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Article(id=1243954933108682924, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1243954925370196248, articleNumber=null, orderNo=null, doi=10.3969/j.issn.0253-4193.2020.04.003, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1559750400000, receivedDateStr=2019-06-06, revisedDate=1567872000000, revisedDateStr=2019-09-08, acceptedDate=null, acceptedDateStr=null, onlineDate=1774511567618, onlineDateStr=2026-03-26, pubDate=1587744000000, pubDateStr=2020-04-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1774511567618, onlineIssueDateStr=2026-03-26, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1774511567618, creator=13701087609, updateTime=1774511567618, updator=13701087609, issue=Issue{id=1243954925370196248, tenantId=1146029695717560320, journalId=1149651085930835976, year='2020', volume='42', issue='4', pageStart='1', pageEnd='136', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1774511565773, creator=13701087609, updateTime=1774511565773, updator=13701087609, preIssue=null, nextIssue=null, ext=null, issueFiles=null}, startPage=20, endPage=28, ext={EN=ArticleExt(id=1243954933465198774, articleId=1243954933108682924, tenantId=1146029695717560320, journalId=1149651085930835976, language=EN, title=The distribution of fatty acids in tissues of rock cod (Patagonotothen ramsayi) in the Argentina Patagonian Shelf and their implications to feeding habit, columnId=1243954927383462170, journalTitle=Haiyang Xuebao, columnName=Marine Biology, runingTitle=null, highlight=null, articleAbstract=

As one of the most abundant species of family Notothenioidei, rock cod (Patagonotothen ramsayi) plays an important role in energy transfer in the food web in the Patagonian Shelf. Currently, the work on feeding ecology of this species mainly bases on the conventional stomach content analysis, which only reflects short-term variation in feeding. Therefore, this study analyzed the distribution of fatty acids in three tissues (muscle, liver and gonad) of rock cod, and analyzed whether fatty acids in these tissues can characterize their feeding habits and transfer of diet. The results indicate that 27 kinds of fatty acids were detected in the tissues of rock cod. The storage and use of fatty acids in liver and gonad tissues are not suitable for indicating the diet of rock cod because they are involved in growth and reproduction. Compared to liver and gonad tissues, the longer renewal cycle of muscle tissue may better reflect its absorption of fatty acids from food. For the analysis of fatty acids in the muscle tissue of the rock cod in different size groups, it is inferred that the small-size rock cod (immature individual, 100−240 mm) mainly fed on planktonic organisms. The benthic feeding characteristics of rock cod increase with the increasing of size, and the feeding pattern transfer from planktonic feeding to planktonic-benthic feeding. In addition, due to the consumption of a certain amount of fishery discards by rock cod, the contribution rate of saprophytic food chain to predatory food chain is weakened. The results derive from this study further demonstrate that, compare to tissues of liver and gonad, the fatty acids of muscle tissue can be more suitable to indicate the food source of oceanic fish species.

, correspAuthors=Guoping Zhu, authorNote=null, correspAuthorsNote=null, copyrightStatement=Haiyang Xuebao, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Qingyuan Yang, Guoping Zhu), CN=ArticleExt(id=1243954935008702729, articleId=1243954933108682924, tenantId=1146029695717560320, journalId=1149651085930835976, language=CN, title=阿根廷巴塔哥尼亚陆架拉氏南美南极鱼（Patagonotothen ramsayi）不同组织中脂肪酸分布及食物来源指示, columnId=1243954927517679901, journalTitle=海洋学报, columnName=海洋生物, runingTitle=null, highlight=null, articleAbstract=

作为南美南极鱼科中数量最为丰富的一个种，拉氏南美南极鱼(Patagonotothen ramsayi)在巴塔哥尼亚海域食物网能量流动中起着重要的传递作用。目前针对该鱼种的营养及摄食生态学研究主要集中于反映短期摄食的传统胃含物分析上。为此，本研究分析了拉氏南美南极鱼3种组织(肌肉、肝脏和性腺)中脂肪酸含量及分布情况，并就3种组织中的脂肪酸是否能表征其食性及食性转换进行了探究。结果表明，拉氏南美南极鱼体内共检测出27种脂肪酸；由于涉及生长、繁殖等因素影响，肝脏、性腺组织对于脂肪酸的储存及使用情况并不适用于指征食性，而肌肉组织更新时间相对较长，故能够较好地反映其对食物中脂肪酸的吸收。对不同体长组拉氏南美南极鱼肌肉组织中脂肪酸分析可知，未成熟拉氏南美南极鱼(100～240 mm)主要摄食浮游性生物；随着体长增加对底栖生物的摄食也随之增加，由浮游性摄食方式转变为浮游−底栖性摄食。另外，因拉氏南美南极鱼摄食一定量的渔业丢弃物，导致腐生食物链和捕食食物链的贡献率特征弱化。研究结果进一步显示，针对大洋性鱼类，与肝脏和性腺相比，肌肉组织脂肪酸更适用于表征食物来源。

, correspAuthors=朱国平, authorNote=null, correspAuthorsNote=
*朱国平（1976-），男，教授，从事极地生物学与渔业生态学方面的研究。E-mail：
, copyrightStatement=版权所有©《海洋学报》编辑部 2023
杨清源，朱国平. 阿根廷巴塔哥尼亚陆架拉氏南美南极鱼（Patagonotothen ramsayi）不同组织中脂肪酸分布及食物来源指示[J]. 海洋学报，2020，42(4)：20–28，doi:10.3969/j.issn.0253−4193.2020.04.003Yang Qingyuan，Zhu Guoping. The distribution of fatty acids in tissues of rock cod (Patagonotothen ramsayi) in the Argentina Patagonian Shelf and their implications to feeding habit[J]. Haiyang Xuebao，2020, 42(4)：20–28，doi:10.3969/j.issn.0253−4193.2020.04.003
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杨清源（1994-），女，山东省青岛市人，研究方向为海洋生物学。E-mail：

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杨清源（1994-），女，山东省青岛市人，研究方向为海洋生物学。E-mail：

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Potential dietary effects on the fatty acid composition of the common jellyfish Aurelia aurita[J]. 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PC1 (PC2) represents the contribution ratio of the first (second) principal component; the vertical coordinate on the right and the horizontal coordinate on the top are the scores of sample points on PC1 and PC2, respectively

, figureFileSmall=88hyWfL3T8VRWMw1eZOdwA==, figureFileBig=rkvMvTNmJuT59CkFCfQJiw==, tableContent=null), ArticleFig(id=1246538013254242926, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=CN, label=图2, caption=拉氏南美南极鱼不同组织脂肪酸含量主成分分析

PC1为第一主成分贡献率，PC2为第二主成分贡献率；右方纵坐标与上方横坐标分别为样品点在PC1及PC2上的得分情况

, figureFileSmall=88hyWfL3T8VRWMw1eZOdwA==, figureFileBig=rkvMvTNmJuT59CkFCfQJiw==, tableContent=null), ArticleFig(id=1246538013388460661, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=EN, label=Table 1, caption=

The main fatty acids found in the rock cod and their corresponding potential food sources

, figureFileSmall=null, figureFileBig=null, tableContent=
脂肪酸直接与间接食物源参考文献
C14:0硅藻纲[21]
C16:0硅藻纲[21]
C15:0+C17:0浮游细菌[22]
C16:1n7硅藻纲[23]
C18:1n9褐藻/浮游动物[24]
C18:2n6陆源脂肪酸[25]
C20:1+C22:1桡足类[26]
C20:4n6底栖生物[26]
C20:5n3硅藻纲[21]
C22:6n3甲藻纲[21]
), ArticleFig(id=1246538013480735355, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=CN, label=表1, caption=

在拉氏南美南极鱼体内发现的主要脂肪酸及直接与间接食物源

, figureFileSmall=null, figureFileBig=null, tableContent=
脂肪酸直接与间接食物源参考文献
C14:0硅藻纲[21]
C16:0硅藻纲[21]
C15:0+C17:0浮游细菌[22]
C16:1n7硅藻纲[23]
C18:1n9褐藻/浮游动物[24]
C18:2n6陆源脂肪酸[25]
C20:1+C22:1桡足类[26]
C20:4n6底栖生物[26]
C20:5n3硅藻纲[21]
C22:6n3甲藻纲[21]
), ArticleFig(id=1246538013556232830, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=EN, label=Table 2, caption=

The size group of the rock cod

, figureFileSmall=null, figureFileBig=null, tableContent=
体长组体长范围数量
小体长100 mm≤L<240 mm26
大体长240 mm≤L10
), ArticleFig(id=1246538013631730303, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=CN, label=表2, caption=

拉氏南美南极鱼体长分组

, figureFileSmall=null, figureFileBig=null, tableContent=
体长组体长范围数量
小体长100 mm≤L<240 mm26
大体长240 mm≤L10
), ArticleFig(id=1246538013740782210, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=EN, label=Table 3, caption=

Fatty acid contents in different tissues of the rock cod

, figureFileSmall=null, figureFileBig=null, tableContent=
脂肪酸平均值/mg·g−1
肌肉肝脏性腺
小体长大体长总体小体长大体长总体小体长大体长总体
　　注：SFA为饱和脂肪酸，MUFA为单不饱和脂肪酸，PUFA为多不饱和脂肪酸，TFA为总脂肪酸；±后为标准偏差。
C14:01.09±0.631.71±1.431.26±0.9415.09±12.0717.39±7.3015.73±10.908.81±16.0814.31±18.5910.34±16.73
C15:00.23±0.100.33±0.100.26±0.111.27±0.860.93±0.181.18±0.751.66±1.601.87±2.111.72±1.72
C16:07.54±3.6311.50±4.588.64±4.2562.57±49.0765.78±23.7963.46±43.2142.59±51.5559.26±64.7947.22±55.09
C17:01.37±0.801.64±0.691.45±0.773.14±1.032.67±0.193.01±0.907.94±7.374.07±3.106.86±6.66
C18:02.84±0.873.28±0.622.96±0.8212.83±9.0011.95±6.3012.58±8.2613.69±9.8010.08±8.4012.69±9.46
C20:01.33±0.821.56±0.781.40±0.812.43±0.652.29±0.142.39±0.567.57±7.723.34±2.786.39±6.95
C21:01.09±0.681.26±0.651.14±0.671.87±0.541.67±0.011.82±0.466.17±6.402.54±2.215.17±5.77
C22:00.47±0.290.55±0.270.49±0.290.82±0.230.74±0.010.80±0.202.21±1.941.15±0.981.91±1.78
C23:01.02±0.700.92±0.780.99±0.711.75±0.731.68±0.031.73±0.626.06±6.392.37±2.325.04±5.78
C24:00.96±0.601.11±0.571.00±0.591.64±0.481.47±0.011.60±0.415.38±5.712.09±2.054.46±5.15
SFA17.95±6.7423.86±5.6919.59±6.93103.41±70.60106.55±36.18104.28±62.44102.08±83.38101.09±100.31101.80±86.91
C15:1n50.61±0.460.80±0.420.66±0.451.16±0.411.07±0.011.14±0.353.77±4.151.62±1.403.18±3.71
C16:1n72.40±1.243.32±1.412.65±1.3453.82±50.0970.35±44.1858.41±48.4818.61±20.9722.26±25.2019.62±21.91
C17:1n71.49±1.061.27±1.251.43±1.100.95±1.410.25±0.790.76±1.307.83±10.123.23±3.606.55±8.99
C18:1n95.11±3.317.58±4.335.80±3.73133.75±146.03151.35±86.67138.64±131.2538.17±49.9154.61±66.5942.74±54.54
C20:11.76±0.852.44±0.891.95±0.907.51±5.048.46±4.997.77±4.979.78±7.887.46±5.959.14±7.39
C22:1n91.49±0.832.07±0.791.65±0.853.79±1.414.43±1.833.97±1.538.63±7.807.82±8.858.41±7.98
C24:1n91.25±0.741.52±0.671.33±0.722.62±0.682.38±0.322.55±0.607.21±6.883.90±3.186.29±6.22
MUFA14.11±5.7419.00±6.0615.47±6.16203.60±197.94238.29±130.78213.23±180.6494.01±81.53100.90±107.7195.92±87.98
C18:2n66.01±3.616.85±3.256.24±3.4911.94±3.529.98±0.6211.40±3.1232.94±32.7414.94±12.1727.94±29.50
C18:3n61.49±0.921.73±0.871.56±0.902.64±0.732.36±0.062.56±0.638.43±8.643.60±3.117.09±7.78
C18:3n31.80±1.032.16±0.931.90±1.013.42±0.893.16±0.233.34±0.778.33±6.925.03±4.237.41±6.40
C20:21.46±0.851.71±0.771.53±0.823.04±0.772.74±0.162.96±0.678.14±7.973.80±3.056.94±7.19
C20:3n61.08±0.691.27±0.641.14±0.681.93±0.531.75±0.061.88±0.464.55±4.762.61±2.214.01±4.27
C20:3n33.17±1.713.77±1.553.34±1.676.29±2.105.39±0.836.04±1.8716.54±16.097.44±5.2614.01±14.46
C20:4n65.57±3.126.50±2.975.83±3.0710.63±3.139.35±1.0210.28±2.7629.82±29.5913.69±10.4525.34±26.59
C20:5n38.86±4.8611.24±4.769.52±4.8921.04±24.7618.50±8.5920.33±21.4121.35±27.6441.97±45.1027.08±34.00
C22:2n61.12±1.371.01±1.611.09±1.422.40±1.192.18±1.152.34±1.167.02±11.243.02±3.985.91±9.88
C22:6n326.48±12.8432.61±12.3828.18±12.8446.06±64.4043.11±24.9945.24±55.9036.19±47.7182.41±85.9849.03±62.99
PUFA57.05±18.2568.85±13.7660.33±17.76109.40±92.2398.51±34.06106.37±79.99173.30±138.01178.51±157.40174.75±141.35
C15:0+C17:01.60±0.881.97±0.711.71±0.844.42±1.773.59±0.364.19±1.559.60±8.055.94±4.818.58±7.42
C20:1+C22:13.26±1.674.51±1.653.61±1.7411.30±6.3012.89±6.7911.74±6.3818.41±15.5615.28±14.4217.54±15.11
C20:5n3 / C22:6n30.35±0.160.36±0.130.35±0.150.59±0.200.47±0.110.55±0.190.62±0.200.46±0.150.58±0.20
n-3 PUFA40.31±16.3749.78±14.4942.95±16.2576.80±89.3770.16±33.6674.95±77.5082.41±79.43136.86±135.0797.53±99.04
n-6 PUFA15.28±9.0217.36±6.8315.85±8.4229.55±7.3725.62±2.0728.46±6.5782.75±83.9037.85±31.4970.28±75.49
TFA89.12±28.89111.70±23.9495.39±29.14416.40±322.22443.36±178.00423.89±287.16369.39±285.84380.50±359.57372.48±302.71
), ArticleFig(id=1246538013937914505, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=CN, label=表3, caption=

拉氏南美南极鱼不同组织中脂肪酸含量

, figureFileSmall=null, figureFileBig=null, tableContent=
脂肪酸平均值/mg·g−1
肌肉肝脏性腺
小体长大体长总体小体长大体长总体小体长大体长总体
　　注：SFA为饱和脂肪酸，MUFA为单不饱和脂肪酸，PUFA为多不饱和脂肪酸，TFA为总脂肪酸；±后为标准偏差。
C14:01.09±0.631.71±1.431.26±0.9415.09±12.0717.39±7.3015.73±10.908.81±16.0814.31±18.5910.34±16.73
C15:00.23±0.100.33±0.100.26±0.111.27±0.860.93±0.181.18±0.751.66±1.601.87±2.111.72±1.72
C16:07.54±3.6311.50±4.588.64±4.2562.57±49.0765.78±23.7963.46±43.2142.59±51.5559.26±64.7947.22±55.09
C17:01.37±0.801.64±0.691.45±0.773.14±1.032.67±0.193.01±0.907.94±7.374.07±3.106.86±6.66
C18:02.84±0.873.28±0.622.96±0.8212.83±9.0011.95±6.3012.58±8.2613.69±9.8010.08±8.4012.69±9.46
C20:01.33±0.821.56±0.781.40±0.812.43±0.652.29±0.142.39±0.567.57±7.723.34±2.786.39±6.95
C21:01.09±0.681.26±0.651.14±0.671.87±0.541.67±0.011.82±0.466.17±6.402.54±2.215.17±5.77
C22:00.47±0.290.55±0.270.49±0.290.82±0.230.74±0.010.80±0.202.21±1.941.15±0.981.91±1.78
C23:01.02±0.700.92±0.780.99±0.711.75±0.731.68±0.031.73±0.626.06±6.392.37±2.325.04±5.78
C24:00.96±0.601.11±0.571.00±0.591.64±0.481.47±0.011.60±0.415.38±5.712.09±2.054.46±5.15
SFA17.95±6.7423.86±5.6919.59±6.93103.41±70.60106.55±36.18104.28±62.44102.08±83.38101.09±100.31101.80±86.91
C15:1n50.61±0.460.80±0.420.66±0.451.16±0.411.07±0.011.14±0.353.77±4.151.62±1.403.18±3.71
C16:1n72.40±1.243.32±1.412.65±1.3453.82±50.0970.35±44.1858.41±48.4818.61±20.9722.26±25.2019.62±21.91
C17:1n71.49±1.061.27±1.251.43±1.100.95±1.410.25±0.790.76±1.307.83±10.123.23±3.606.55±8.99
C18:1n95.11±3.317.58±4.335.80±3.73133.75±146.03151.35±86.67138.64±131.2538.17±49.9154.61±66.5942.74±54.54
C20:11.76±0.852.44±0.891.95±0.907.51±5.048.46±4.997.77±4.979.78±7.887.46±5.959.14±7.39
C22:1n91.49±0.832.07±0.791.65±0.853.79±1.414.43±1.833.97±1.538.63±7.807.82±8.858.41±7.98
C24:1n91.25±0.741.52±0.671.33±0.722.62±0.682.38±0.322.55±0.607.21±6.883.90±3.186.29±6.22
MUFA14.11±5.7419.00±6.0615.47±6.16203.60±197.94238.29±130.78213.23±180.6494.01±81.53100.90±107.7195.92±87.98
C18:2n66.01±3.616.85±3.256.24±3.4911.94±3.529.98±0.6211.40±3.1232.94±32.7414.94±12.1727.94±29.50
C18:3n61.49±0.921.73±0.871.56±0.902.64±0.732.36±0.062.56±0.638.43±8.643.60±3.117.09±7.78
C18:3n31.80±1.032.16±0.931.90±1.013.42±0.893.16±0.233.34±0.778.33±6.925.03±4.237.41±6.40
C20:21.46±0.851.71±0.771.53±0.823.04±0.772.74±0.162.96±0.678.14±7.973.80±3.056.94±7.19
C20:3n61.08±0.691.27±0.641.14±0.681.93±0.531.75±0.061.88±0.464.55±4.762.61±2.214.01±4.27
C20:3n33.17±1.713.77±1.553.34±1.676.29±2.105.39±0.836.04±1.8716.54±16.097.44±5.2614.01±14.46
C20:4n65.57±3.126.50±2.975.83±3.0710.63±3.139.35±1.0210.28±2.7629.82±29.5913.69±10.4525.34±26.59
C20:5n38.86±4.8611.24±4.769.52±4.8921.04±24.7618.50±8.5920.33±21.4121.35±27.6441.97±45.1027.08±34.00
C22:2n61.12±1.371.01±1.611.09±1.422.40±1.192.18±1.152.34±1.167.02±11.243.02±3.985.91±9.88
C22:6n326.48±12.8432.61±12.3828.18±12.8446.06±64.4043.11±24.9945.24±55.9036.19±47.7182.41±85.9849.03±62.99
PUFA57.05±18.2568.85±13.7660.33±17.76109.40±92.2398.51±34.06106.37±79.99173.30±138.01178.51±157.40174.75±141.35
C15:0+C17:01.60±0.881.97±0.711.71±0.844.42±1.773.59±0.364.19±1.559.60±8.055.94±4.818.58±7.42
C20:1+C22:13.26±1.674.51±1.653.61±1.7411.30±6.3012.89±6.7911.74±6.3818.41±15.5615.28±14.4217.54±15.11
C20:5n3 / C22:6n30.35±0.160.36±0.130.35±0.150.59±0.200.47±0.110.55±0.190.62±0.200.46±0.150.58±0.20
n-3 PUFA40.31±16.3749.78±14.4942.95±16.2576.80±89.3770.16±33.6674.95±77.5082.41±79.43136.86±135.0797.53±99.04
n-6 PUFA15.28±9.0217.36±6.8315.85±8.4229.55±7.3725.62±2.0728.46±6.5782.75±83.9037.85±31.4970.28±75.49
TFA89.12±28.89111.70±23.9495.39±29.14416.40±322.22443.36±178.00423.89±287.16369.39±285.84380.50±359.57372.48±302.71
), ArticleFig(id=1246538014034383502, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1243954933108682924, language=EN, label=Table 4, caption=

The relationship between standard length and percentage of specific fatty acids in the different tissues of rock cod

, figureFileSmall=null, figureFileBig=null, tableContent=
脂肪酸肌肉肝脏性腺
R2pR2pR2p
　　注：*表示两者关系呈线性关系（p<0.05）。
C14:00.0200.4120.0000.9760.0000.915
C16:00.1290.031*0.0090.5750.0000.917
C15:0+C17:00.1250.035*0.1380.026*0.2300.003*
C16:1n70.0320.2950.0100.5540.0010.848
C18:1n90.0210.3950.0040.7200.0020.797
C18:2n60.0950.0670.2350.003*0.2510.002*
C20:1+C22:10.2320.003*0.0190.4170.1520.019*
C20:4n60.0970.0640.2250.003*0.2400.002*
C20:5n30.0160.4620.0010.8560.0250.362
C22:6n30.0110.5510.0020.7950.0610.148
C20:5n3/ C20:5n30.0100.5530.0890.0780.3260.000*
n-6 PUFA/n-3 PUFA0.0040.7060.0340.2800.2520.002*
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拉氏南美南极鱼体长与特征脂肪酸含量之间的关系

, figureFileSmall=null, figureFileBig=null, tableContent=
脂肪酸肌肉肝脏性腺
R2pR2pR2p
　　注：*表示两者关系呈线性关系（p<0.05）。
C14:00.0200.4120.0000.9760.0000.915
C16:00.1290.031*0.0090.5750.0000.917
C15:0+C17:00.1250.035*0.1380.026*0.2300.003*
C16:1n70.0320.2950.0100.5540.0010.848
C18:1n90.0210.3950.0040.7200.0020.797
C18:2n60.0950.0670.2350.003*0.2510.002*
C20:1+C22:10.2320.003*0.0190.4170.1520.019*
C20:4n60.0970.0640.2250.003*0.2400.002*
C20:5n30.0160.4620.0010.8560.0250.362
C22:6n30.0110.5510.0020.7950.0610.148
C20:5n3/ C20:5n30.0100.5530.0890.0780.3260.000*
n-6 PUFA/n-3 PUFA0.0040.7060.0340.2800.2520.002*
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阿根廷巴塔哥尼亚陆架拉氏南美南极鱼（Patagonotothen ramsayi）不同组织中脂肪酸分布及食物来源指示
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杨清源 1, 2, 3 , 朱国平 1, 2, 3, 4, *
海洋学报 | 海洋生物 2020,42(4): 20-28
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海洋学报 | 海洋生物 2020, 42(4): 20-28
阿根廷巴塔哥尼亚陆架拉氏南美南极鱼（Patagonotothen ramsayi）不同组织中脂肪酸分布及食物来源指示
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杨清源1, 2, 3 , 朱国平1, 2, 3, 4, *
作者信息
  • 1 上海海洋大学 海洋科学学院，上海 201306
  • 2 上海海洋大学 极地研究中心，上海 201306
  • 3 大洋渔业资源可持续开发教育部重点实验室 极地海洋生态系统研究室，上海 201306
  • 4 国家远洋渔业工程技术研究中心，上海 201306

    • 杨清源（1994-），女，山东省青岛市人，研究方向为海洋生物学。E-mail：

通讯作者:

*朱国平（1976-），男，教授，从事极地生物学与渔业生态学方面的研究。E-mail：
The distribution of fatty acids in tissues of rock cod (Patagonotothen ramsayi) in the Argentina Patagonian Shelf and their implications to feeding habit
Qingyuan Yang1, 2, 3 , Guoping Zhu1, 2, 3, 4, *
Affiliations
  • 1 College of Marine Sciences, Shanghai Ocean University, Shanghai 201306, China
  • 2 Center for Polar Research, Shanghai Ocean University, Shanghai 201306, China
  • 3 Polar Marine Ecosystem Laboratory, Key Laboratory of Sustainable Exploitation of Oceanic Fisheries Resources, Ministry of Education, Shanghai 201306, China
  • 4 National Engineering Research Center for Oceanic Fisheries, Shanghai 201306, China
出版时间: 2020-04-25 doi: 10.3969/j.issn.0253-4193.2020.04.003
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摘要
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作为南美南极鱼科中数量最为丰富的一个种，拉氏南美南极鱼(Patagonotothen ramsayi)在巴塔哥尼亚海域食物网能量流动中起着重要的传递作用。目前针对该鱼种的营养及摄食生态学研究主要集中于反映短期摄食的传统胃含物分析上。为此，本研究分析了拉氏南美南极鱼3种组织(肌肉、肝脏和性腺)中脂肪酸含量及分布情况，并就3种组织中的脂肪酸是否能表征其食性及食性转换进行了探究。结果表明，拉氏南美南极鱼体内共检测出27种脂肪酸；由于涉及生长、繁殖等因素影响，肝脏、性腺组织对于脂肪酸的储存及使用情况并不适用于指征食性，而肌肉组织更新时间相对较长，故能够较好地反映其对食物中脂肪酸的吸收。对不同体长组拉氏南美南极鱼肌肉组织中脂肪酸分析可知，未成熟拉氏南美南极鱼(100～240 mm)主要摄食浮游性生物；随着体长增加对底栖生物的摄食也随之增加，由浮游性摄食方式转变为浮游−底栖性摄食。另外，因拉氏南美南极鱼摄食一定量的渔业丢弃物，导致腐生食物链和捕食食物链的贡献率特征弱化。研究结果进一步显示，针对大洋性鱼类，与肝脏和性腺相比，肌肉组织脂肪酸更适用于表征食物来源。

拉氏南美南极鱼  /  脂肪酸  /  食性  /  巴塔哥尼亚陆架

As one of the most abundant species of family Notothenioidei, rock cod (Patagonotothen ramsayi) plays an important role in energy transfer in the food web in the Patagonian Shelf. Currently, the work on feeding ecology of this species mainly bases on the conventional stomach content analysis, which only reflects short-term variation in feeding. Therefore, this study analyzed the distribution of fatty acids in three tissues (muscle, liver and gonad) of rock cod, and analyzed whether fatty acids in these tissues can characterize their feeding habits and transfer of diet. The results indicate that 27 kinds of fatty acids were detected in the tissues of rock cod. The storage and use of fatty acids in liver and gonad tissues are not suitable for indicating the diet of rock cod because they are involved in growth and reproduction. Compared to liver and gonad tissues, the longer renewal cycle of muscle tissue may better reflect its absorption of fatty acids from food. For the analysis of fatty acids in the muscle tissue of the rock cod in different size groups, it is inferred that the small-size rock cod (immature individual, 100−240 mm) mainly fed on planktonic organisms. The benthic feeding characteristics of rock cod increase with the increasing of size, and the feeding pattern transfer from planktonic feeding to planktonic-benthic feeding. In addition, due to the consumption of a certain amount of fishery discards by rock cod, the contribution rate of saprophytic food chain to predatory food chain is weakened. The results derive from this study further demonstrate that, compare to tissues of liver and gonad, the fatty acids of muscle tissue can be more suitable to indicate the food source of oceanic fish species.

rock cod  /  fatty acid  /  feeding habit  /  Patagonian Shelf
杨清源, 朱国平. 阿根廷巴塔哥尼亚陆架拉氏南美南极鱼（Patagonotothen ramsayi）不同组织中脂肪酸分布及食物来源指示. 海洋学报, 2020 , 42 (4) : 20 -28 . DOI: 10.3969/j.issn.0253-4193.2020.04.003
Qingyuan Yang, Guoping Zhu. The distribution of fatty acids in tissues of rock cod (Patagonotothen ramsayi) in the Argentina Patagonian Shelf and their implications to feeding habit[J]. Haiyang Xuebao, 2020 , 42 (4) : 20 -28 . DOI: 10.3969/j.issn.0253-4193.2020.04.003
正文
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1 引言
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南美洲南部大陆架水域共有14种南美南极鱼[1]，其中位于巴塔哥尼亚海域的拉氏南美南极鱼（Patagonotothen ramsayi）数量最为丰富[2]。尽管拉氏南美南极鱼具有重要的生物学意义，但对于其生物学的研究仍不够深入。Ekau[2]发现这一物种明显表现出性别的二态性，雄性的臀鳍、腹鳍和身体前部呈深黑色，而雌性则呈棕色至绿色。该文还指出，拉氏南美南极鱼产卵发生在南半球秋季的阿根廷陆架以及春季的布尔德伍德浅滩。Sosiński和Janusz[3]也证实了两个区域产卵时间的差异。Brickle等[4]对拉氏南美南极鱼的年龄与生长进行了研究，发现它是一种生长速度相对较慢的鱼类，最大年龄可达14龄。另外，Hart[5]还提供了一些关于其摄食及捕食者的信息。因拉氏南美南极鱼存在昼夜垂直移动的特性[6]，所以它不仅可以捕食一些胶状及甲壳类的浮游生物，还会捕食部分底栖生物[2, 7]。近期研究还发现，拉氏南美南极鱼偶尔也会捕食一些渔业丢弃物，尤其是鱿鱼[8-9]。与此同时，拉氏南美南极鱼也是许多大型鱼类的摄食对象[7]，甚至被巴布亚企鹅（Pygoscelis papua）所捕食[10]。该过程对于巴塔哥尼亚海域能量由海表至海底间的传递有着重要作用。因此，开展拉氏南美南极鱼摄食生态学的研究可为了解由低营养级至高营养级之间的能量流动以及有机质由上层至中下层的垂直运输过程提供依据。
使用传统的胃含物分析法可以知晓拉氏南美南极鱼的主要摄食对象、摄食的时空差异[6-7, 9, 11]。但该方法只能反映短期的摄食情况，对于食物同化情况以及长期的摄食趋势不能够反映。为了解决这个问题，已有学者提出利用肌肉组织中稳定同位素及脂类（尤其是脂肪酸和脂肪醇）作为食性指标[12-15]。对于脂类来说，食物中的脂肪酸组成会影响肝脏、性腺、肌肉等组织的脂质成分[16]。dos Santos等[17]曾分析不同摄食条件下鳕鱼不同组织内脂肪酸组成情况，Beckmann等[18]则分析了不同喂养条件下，鲨鱼肝脏和肌肉组织中脂肪酸分布情况。因此，若利用脂肪酸作为生物标志物探究拉氏南美南极鱼的摄食生态学，需要了解各组织中脂肪酸的分布情况。目前，关于拉氏南美南极鱼脂肪酸组成的研究仅见于肌肉组织，且仅对其营养价值进行了评估[19]，尚未有人将其脂肪酸组成应用于摄食生态学研究。鉴于此，本研究旨在：（1）描述拉氏南美南极鱼3种组织中脂肪酸分布情况，（2）分析3种组织中的脂肪酸是否能表征其摄食情况，以及（3）研究拉氏南美南极鱼的食性及其能量转移情况。
2 方法
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2.1 样品收集
拉氏南美南极鱼样本由中国水产有限公司“龙腾”号于西南大西洋水域进行中层拖网作业时所收集，采样深度范围为20～160 m（即阿根廷巴塔哥尼亚大陆架），取样时间为2014年12月至2015年4月，取样站点如图1所示。所有海上随机采集的样本立即放入−20℃冷冻条件下保存，运回国内实验室后放入−80℃的超低温冰箱内保存，以便进行脂类分析。
2.2 方法
2.2.1 脂肪酸提取及测定
在实验室中，将样本解冻后首先测定其体长和体质量等，其中体长测量使用精度为1 mm的钢尺，体质量测定使用精度为1 g的电子秤；测量完基础生物学后，取样本背鳍附近肌肉、肝脏及性腺部分，用超纯水冲洗干净后装入离心管，然后放入冷冻干燥机（Christ Alpha 1−4），在−55°C下冷冻干燥24 h，所有样品使用混合型球磨仪（Retsch - MM440）粉碎，装入离心管待测。根据Folch法进行粗酯的提取[20]。依据GB/T 17376—2008《动植物油脂脂肪酸甲酯制备》三氟化硼法进行脂肪酸甲酯化。以37种脂肪酸混标及内标十九烷酸甲酯标品作为标准，利用气相色谱−质谱联用仪（Agilent，GC-MS 7890B）进行测试。
2.2.2 指标的测定与计算
以37种脂肪酸混标及内标十九烷酸甲酯标品作为标准，通过比对保留时间对脂肪酸进行定性分析，采用内标法对脂肪酸进行定量分析。脂肪酸含量使用以下内标法公式得出其质量分数，并用单位干质量的拉氏南美南极鱼肌肉组织中每种脂肪酸的含量（单位：mg/g）表示，即
${X_{i}} = {m_{\rm{s}}} \times {A_{i}} \times {f_{{\rm{s}},{i}}}/\left( {m \times {A_{\rm{s}}}} \right),$
式中，Xi为待测样品中组分i的含量，单位为mg/g；ms为待测样品中加入内标物的质量，单位为mg；Ai为组分i的峰面积，无单位；$ {f_{{\rm{s}},{i}}}$为组分i与内标物的校正因子之比，称为相对校正因子，无单位；m为样品的质量，单位为g；As为待测样品中内标物的峰面积，无单位。
2.2.3 指示食性的特征脂肪酸
特征脂肪酸可以指示生物食性，结合已有研究（表1），分析不同组织中脂肪酸组成与食性关系。
2.3 数据处理
首先利用Shapiro-Wilk检验分析脂肪酸含量数据是否符合正态分布，若不符合，则将数据转换成对数形式后再进行检验，以使得数据符合正态分布。利用配对t检验分析两两脂肪酸含量或类型之间是否存在显著性差异（显著性水平设为0.05）。使用主成分分析（Principal Components Analysis，PCA）以研究3种组织间脂肪酸特征的变化，并鉴定对这种变异影响最大的脂肪酸类型。所有平均值均以平均值±标准差表示。PCA及统计分析均采用R程序（R Core Team 2017）处理。
3 结果
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3.1 拉氏南美南极鱼生物学信息
拉氏南美南极鱼体长范围为111.0～288.0 mm，平均体长为（209.7±40.9）mm；体质量范围为20.8～451.3 g，平均体质量为（177.12±115.90）g。据Sosiński和Janusz[3]的研究，拉氏南美南极鱼于体长为240 mm时达到性成熟。据此，本研究将样品体长分为2组：100 mm≤L<240 mm属小体长组，240 mm≤L归大体长组（表2）。
3.2 3种组织中脂肪酸组成
由西南大西洋拉氏南美南极鱼3种组织中共检测出27种脂肪酸，饱和脂肪酸（Saturated Fatty Acid，SFA）包括10种，单不饱和脂肪酸（Monounsaturated Fatty Acid, MUFA）和多不饱和脂肪酸（Polyunsaturated Fatty Acid, PUFA）分别含7种和10种（表3）。对于3个组织而言，肌肉组织中脂肪酸含量最低为（95.39±29.14）mg/g，显著低于肝脏（（423.89±287.16）mg/g）和性腺（（372.48±302.71）mg/g）组织（配对t检验：肌肉组织与肝脏组织，t=−6.99，p<0.001；肌肉组织与性腺组织，t=−5.62，p<0.01）。对于SFA，肝脏组织与性腺组织中含量均较高，显著高于肌肉组织（肝脏组织与肌肉组织，t=−8.14，p<0.001；性腺组织与肌肉组织，t=−5.77，p<0.001）。对于MUFA，肝脏组织中含量最高，显著高于性腺组织和肌肉组织（肝脏组织与性腺组织，t=3.54，p<0.001；肝脏组织与肌肉组织，t=6.63，p<0.001）。就PUFA而言，性腺组织中含量最高，显著高于肝脏和肌肉组织（性腺组织与肝脏组织，t=2.56，p<0.05；性腺组织与肌肉组织，t=4.99，p<0.001）。对于n-3 MUFA，性腺组织中含量最高，肝脏组织次之，且均显著高于肌肉组织（性腺组织与肌肉组织，t=−3.24，p=0.001<0.05；肝脏组织与肌肉组织，t=−2.47，p=0.009<0.05）；而在n-6 PUFA中，同样的性腺组织含量最高，肝脏组织次之，且性腺组织含量显著高于肝脏组织；另外，两者含量均显著高于肌肉组织（性腺组织与肝脏组织，t=−3.66，p<0.001；性腺组织与肌肉组织，t=−4.78，p<0.001；肝脏组织与肌肉组织，t=−6.35，p<0.001）。
在C20:5n3与C22:6n3中，性腺组织中含量最高，肝脏组织次之，均显著高于肌肉组织（C20:5n3∶性腺组织与肌肉组织，t=−3.07，p=0.002<0.05；肝脏组织与肌肉组织，t=−3.00，p=0.003<0.05；C22:6n3:性腺组织与肌肉组织，t=−2.03，p=0.025<0.05；肝脏组织与肌肉组织，t=−1.84，p=0.037<0.05）。对于C20:4n6来说，性腺组织中含量最高，肝脏组织次之，且性腺组织显著高于肝脏组织，两者均显著高于肌肉组织（性腺组织与肌肉组织，t=−4.96，p<0.001；肝脏组织与肌肉组织，t=−6.10，p<0.001；性腺组织与肝脏组织，t=−3.87，p<0.001）。
3.3 脂肪酸组成主成分分析
对拉氏南美南极鱼3个组织脂肪酸组成进行主成分分析后发现，主成分1和主成分2分别解释了3个组织脂肪酸组成总变异的71.61%和11.25%（图2）。由因子载荷图可以看出，主成分1维度负向上有C15:0+C17:0、C20:1+C22:1、C18:2n6以及C20:4n6；主成分2维度正向上有C14:0、C16:0、C16:1n7和C18:1n9；负向上有C20:5n3及C22:6n3。
由因子得分图可知，3个组织之间表现出明显的差异。肝脏组织主要分布在主成分1正向及主成分2的正向上，该组织中C14:0、C16:0、C16:1n7和C18:1n9这4种脂肪酸对其影响较大；而性腺组织主要分布于主成分1的负向上，C15:0+C17:0、C18:2n6、C20:4n6及C20:1+C22:1对其脂肪酸组成的贡献较大；肌肉组织主要分布于主成分1的负向和主成分2的负向上，除与性腺组织的脂肪酸具有同源脂肪酸外，C20:5n3和C22:6n3对其的贡献也较大。
3.4 体长与特征脂肪酸含量的关系
表4显示了拉氏南美南极鱼3种组织脂肪酸含量与体长之间的相关性。由表4可见，肌肉组织的特征脂肪酸中，有3种脂肪酸含量与体长之间存在着明显的线性关系，分别为C15:0+C17:0、C16:0及C20:1+C22:1，且随着体长的增加而增加；就肝脏组织而言，有3种特征脂肪酸含量与体长之间存在着显著的线性关系，分别为C15:0+C17:0、C18:2n6、C20:4n6，随着体长的增加而降低。性腺组织中，有4种特征脂肪酸与体长之间存在显著的线性关系，分别为C15:0+C17:0、C18:2n6、C20:1+C22:1以及C20:4n6，同样随着体长的增加而降低。
4 讨论
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4.1 3种组织对脂肪酸的吸收与分配
鱼类在其不同生活史阶段可能会转变食性，且随着鱼类的生长，身体各组织部位对脂肪酸的吸收以及利用均有所不同。在鱼类未达性成熟之前，鱼类由外界食物来源获取的能量（脂肪酸）用于其身体及各组织的生长；而当鱼类达到性成熟之后，性腺部位获取的脂肪酸在繁殖阶段则更多地用于繁殖。基于Ekau[2]的研究，本研究中的拉氏南美南极鱼样本均未达到性成熟。因此，本文中仅由3种组织的能量分配用于生长这个角度进行分析。
脂肪酸是鱼体的重要组成成分。有研究表明，肝脏是脂肪合成与转化的主要场所，肝脏中有丰富的脂肪积累，可为性腺、肌肉等组织的发育提供能量[27-29]。拉氏南美南极鱼3种组织中，肝脏中总脂肪酸含量为（423.89±287.16）mg/g，远高于性腺中的（372.48±302.71）mg/g及肌肉的（95.39±29.14）mg/g。由此可见，拉氏南美南极鱼更倾向于在肝脏中存储及合成脂类，之后通过血液、淋巴系统等将脂肪酸转移到其他组织[21]。此外，肝脏中SFA和MUFA含量均为3个组织中最高，而PUFA含量却相对较低，这可能由于食物中的PUFA被优先输送到性腺和肌肉等组织中，以供应生长和发育的需求。类似的结果在贝类中也得到证实[22]，这种差异的存在也因各组织具有的不同生理功能所致。性腺的PUFA含量显著高于肝脏和肌肉，这是因为PUFA在卵巢中选择性地积累规律所致，丰富的PUFA可以为卵子和胚胎的发育提供相应的营养，如形成细胞膜结构、调节细胞膜的离子通道和促进发育等[30]；其中n-3 PUFA和n-6 PUFA最为重要，特别是C22:6n3（DHA）、C20:5n3（EPA）和C20:4n6（ARA）。DHA和EPA是海水鱼类不能完全依靠自身合成以满足其生长发育所需的长链不饱和脂肪酸，必须由食物中获取，ARA在某些海水鱼类的正常发育中也起着重要作用[31]。由本文结果来看，拉氏南美南极鱼3个组织中DHA、EPA和ARA的含量均以性腺最高，肝脏次之，肌肉最低。这表明拉氏南美南极鱼性腺组织在生长过程中选择性地储存这些脂肪酸。由于本研究所使用的样本体长多低于27.5 cm，未达到性成熟体长[2, 32]。所以，可推测现阶段对于脂肪酸的积累为后期性腺发育储备了条件，如ARA可以提升精子的生存能力[33]，DHA和EPA可保障幼体的视觉系统发育等[34-35]。
4.2 哪种组织脂肪酸组成最适合用于指征食性？
Jeffries[36]发现，某一海区生态系统中脂肪酸组成特征由该海区存在的浮游植物所决定，而每一个门类的浮游植物均由其特征脂肪酸指示。于此，本研究总结了拉氏南美南极鱼体内脂肪酸潜在的食物来源（表1）。
对于肝脏组织来说，在两个体长组的样品中，C14:0、C16:0、C16:1n7及C18:1n9这4种脂肪酸含量均较高，而PUFA并未体现出明显特征，这与肝脏本身的功能有关，其通常储存大量的SFA和MUFA，而PUFA优先供给其他组织。此结果似乎暗示拉氏南美南极鱼可能会大量摄食硅藻、甲藻以及褐藻等，然而对比已有研究[2, 7, 9]可知，其胃含物中并未发现这些藻类的存在。另外，C15:0+C17:0、C18:2n6、C20:4n6的含量随体长增加而降低，暗示随着拉氏南美南极鱼的生长，其对于浮游细菌、海藻的摄食减少，且底栖摄食特性减弱。而基于已有胃含物分析[2]发现，相对于小体长，大体长鱼类的摄食范围较广，且更多地摄食底栖鱼类。Rossi等[37]曾指出，DHA/EPA的比值越高，其指征的营养级越高，在两个体长组中比值分别为0.59±0.20及0.47±0.11，说明大体长拉氏南美南极鱼呈现出较低的营养级水平。此外，本研究水域处于西南大西洋重要的鱿鱼渔场。已有多项研究显示，拉氏南美南极鱼胃中发现了渔业丢弃物的存在，如鱿鱼和各种经济鱼种等[2, 7-9]。综合这些证据可知，该海域的拉氏南美南极鱼应处于一个较高的营养级水平。据此，以肝脏组织中脂肪酸组成指示其食性会存在一定的误差。
在两个体长组样品的性腺组织中，C15:0+C17:0、C18:2n6、C20:1+C22:1及C20:4n6的贡献率均较大，表现出大量摄食浮游细菌、陆源有机质以及浮游动物的情况，同时也表现出底栖性摄食特征。考虑到本研究的样本基本未达性成熟，性腺内储存的脂肪酸大部分用于正常生长。但性腺组织中DHA与EPA含量较低，且小体长组中DHA/EPA为0.62±0.20，大体长组中为0.46±0.15，对于食性的指征体现出大体长的拉氏南美南极鱼营养级低于小体长鱼类，这与胃含物分析结果存在偏差。考虑到处于未成熟阶段的个体性腺组织还未发育，对于DHA和EPA并未进行选择性储存。此外，性腺组织在不同的发育阶段对于不同类型脂肪酸的需求程度有异[16]。因此，利用性腺组织对其食性进行指征也会出现一定的差异。
综上分析，肝脏和性腺组织呈现的结果稍显偏差，这与各组织对于脂肪酸的储存及使用情况有关。对于拉氏南美南极鱼来说，肌肉组织更新时间长，能够反映较长时间，甚至一生对食物中脂肪酸的吸收情况。
4.3 拉氏南美南极鱼在生长过程中如何转换食性？
通过胃含物分析发现，拉氏南美南极鱼整个生活史过程中存在着食性的转换[2, 6-7, 9]。在肌肉组织中，DHA与EPA的贡献率较高，且小、大体长组DHA/EPA值分别为0.35±0.16和0.36±0.13，说明大体长的拉氏南美南极鱼体现出稍高的营养级[37]。此外，Budge和Parrish[38]指出，在捕食者中只有植食性桡足类能够通过其特有的途径将C18:1和C20:1通过碳链延长大量合成C20:1和C22:1。因此，C20:1和C22:1可作为桡足类摄食其他浮游植物的指示。Falk-Petersen等[26]提出，这两种脂肪酸也可以作为高营养级捕食者摄食植食性桡足类的指示。在本研究中，大体长拉氏南美南极鱼体内C20:1+C22:1含量高于小体长鱼类，且随着体长的增加，其含量呈显著性增加趋势。由此推测，随着拉氏南美南极鱼的生长，其摄食以植食性桡足类为食的生物的程度有所增加。就C18:2n6来说，曾有研究提出其为一种典型的陆源脂肪酸[25]，其在肌肉组织脂肪酸中贡献率较大。这表明，拉氏南美南极鱼在巴塔哥尼亚陆架区对于陆源有机物或以其为食物的生物有一定的摄食。考虑到拉氏南美南极鱼在近岸和大洋水域均有分布，而本研究区域位于陆架区。因此，这种陆源有机物可能由东福克兰流由福克兰群岛或阿根廷漂流由阿根廷近岸携带至本研究海区[34]，也有可能是本研究区域的拉氏南美南极鱼在近岸水域摄食，然后洄游至陆架区所致[39]。此外，大体长拉氏南美南极鱼体内C18:2n6含量高于小体长鱼类，说明大体长鱼类洄游能力高于小体长鱼类，摄食范围也随着增加。饱和脂肪酸中，C15:0和C17:0可用来指示食物中细菌的贡献[22]，肌肉组织中C15:0+C17:0也表现出明显的特征，其大体长拉氏南美南极鱼体内含量较高，由此推测拉氏南美南极鱼在摄食浮游生物过程中可能顺带摄入浮游细菌。总的来讲，肌肉组织脂肪酸表现出的特征表明拉氏南美南极鱼呈现出浮游生物食性[40]。另一方面，肌肉组织中C20:4n6含量较高，贡献率较大，是能够指征底栖生物食性的脂肪酸[26]。由此可见，拉氏南美南极鱼能够较易地在浮游及底栖生活之间转换。该结果与Kock[41]的分析一致。这也是为什么拉氏南美南极鱼肌肉中δ15N和δ13C含量与体长之间无线性关系存在的原因[39]。针对PUFA，主要由n-3系列和n-6系列脂肪酸组成，其中n-3系列脂肪酸可以指示浮游植物通过捕食食物链向更高营养级捕食者传递[42]，而n-6系列脂肪酸中，C20:4n6所占比重最高。所以，n-6 PUFA/n-3 PUFA的比值被用来估计腐生食物链和捕食食物链对于高营养级捕食者的贡献程度[42]。在拉氏南美南极鱼体内，n-6 PUFA/n-3 PUFA的比值与体长之间并无明显的线性关系。这可能是因为大、小个体拉氏南美南极鱼胃含物中均发现了渔业丢弃物，从而导致腐生食物链和捕食食物链的贡献率特征弱化。此外，该现象也可能是因为随着生长，其在浮游与底栖环境之间频繁转换所致。
在本研究中，随着拉氏南美南极鱼的生长，其对于浮游细菌、浮游动物以及底栖鱼类的摄食均会增加，表明其摄食特性由浮游性摄食方式转变为浮游–底栖性摄食，且增强对于陆源有机质的摄食，推测其洄游能力有所增强。但由于渔业丢弃物的影响，导致其营养级水平升高，腐生食物链和捕食食物链的贡献率特征弱化。从这个角度来看，人类活动，尤其是渔业活动，可能会影响或干扰大洋性生态系统结构。
致谢：本研究所使用样品均为中国水产有限公司所属渔船“龙腾”号船长及全体船员协助收集，实验过程在上海海洋大学极地研究中心和大洋渔业资源可持续开发教育部重点实验室完成，在此深表谢忱！
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  • 国家自然科学基金（41776185）；国家重点研发计划（2018YFC1406801）。
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2020年第42卷第4期
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doi: 10.3969/j.issn.0253-4193.2020.04.003
  • 接收时间：2019-06-06
  • 首发时间：2026-03-26
  • 出版时间：2020-04-25
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  • 收稿日期：2019-06-06
  • 修回日期：2019-09-08
基金
国家自然科学基金（41776185）；国家重点研发计划（2018YFC1406801）。
作者信息
    1 上海海洋大学 海洋科学学院，上海 201306
    2 上海海洋大学 极地研究中心，上海 201306
    3 大洋渔业资源可持续开发教育部重点实验室 极地海洋生态系统研究室，上海 201306
    4 国家远洋渔业工程技术研究中心，上海 201306

通讯作者:

*朱国平（1976-），男，教授，从事极地生物学与渔业生态学方面的研究。E-mail：
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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