微藻无机碳利用在岩石风化及碳循环过程中的作用

赵丽华; 吴沿友; 谢腾祥; 李海涛

doi:10.11932/karst20230101

微藻无机碳利用在岩石风化及碳循环过程中的作用

doi: 10.11932/karst20230101

赵丽华^{1, 2,},
吴沿友^2, ,,
谢腾祥³,
李海涛²

1.
贵州省分析测试研究院, 贵州贵阳 550000
2.
中国科学院地球化学研究所环境地球化学国家重点实验室, 贵州贵阳 550081
3.
西南科技大学环境与资源学院, 四川绵阳 621000

基金项目: 贵州省高层次创新型人才培养项目“百层次”（黔科合人才（2015）4035号）；国家自然科学基金 (U1612441)；贵州科学院青年科学基金（黔科院J字 (2023) 22号）

详细信息

作者简介:
赵丽华（1992－），女，工程师，硕士研究生，主要从事环境生物地球化学等研究工作。E-mail：18798012493@163.com

通讯作者:
吴沿友（1966－），男，博士，研究员，博士生导师，主要从事生态环境和环境地球化学等研究工作。E-mail：wuyanyou@mail.gyig.ac.cn

中图分类号: P642.25
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出版历程
- 收稿日期: 2021-10-12
- 刊出日期: 2023-02-25

Role of carbonic utilization of microalgae on rock weathering and carbon cycle

ZHAO Lihua^{1, 2
,},
WU Yanyou^{2
, ,},
XIE Tengxiang³,
LI Haitao²

1.
Guizhou Academy of Testing and Analysis, Guiyang, Guizhou 550000, China
2.
State Key Laboratory of Environmental Geochemistry, Institute of Geochemistry, Chinese Academy of Sciences, Guiyang, Guizhou 550081, China
3.
School of Environment and Resource, Southwest University of Science and Technology, Mianyang, Sichuan 621000, China

摘要

摘要: 岩溶碳汇呈现两种不同观点：（1）岩溶碳汇巨大，其机理在于岩溶区藻类及光合细菌利用碳酸氢根离子（${\rm{HCO}}_3^{-}$）实现光合作用，从动力学上加速了岩溶风化过程，促进大气CO₂的溶解。（2）岩溶区碳酸盐岩的风化作用，产生${\rm{HCO}}_3^{-}$，随后产生等量的阳离子在海洋中进行碳酸盐岩的沉积作用，这仅仅体现的是碳酸盐岩的搬运作用，不能体现碳汇，在长期尺度上仅仅有硅酸盐岩风化产生净碳汇。文章抓住岩石风化产生${\rm{HCO}}_3^{-}$与微藻光合作用利用${\rm{HCO}}_3^{-}$的耦合点，分析了典型代表性水生生物——微藻在无机碳利用上对岩石风化及碳汇的影响。从微藻光合无机碳利用机制以及光合作用关键性酶－碳酸酐酶（CA）作用两方面，论证了微藻生长对岩石风化及其碳汇的的促进作用；同时论述高pH、高${\rm{HCO}}_3^{-}$的风化环境对微藻生长影响。获得以下新认识：（1）微藻通过胞外碳酸酐酶（CAex）利用了大量${\rm{HCO}}_3^{-}$，加速岩石风化，并促使风化朝着形成${\rm{HCO}}_3^{-}$的方向进行；（2）微藻加速钙镁硅酸盐岩风化，风化溶出的Ca²⁺、Mg²⁺会促使碳酸盐岩的沉积，因此微藻加速硅酸盐岩风化形成净碳汇；（3）长时间尺度下，单纯的碳酸盐岩化学风化并不能直接产生净碳汇，但微藻对${\rm{HCO}}_3^{-}$利用使得碳酸盐岩风化朝着${\rm{HCO}}_3^{-}$转化方向进行，微藻参与碳酸钙沉积作用的同时转化无机碳为惰性有机碳，产生碳汇。故微藻通过CAex的作用，催化加速${\rm{HCO}}_3^{-}$与CO₂之间的转化，形成水体${\rm{HCO}}_3^{-}$消耗的动力基础，微藻无机碳利用对岩石风化具有促进作用，从而调节大气CO₂、浓度变化。基于当前研究，提出三点展望：（1）开展岩溶区区域水体系统的岩石风化、水生生物碳汇评估成为解决当前区域碳收支不平衡问题的关键；（2）查明岩石风化作用中生物作用碳转化机理及转化量，解决单纯的水化学径流法计算岩石风化碳汇精度不够问题；（3）构建光合生物参与下的新的评估方法，评估当前岩石风化在水生生物、水循环作用下的碳汇的时间尺度问题，厘清岩石风化碳汇在碳收支中的贡献。
- 微藻 /
- 风化碳汇 /
- ${\rm{HCO}}_3^{-} $ /
- 碳酸酐酶 /
- CO₂
Abstract: The carbon sink on rock weathering is widely discussed for reducing global atmospheric carbon dioxide (CO₂). Two different views on karst carbon sink are proposed. One is that the karst carbon sink is huge because bicarbonate ion (${\rm{HCO}}_3^{-}$) is used by the photosynthesis of algae and photosynthetic bacteria in karst areas, which dynamically accelerates the process of karst weathering and subsequently promote the dissolution of atmospheric CO₂. Another is that the weathering of carbonate rock generates HCO$_3^{-}$, and then the equivalent calcium ions (Ca²⁺) and Magnesium ions (Mg²⁺) are produced for the deposition of carbonate rock on the sea floor as the river enters the ocean. This process only reflects the transport of carbonate rock instead of the carbon sink because only the weathering of silicate rock may generate the net carbon sink in the long term. By literature review in this paper, the effects of microalgae (a typical aquatic organism) on rock weathering and its carbon sink are discussed based on the coupling between inorganic carbon utilization of microalgae in photosynthesis and ${\rm{HCO}}_3^{-}$ produced from rock-weathering. The facilitation on rock-weathering and its carbon sink by microalgae growth is demonstrated from two aspects, namely, the utilization mechanism of inorganic carbon and the action of carbonic anhydrase (the key enzyme of photosynthesis) in microalgae. Besides, the biomass of microalgae, in turn, is enhanced by the effects of weathering-environment, such as, higher pH value and higher ${\rm{HCO}}_3^{-}$. In this study, the following three arguments are proposed. Firstly, the weathering is accelerated because of the continuous consumption of ${\rm{HCO}}_3^{-}$ utilized by catalysis of extracellular carbonic anhydrase (CAex) in microalgae, which makes the weathering towards the direction on forming ${\rm{HCO}}_3^{-}$. Secondly, the microalgae can accelerate the weathering of calcium-magnesium silicate rocks, and Ca²⁺ and Mg²⁺ dissolved out by weathering may, in turn, facilitate the deposition of carbonate rock, hence a net carbon sink is generated. Thirdly, pure chemical weathering of carbonate rock cannot directly generate a net carbon sink at long time scale, but the ${\rm{HCO}}_3^{-}$ utilization from CO₂ in microalgae makes the weathering of carbonate rock proceed in the direction of HCO$_3^{-}$ conversion. In the process of calcium carbonate deposition involved by microalgae, inorganic carbon is converted into recalcitrant organic carbon and thus the carbon sink is generated.Research findings can be concluded that through the CAex effect, the catalysis and acceleration of conversion of HCO$_3^{-}$ to CO₂by microalgae will form the dynamic basis of water HCO$_3^{-}$ consumption. The utilization of inorganic carbon in microalgae can facilitate rock weathering, and hence the concentration of atmospheric CO₂ will be regulated. In this study, three aspects of prospect are also put forward. Firstly, to address the regional unbalance of carbon budget, it is crucial to assess the carbon sink of rock weathering under aquatic organism in karst areas. Besides, to improve the precision of calculating carbon sink of rock-weathering by hydrochemical runoff method, the mechanism and amount of biological carbon conversion in rock weathering should be determined. Finally, it is urgent to establish a new method to assess the time scale of carbon sink of rock weathering under the effects of aquatic organisms by water-cycle, which can clarify the contribution of carbon sink of rock weathering to the carbon budget.
- microalgae /
- carbon sink of rock weathering /
- ${\rm{HCO}}_3^{-} $ /
- carbonic anhydrase /
- CO₂

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图 1 地表温度（a）、大气CO₂浓度（b）、大气CO₂浓度增长率（c）随时间的变化（据IPCC AR6，2021^[1]修编）

Figure 1. Changes of surface temperature (a), atmospheric CO₂ concentration (b) and the growth rate of atmospheric CO₂ concentration (c) over time (revised from IPCC AR6, 2021^[1])

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图 2 长期碳循环模型（a）、短期碳循环模型（b）（据Berner，1999修编^[13]）

Figure 2. Long-term carbon cycle (a) and short-term carbon cycle (b) (revised from Berner, 1999^[13])

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图 3 不同pH条件下水体中各无机碳分量的百分含量（ Larkum等，1989^[76]）

Figure 3. Percentage of the inorganic carbon component in water under different pH values (Larkum et al., 1989^[76])

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图 4 微藻CO₂浓缩机制（ CCM）模型（据Sültemeyer，1998^[87]；Li Haitao等，2019^[96]修编）

Figure 4. Model of CO₂concentrating mechanism (CCM) in microalgae (revised from Sültemeyer, 1989^[87]; Li Haitao, 2019^[96])

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图 5 不同时间下方解石的镁离子释放量（mg·L⁻¹）和叶绿素a的含量（μg·L⁻¹）（a）对照组；（b、e）莱茵衣藻；（c、f）蛋白核小球藻；（d、g）铜绿微囊藻（据Xie Tengxiang等，2017^[67]）

Figure 5. Concentrations of Mg²⁺ from calcite (mg·L⁻¹) and chl-a (μg·L⁻¹) at different pH values after different hours of incubation (a) the treatment without microalgae (control group); (b, e) the treatment with C. reinhardtii (C.R.); (c, f) the treatment with C. pyrenoidosa (C.P.); and (d, g) the treatment with M. aeruginosa (M.A.) (revised from Xie Tengxiang et al., 2017^[67] )

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图 6 微藻无机碳利用与岩石风化、大气碳之间的动力关系

Figure 6. Relationship between inorganic carbon utilization by microalgae, rock weathering and atmospheric carbon

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图 7 微藻对水环境中${\rm{HCO}}_3^{-}$的响应

Figure 7. Response of microalgae to ${\rm{HCO}}_3^{-}$ in water

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图 8 流域岩石风化与微藻生长的耦合关系

Figure 8. Coupling relationship between rock weathering and microalgae growth in the watershed

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表 1 1850-2019年全球人为累计碳收支情况（引自AR6^[1,10]）

Table 1. Global accumulated anthropogenic CO₂budget from 1850 to 2019(revised from AR6^[1,10])

		碳排放量/PgC	总量/PgC	收支不平衡量/PgC
排放（源）	化石燃料燃烧及水泥生产	445±20	685±65	20
排放（源）	净通量土地使用	240±60	685±65
分配（汇）	大气增加CO₂	265±5	635±80
	海洋碳汇	160±20
	陆地碳汇	210±55

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表 2 全球碳酸盐岩风化碳汇一些数据对比（据张之淦，2012^[35]）

Table 2. Comparison of global weathering carbon sink from carbonate rock (revised from Zhang Zhigan, 2012^[35])

通量/PgC·yr⁻¹	数据来源	计算方法
0.142	Berner 1983^[23]；	水化学径流法
0.117	数据引自徐胜友等，1997；据张之淦，2012外推^[32,35]	水化学径流法
0.131	数据引自蒋忠诚等，2011；据张之淦，2012外推^[33,35]	水化学径流法
0.165	数据引自邱冬生等，2004）；据张之淦，2012外推^[34-35]	水化学径流法
0.608	Yuan，1997^[36]	定点磨片

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表 3 微藻的碳酸氢根离子利用途径份额

Table 3. Proportion of bicarbonate utilization pathway to the whole carbon utilization pathway of microalgae

NaHCO₃/mmol·L⁻¹	莱茵衣藻	蛋白核小球藻	野外混合微藻	数据来源
\	1（1.067）	0.81	0.98	赵丽华等，2016^[98]
\	0.96	0.77	/	Wu等，2015^[100]
0.50	0.97	0.81	/	Wu等，2015^[100]
2.00	1（1.05）	0.92	/	Wu等，2015^[100]
4.00	1（1.02）	1.00（1.10）	/	Wu等，2015^[100]
8.00	1（1.10）	1.00（1.29）	/	Wu等，2015^[100]

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表 4 微藻对碳酸钙碳源的利用份额(f_B)(谢腾祥等，2014^[104])

Table 4. Proportion of calcium carbonate-carbon source utilized by microalgae (Xie Tengxiang et al., 2014^[104])

藻种	AZ/mmol·L⁻¹	f_B
莱茵衣藻	0	0.02
	0.1	0.03
	1	0.08
	10	0.30
蛋白核小球藻	0	0.02
	0.1	0.02
	1	0.10
	10	0.15

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表 5 野外湖泊微藻利用添加的碳酸氢钠与总无机碳碳源的占比（Li Haitao等，2018^[96]）

Table 5. Proportion of NaHCO₃-carbon source utilized by lake microalgae (Li Haitao et al., 2018^[96])

NaHCO₃/ mmol·L⁻¹	AZ /mmol·L⁻¹
NaHCO₃/ mmol·L⁻¹	0	1.0	10.0
1.0	0.06±0.03	0.06±0.04	0.03±0.03
2.5	0.08±0.02	0.08±0.05	0.09±0.03
5.0	0.09±0.04	0.17±0.05	0.12±0.05

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表 6 碳酸酐酶加速岩石风化的数据统计

Table 6. Statistics of the acceleration of rock weathering by carbonic anhydrase

岩石种类	岩石风化变化情况	生物种类	CA种类	数据来源
白云岩	在CO₂分压低于5 000 Pa实验条件下，对白云岩的溶解速率促进倍数在1.29~3.07之间	体外实验	高分子催化剂－牛碳酸酐酶	刘再华，2001^[117]
灰岩	加入CA后，在高CO₂ 分压时，其溶解速率可增加 10倍	体外实验	高分子催化剂－牛碳酸酐酶	刘再华，2001^[117]
灰岩	在加入AZ后，单位时间单位藻体的镁离子释放量从3.37×10⁻⁴ mg·(μg·day)⁻¹降低至1.99×10⁻⁴ mg·(μg·day)⁻¹	莱茵衣藻	胞外碳酸酐酶	Xie等，2014^[66]
灰岩	在加入AZ后，单位时间单位藻体的镁离子释放量从2.44×10⁻⁴ mg·(μg·day)⁻¹降低至2.19×10⁻⁴ mg·(μg·day)⁻¹	蛋白核小球藻	胞外碳酸酐酶	Xie等，2014^[66]

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表 7 不同处理下微藻的增殖倍数（摘自Wu等，2015^[103]）

Table 7. Microalgae growth with different treatments ( Wu et al., 2015^[103])

NaHCO₃/ mmol·L⁻¹	增值倍数
NaHCO₃/ mmol·L⁻¹	0 mmol·L⁻¹ AZ	1.0 mmol·L⁻¹ AZ	10.0 mmol·L⁻¹ AZ
1	4.89±0.07	3.75±0.25	1.59±0.24
2.5	5.27±0.52	3.67±0.39	1.93±0.30
5	6.28±0.44	3.93±0.19	2.16±0.19
20	6.55±0.32	4.46±0.28	2.53±0.47

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