Exploring biomineralization in the tropical western Pacific sediments based on phospholipid fatty acid analysis

doi:10.13745/j.esf.sf.2022.1.3

Abstract

Abstract:

Identification of marine sediment mineralization processes plays an important role in the study of biogeochemical cycling of biogenic elements, although such processes—including aerobic respiration, nitrate reduction, iron and manganese reduction, sulfate reduction, etc. —can be difficult to distinguish one from another. In this study, gas chromatography-mass spectrometry (GC-MS) was used to analyze the phospholipid fatty acid (PLFA) profiles and their influencing factors in different mineralization processes, and further explore the use of PLFAs as sediment mineralization indicator in the tropical western Pacific. The total PLFA content was found to be highest in aerobic respiration and lowest in nitrate reduction. The obvious decrease of the sum of 14:0, i14:0, i15:0 and i19:0—the main components of microbial PLFA in aerobic respiration—could have certain indication for the occurrence of nitrate reduction; whilst the decrease of the sum of 10:0, 17:0, 20:0 and 22:0 could indicate the occurrence of sulfate reduction. In the sediments of the tropical western Pacific, the total organic carbon (TOC), total organic nitrogen (TON) and NO₃-N content in porewater were important factors influencing PLFA. as the total PLFA content decreased with decreasing TOC and TON, whilst the degradation of TOC and TON promoted the degradation of PLFA, which had a more direct impact on the PLFA composition.

Key words: phospholipid fatty acid, mineralization, indicative function, marine sediment, tropical western Pacific

CLC Number:

P736.2
P593

YANG Ziyang, REN Denglong, HE Zhipeng, LI Xuegang, SONG Jinming, YUAN Huamao, DUAN Liqin, LI Ning, ZHANG Qian. Exploring biomineralization in the tropical western Pacific sediments based on phospholipid fatty acid analysis[J]. Earth Science Frontiers, 2022, 29(4): 93-102.

Figures/Tables 12

Fig.1 Location of the sampling station in the tropical western Pacific

Table 1 Detection limits, RSD and characteristic ions of fatty acid methyl esters

脂肪酸种类	检出限/(μg·mL^-1)	RSD/%	特征离子
8:0	0.032	2%	74 115 158
10:0	0.027	2%	74 143 186
12:0	0.023	2%	74 171 214
14:0	0.034	2%	74 199 242
i14:0	0.030	4%	74 211 242
15:0	0.044	3%	74 213 256
i15:0	0.033	4%	74 213 256
a15:0	0.032	4%	74 199 256
16:0	0.022	1%	74 227 270
i16:0	0.028	4%	74 227 270
17:0	0.031	2%	74 284 241
i17:0	0.033	4%	74 241 284
17:1ω7	0.039	3%	55 250 282
18:0	0.068	5%	74 298 255
i19:0	0.035	4%	143 269 312
20:0	0.053	4%	74 326 283
i20:1	0.035	4%	55 292 324
24:0	0.110	7%	74 382 339
13:0	0.022	1%	74 185 228
14:1	0.028	2%	55 208 240
16:1ω7	0.033	2%	55 236 268
i16:1	0.034	4%	55 194 236
18:1	0.071	5%	55 264 296
18:2	0.087	6%	67 294 263
20:3	0.077	5%	79 320 289
20:5	0.250	17%	79 287 316
22:1	0.080	5%	55 320 352
11:0	0.024	2%	74 157 200
20:4	0.093	6%	79 287 318

Fig.2 Chromatogram of fatty acid methyl esters

Table 2 Corresponding relationships between microbial groups and PLFA biomarkers [23⇓-25]

微生物	PLFA标志物
细菌	一些饱和或单不饱和脂肪酸:15:0,16:0,17:0,18:1ω5,18:1ω7,i19:0,a19:0
好氧细菌	16:1ω7,i14:0,a14:0,15:0 2OH,15:0 3OH,a15:0,i15:0
厌氧细菌	cy17:0,cy19:0
革兰氏阳性菌	a16:0,i16:0,a17:0,i17:0,i18:0
革兰氏阴性菌	i15:0 3OH,16:1ω9,i17:0 3OH
硫酸盐还原菌	17:1ω6,17:1ω7
放线菌	10Me16:0,10Me17:0,10Me18:0
假单胞菌	18:1ω7
真菌	18:3ω6 (6,9,12),18:1ω9

Fig.3 Vertical changes of sediment TOC, TON, TIN, TOC/TON, pH and δ13C

Fig.4 Vertical changes of nutrient concentration in sediment porewater

Fig.5 Relative percentages of various PLFAs in sediment

Table 3 Content of various PLFAs in the sediment

深度/ cm	总 PLFA	各种PLFA含量/(μg·g^-1)
深度/ cm	总 PLFA	8:0	10:0	12:0	14:0	i14:0	i15:0	15:0	16:0	17:0	17:1ω7	18:0	i19:0	20:0	22:0	24:0	i16:1	i16:0	13:0	16:1ω7	i20:1	14:1	a15:0	i17:0	18:1	20:3	20:5	22:4	20:4	11:0	18:2	22:1
0~<1	0.27	0.002 7	0.001 2	0.001 5	0.011	0.024	0.007 1	0.004 4	0.046	0.005 0	0.013	0.012	0.012	0.003 7	0.005 6	0.007 2	-	0.039	0.001 3	0.038	0.005 0	0.001 4	0.001 7	0.004 0	0.008 0	0.002 3	0.002 7	0.003 2	-	0.000 64	0.002 0	0.004 0
1~<2	0.22	0.002 3	0.001 2	0.001 4	0.007 6	0.019	0.005 4	0.003 8	0.035	0.005 2	0.008 4	0.011	0.009 8	0.003 8	0.005 9	-	0.021	0.031	0.001 4	0.015	0.004 3	0.001 6	0.001 3	0.003 2	0.007 7	0.002 6	0.003 1	0.004 3	-	-	-	-
2~<3	0.20	0.002 3	0.001 1	0.001 2	0.007 1	0.017	0.004 9	0.003 4	0.032	0.004 6	0.007 0	0.009 1	0.003 6	0.003 4	0.005 4	0.007 0	0.018	0.027	0.001 2	0.025	-	0.001 4	0.001 2	0.002 8	0.006 7	0.002 3	0.002 8	0.003 9	0.003 2	-	-	-
3~<4	0.087	0.001 7	0.001 1	0.001 1	0.003 4	0.005 3	0.001 5	0.002 4	0.014	0.004 0	-	0.006 9	0.002 8	0.003 3	0.005 2	0.007 0	0.005 6	0.008 4	0.001 1	0.011	0.001 3	-	-	-	-	-	-	-	-	-	-	-
4~<5	0.028	0.001 4	0.000 89	0.000 89	0.001 6	0.000 54	0.000 17	0.001 7	0.004 8	0.003 3	-	0.004 7	0.000 29	0.002 7	0.004 5	-	-	0.000 59
5~<6	0.028	0.001 5	0.000 91	0.000 91	0.001 5	0.000 39	0.000 13	0.001 7	0.004 6	0.003 4	-	0.004 9	0.000 23	0.002 8	0.004 6	-	-	0.000 44
6~<7	0.032	0.001 4	0.000 86	0.000 86	0.001 5	0.000 38	0.000 11	0.001 6	0.004 5	0.003 1	-	0.004 5	0.000 13	0.002 6	0.004 3	0.005 9	-	0.000 32
7~<8	0.035	0.001 9	0.000 92	0.000 97	0.001 6	0.000 30	8.1E-05	0.001 8	0.004 5	0.003 5	-	0.005 0	0.000 11	0.002 9	0.004 8	0.006 6	-	-
8~<9	0.022	0.001 4	0.000 74	-	0.001 2	0.000 10	2.2E-05	0.001 4	0.003 4	0.002 9	-	0.004 0	2.2E-05	0.002 3	0.003 9	-	-	-
9~<10	0.021	0.001 2	0.000 68	-	0.001 2	0.000 13	3.3E-05	0.001 4	0.003 3	0.002 9	-	0.004 0	4.4E-05	0.002 3	0.003 8	-	-	-
10~<11	0.056	0.004 6	0.002 2	0.001 5	0.002 5	0.000 98	0.000 20	0.002 8	0.008 4	0.005 3	0.007 5	0.008 0	0.000 14	0.004 3	0.007 2	-	0.000 84	-
11~<12	0.039	0.003 0	0.001 3	0.001 4	0.002 1	0.000 27	5.7E-05	0.002 5	0.005 9	0.004 9	-	0.006 8	-	0.004 0	0.006 7	-	-	-
12~<13	0.043	0.002 3	0.001 5	-	0.001 9	0.000 21	5.2E-05	0.002 2	0.005 4	0.004 5	-	0.006 2	-	0.003 7	0.006 1	0.008 4	0.000 14	-
13~<14	0.044	0.002 2	0.001 2	0.001 1	0.001 7	0.000 27	6.0E-05	0.002 0	0.005 2	0.003 9	0.004 1	0.005 6	7.5E-05	0.003 3	0.005 4	0.007 4	-	-
14~<15	0.042	0.002 2	0.001 1	0.001 2	0.001 9	0.000 38	9.8E-05	0.002 2	0.005 5	0.004 2	-	0.006 1	9.8E-05	0.003 5	0.005 8	0.007 9	0.000 18	-
15~<16	0.043	0.002 0	0.001 1	0.001 1	0.001 8	0.000 22	6.2E-05	0.002 0	0.005 4	0.004 0	0.003 4	0.005 8	9.2E-05	0.003 4	0.005 6	0.007 6	-	-
16~<17	0.050	0.002 9	0.001 3	0.001 4	0.002 2	0.000 20	4.0E-05	0.002 6	0.006 3	0.005 1	-	0.007 2	3.9E-05	0.004 2	0.007 0	0.009 6	5.9E-05	-
17~<18	0.043	0.002 5	0.001 6	-	0.002 1	0.000 30	3.8E-05	0.002 5	0.005 8	0.004 9	0.005 7	0.006 8	3.8E-05	0.004 0	0.006 7	-	-	-
18~<19	0.046	0.002 8	0.001 4	0.001 3	0.002 0	0.000 27	5.4E-05	0.002 3	0.006 0	0.004 7	-	0.006 6	-	0.003 8	0.006 4	0.008 7	0.000 13	-
19~≤20	0.028	0.001 7	0.000 80	0.000 78	0.001 2	0.000 14	3.2E-05	0.001 4	0.003 5	0.002 8	-	0.004 1	-	0.002 3	0.003 8	0.005 3	-	-

Table 3 Content of various PLFAs in the sediment

深度/ cm	总 PLFA	各种PLFA含量/(μg·g^-1)
深度/ cm	总 PLFA	8:0	10:0	12:0	14:0	i14:0	i15:0	15:0	16:0	17:0	17:1ω7	18:0	i19:0	20:0	22:0	24:0	i16:1	i16:0	13:0	16:1ω7	i20:1	14:1	a15:0	i17:0	18:1	20:3	20:5	22:4	20:4	11:0	18:2	22:1
0~<1	0.27	0.002 7	0.001 2	0.001 5	0.011	0.024	0.007 1	0.004 4	0.046	0.005 0	0.013	0.012	0.012	0.003 7	0.005 6	0.007 2	-	0.039	0.001 3	0.038	0.005 0	0.001 4	0.001 7	0.004 0	0.008 0	0.002 3	0.002 7	0.003 2	-	0.000 64	0.002 0	0.004 0
1~<2	0.22	0.002 3	0.001 2	0.001 4	0.007 6	0.019	0.005 4	0.003 8	0.035	0.005 2	0.008 4	0.011	0.009 8	0.003 8	0.005 9	-	0.021	0.031	0.001 4	0.015	0.004 3	0.001 6	0.001 3	0.003 2	0.007 7	0.002 6	0.003 1	0.004 3	-	-	-	-
2~<3	0.20	0.002 3	0.001 1	0.001 2	0.007 1	0.017	0.004 9	0.003 4	0.032	0.004 6	0.007 0	0.009 1	0.003 6	0.003 4	0.005 4	0.007 0	0.018	0.027	0.001 2	0.025	-	0.001 4	0.001 2	0.002 8	0.006 7	0.002 3	0.002 8	0.003 9	0.003 2	-	-	-
3~<4	0.087	0.001 7	0.001 1	0.001 1	0.003 4	0.005 3	0.001 5	0.002 4	0.014	0.004 0	-	0.006 9	0.002 8	0.003 3	0.005 2	0.007 0	0.005 6	0.008 4	0.001 1	0.011	0.001 3	-	-	-	-	-	-	-	-	-	-	-
4~<5	0.028	0.001 4	0.000 89	0.000 89	0.001 6	0.000 54	0.000 17	0.001 7	0.004 8	0.003 3	-	0.004 7	0.000 29	0.002 7	0.004 5	-	-	0.000 59
5~<6	0.028	0.001 5	0.000 91	0.000 91	0.001 5	0.000 39	0.000 13	0.001 7	0.004 6	0.003 4	-	0.004 9	0.000 23	0.002 8	0.004 6	-	-	0.000 44
6~<7	0.032	0.001 4	0.000 86	0.000 86	0.001 5	0.000 38	0.000 11	0.001 6	0.004 5	0.003 1	-	0.004 5	0.000 13	0.002 6	0.004 3	0.005 9	-	0.000 32
7~<8	0.035	0.001 9	0.000 92	0.000 97	0.001 6	0.000 30	8.1E-05	0.001 8	0.004 5	0.003 5	-	0.005 0	0.000 11	0.002 9	0.004 8	0.006 6	-	-
8~<9	0.022	0.001 4	0.000 74	-	0.001 2	0.000 10	2.2E-05	0.001 4	0.003 4	0.002 9	-	0.004 0	2.2E-05	0.002 3	0.003 9	-	-	-
9~<10	0.021	0.001 2	0.000 68	-	0.001 2	0.000 13	3.3E-05	0.001 4	0.003 3	0.002 9	-	0.004 0	4.4E-05	0.002 3	0.003 8	-	-	-
10~<11	0.056	0.004 6	0.002 2	0.001 5	0.002 5	0.000 98	0.000 20	0.002 8	0.008 4	0.005 3	0.007 5	0.008 0	0.000 14	0.004 3	0.007 2	-	0.000 84	-
11~<12	0.039	0.003 0	0.001 3	0.001 4	0.002 1	0.000 27	5.7E-05	0.002 5	0.005 9	0.004 9	-	0.006 8	-	0.004 0	0.006 7	-	-	-
12~<13	0.043	0.002 3	0.001 5	-	0.001 9	0.000 21	5.2E-05	0.002 2	0.005 4	0.004 5	-	0.006 2	-	0.003 7	0.006 1	0.008 4	0.000 14	-
13~<14	0.044	0.002 2	0.001 2	0.001 1	0.001 7	0.000 27	6.0E-05	0.002 0	0.005 2	0.003 9	0.004 1	0.005 6	7.5E-05	0.003 3	0.005 4	0.007 4	-	-
14~<15	0.042	0.002 2	0.001 1	0.001 2	0.001 9	0.000 38	9.8E-05	0.002 2	0.005 5	0.004 2	-	0.006 1	9.8E-05	0.003 5	0.005 8	0.007 9	0.000 18	-
15~<16	0.043	0.002 0	0.001 1	0.001 1	0.001 8	0.000 22	6.2E-05	0.002 0	0.005 4	0.004 0	0.003 4	0.005 8	9.2E-05	0.003 4	0.005 6	0.007 6	-	-
16~<17	0.050	0.002 9	0.001 3	0.001 4	0.002 2	0.000 20	4.0E-05	0.002 6	0.006 3	0.005 1	-	0.007 2	3.9E-05	0.004 2	0.007 0	0.009 6	5.9E-05	-
17~<18	0.043	0.002 5	0.001 6	-	0.002 1	0.000 30	3.8E-05	0.002 5	0.005 8	0.004 9	0.005 7	0.006 8	3.8E-05	0.004 0	0.006 7	-	-	-
18~<19	0.046	0.002 8	0.001 4	0.001 3	0.002 0	0.000 27	5.4E-05	0.002 3	0.006 0	0.004 7	-	0.006 6	-	0.003 8	0.006 4	0.008 7	0.000 13	-
19~≤20	0.028	0.001 7	0.000 80	0.000 78	0.001 2	0.000 14	3.2E-05	0.001 4	0.003 5	0.002 8	-	0.004 1	-	0.002 3	0.003 8	0.005 3	-	-

Fig.6 Vertical variations of total PLFA and microbial PLFAs abundances in three types of sediment mineralization processes

Table 4 Sediment mineralization and related chemical reactions. Adapted from [37].

矿化过程	化学反应
有氧呼吸	(CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+138O₂→106CO₂+16HNO₃+H₃PO₄+122H₂O
硝酸盐还原	① (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+94.4HNO₃→H₃PO₄+177.2H₂O+106CO₂+55.2N₂ ② (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+84.8HNO₃→H₃PO₄+148.4H₂O+106CO₂+42.4N₂+16NH₃
铁还原	① (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+212Fe₂O₃+848H⁺→H₃PO₄+530H₂O+106CO₂+16NH₃+424Fe²⁺ ② (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+424FeOOH+848H⁺→424Fe²⁺+106CO₂+16NH₃+H₃PO₄+742H₂O
锰还原	(CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+472H⁺+236MnO₂→H₃PO₄+106CO₂+236Mn²⁺+8N₂+366H₂O
硫酸盐还原	(CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄) +53S $O 4 2 -$ →H₃PO₄+106H₂O+106CO₂+16NH₃+53S^2-

Table 4 Sediment mineralization and related chemical reactions. Adapted from [37].

矿化过程	化学反应
有氧呼吸	(CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+138O₂→106CO₂+16HNO₃+H₃PO₄+122H₂O
硝酸盐还原	① (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+94.4HNO₃→H₃PO₄+177.2H₂O+106CO₂+55.2N₂ ② (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+84.8HNO₃→H₃PO₄+148.4H₂O+106CO₂+42.4N₂+16NH₃
铁还原	① (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+212Fe₂O₃+848H⁺→H₃PO₄+530H₂O+106CO₂+16NH₃+424Fe²⁺ ② (CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+424FeOOH+848H⁺→424Fe²⁺+106CO₂+16NH₃+H₃PO₄+742H₂O
锰还原	(CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄)+472H⁺+236MnO₂→H₃PO₄+106CO₂+236Mn²⁺+8N₂+366H₂O
硫酸盐还原	(CH₂O)₁₀₆(NH₃)₁₆(H₃PO₄) +53S $O 4 2 -$ →H₃PO₄+106H₂O+106CO₂+16NH₃+53S^2-

Table 5 Correlation coefficients between sediment physio-chemical properties and PLFAs

PLFA种类	TON	TOC	TOC/TON	NO₃-N	NO₂-N	NH₄-N	pH值
总PLFA	0.933^**	0.942^**	0.769^**	0.698^**	0.464	0.198	-0.334
SSFA	0.822^**	0.840^**	0.692^**	0.622^**	0.346	0.162	-0.444
BSFA	0.955^**	0.956^**	0.774^**	0.674^**	0.515	0.179	-0.272
MUFA	0.951^**	0.958^**	0.779^**	0.740^**	0.485	0.228	-0.298
PUFA	0.910^**	0.945^**	0.796^**	0.747^**	0.433	0.257	-0.247

Fig.7 PCA analysis of sediment PLFA

References 47

[1]	PROVOOST P, BRAECKMAN U, VAN GANSBEKE D, et al. Modelling benthic oxygen consumption and benthic-pelagic coupling at a shallow station in the southern North Sea[J]. Estuarine, Coastal and Shelf Science, 2013, 120: 1-11.
[2]	GUO J, YUAN H, SONG J, et al. Hypoxia, acidification and nutrient accumulation in the Yellow Sea Cold Water of the South Yellow Sea[J]. Science of the Total Environment, 2020, 745: 141050.
[3]	宋金明, 李学刚. 海洋沉积物/颗粒物在生源要素循环中的作用及生态学功能[J]. 海洋学报, 2018, 40(10): 1-13.
[4]	朱茂旭, 史晓宁, 杨桂朋, 等. 海洋沉积物中有机质早期成岩矿化路径及其相对贡献[J]. 地球科学进展, 2011, 26(4): 355-364.
[5]	TOUSSAINT E, DE BORGER E, BRAECKMAN U, et al. Faunal and environmental drivers of carbon and nitrogen cycling along a permeability gradient in shallow North Sea sediments[J]. Science of the Total Environment, 2021, 767: 144994.
[6]	GUO J, YUAN H, SONG J, et al. Evaluation of sedimentary organic carbon reactivity and burial in the Eastern China Marginal Seas[J]. Journal of Geophysical Research: Oceans, 2021, 126(4): e2021JC017207.
[7]	尹美玲, 段丽琴, 宋金明, 等. 长江口邻近海域表层沉积物中的细菌藿多醇及对低氧区的响应判别[J]. 环境科学, 2021, 42(3): 1343-1353.
[8]	SINKKO H, LUKKARI K, SIHVONEN L M, et al. Bacteria contribute to sediment nutrient release and reflect progressed eutrophication-driven hypoxia in an organic-rich continental sea[J]. PLoS One, 2013, 8(6): e67061.
[9]	WUNDERLICH A, HEIPIEPER H J, ELSNER M, et al. Solvent stress-induced changes in membrane fatty acid composition of denitrifying bacteria reduce the extent of nitrogen stable isotope fractionation during denitrification[J]. Geochimica et Cosmochimica Acta, 2018, 239: 275-283.
[10]	ATASHGAHI S, SANCHEZ-ANDREA I, HEIPIEPER H J, et al. Prospects for harnessing biocide resistance for bioremediation and detoxification[J]. Science, 2018, 360(6390): 743-746.
[11]	FOSTER A L, MUNK L, KOSKI R A, et al. Relationships between microbial communities and environmental parameters at sites impacted by mining of volcanogenic massive sulfide deposits, Prince William Sound, Alaska[J]. Applied Geochemistry, 2008, 23(2): 279-307.
[12]	李冬梅, 施雪华, 孙丽欣, 等. 磷脂脂肪酸谱图分析方法及其在环境微生物学领域的应用[J]. 科技导报, 2012, 30(2): 65-69.
[13]	张一鸣, 黄咸雨, 谢树成. 微生物磷脂脂肪酸单体碳同位素示踪碳循环过程[J]. 第四纪研究, 2021, 41(4): 877-892.
[14]	郭景腾. 15万年来热带西太平洋表层pH和pCO₂演化及其影响因素[D]. 青岛: 中国科学院研究生院(海洋研究所), 2015.
[15]	马骏, 宋金明, 李学刚, 等. 西太平洋Y3海山对营养盐的影响及其生态环境效应[J]. 地学前缘, 2020, 27(4): 322-331. DOI
[16]	李学刚, 宋金明, 牛丽凤, 等. 近海沉积物中氮磷的同时测定及其在胶州湾沉积物中的应用[J]. 岩矿测试, 2007, 2: 87-92.
[17]	王丽莎, 石晓勇, 张传松. 东海赤潮高发区沉积物中有机碳、有机氮的分布及其来源[J]. 海洋环境科学, 2010, 29: 165-169.
[18]	卢凤艳, 安芷生. 鹤庆钻孔沉积物总有机碳、氮含量测定的前处理方法及其环境意义[J]. 地质力学学报, 2010, 16(4): 393-401.
[19]	HU B, LI J, ZHAO J, et al. Late Holocene elemental and isotopic carbon and nitrogen records from the East China Sea inner shelf: implications for monsoon and upwelling[J]. Marine Chemistry, 2014, 162: 60-70.
[20]	LUPWAYI N Z, LARNEY F J, BLACKSHAW R E, et al. Phospholipid fatty acid biomarkers show positive soil microbial community responses to conservation soil management of irrigated crop rotations[J]. Soil and Tillage Research, 2017, 168: 1-10.
[21]	PETERSEN S O, KLUG M J. Effects of sieving, storage, and incubation temperature on the phospholipid Fatty Acid profile of a soil microbial community[J]. Applied Environmental Microbiology, 1994, 60(7): 2421-2430.
[22]	VESTAL J R, WHITE D C. Lipid analysis in microbial ecology[J]. BioScience, 1989, 39(8): 535-541.
[23]	孙和泰, 华伟, 祁建民, 等. 利用磷脂脂肪酸(PLFAs)生物标记法分析人工湿地根际土壤微生物多样性[J]. 环境工程, 2020, 38(11): 103-109.
[24]	颜慧, 蔡祖聪, 钟文辉. 磷脂脂肪酸分析方法及其在土壤微生物多样性研究中的应用[J]. 土壤学报, 2006, 5: 851-859.
[25]	陈振翔, 于鑫, 夏明芳, 等. 磷脂脂肪酸分析方法在微生物生态学中的应用[J]. 生态学杂志, 2005(7): 828-832.
[26]	SHI Y, XIANG X, SHEN C, et al. Vegetation-associated impacts on arctic tundra bacterial and microeukaryotic communities[J]. Applied Environmental Microbiology, 2015, 81(2): 492-501.
[27]	WU H, LI Y, ZHANG J, et al. Sediment bacterial communities in a eutrophic lake influenced by multiple inflow-rivers[J]. Environmental Science and Pollution Research, 2017, 24(24): 19795-19806.
[28]	邓延慧, 丁润楠. 湖泊沉积物氮矿化及其影响因素研究进展[J]. 环境生态学, 2020, 2(11): 91-95.
[29]	欧阳媛, 王圣瑞, 金相灿, 等. 外加氮源对滇池沉积物氮矿化影响的研究[J]. 中国环境科学, 2009, 29(8): 879-884.
[30]	张嘉雯, 魏健, 刘利, 等. 衡水湖沉积物营养盐形态分布特征及污染评价[J]. 环境科学, 2020, 41(12): 5389-5399.
[31]	田东凡, 李学刚, 宋金明, 等. 海洋最小含氧带氮流失过程与机制[J]. 应用生态学报, 2019, 30(3): 1047-1056. DOI
[32]	TIAN D, WANG Y, XING J, et al. Nitrogen loss process in hypoxic seawater based on the culture experiment[J]. Marine Pollution Bulletin, 2020, 152: 110912.
[33]	WAKEHAM S G. Organic biogeochemistry in the oxygen-deficient ocean: a review[J]. Organic Geochemistry, 2020, 149: 104096.
[34]	ZHANG X, CHEN Q, WANG C, et al. Characteristic analysis of phospholipid fatty acids (PLFAs) in typical nutrient polluted lake sediment in Wuhan[J]. International Journal of Sediment Research, 2021, 36(2): 221-228.
[35]	WIESENBERG G L B, DORODNIKOV M, KUZYAKOV Y. Source determination of lipids in bulk soil and soil density fractions after four years of wheat cropping[J]. Geoderma, 2010, 156(3/4): 267-277.
[36]	ESPINOSA L F, PANTOJA S, PINTO L A, et al. Water column distribution of phospholipid-derived fatty acids of marine microorganisms in the Humboldt Current system off northern Chile[J]. Deep Sea Research Part Ⅱ: Topical Studies in Oceanography, 2009, 56(16): 1063-1072.
[37]	FROELICH P N, KLINKHAMMER G P, BENDER M L, et al. Early oxidation of organic matter in pelagic sediments of the eastern equatorial Atlantic: suboxic diagenesis[J]. Geochimica et Cosmochimica Acta, 1979, 43(7): 1075-1090.
[38]	吴雪停, 刘丽华, 吴能友, 等. 海洋沉积物中早期成岩作用地球化学研究进展[J]. 海洋地质前沿, 2015, 31(12): 17-26.
[39]	PIMENOV N V, LUNINA O N, PRUSAKOVA T S, et al. Biological fractionation of stable carbon isotopes at the aerobic/anaerobic water interface of meromictic water bodies[J]. Microbiology, 2008, 77(6): 751-759.
[40]	LEHMANN M F, BERNASCONI S M, BARBIERI A, et al. Preservation of organic matter and alteration of its carbon and nitrogen isotope composition during simulated and in situ early sedimentary diagenesis[J]. Geochimica et Cosmochimica Acta, 2002, 66: 3573-3584.
[41]	RAVENSCHLAG K, SAHM K, AMANN R. Quantitative molecular analysis of the microbial community in marine arctic sediments (Svalbard)[J]. Applied Environmental Microbiology, 2001, 67(1): 387-395.
[42]	高爱国, 陈皓文, 林学政. 加拿大海盆与楚科奇海柱状沉积物中硫酸盐还原菌的分布状况[J]. 环境科学学报, 2008(5): 1014-1020.
[43]	WILLERS C, JANSEN VAN RENSBURG P J, CLAASSENS S. Phospholipid fatty acid profiling of microbial communities: a review of interpretations and recent applications[J]. Journal of Applied Microbiology, 2015, 119(5): 1207-1218.
[44]	MILLS C T, DIAS R F, GRAHAM D, et al. Determination of phospholipid fatty acid structures and stable carbon isotope compositions of deep-sea sediments of the Northwest Pacific, ODP site 1179[J]. Marine Chemistry, 2006, 98(2/3/4): 197-209.
[45]	SUN H, WU Y, BING H, et al. Available forms of nutrients and heavy metals control the distribution of microbial phospholipid fatty acids in sediments of the Three Gorges Reservoir, China[J]. Environmental Science and Pollution Research, 2018, 25(6): 5740-5751.
[46]	ZHAO J, WANG P, YU X, et al. Structure and composition of sediment: associated bacterial and eukaryotic communities in the river-lake system of Poyang Lake, China[J]. Geomicrobiology Journal, 2019, 36(8): 727-736.
[47]	LI S, XIAO X, YIN X, et al. Bacterial community along a historic lake sediment core of Ardley Island, west Antarctica[J]. Extremophiles, 2006, 10(5): 461-467.