一次, 共12個(gè)月。退潮時(shí)采取表層1cm沉積物, 保存于采樣瓶中, 并加入適量海水。采樣時(shí)用水銀溫度計(jì)測(cè)量氣溫、水溫和泥溫, 用手持式鹽度計(jì)測(cè)量海水鹽度。樣品帶回實(shí)驗(yàn)室后, 去掉海水, 立即加入虎紅和酒精的混合溶液對(duì)活體進(jìn)行染色。放置48 h后, 將沉積物用300目(約48 μm)的篩絹過水沖洗, 粗粒部分放置于50℃烘箱中烘干。從烘干的樣品中挑出被虎紅染色的活體(圖1d)用于進(jìn)一步分析。

a.青島灣采樣點(diǎn); b. 潮間帶現(xiàn)場(chǎng)照片; c. 活體(見黃色原生質(zhì)); d. 經(jīng)虎紅染色后用于測(cè)試殼體18O的。圖中標(biāo)尺為200 μm

a. Sampling site in Qingdao Bay, Yellow Sea, China; b. Photograph of the intertidal flats; c. Livingwith yellow protoplasm; d. Individuals stained for stable oxygen isotope measurement. The scale bar is 200 μm

1.2 殼體18O測(cè)試

有孔蟲按殼徑每增加50 μm間隔被劃分成7個(gè)組: 200~250 μm, 250~300 μm, 300~350 μm, 350~400 μm, 400~450 μm, 450~500 μm和500~550 μm進(jìn)行18O測(cè)試。殼徑測(cè)量由實(shí)體顯微鏡LEICA S8AP0利用cellSens Standard軟件完成。從各個(gè)殼徑范圍分別挑取4~32枚。將每份樣品放入0.3 mL的玻璃小管內(nèi), 用解剖針扎破殼體。先加入適量3%的次氯酸鈉溶液, 用于除去被虎紅染色的原生質(zhì)。兩小時(shí)后吸掉廢液, 用去離子水清洗3次。再加入適量5%的H2O2溶液浸泡殼體約2 h, 期間可輕輕敲打玻璃小管管壁使H2O2溶液和殼體充分反應(yīng), 去除有機(jī)雜質(zhì)。再加入丙酮溶液洗去殼體表面粘附的油脂, 超聲30 s后用針管吸去濁液, 加水再洗3次, 放入50℃的烘箱中干燥之后待測(cè)試。氧同位素測(cè)試在中國(guó)科學(xué)院海洋研究所海洋地質(zhì)與環(huán)境重點(diǎn)實(shí)驗(yàn)室GV IsoPrime質(zhì)譜儀上進(jìn)行, 通過NBS18標(biāo)準(zhǔn)校正為PDB標(biāo)準(zhǔn), 本文數(shù)據(jù)測(cè)試標(biāo)準(zhǔn)誤差是0.07‰。

2 實(shí)驗(yàn)結(jié)果

表1顯示了2014年6月至2015年5月不同殼徑范圍的活體的殼體18O值和月平均值, 以及測(cè)量的氣溫、水溫、泥溫和鹽度值。野外測(cè)量的氣溫、水溫和泥溫的變化基本一致(圖2), 表現(xiàn)為夏季溫度高(8月最高), 冬天溫度低(2月最低)。測(cè)量氣溫的年變化范圍為4.2~32.5℃, 水溫的年變化范圍為5.8~33℃, 泥溫的年變化范圍為3.8~30℃。海水鹽度的季節(jié)變化范圍相對(duì)溫度的變化要窄很多, 為29~31, 體現(xiàn)為夏季月份最低, 而冬季月份最高, 全年的海水鹽度波動(dòng)為2個(gè)鹽度單位。

表1 青島灣潮間帶2014年6月至2015年5月各月活體A. aomoriensis的殼體δ18O值, 以及測(cè)量的環(huán)境因子

“/”表示沒有數(shù)據(jù)

殼體的月平均18O表現(xiàn)出在9月和10月最輕, 在3月和4月最重的特點(diǎn)。全年變化范圍為–0.75‰到0.72‰, 變化幅度為1.47‰。從圖2可以看到,殼體18O的季節(jié)變化模式與溫度和鹽度的變化模式相同, 但是其變化要滯后于環(huán)境變化約2個(gè)月。

3 討論

3.1 溫度對(duì)殼體18O的影響

Wefer和Berger對(duì)采自于1 ~ 3 m水深(年最低溫度26℃在2月, 年最高溫度30℃在7月)的熱帶大型底棲有孔蟲和10 m水深(年最低溫度16℃在2月, 年最高溫度29.5℃在8月)的進(jìn)行了殼體同位素分析, 結(jié)果顯示其殼體18O具有很好的季節(jié)變化[20]。放置于Cariaco海盆3 a的沉積物捕獲器顯示,(pink)的殼體18O基本精確地記錄了該海區(qū)表層水溫度的季節(jié)變化[28]。對(duì)加利福尼亞灣7 a的沉積物捕獲器研究也顯示的殼體18O精確地記錄了表層水溫度的季節(jié)變化[29]。但是由于自然環(huán)境的復(fù)雜性, 有孔蟲殼體18O對(duì)溫度的響應(yīng)可能會(huì)受到干擾。Fontanier等[18]對(duì)法國(guó)Biscay灣約550 m水深的和等活體底棲有孔蟲進(jìn)行研究, 沒有觀察到殼體18O的季節(jié)變化。Filipsson等[19]對(duì)Havstens灣20 m水深和Gullmar灣119 m水深的進(jìn)行了16個(gè)月調(diào)查, 結(jié)果表明溫度和鹽度的季節(jié)變化與殼體18O沒有顯著關(guān)系, 認(rèn)為海水18O對(duì)溫度的作用有一定的抵消。另外, 相對(duì)于野外調(diào)查, 能更好地控制變量的培養(yǎng)實(shí)驗(yàn)也證實(shí)了溫度對(duì)有孔蟲殼體18O的控制作用。例如將和分別放置于4℃, 7℃, 14℃和21℃下培養(yǎng), 實(shí)驗(yàn)表明其殼體18O隨溫度升高而變輕[30]。另一組在4~19℃的培養(yǎng)實(shí)驗(yàn)表明, 在控制殼徑的前提下, 溫度對(duì)殼體18O的控制作用為–0.22‰/℃[31]。將淺水底棲有孔蟲在10~27℃, 鹽度18~33范圍內(nèi)培養(yǎng), 觀察到殼體18O隨溫度升高而線性變輕[32]。

近岸淺水環(huán)境的季節(jié)性波動(dòng)比深海大, 如本研究的采樣點(diǎn)青島灣潮間帶季節(jié)溫差變化非常明顯, 冬夏差異高達(dá)約27℃(表1)。殼體18O值在變化模式上與溫度和鹽度的年際周期性變化一致, 顯示對(duì)溫度和鹽度變化響應(yīng)顯著。但是, 其最輕值出現(xiàn)在2014年9~10月, 而最重值出現(xiàn)在2015年3~4月, 比溫、鹽的變化滯后了約2個(gè)月。

根據(jù)Shackleton[10], Lynch-Stieglitz等[12]和Toyofuku等[32]所得的有孔蟲殼體18O的分餾平衡方程, 假設(shè)海水18O不變且分餾平衡, 溫度升高1℃, 殼體18O將偏輕0.21‰~0.25‰, 青島灣潮間帶冬季和夏季泥溫的最大差值約為26℃, 則殼體18O的年變化幅度應(yīng)該有5.46‰~6.5‰, 而實(shí)際的變化幅度只有1.47‰。其可能是因?yàn)檎鎸?shí)的鈣化溫度差沒有測(cè)量的那么大, 夏季的豐度非常低, 可能高溫不適合這個(gè)種的生長(zhǎng), 因此它們?cè)跍囟容^低的時(shí)間內(nèi)鈣化。另外, 也有可能是受到了生命效應(yīng)的影響。

3.2 鹽度對(duì)殼體18O的影響

有孔蟲殼體18O除了受溫度的控制以外, 海水18O也會(huì)對(duì)其產(chǎn)生影響[33-34]。海水18O值與水循環(huán)中的分餾密切相關(guān), 主要受蒸發(fā)、大氣水汽交換、降雨以及淡水輸入(降雨、河流和冰蓋融化)的影響[35-36]。由于這些過程最終可以在鹽度上體現(xiàn)出來, 因此海水18O與鹽度之間存在很好的線性正相關(guān)關(guān)系[37], 但是18O-鹽度關(guān)系式存在海區(qū)差異[12, 38]。另外, 海水18O的變化比鹽度更復(fù)雜, 顯著的季節(jié)和年際變化可能給海水18O或海水18O-鹽度的關(guān)系帶來偏差, 其原因有大氣中的附加分餾或海冰的形成[38]。

在中國(guó)邊緣海, 吳世迎[39]首先建立了黃河三角洲海區(qū)海水18O與鹽度的關(guān)系:18Ow= 0.25– 8.39。隨后, Kim等[40-41]將對(duì)馬暖流的數(shù)據(jù)與前人在長(zhǎng)江口、黃河口和黑潮區(qū)的數(shù)據(jù)相結(jié)合, 得到18Ow= 0.24– 8.13[42]。杜金秋等于夏季對(duì)黃、東海8個(gè)斷面57個(gè)站位的海水18O值進(jìn)行了測(cè)定, 得到關(guān)系式:18Ow= 0.29– 9.85(2= 0.67)[43]。Ye等[36]在夏季和冬季對(duì)黃、東海的表層海水18O做了調(diào)查, 海水18O值與鹽度呈正相關(guān)但存在季節(jié)差異:18Ow= 0.27– 10.68(2= 0.68,<0.001, 夏季);18Ow= 0.24– 8.66(2= 0.89,<0.001, 冬季), 大氣強(qiáng)迫的季節(jié)差異(潮濕溫和的夏季風(fēng)和強(qiáng)勁干燥的冬季風(fēng))和區(qū)域降水會(huì)給海水18O帶來季節(jié)和空間上的差異。

在本研究中, 青島灣的季節(jié)性非常明顯,殼體18O的變化與鹽度的變化呈正相關(guān), 由上述公式推算, 青島灣全年的鹽度變化為2, 相應(yīng)的海水18O的變化幅度為0.48‰~0.58‰, 可能會(huì)對(duì)殼體18O的變化帶來一定的影響。

3.3殼體18O季節(jié)變化的滯后

根據(jù)有孔蟲殼體的生長(zhǎng)特征, 出現(xiàn)滯后的原因可能來自于以下幾個(gè)方面。首先, 在野外采得的有孔蟲殼體記錄的信息是它在生長(zhǎng)期間的信息, 而我們現(xiàn)場(chǎng)測(cè)得的環(huán)境因子數(shù)據(jù)只是一瞬間的[44]。屬的實(shí)驗(yàn)室培養(yǎng)結(jié)果顯示, 有孔蟲個(gè)體從繁殖幼體到成體, 大概需要幾個(gè)月的時(shí)間。在10~27℃、鹽度18~33的培養(yǎng)條件下在2個(gè)月內(nèi)才長(zhǎng)到300 μm以上[32];在20℃, 鹽度32的條件下, 2個(gè)月最大長(zhǎng)到700 μm[45];sp.在26℃下2~3個(gè)月內(nèi)只長(zhǎng)到247~288 μm[46];在25℃條件下1個(gè)月生長(zhǎng)了10~21個(gè)房室[47];在18℃中6周最多只生長(zhǎng)了100 μm[27];而在1個(gè)月(25℃, 鹽度32)的時(shí)間內(nèi)只生長(zhǎng)了不到200 μm[26]。目前唯一關(guān)于淺水底棲有孔蟲殼體18O的季節(jié)變化的報(bào)道來自于熱帶大型底棲有孔蟲和, 它們分別采自于1 ~ 3 m水深和10 m水深, 由于個(gè)體較大, 可用解剖針進(jìn)行分房室測(cè)試, 每份樣品1~10個(gè)房室, 從而避免與早期房室的混合, 精確地恢復(fù)了殼體18O變化時(shí)間序列, 沒有出現(xiàn)滯后現(xiàn)象[20]。本文用來測(cè)試的殼徑范圍為200~550 μm, 由此推測(cè), 實(shí)驗(yàn)得到的殼體18O月平均值可能記錄了若干個(gè)月的環(huán)境信息, 而且不同季節(jié)有孔蟲殼體的生長(zhǎng)速度差異較大。的個(gè)體相對(duì)于大型有孔蟲要小, 為了滿足上機(jī)要求的樣品用量, 不可能對(duì)殼體進(jìn)行分房室測(cè)試, 從而無法剔除不同時(shí)間段、不同溫度條件下生長(zhǎng)的殼室的影響。

其次, 有孔蟲的生長(zhǎng)情況非常復(fù)雜。有孔蟲早期的房室小且生長(zhǎng)快, 后期房室大且生長(zhǎng)慢[34, 48-50]。的生長(zhǎng)速率從最開始的一天一個(gè)房室降低到一周一個(gè)房室[47]。Bradshaw[48]在實(shí)驗(yàn)室觀察了var.的生長(zhǎng)情況, 發(fā)現(xiàn)其成體在1個(gè)月內(nèi)只生長(zhǎng)了5~8個(gè)房室(10~35℃)。另外, 有孔蟲殼體的形成可能是集中在某個(gè)時(shí)間段(幾小時(shí))完成的[46]。利用微計(jì)算機(jī)斷層掃描技術(shù)(顯微CT)對(duì)野外共生大型有孔蟲進(jìn)行房室體積掃描, 發(fā)現(xiàn)其生長(zhǎng)出現(xiàn)潮汐周期(約14.8 d)和月相周期(約29.5 d), 可能是因?yàn)槌毕a(chǎn)生的強(qiáng)烈的潮流影響到了半附著的底棲有孔蟲[51-52], 細(xì)粒沉積物被懸浮而減弱了光的強(qiáng)度, 但增加了無機(jī)營(yíng)養(yǎng)物質(zhì), 從而影響了內(nèi)共生體, 且內(nèi)共生的微藻在滿月的時(shí)候光合作用速率更高[53]。月光的增強(qiáng)和潮汐濁流引起的水體中光的減弱這兩個(gè)因素之間的相互作用, 導(dǎo)致時(shí)而相互助益時(shí)而相互削弱[54]。這種潮汐和月亮周期在浮游有孔蟲[55-59]和其它大型底棲有孔蟲也有觀察到。

4 結(jié)論

本文對(duì)青島灣潮間帶淺水底棲有孔蟲進(jìn)行了連續(xù)12個(gè)月的活體采樣, 現(xiàn)場(chǎng)測(cè)量氣溫, 水溫, 泥溫和鹽度, 對(duì)虎紅染色的活體進(jìn)行分殼徑(200~550 μm, 50 μm間隔)殼體18O測(cè)試。研究顯示, 青島灣殼體18O值受溫度與海水18O的影響, 其殼體18O值表現(xiàn)出了很好的季節(jié)性波動(dòng)。但是,殼體18O滯后于溫度和鹽度的變化約2個(gè)月, 有孔蟲殼體記錄的是其在生長(zhǎng)期間, 在真實(shí)鈣化溫度下, 分餾所得的18O值。

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Seasonal oxygen isotope variation of a living benthic foraminifera(Asano, 1951) in the intertidal area of Qingdao Bay, Yellow Sea, China

YU Zhou-fei1, 2, LEI Yan-li1, LI Tie-gang1, 3, 4

(1. Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China; 2. University of Chinese Academy of Sciences, Beijing 100049, China; 3. Key Laboratory of Marine Sedimentology and Environmental Geology, First Institute of Oceanography, State Oceanic Administration People’s Republic of China, Qingdao 266061, China; 4. Function Laboratory of Marine Geology and Environment, National Laboratory for Marine Science and Technology, Qingdao 266061, China)Received：Mar. 23, 2016

intertidal flats;; stable oxygen isotope; seasonal variation

A successive 12-month study (June 2014 to May 2015) was conducted on the oxygen isotope composition (18O) of living benthic foraminifera, from the intertidal flats of Qingdao Bay, Yellow Sea, China.18O values were measured in different diameter test groups, ranging from 200 μm to 550 μm, at 50 μm intervals, from Rose Bengal stained specimens. The results showed that the fluctuant pattern of18O was remarkably similar to the temperature and salinity changes, negative and positive, respectively. The variation in18O in livingwas influenced by temperature and seawater18O, and exhibited a seasonal pattern which showed a time-lag of about two months. This result implies that the large shell ofreflects the environmental18O conditions during the months when the chambers form, and probably record the oxygen isotopes, obtained by fractionation under actual calcification temperatures, over the whole life cycle.

P67

A

1000-3096(2016)07-0132-08

10.11759/hykx20160323001

2016-03-23;

2016-05-26

國(guó)家自然科學(xué)基金(41476043, 41230959, 41176132); 中國(guó)科學(xué)院戰(zhàn)略性先導(dǎo)科技專項(xiàng)(A類)(XDA11030104); 大陸架科學(xué)鉆探項(xiàng)目(GZH201100202); 海洋生物群落結(jié)構(gòu)對(duì)氣候變化的響應(yīng)(GASI-03- 01-03-01; DOMEP(MEA)-01-01-E; No. 201303; MGK1210)

[Foundation: National Natural Science Foundation of China, No. 41476043, No. 41230959, No. 41176132; Strategic Priority Research Program of the Chinese Academy of Sciences, No. XDA11030104; Continental Shelf Drilling Program of China, No. GZH201100202; Response of marine biotic community structure to climate change, National Program on Global Change and Air-Sea Interaction, GASI-03-01-03-01; DOMEP (MEA)-01-01-E; No. 201303; MGK1210]

俞宙菲(1987-), 女, 江蘇常州人, 博士研究生, 研究方向: 海洋地質(zhì)學(xué), E-mail: yuzhoufei5@126.com;李鐵剛，通信作者，E-mail: tgli@fio.org.cn;

(本文編輯: 劉珊珊)