海洋环境中的氨基糖及其在有机质循环过程中的指示作用

doi:10.11867/j.issn.1001-8166.2017.09.0959

地球科学进展 ›› 2017, Vol. 32 ›› Issue (9): 959 -971. doi: 10.11867/j.issn.1001-8166.2017.09.0959

任成喆(

), 袁华茂 ^*(

), 宋金明, 李学刚, 李宁, 段丽琴

1.中国科学院海洋研究所海洋生态与环境科学重点实验室,山东青岛 266071
2.中国科学院大学, 北京 100049
3.青岛海洋科学与技术国家实验室海洋生态与环境科学功能实验室,山东青岛 266237

收稿日期:2017-03-03 修回日期:2017-07-17 出版日期:2017-09-20
通讯作者: 袁华茂 E-mail:renchengzhe@outlook.com;yuanhuamao@qdio.ac.cn
基金资助:
国家自然科学基金委员会—山东省联合基金项目“海洋生态环境变化的生物地球化学机制”(编号:U1406403);国家重点基础研究发展计划项目“海湾营养物质迁移转化规律及其环境效应”(编号:2015CB452902)资助

Amino Sugars and Their Indicating Role in the Cycling of Organic Matter in Marine Environment

Chengzhe Ren( ), Huamao Yuan ^*( ), Jinming Song, Xuegang Li, Ning Li, Liqin Duan

1.Key Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China
2.University of Chinese Academy of Sciences, Beijing 100049, China
3.Function Laboratory of Marine Ecology and Environmental Sciences, Qingdao National Laboratory for Marine Science and Technology, Qingdao 266237, China

Received:2017-03-03 Revised:2017-07-17 Online:2017-09-20 Published:2017-09-20
Contact: Huamao Yuan E-mail:renchengzhe@outlook.com;yuanhuamao@qdio.ac.cn
About author:
First author:Ren Chengzhe (1991-), male, Baicheng County, Jilin Province, Ph.D studernt. Research areas include marine biogeochemistry.E-mail:renchengzhe@outlook.com
Supported by:
*Project supported by the National Natural Science Foundation of China and Shandong Province Joint Fund “Biogeochemical mechanism of marine ecological environment changes” (No.U1406403);The National Basic Research Program of China “Transportation and transformation of nutrients and theirenvironmental effects in the gulf”(No.2015CB452902)

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氨基糖作为海洋环境中一类具有重要地球化学特征的有机质,其在海水、颗粒物和沉积物中的含量和组成等信息能够有效反映有机质的来源、降解过程及成岩状态。从氨基糖的来源与组成、海洋环境中的分布特征和影响因素,以及其作为生物标志物对有机质来源和降解状态的指示作用等方面,系统总结了海洋环境中氨基糖的研究进展。结果表明,氨基糖的活性受其大分子形态、环境中溶解氧、营养盐水平和沉积环境的影响。葡萄糖胺/半乳糖胺(GlcN/GalN)和总可水解氨基酸/总可水解氨基糖(THAA/THAS)对有机质来源和降解状态的指示具有一致性,较高的GlcN/GalN和THAA/THAS值可反映浮游生物来源的新鲜有机质,其比值的降低表明有机质逐渐向细菌有机质转化。氨基糖的碳、氮归一化含量对二者的指示具有差异性,其比例的升高和降低取决于有机质降解程度和来源影响的相对贡献大小。胞壁酸可用于估算较为新鲜的细菌有机质对总有机质的贡献,但由于其快速循环而导致在溶解有机质中的含量极低,不适合应用在溶解有机质中。今后的工作应进一步加强不同微生物对海洋环境中氨基糖的贡献,区分有机质来源和降解对氨基糖的影响以及转化和归宿研究。

关键词: 氨基糖, 胞壁酸, 有机质来源, 成岩状态, 细菌降解

As a kind of marine organic matter with important geochemical characteristics, amino sugars can effectively reflect the source, diagenetic state and mineralization process of organic matter by their concentration and composition in marine environment. This article systematically concluded the research progresses of amino sugars from the aspects of their source, composition and distribution characteristics in marine environment, and the role as a biomarker indicating source and diagenetic state of marine organic matter. The result showed that the macromolecular morphology, the oxygen and nutrient level and the sedimentary environment could affect the reactivity of amino sugars. The higher ratios of glucosamine to galactosamine (GlcN/GalN) and the Total Hydrolysable Amino Acids to Total Hydrolysable Amino Sugars (THAA/THAS) can reflect the fresh planktonic organic matter source and the lower ratios can reflect the conversion from planktonic to bacterial organic matter. The carbon and nitrogen normalized yield of total hydrolysable amino sugars, however, could give contradictory results depending on the relative contribution of the source and degradation degree of organic matter. Muramic acid is suitable to estimate the contribution of relatively fresh bacteria organic matter to particulate and sediment organic matter, but it is not suitable for applying in the dissolved organic matter because of its very low concentration leading from its rapid recycle. It is critical to enhance the research on the contribution of different microorganisms to amino sugars and differentiate the influence of organic matter source and degradation on amino sugars in marine environment. The research on the conversion and fate of amino sugars in marine environment is also needed.

Key words: Amino sugars, Muramic acid, Organic matter source, Diagenetic state, Bacteria degradation.

中图分类号:

P734

表1 海洋DOM,UDOM,POM,SOM和沉降颗粒物中氨基糖的分布

Table 1 The distribution of AS in marine DOM, UDOM, POM, SOM and settling particulate matter

	%THAS-TOC	%THAS-TN	GlcN/GalN	THAS /(μmol/(g dw))	THAS /(nmol/L)	海域	参考文献
DOM	0.6	1.5(DON)	1.85		49	西北冰洋(0~100 m)	[7]
DOM	0.2	0.7(DON)	1.4		11	西北冰洋(>1 000 m)	[7]
UDOM	1.2±0.6	3.3±1.7	1.6±0.1		34.9±28.8	太平洋(0~4 000 m)	[8]
	1.1±0.6	3.1±1.7	1.7±0.2		23.6±21.7	大西洋(0~2 400 m)	[8]
	0.6±0.1	1.7±0.4	1.3±0.3		17.8±7.9	北冰洋(10~1 600 m)	[8]
POM	0.6±0.3	0.8±0.4	1.9±0.2		0.7±0.6	太平洋(0~4 000 m)	[8]
	0.2±0.2	0.5±0.4	0.4±0.3		4.3±3.7	孟加拉湾(0~1 000 m)	[9]
	0.6	0.6	2.2		8.35	西北冰洋(0~100 m)	[7]
	2.1±0.9	2.8±0.8	1.1±0.3		127.2±48.1	曼多维河口雨季	[10]
	0.7±0.2	0.6±0.2	1.8±0.4		64.4±12.8	曼多维河口雨季前期	[10]
沉降颗粒物	2.0±0.9	1.9±0.9	5.8±2.0	17.8±9.5		赤道太平洋(1 357 m)	[11]
	1.9±0.4	1.9±0.4	4.2±0.8	10.8±3.5		赤道太平洋(4 363 m)	[11]
	0.8±0.4	1.9±0.8		6.9±3.0		马尾藻海(3 200 m)	[12]
	1.8±1.3	2.9±1.8		12.0±10.6		巴拿马海盆(890~3 560 m)	[13]
	1.9±0.3		3~4.5	12.9±2.4		中国南海(1 000~1 200 m)	[14]
	1.3	1.9		13.4		楚科奇海(<40 m)	[15]
	1.0	1.7		17.3		日本大槌湾(<30 m)	[15]
	1.1	1.7		27.4		日本喷火湾(<70 m)	[15]
SOM	1.6±0.3	2.7±1.1		2.4±2.7		北海东部表层沉积物	[16]
	2.1±0.2	3.7±0.6	1.6±0.1	35.4±11.8		秘鲁沿海表层沉积物(OMZ区)	[17]
	2.1±0.2	3.7±0.6	1.6±0.1	14.8±4.3		秘鲁沿海表层沉积物(非OMZ区)	[17]
	0.8	1.2	1.2			孟加拉湾表层沉积物	[18]
	1.7±0.3	3.3±0.6	1.4±0.1	20.0±14.8		秘鲁沿海沉积物柱状样	[19]
	0.4~1.5	n.a.	1.2±0.3			南极沉积物柱状样	[20]
	2.1~3.2(2.6)	3.9	1.1~1.3(1.2)			巴西沿海表层沉积物	[21]

表1 海洋DOM,UDOM,POM,SOM和沉降颗粒物中氨基糖的分布

Table 1 The distribution of AS in marine DOM, UDOM, POM, SOM and settling particulate matter

	%THAS-TOC	%THAS-TN	GlcN/GalN	THAS /(μmol/(g dw))	THAS /(nmol/L)	海域	参考文献
DOM	0.6	1.5(DON)	1.85		49	西北冰洋(0~100 m)	[7]
DOM	0.2	0.7(DON)	1.4		11	西北冰洋(>1 000 m)	[7]
UDOM	1.2±0.6	3.3±1.7	1.6±0.1		34.9±28.8	太平洋(0~4 000 m)	[8]
	1.1±0.6	3.1±1.7	1.7±0.2		23.6±21.7	大西洋(0~2 400 m)	[8]
	0.6±0.1	1.7±0.4	1.3±0.3		17.8±7.9	北冰洋(10~1 600 m)	[8]
POM	0.6±0.3	0.8±0.4	1.9±0.2		0.7±0.6	太平洋(0~4 000 m)	[8]
	0.2±0.2	0.5±0.4	0.4±0.3		4.3±3.7	孟加拉湾(0~1 000 m)	[9]
	0.6	0.6	2.2		8.35	西北冰洋(0~100 m)	[7]
	2.1±0.9	2.8±0.8	1.1±0.3		127.2±48.1	曼多维河口雨季	[10]
	0.7±0.2	0.6±0.2	1.8±0.4		64.4±12.8	曼多维河口雨季前期	[10]
沉降颗粒物	2.0±0.9	1.9±0.9	5.8±2.0	17.8±9.5		赤道太平洋(1 357 m)	[11]
	1.9±0.4	1.9±0.4	4.2±0.8	10.8±3.5		赤道太平洋(4 363 m)	[11]
	0.8±0.4	1.9±0.8		6.9±3.0		马尾藻海(3 200 m)	[12]
	1.8±1.3	2.9±1.8		12.0±10.6		巴拿马海盆(890~3 560 m)	[13]
	1.9±0.3		3~4.5	12.9±2.4		中国南海(1 000~1 200 m)	[14]
	1.3	1.9		13.4		楚科奇海(<40 m)	[15]
	1.0	1.7		17.3		日本大槌湾(<30 m)	[15]
	1.1	1.7		27.4		日本喷火湾(<70 m)	[15]
SOM	1.6±0.3	2.7±1.1		2.4±2.7		北海东部表层沉积物	[16]
	2.1±0.2	3.7±0.6	1.6±0.1	35.4±11.8		秘鲁沿海表层沉积物(OMZ区)	[17]
	2.1±0.2	3.7±0.6	1.6±0.1	14.8±4.3		秘鲁沿海表层沉积物(非OMZ区)	[17]
	0.8	1.2	1.2			孟加拉湾表层沉积物	[18]
	1.7±0.3	3.3±0.6	1.4±0.1	20.0±14.8		秘鲁沿海沉积物柱状样	[19]
	0.4~1.5	n.a.	1.2±0.3			南极沉积物柱状样	[20]
	2.1~3.2(2.6)	3.9	1.1~1.3(1.2)			巴西沿海表层沉积物	[21]

表2 氨基糖在生物体中的组成及含量

Table 2 Concentration and composition of AS in some organisms

微生物种类	GlcN	GalN	ManN	MurA	THAS	GlcN/GalN	%THAS-TOC	%THAS-TN	物种数	参考文献
藻类	5.9~63.7(23.3)	0~13.5(5.4)	0.2~4.7(2.8)	0.0	14.4~69.9(31.5)	0.4~20.5(7.6)	0.1~0.5	0.2~0.6	4	[8,27]
桡足类	457.4~1 210.4(833.9)	32.3~58.0(45.2)	0.0	489.7~1 268.4(879.1)	14.2~20.9(17.5)		约12	约7	2
光合自养细菌	25~243.2(127.1)	3.1~14.1(7.2)	3.3~24.9(11.8)	11.0~33.9(20.0)	42.4~306.3(166.1)	8.0~56.6(24.2)	0.3~3.9	0.3~2.5	3
淡水&土壤异养G-细菌	97.4	30.9		15.7~44.6(28.4)	172.9	3.2			4
G+细菌	251.2	30.5	4.7	66.7	353.1	8.2			6
海洋异养 G-细菌				15.2~52.5(29.8)					1
海洋天然细菌种群	23.1~58.8(39.8)	10.0~35.8(18.2)	2.4~4.3(3.2)	3~7.7(5.1)	40.3~97.9(65.0)	1.6~3.4(2.5)			5
细菌DOM				trace					3
土壤G-细菌				30.0~40.0(37.8)					5	[28]
土壤G+细菌				159.2~226.9(195.0)					2
海洋G-细菌				20.0~40.0(29.0)					8
海洋G-细菌和弱G+细菌			47.8				不详	[28]
海洋G+细菌				159.2					不详
蓝细菌				43.8					1
淡水湖泊藻类	178.8	22.3	2.4	0.0	200.0	8.0			2	[29]

表2 氨基糖在生物体中的组成及含量

Table 2 Concentration and composition of AS in some organisms

微生物种类	GlcN	GalN	ManN	MurA	THAS	GlcN/GalN	%THAS-TOC	%THAS-TN	物种数	参考文献
藻类	5.9~63.7(23.3)	0~13.5(5.4)	0.2~4.7(2.8)	0.0	14.4~69.9(31.5)	0.4~20.5(7.6)	0.1~0.5	0.2~0.6	4	[8,27]
桡足类	457.4~1 210.4(833.9)	32.3~58.0(45.2)	0.0	489.7~1 268.4(879.1)	14.2~20.9(17.5)		约12	约7	2
光合自养细菌	25~243.2(127.1)	3.1~14.1(7.2)	3.3~24.9(11.8)	11.0~33.9(20.0)	42.4~306.3(166.1)	8.0~56.6(24.2)	0.3~3.9	0.3~2.5	3
淡水&土壤异养G-细菌	97.4	30.9		15.7~44.6(28.4)	172.9	3.2			4
G+细菌	251.2	30.5	4.7	66.7	353.1	8.2			6
海洋异养 G-细菌				15.2~52.5(29.8)					1
海洋天然细菌种群	23.1~58.8(39.8)	10.0~35.8(18.2)	2.4~4.3(3.2)	3~7.7(5.1)	40.3~97.9(65.0)	1.6~3.4(2.5)			5
细菌DOM				trace					3
土壤G-细菌				30.0~40.0(37.8)					5	[28]
土壤G+细菌				159.2~226.9(195.0)					2
海洋G-细菌				20.0~40.0(29.0)					8
海洋G-细菌和弱G+细菌			47.8				不详	[28]
海洋G+细菌				159.2					不详
蓝细菌				43.8					1
淡水湖泊藻类	178.8	22.3	2.4	0.0	200.0	8.0			2	[29]

图1 北冰洋和太平洋中POM和DOM,UDOM中AS含量和组成的差异(数据来自参考文献[7,8])

Fig.1 Differences of AS concentration and composition between POM, DOM and UDOM in Arctic and Pacific Ocean(data from references[7,8])

图2 太平洋POM(0.1~60 μm)中AS的垂直变化(数据来自参考文献[8])

Fig.2 Vertical changes of AS in POM(0.1~60 μm)of Pacific Ocean(data from reference[8])

图3 太平洋UDOM(<0.1 μm)中AS的垂直变化(数据来自参考文献[8])

Fig.3 Vertical changes of AS in UDOM(<0.1 μm) of Pacific Ocean(data from reference[8])

图4 近海及海湾沉降颗粒物中氨基糖的垂直变化(数据来自参考文献[15])

Fig.4 Vertical changes of AS in settling particulate matter of coastal area and bays (data from reference[15])

图5 赤道太平洋沉降颗粒物中氨基糖的垂直变化(数据来自参考文献[11])

Fig.5 Vertical changes of AS in settling particulate matter of equatorial pacific(data from reference[11])

图6 秘鲁沉积物中氨基糖随沉积物深度的变化(数据来自参考文献[18])

Fig.6 Vertical changes of AS in Peru sediment(data from reference[18])

表3 利用MurA和D-Ala估算细菌有机质对总有机碳、总氮的贡献

Table 3 Estimating the contribution of bacteria organic matter to TOC, TN by using MurA and D-Ala

研究对象	细菌MurA /(nmol/(mg C)))	细菌D-Ala /(nmol/(mg C))	生物标志物含量注释	%Bacteria-OC (基于MurA)	%Bacteria-OC (基于D-Ala)	%Bacteria-N (基于MurA)	%Bacteria-N (基于D-Ala)	参考文献
太平洋POM	12.6	-	应用于海水环境	35	-	-	-	[8]
太平洋POM	28.1	50.3	应用于海水环境,数据通过80%异养菌+20%自养菌计算得到		12~32		20~64	[27]
太平洋DOM	-	50.3		-	24~29	-	45~54
秘鲁沿岸SOM	60	-	数据来源:Moriarty(1977)	3~17	-	-	-	[19]
圣劳伦斯河口POM	42.3,21,28.1	95.2,36.9,50.3	分别用于土壤和淡水, 河口,海水环境	0.1~6.7	2.7~42	-	-	[4]
圣劳伦斯河口SOM				4.3~22	12~38	-	-
曼多维河口POM (雨季)	42.3, 21	-	分别用于土壤和淡水, 河口环境	35±10	-	-	-	[10]
曼多维河口POM (雨季前期)				24±9	-	-	-
布里安兹湖SOM	42.3	92.2	用于土壤和淡水环境	0.2~11	0.4~14.4	-	-	[29]
楚格湖SOM				0.1~4.7	0.1~3.1	-	-
亚马逊河CPOM (>63 μm)	49.1	111.2	用于淡水环境,数据通过 85%G-细菌和15%G+细菌计算得到	1~2	4~8	8~10	20~37	[57]
亚马逊河FPOM (0.1~63 μm)				3~4	11~17	4~6	17~26

表3 利用MurA和D-Ala估算细菌有机质对总有机碳、总氮的贡献

Table 3 Estimating the contribution of bacteria organic matter to TOC, TN by using MurA and D-Ala

研究对象	细菌MurA /(nmol/(mg C)))	细菌D-Ala /(nmol/(mg C))	生物标志物含量注释	%Bacteria-OC (基于MurA)	%Bacteria-OC (基于D-Ala)	%Bacteria-N (基于MurA)	%Bacteria-N (基于D-Ala)	参考文献
太平洋POM	12.6	-	应用于海水环境	35	-	-	-	[8]
太平洋POM	28.1	50.3	应用于海水环境,数据通过80%异养菌+20%自养菌计算得到		12~32		20~64	[27]
太平洋DOM	-	50.3		-	24~29	-	45~54
秘鲁沿岸SOM	60	-	数据来源:Moriarty(1977)	3~17	-	-	-	[19]
圣劳伦斯河口POM	42.3,21,28.1	95.2,36.9,50.3	分别用于土壤和淡水, 河口,海水环境	0.1~6.7	2.7~42	-	-	[4]
圣劳伦斯河口SOM				4.3~22	12~38	-	-
曼多维河口POM (雨季)	42.3, 21	-	分别用于土壤和淡水, 河口环境	35±10	-	-	-	[10]
曼多维河口POM (雨季前期)				24±9	-	-	-
布里安兹湖SOM	42.3	92.2	用于土壤和淡水环境	0.2~11	0.4~14.4	-	-	[29]
楚格湖SOM				0.1~4.7	0.1~3.1	-	-
亚马逊河CPOM (>63 μm)	49.1	111.2	用于淡水环境,数据通过 85%G-细菌和15%G+细菌计算得到	1~2	4~8	8~10	20~37	[57]
亚马逊河FPOM (0.1~63 μm)				3~4	11~17	4~6	17~26

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