海洋微微型蓝细菌分子生态学研究进展

doi:10.1167/j.issn.1001-8166.2015.04.0477

微微型蓝细菌是迄今发现最小也是最古老的光合自养生物,在海洋中分布极广且丰度较高,作为重要的初级生产者对全球碳循环和海洋食物网具有重要贡献。在长期进化中微微型蓝细菌形成简并基因组和高度多样性,其分子生态学的研究能够为理解生物的基因型、表型与生态型的关系以及生物的适应性进化提供关键科学依据。借助分子生物技术的发展,海洋微微型蓝细菌分子生态学近年来的研究揭示了其分布的时空变化特征、不同亚型微尺度分布差异及影响分布的主要环境因子,探讨了环境因子影响下基因组在功能和表达上传递的适应机制信息。通过进一步扩充和挖掘基因组信息,结合分布规律论证微微型蓝细菌在分子水平上的环境适应机制,是实现生态系统微微型蓝细菌功能模块精细化数值模拟的关键,也是未来研究的主要方向和重要内容。

关键词: 微微型浮游植物, 聚球藻, 原绿球藻, 生态分布, 适应机制

Picocyanobacteria are the smallest and most ancient photosynthetic autotroph known to date with wide distribution and abundant density in the oceans. As the major contributor to primary production, marine picocyanobacteria have significant impacts on global carbon cycles and ocean food web. Given the picocyanobacterial genome streamlining and remarkable diversity during the long process of evolution, studies of their molecular ecology provide key scientific basis for relationships among genotypic, phenotypic and ecological variations, as well as for insights of biotic adaptation evolution in the context of environmental requirements. With the application of newly molecular biotechniques over recent years, our reports on the molecular ecology of marine picocyanobacteria have not only revealed their features of temporal-spatial distribution, fine distribution patterns of genetical subgroups and environmental factors controlling their variations, but also explored their adaptation mechanism derived from information about genomic functions and expressions. Accessing the further progresses in genomic information of these organism, coupled with the advances in the environmental adaptation mechanisms on molecular level given valuable information on distribution characteristics, it would be able to simulate the regulatory and metabolic network of picocyanobacteria in high resolution integrated into the framework of ecosystem model which is the main focus on the future perspectives in molecular ecology of picocyanobacteria.

Key words: Picophytoplankton, Synechococcus, Prochlorococcus, Ecological distribution, Adaptation mechanisms.

中图分类号:

P735
Q938

图1 海洋微微型蓝细菌的16S rDNA系统发育树(根据文献[12,19,25]序列信息构建)

Fig.1 Phylogenetic tree of marine picocyanobacteria based on 16S rDNA (sequences derived from references[12, 19, 25])

表1 海洋微微型蓝细菌系统发育分析常用目标基因

Table 1 Target genes used in phylogenetic analysis of marine picocyanobacteria

目标基因	编码蛋白	聚合酶链式反应前引物(5′-3′)	片段长度/bp	退火温度/°C	参考文献
目标基因	编码蛋白	聚合酶链式反应后引物(5′-3′)	片段长度/bp	退火温度/°C	参考文献
16S	蓝细菌：16S核糖体DNA	106F：CGGACGGGTGAGTAACGCGTGA 1509R：GGTTACCTTGTTACGACTT-3’	1363	45	[43]
ITS	蓝细菌：16S-23S rDNA内转录间隔区	16S-322F：TGTACACACCGCCCGTC 23S-340R：CTCTGTGTGCCTAGGTATCC	不等	55	[44]
ITS	原绿球藻：16S-23S rDNA内转录间隔区	16S-2F：GAAGTCGTTACTYYAACCC 23S-3R：TCATCGCCTCTGTGTGCC	不等	58	[41]
rbcL	1,5-二磷酸核酮糖羧化酶/加氧酶I A/B型	DJ-rbcL-F：CTGAGIGGIAARAACTACGG DJ-rbcL-R：GGCATRTGCCANACRTGRAT	615	46	[45]
psbA	光系统II D1蛋白	psbA-F：GTNGAYATHGAYGGNATHMGNGARCC psbA-R：GGRAARTTRTGNGCRTTNCKYTCRTGCAT	849	55	[46]
atpB	ATP合成酶β亚基	atpBF：CCGGTTCTGGACGTKGAATT atpBR：GGCTCRTTCATCTGGCCGTA	641	56	[25]
petB	细胞色素b₆/f复合体亚基b₆	petBF：TACGACTGGTTCCAGGAACG petBR：GAAGTGCATGAGCATGAA	597	55	[25]
cpcBA	藻蓝蛋白基因β亚基和α亚基	SyncpcB-Fw：ATGGCTGCTTGCCTGCG SyncpcA-Rev：ATCTGGGTGGTGTAGGG	500	55	[22]
cpeBA	藻红蛋白基因间隔区和α亚基	B3FW：TCAAGGAGACCTACATCG SynA1R：CAGTAGTTGATCAGRCGCAGGT	521	58	[22]
rpoC1	聚球藻：依赖DNA的RNA聚合酶	SAN-157：YTNAARCCNGARATGGAYGG SAC-1039：CYTGYTTNCCYTCDATDATRTC	841	52	[47]
narB	硝酸盐还原酶	narB1F：GAAGGCAAYCCGATGTGGA narBdegR1：GCCCCTACTGCGGTGTNGGCTGYGG	1837	58	[38]

表1 海洋微微型蓝细菌系统发育分析常用目标基因

Table 1 Target genes used in phylogenetic analysis of marine picocyanobacteria

目标基因	编码蛋白	聚合酶链式反应前引物(5′-3′)	片段长度/bp	退火温度/°C	参考文献
目标基因	编码蛋白	聚合酶链式反应后引物(5′-3′)	片段长度/bp	退火温度/°C	参考文献
16S	蓝细菌：16S核糖体DNA	106F：CGGACGGGTGAGTAACGCGTGA 1509R：GGTTACCTTGTTACGACTT-3’	1363	45	[43]
ITS	蓝细菌：16S-23S rDNA内转录间隔区	16S-322F：TGTACACACCGCCCGTC 23S-340R：CTCTGTGTGCCTAGGTATCC	不等	55	[44]
ITS	原绿球藻：16S-23S rDNA内转录间隔区	16S-2F：GAAGTCGTTACTYYAACCC 23S-3R：TCATCGCCTCTGTGTGCC	不等	58	[41]
rbcL	1,5-二磷酸核酮糖羧化酶/加氧酶I A/B型	DJ-rbcL-F：CTGAGIGGIAARAACTACGG DJ-rbcL-R：GGCATRTGCCANACRTGRAT	615	46	[45]
psbA	光系统II D1蛋白	psbA-F：GTNGAYATHGAYGGNATHMGNGARCC psbA-R：GGRAARTTRTGNGCRTTNCKYTCRTGCAT	849	55	[46]
atpB	ATP合成酶β亚基	atpBF：CCGGTTCTGGACGTKGAATT atpBR：GGCTCRTTCATCTGGCCGTA	641	56	[25]
petB	细胞色素b₆/f复合体亚基b₆	petBF：TACGACTGGTTCCAGGAACG petBR：GAAGTGCATGAGCATGAA	597	55	[25]
cpcBA	藻蓝蛋白基因β亚基和α亚基	SyncpcB-Fw：ATGGCTGCTTGCCTGCG SyncpcA-Rev：ATCTGGGTGGTGTAGGG	500	55	[22]
cpeBA	藻红蛋白基因间隔区和α亚基	B3FW：TCAAGGAGACCTACATCG SynA1R：CAGTAGTTGATCAGRCGCAGGT	521	58	[22]
rpoC1	聚球藻：依赖DNA的RNA聚合酶	SAN-157：YTNAARCCNGARATGGAYGG SAC-1039：CYTGYTTNCCYTCDATDATRTC	841	52	[47]
narB	硝酸盐还原酶	narB1F：GAAGGCAAYCCGATGTGGA narBdegR1：GCCCCTACTGCGGTGTNGGCTGYGG	1837	58	[38]

图2 海洋微微型蓝细菌的生态分布特征及其主要影响因子示意图(根据文献 ^{[

13
]}修改)

Fig.2 Conceptual diagram showing ecological patterns of marine picocyanobacteria, and representative environmental factors influencing their distribution (modified after reference ^{[

13
]})

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[1]	张琳琳, 赵晓英, 原慧. 风对植物的作用及植物适应对策研究进展[J]. 地球科学进展, 2013, 28(12): 1349-1353.
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Advances in Molecular Ecology of Marine Picocyanobacteria