大气圈氧气上升与生物进化:一个重要的地球生物学过程

摘要/Abstract

摘要：

以生氧光合作用为主造成的大气圈氧气上升,与生物进化存在着密切的成因联系。在大气圈氧气含量明显上升之前,微生物在地球大气圈演化中可能起着主要作用,形成了埃迪卡拉纪之前的微生物世界;甚至到今天,这些细菌以及其他的微体藻类,一直在向地球的大气圈提供氧气,而且在海洋中进行着艰苦的固氮作用。早期很少受到动物影响的生物圈,与现代动物所占据的生物圈(后生动物世界)明显不同,这个转变随着埃迪卡拉纪—寒武纪器官级别的动物辐射而发生,动物辐射造成了宏观生态和宏观进化表现的根本性变化。因此,地球大气圈的氧气上升,实际上是一个复杂的地球生物学过程;追索这个复杂的地球生物学过程所产生的环境变化及其与生物进化与革新之间的成因联系,对深入了解地球复杂的演变历史将提供一些重要线索和思考途径。

关键词: 大气圈氧气上升, 生物进化与革新, 地球生物学, 研究进展

Abstract:

The rise of atmosphere oxygen that is chiefly resulted from the oxygenic photosynthesis illustrates a genetically consanguineous relation with the biological evolution. Microorganisms may also have played a major role in atmosphere evolution before the rise of oxygen, which shapes a microbial world before the Ediacaran. Even in today, these bacteria, as well as microscopic algae, supply oxygen to the atmosphere and churn out fixed nitrogen in Earth’s vast oceans. Earlier, less animal-influenced biospheres worked quite differently from the one currently occupied, with the Ediacaran-Cambrian radiation of organ-grade animals marking a fundamental shift in macroecological and macroevolutionary expression. Therefore, the rise of the atmosphere oxygen is actually a sophisticated geobiological process. Tracing the complex relationship between evolution and environmental change that is caused by the rise of the atmosphere oxygen will provide some important thinking approaches and researching clues for the further understanding of the sophisticatedly evolutionary history of the Earth.

Key words: rise of atmosphere oxygen, biological evolution and innovation, geobiology, research advancement

中图分类号:

P512.5

梅冥相, 孟庆芬. 大气圈氧气上升与生物进化:一个重要的地球生物学过程[J]. 现代地质, 2017, 31(05): 1022-1038.

MEI Mingxiang, MENG Qingfen. The Rise of Atmosphere Oxygen and the Biological Evolution: An Important Geobiological Process[J]. Geoscience, 2017, 31(05): 1022-1038.

图/表 4

图1 太古宙以来主要的地质过程 A.碳同位素曲线,整个地球历史的δ13C记录,其中虚线代表了有限的资料,Campbell and Allen[40]运用了许多学者的资料综合而成,注意被定为巨型氧化作用事件的GOE、NOE和POE所代表的主要的大气圈氧气上升及其所代表的变冷过程,对应着明显的碳同位素异常事件代表的含氧碳循环的剧烈波动;B.冰川作用期,包括(1)地球上最古老的冰川作用(2.9 Ga),(2)休伦冰川作用(2 420~2 250 Ma),(3)1.8 Ga的King Leopold冰川作用,(4)720~658 Ma的Sturtian冰川作用,(5)655~635 Ma的Marinoan冰川作用,(6)584~582 Ma的Gaskiers冰川作用,(7)晚奥陶世冰川作用,(8)石炭纪—二叠纪冰川作用,(9)第四纪冰川作用;太古宙和古元古代冰川作用的作用时期是不确定的,所以用虚线表示;C.超大陆形成的时期(修改自文献[40]):[1] 地球上最古老的超大陆即基诺超大陆,[2] 奴纳乌提亚超大陆,[3] 哥伦比亚超大陆,[4] 罗迪利亚超大陆,[5] 冈瓦纳大陆,[6] 泛大陆;D.大气圈氧气含量变化曲线(修改自文献[27]),箭头所指代表相对较低的氧气含量周期,大气圈氧气含量上升的步骤表示为(1)至(7),说明见正文

Fig.1 Diagram showing the main geological processes since the Archean to the present

图2 展示了后生动物前与显生宙海洋生物圈的宏观生态和宏观进化差异性的概略性图解大约100 Ma(635~541 Ma)的埃迪卡拉纪,标定了本质上存在差异的生命世界之间的回转;恒定不变的海洋生物量(较为宽泛地与生物大小相关),随着埃迪卡拉纪—奥陶纪三重营养结构的确立而急剧上升;在埃迪卡拉纪之前和之后,生态系均是“稳定的”,但是,稳定性的基础来源于生物圈本质上存在差异的动力;形态差异性曲线来源于微化石资料和多细胞生物中的细胞类型的数量;修改自Butterfield[20]

Fig.2 A schematic depiction of macroecological and macroevolutionary differences between the pre-metazoan and Phanerozoic marine biospheres

图3 天津蓟县剖面形态各异的宏观藻类化石 (A)箭头所指为类似于丘尔藻的化石;(B)箭头所指为球形丘尔藻化石的铸模;(C)新鲜的岩石破裂面上的类似于拟丘尔藻的化石;(D)新鲜的岩石破裂面上椭圆形的类似于寿县藻的化石:a为中心的髓部,b为边部,具有可能的原始生物组织分异;(E)新鲜的岩石破裂面上似球形的类似于丘尔藻的化石,涵义同照片(D);(F)各种形态的宏观藻类化石:a为类似于丘尔藻的化石,b为类似于寿县藻的化石;c为类似于荚藻的化石,d为类似于卵圆藻的化石,MT代表臼齿状构造

Fig.3 Various categories of fossils of megascopic algae reflected by different morphological features at the Jixian section in Tianjin

图4 显生宙大气圈氧气浓度变化与生物进化与革新之间的一个可能的联系其上部和下部边界是模拟大气圈氧气浓度的边界误差,标有数字的间隔标注了重要的进化事件,这些事件可能与大气圈氧气浓度的变化存在联系。其中,(1)至(7)代表与氧气浓度上升存在关联的生物进化事件,(1)至(5)则代表与大气圈氧气浓度减小相关联的进化事件,修改自Berner et al.[24]

Fig.4 A possible link between the variation in atmospheric O2 concentration through the Phanerozoic and the evolution and development of life on Earth

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