Evolution of the Marine Biological Carbon Pump and geological events in the Ross Sea during Modern and the Last Glacial Maximum Period — Stratigraphic Records of Molecular Biomarkers
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摘要: 海洋生物碳泵（BCP）与古气候的关系是碳循环研究的一个核心科学问题。 本文利用罗斯海沉积物总有机碳（TOC）和一些重要的类脂化合物所隐含的生态学特性，即那些可引起海底沉积碳库组成和海洋BCP（包括生物泵BP和微型生物碳泵MCP）效率改变的分子生物标志物，从罗斯海现代海洋出发，追溯到末次冰期以来（27 ka BP～0.6 ka BP）古海洋生物碳泵演化（重点讨论BP和MCP）及其与一些重大地质事件的联系。研究结果表明；（1）罗斯海表层沉积物中相对高的TOC和较低C/N比值、以及正构烷烃色谱峰型、主峰碳、分子组合特征低碳数和高碳数比值（L/H）和(nC21+nC22)/(nC28+nC29)比值、低碳烃（nC15+nC17+nC19）、中碳烃（nC21+nC23+nC25）和高碳烃（nC27+nC29+nC31）、细菌（BrGDGT）和较低的陆源土壤指标（BIT）一致性表征海源有机质贡献占主导地位，通过海洋BP和MCP效应将内源有机碳输送到海底进行长期储存；Pr/Ph比值结合Ph/nC18和Pr/C17比值均表明表层沉积物为还原—强还原性的缺氧环境，有利于海底储存有机碳，同时还与海冰消融关系密切。（2）利用低温校正公式
$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ -SST（2式）反演现代的海洋表层温度（SST），并与WOA-SST（3月)表层水体温度较接近（R2=0.78、P<0.01，n=15）；同时利用$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ -SST-2重建罗斯海JB03岩芯末次冰期以来的古海洋温度在−0.74～2.86℃范围（平均为1.03℃），接近现代南极罗斯海夏季温度。（3）JB03岩芯记录年代为27.27～0.6 ka BP，分为末次冰盛期（27.3～21 ka BP）、末次冰消期（21～11.7 ka BP）和全新世（11.7～0.6 ka BP）三个地质历史时期。末次冰期古海洋受冰盖和海冰限制作用的影响，初级生产力低下、沉积速率仅为0.45 cm/ka、Pr/Ph比值、Ph/nC18和Pr/nC17反映沉积环境氧化性较强、不利于MCP和BP储碳，在寒冷气候时段碳储量潜力降低；进入全新世暖期，冰架退缩解体，温暖的气候条件有利于浮游植物生产力和硅藻增加、浮游动物增加及影响粪便物质组成和提高沉降速率，促进有机碳向深海的输送加快，海底沉积为弱还原—弱氧化环境，有利于碳保存；硅藻生物量提高就意味着硅质泵加强，进而微生物活性增强、促进古菌和细菌生长，因而微生物总量-GDGTs、产甲烷古菌或广古菌-GDGT-0、奇古菌-Crenarchaeol生物量大大提高，显示罗斯海全新世以来古海洋BP和MCP作用大大加强，且古海洋BP与现代海洋均以硅藻/硅质泵为绝对主导。研究还发现，罗斯海末次冰期和全新世以来的古海洋BP和MCP储碳效率变化均与古海洋地质事件，即全球性大尺度的气候变化有关，这个碳库的大小与气候冷暖之间存在对应关系，罗斯海古海洋调节大气CO2的能力——尤其在全新世暖期最强，这对于认识全球气候变化的海洋调控机制具有重要的科学意义。Abstract: The relationship between the marine biological carbon pump(BCP) and paleoclimate is a central scientific issue in carbon cycle research. This study utilizes the ecological characteristics implied by total organic carbon (TOC) and key lipid compounds in Ross Sea sediments, specifically molecular biomarkers that can influence the composition of the seabed carbon reservoir and the efficiency of the marine BCP (including the biological pump BP and the microbial carbon pump MCP), tracing the evolution of the ancient marine biological carbon pump (with a focus on BP and MCP) since the Last Glacial Maximum (27ka BP～0.6ka BP) and its connection to significant geological events. The research findings indicate: (1) the relatively high TOC and low C/N ratios in surface sediments of the Ross Sea, along with characteristics such as normal alkane chromatographic peak shapes, dominant carbon, molecular composition features with low to high carbon number ratios (L/H) and(nC21+nC22)/(nC28+nC29)ratios, low n-alkanes(nC15+nC17+nC19), mid-chain alkanes(nC21+nC23+nC25), high-chain alkanes (nC27+nC29+nC31), bacterial (BrGDGT), and lower terrigenous input index (BIT), collectively indicate a predominance of marine-derived organic matter contributing to the seabed through the effects of the marine BP and MCP, facilitating long-term carbon storage. The Pr/Ph ratio combined with Ph/nC18 and Ph/nC17 ratios suggest that surface sediments represent a reducing to strongly reducing anoxic environment conducive to organic carbon sequestration on the seafloor, closely linked to sea ice melt. (2) By employing a low-temperature correction formula$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ -SST (Equation 2) to infer modern sea surface temperatures (SST) in the ocean and comparing them to WOA-SST (March) surface water temperatures, a close match is observed (R2=0.78, P<0.01, n=15). Additionally, using$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ -SST-2 to reconstruct paleoceanic temperatures in the Ross Sea JB03 core since the Last Glacial Maximum yields a range of −0.74 to 2.86°C (average 1.03°C), approximating modern summer temperatures in the Antarctic Ross Sea. (3) The JB03 core record spans 27.27 to 0.6 ka BP, divided into three geological periods: the Last Glacial Maximum (27.3～21 ka BP), the deglaciation period (21～11.7 ka BP), and the Holocene (11.7～0.6 ka BP). During the Last Glacial Maximum, the ancient ocean was influenced by ice cover and sea ice constraints, resulting in reduced primary productivity, sedimentation rates of only 0.45 cm/ka, and indicators such as Pr/Ph ratios, Ph/nC18, and Ph/nC17 reflecting a strongly oxidative sedimentary environment unfavorable for MCP and BP carbon sequestration, leading to diminished carbon storage potential during cold climate periods. As the Holocene warm period commenced, ice shelves retreated and disintegrated, creating favorable warm climate conditions for increased phytoplankton productivity, diatom abundance, zooplankton proliferation, alterations in fecal matter composition, and enhanced sedimentation rates, accelerating the transfer of organic carbon to the deep sea. Seabed sediments transitioned to weakly reducing to weakly oxidizing conditions, conducive to carbon preservation. Elevated diatom biomass signifies strengthened siliceous pump activity, subsequently enhancing microbial activity, promoting the growth of archaea and bacteria, leading to significant increases in total microbial-GDGTs, methane-producing archaea or widespread archaea-GDGT-0, and crenarchaeol archaeal biomass, indicating a substantial enhancement in ancient marine BP and MCP activity in the Ross Sea since the Holocene, with diatoms/siliceous pumps dominating both ancient and modern marine environments. The study further reveals that changes in carbon sequestration efficiency of the ancient marine BP and MCP in the Ross Sea during the Last Glacial Maximum and the Holocene are linked to geological events, specifically large-scale global climate variations, where the size of this carbon reservoir corresponds to climatic fluctuations. The Ross Sea’s ability to regulate atmospheric CO2 through ancient marine processes, particularly during the Holocene warm period, holds significant scientific implications for understanding the oceanic regulatory mechanisms of global climate change.-
Key words:
- Marine primary productivity /
- Biological pump /
- Microbial carbon pump /
- Molecular biomarkers /
- TEXL86.
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图 1 罗斯海表层沉积物和岩芯采样站位
箭头代表流向，灰色为表层环流，蓝色为变性绕极深层水（MCDW）涌入陆架
Fig. 1 Ross Sea surface sediment and core sampling stations
Arrows represent flow direction, Grey represent the surface gyre, and Blue represent the modified Circumpolar Deep Water (MCDW) spilling onto the shelf
图 2 表层沉积物（a）正构烷烃（RB08B，JB03和RB06B）和（b）甾醇(JB01)气相及（c）GDGTs（JB01）液相色谱图
Fig. 2 Gas (a) and liquid (c) chromatograms of the surface sediments for (a) n-alkanes (08B, JB03, and RB06B) and (b) sterols (JB01) and (c) GDGTs (01)
图 3 (a)初级生产力-TOC(%)，（b）总有机氮-TN(%)，（c）T/N比值
Fig. 3 (a) Primary Productivity-TOC(%), (b) Total Organic Nitrogen-TN(%), (c) T/N Ratio
图 4 (a)TOC(%)与TN(%)和(b) TOC(%)与黏土含量相关性
Fig. 4 (a) TOC(%) vs TN(%) and (b) TOC(%) vs Clay Content Correlation
图 5 现代表层沉积物中TOC和正构烷烃总量及其分子组合特征与海洋生物泵联系
（a）正构烷烃总量（∑n-ALK）ng/g，（b）菌藻类（nC15+nC17+nC19）ng/g，（c）大型浮游植物（nC21+nC23+nC25）ng/g，（d）陆源高等植物（nC27+nC29+nC31）ng/g，（e）姥鲛烷/植烷（Pr/Ph），（f）Ph/nC17值，（g）Pr/nC18值，（h）L/H值，（i）（nC21+nC22）/（nC28+nC29）值，（j）CPI值 ，（k）OEP值
Fig. 5 Comparison of TOC and total normal alkane and its molecular composition characteristics in modern sediments with marine biological pump
(a) normal alkane (∑n-ALK) ng/g, (b) Fungi algae (nC15+ nC17 +nC19）ng/g, (c) Large phytoplankton (nC21+nC23+nC25) ng/g, (d)Terrestrial higher plants (nC27+nC29+nC31) ng/g, (e) Cetane/phytane (Pr/Ph, (f) Ph/nC17, (g) Pr/nC18, (h) L/H, (i) (nC21+nC22)/(nC28+nC29) , (j) CPI, (k) OEP
图 6 现代表层沉积物微生物细胞膜脂GDGTs、不同类群的古菌及其与MCP储碳联系
（a）微生物细胞膜脂-GDGTs、（b）古菌-IsoGDGTs、（c）细菌- BrGDGTs、（d）产甲烷古菌- GDGT-0、（e）奇古菌-Crenarchaeol、（f）R0/5值和（g）BIT值
Fig. 6 Modern surface sedimentary microbial cell membrane GDGTs, archaeal groups and their link to MCP carbon storage
(a) Microbial cell membrane lipids-GDGTs, (b) Archaea-IsoGDGTs, (c) Bacteria-BrGDGTs, (d) Methanogenic archaea-GDGT-0, (e) Thaumarchaeota-Crenarchaeol, (f) R0/5 values and (g) BIT values
图 7 （a）罗斯海WOA 3月和（b）WOA夏季表层温度与
$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ —SST的线性关系Fig. 7 Linear relationship between (a) Ross Sea WOA March and (b) WOA summer surface temperature vs
$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ —SST
图 8 现代表层沉积物的浮游动植物分子生物标志物含量平面分布图
A-（a）菜籽甾醇-brassicasterol（硅藻）、（b）甲藻甾醇—dinostero（甲藻）、（c）长链烯酮-Long-chain alkenones（颗石藻）、（d）胆甾醇-Chlesterol(浮游动物总量)、（e）SUM-初级生产力，（f）菜籽甾醇-硅藻占比（bra/sum）、（g）硅藻/甲藻比值（bra/dino）
Fig. 8 Molecular biomarker contents of planktonic algae in modern surface sediments
A- (a) brassasterol (diatoms), (b) dinosterol (dinoflagellates), (c) long-chain alkenones (ccolithophores), (d) cholestanol (total zooplankton), (e) SUM-primary productivity, (f brassicasterol-silica ratio (bra/sum), (g) diatom/dinoflagellate ratio (bra/dino)
图 9 硅藻和甲藻占比与重建表层海温SST的线性关性
B-（h）菜籽甾醇/总量（bra/sum）和B-（i）菜籽甾醇/甲藻甾醇（bra/dino）与SST/℃的相关性
Fig. 9 The linear correlation between the proportions of diatoms and dinoflagellates and the reconstructed SST
B- (h) The correlation of brassicasterol/sum (bra/sum) and B- (i) brassicasterol/dinosterol (bradino) with SST/℃
图 10 AMS14C测年结果和校正后年龄及其沉积速率
Fig. 10 AMS14C dating results and calibrated ages and their depositional rates
图 11 JB03岩芯TOC/TN值和正构烷烃分子组合特征记录的末次冰期来古海洋陆源/海源碳库变化
（a）总有机碳TOC和TOC/TN、（b）低碳烃/高碳烃L/H、（c）TARHC、（d）(nC21+nC22)/(nC28+nC29)、(e)菌藻类（nC15+nC17+nC19）、（f）大型浮游植物（nC21+nC23+nC25）、（g）陆源高等植物（nC27+nC29+nC31）、（h）Pr/Ph、（i）CPI和（J）OEP值
Fig. 11 Last Glacial Marine/Terrestrial Carbon Isotopic Signature Recorded by TOC/TN Ratio and Molecular Composition n-Alkanes from JB03 Core
(a) TOC and TOC/TN, (b) L/H, (c)TARHC, (d) (nC21+nC22)/(nC28+nC29) (e) Fungi algae (nC15+ nC17 +nC19）, (f) Large planktonic plants (nC21+nC23+nC25), (g) Terrestrial higher plants (nC27+nC29+nC31) , (h) Pr/Ph, (i）CPI and (j) OEP values
图 13 JB03沉积地层
$ {\mathrm{T}\mathrm{E}\mathrm{X}}_{86}^{\mathrm{L}} $ 指标计算的SST-1和SST-2Fig. 13 Calculation of JB03 sedimentary stratigraphic index SST-1 and SST-2
图 12 JB03岩芯微生物细胞膜脂GDGTs记录的古海洋微型生物/MCP地质演化
（a）WDC δ18O、（b）含水率和TOC、（c）微生物细胞膜脂-GDGTs、（d）古菌-IsoGDGTs、（e）细菌- BrGDGTs、（f）产甲烷古菌- GDGT-0、（g）奇古菌-Crenarchaeol、（h）R0/5比值、（i）菜籽甾醇- Brassicasterol（硅藻）、（j）甲藻甾醇- Dinosterol（甲藻）、（k）C37长链烯酮- C37-ketene（颗石藻）、（l）总量SUM-初级生产力、（m）胆甾醇-浮游动物总量
Fig. 12 Geological evolution of the microbial/MCP paleocean recorded by microbial cell membrane lipids GDGTs of JB03 core
(a) WDC δ18O, (b) water content and TOC, (c) microbial cell membrane lipids-GDGTs, (d) archaeal-IsoGDGTs, (e) bacterial- BrGDGTs, (f) Methanogen GDGT-0, (g) Thaumarchaeota-Crenarchaeol, (h) R0/5 ratio, (i) brassicol (diatoms), (j) dinosterol (dinoflagellates), (k) C37 long-chain alken-C37-ketene (coccolithophores), (l) Total SUM-primary production, (m) Cholesterol-Total zoplankton
图 14 分子生物标志重建的浮游植物相对生物量与SST-2和的相关性
（a）菜籽甾醇-硅藻（ng/g）与SST-2/℃、（b）brassicasterol/Sum（%）与SST-2/℃、（c）dinosterol/Sum（%）与SST-2/℃、（d）alkenone/Sum（%）与SST-2/℃的相关性
Fig. 14 Correlation between the relative biomass of phytoplankton with molecular biomarker reconstruction and SST-2
(a) brassicasterol-siliceous (ng/g) with SST-2/℃, (b) brassicasterol/ (%) with SST-2/℃, (c) dinosterol/Sum (%) with SST-2/℃, (d) alken/Sum (%) with SST-2/℃
表 1 南极罗斯海JB03柱样的AMS14C年龄及校正年龄与沉积速率
Tab. 1 AMS14C ages and calibrated ages with sedimentation rates of JB03 core from Ross sea of Southern Ocean
深度 14C年龄 化石碳年龄 海洋储层年龄 日历年龄 沉积速率
地质年代cm a BP a a a BP cm/ka 0-2 4470 ±303045 825 600
全新世2-4 4774 ±303045 825 904 6.58 18-20 6950 ±303045 825 3080 7.35 54-56 12655 ±303045 825 8785 6.31 68-70 14730 ±303045 825 10860 6.75 72-74 15720 ±303045 825 11850 4.04 末次冰消期 78-80 28950 ±303045 825 25080 0.45
末次盛冰期102-104 31010 ±303045 825 27140 11.7 110-112 31090 ±303045 825 27220 100 112-114 31095 ±303045 825 27225 400 128-130 31140 ±303045 825 27270 356 
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