中国南海浮游植物光合固碳与阳光紫外辐射关系的研究
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摘要
海洋对大气CO2的吸收,依赖于海洋光合作用所驱动的“CO2生物泵”。阳光紫外辐射(UVR, 280-400 nm)会影响浮游植物的固碳过程及固碳量。然而,传统固碳量的测定方法忽视了UVR的作用,会导致光合固碳量的测定误差。另外,工业活动导致的平流层臭氧减少,使得地球表面阳光UVB (280-315 nm)辐射增加,影响海洋初级生产过程。基于以上两方面研究的必要性,本文探讨了阳光UVR对南海典型海域浮游植物固碳量的影响,分析了UVR的生理效应与环境变化及种群变化的关系,并比较了近海与外海浮游植物群落对UVR响应的差异,主要结果如下:
海洋初级生产力或光合固碳量测定时,因所用培养容器透光特性的差异,导致固碳量测定误差高达60%。以石英容器(透UV)、硼酸盐玻璃(部分透UV)、聚碳酸酯(透少量UV且消减可见光)及普通玻璃容器(透部分UVA),C-14示踪法同时测定浮游植物固碳量,与石英容器的结果相比,普通玻璃容器会高估约30%,而硼酸盐玻璃和聚碳酸酯容器则会分别低估约3%和30%浮游植物的光合固碳量。
阳光UVA (315-400 nm),在光强较高时抑制浮游植物光合固碳;而在光强较低时促进其固碳量,UVB长波段部分(305-315 nm)也有促进作用。在滤除可见光(PAR, 400-700 nm)条件下,UVR可以驱动浮游植物光合固碳。为此,光强较高的晴天时,UVR抑制浮游植物光合作用,降低其固碳量;光强较低的阴天时,UVR对浮游植物固碳有明显的促进作用。浮游植物对UVR利用的能力与细胞粒径大小有关,>20μm细胞利用能力最强,其光合固碳在低光条件下受UVR的促进;5-20μm细胞次之;<5μm细胞则不能利用UVR进行光合固碳。利用UVR进行光合固碳的浮游植物,含有较高的紫外吸收物质。
不同季节或不同海域,水体理化环境不同,浮游植物种群组成不同,阳光UVR对浮游植物光合固碳的影响也存在差异。夏季,虽然阳光辐射强度较高,但UVR对浮游植物固碳量的抑制率比冬季低;UVR对细胞导致的损伤在水温较高的夏季会得到较快的修复。台风袭击后,近岸海域浮游植物优势种群由粒径较大的微型细胞转变为粒径较小的微微型,光合固碳量受UVR的影响较大。而冬季以微微型浮游植物为优势种,且对UVR较敏感。另外,由近岸到远洋,浮游植物优势种群从微型到微微型转变,而UVR对浮游植物固碳量的抑制率也随之升高。近岸富营养海域,浮游植物生物量较高,较大粒径细胞(>20μm)占比例较高,浮游植物固碳量及固碳速率较高,对阳光UVR耐受能力较强;相反,外海寡营养海域,浮游植物生物量较低,较大粒径细胞占比例较低,水体光合固碳量及单位叶绿素固碳速率较低,对UVR也较敏感。河口海域,受陆源输入和潮汐的共同影响,理化环境极为复杂,浮游植物生物量、种群和光合能力随盐度梯度变化较大;表层水体光合固碳量及同化系数,在低盐区(<10)和高盐区(>20)较低,中盐区(10-20)较高、受UVR影响较大。
总之,阳光UVR对浮游植物的影响,因时空变化而不同,也因浮游植物群落的变化而变化,在微微型占优势的水体光合固碳量受UVR影响较大。UVB始终导致负面效应,而UVA在阳光辐射较低的水层或天气下,对浮游植物的固碳量有促进作用。
Atmospheric CO2 uptake by the oceans depends on the“marine biological CO2 pump”driven by marine photosynthesis. Solar ultraviolet radiation (UVR, 280-400 nm) is known to affect the photosynthetic processes and carbon fixation of phytoplankton. Therefore, measurements of marine primary production, without UVR being considered as in traditional methods, can result in significant errors. On the other hand, the depletion of the stratospheric ozone layer has caused an increase of solar UVB (280-315 nm) irradiance reaching the Earth’s surface, which could influence the marine primary productivity. Based on the above significances of UVR-effects, this thesis was designed to investigate UVR effects on carbon fixation of the phytoplankton assemblages from the typical areas of the South China Sea. The relationships of UVR-related physiological impacts with the changes in environments or taxonomical structure were analyzed, and the responses to UVR of phytoplankton were compared from the coasts to pelagic waters. The main results are as follows:
Different features in light transmission of the vessels can lead to the estimation errors on marine photosynthetic carbon fixation as high as 60%. Carbon fixation rates measured with C-14 method using the quartz (UV transmitted), borate glass (partially UV transmitted), polycarbonate (little UV transmitted and little PAR blocked) or common glass vessels (partially UVA transmitted), showed significant differences. Compared with the results from the quartz vessels, the carbon fixation rate could be overestimated by 30% with the common glass vessels, but underestimated by 3% with borate glass or by 30% with polycarbonate vessels, respectively.
Phytoplankton carbon fixation could be enhanced at moderate levels of solar radiation by UVA (315-400 nm), though it brought about inhibition at higher levels. Longer wavelengths of UVB (305-315 nm) were also effective in enhancing the photosynthetic carbon fixation. Under UVR alone treatment (PAR, filtered out), UVR could drive the photosynthetic carbon fixation. Therefore, the carbon fixation by phytoplankton assemblages exhibited higher rates in the presence of UVR on cloudy days, but lower rates on sunny days. The capacity of UVR-utilization was related to the size of phytoplankton cells. Micro-cells (>20μm) were able to use UVR more efficiently than nano-cells (5-20μm), and smaller nano- or pico-cells (<5μm) were unable to use UVR for the photosynthetic carbon fixation. Additionally, more UV-absorbing compounds were detected in the cells with UVR-utilized capacities.
UVR-induced impacts varied among different seasons and spatially. UVR-induced inhibition was lower in summer than that in winter, although solar radiation was high in the former season when the repair rate of UVR-induced damage was faster in the higher temperature. During the summer period, typhoon’s passage influenced the UVR-induced effects on phytoplankton community structure and carbon fixation. The larger cells were dominant in pre-typhoon; while pico-cells were dominant in post-typhoon periods, and UVR decreased the photosynthesis to more extent after typhoon. In winter, the phytoplankton assemblages were dominated by piconano-cells, and more sensitive to UVR. On the other hand, the dominant phytoplankton species changed from micro- to pico-cells from the coastal to off-shore waters, and UVR-induced inhibition on photosynthetic carbon fixation increased with increased distance from the shore or with decreased cell sizes. In the estuaries, physic-chemical environments were very complex because of the influence by the freshwater inputs and the water-exchange with the adjacent open sea; therefore, dramatic changes were found in phytoplankton biomass, taxonomical structure and photosynthetic capacity. Carbon fixation per Chl a or per volume of seawater were lower in the low (<10) or high (>20) salinity zones, but higher in the middle salinity zone (10-20) where more reduction caused by solar UVR was observed on the photosynthetic rates of phytoplankton assemblages.
In summary, solar UVR effects on phytoplankton varied spatially and temporally; and also changed according to the changes in community structure. Solar UVB always led to the negative effects on the phytoplankton photosynthesis; while UVA could promote the carbon fixation at low or moderate levels of sunlight on cloudy days or at deeper depths.
海洋初级生产力或光合固碳量测定时,因所用培养容器透光特性的差异,导致固碳量测定误差高达60%。以石英容器(透UV)、硼酸盐玻璃(部分透UV)、聚碳酸酯(透少量UV且消减可见光)及普通玻璃容器(透部分UVA),C-14示踪法同时测定浮游植物固碳量,与石英容器的结果相比,普通玻璃容器会高估约30%,而硼酸盐玻璃和聚碳酸酯容器则会分别低估约3%和30%浮游植物的光合固碳量。
阳光UVA (315-400 nm),在光强较高时抑制浮游植物光合固碳;而在光强较低时促进其固碳量,UVB长波段部分(305-315 nm)也有促进作用。在滤除可见光(PAR, 400-700 nm)条件下,UVR可以驱动浮游植物光合固碳。为此,光强较高的晴天时,UVR抑制浮游植物光合作用,降低其固碳量;光强较低的阴天时,UVR对浮游植物固碳有明显的促进作用。浮游植物对UVR利用的能力与细胞粒径大小有关,>20μm细胞利用能力最强,其光合固碳在低光条件下受UVR的促进;5-20μm细胞次之;<5μm细胞则不能利用UVR进行光合固碳。利用UVR进行光合固碳的浮游植物,含有较高的紫外吸收物质。
不同季节或不同海域,水体理化环境不同,浮游植物种群组成不同,阳光UVR对浮游植物光合固碳的影响也存在差异。夏季,虽然阳光辐射强度较高,但UVR对浮游植物固碳量的抑制率比冬季低;UVR对细胞导致的损伤在水温较高的夏季会得到较快的修复。台风袭击后,近岸海域浮游植物优势种群由粒径较大的微型细胞转变为粒径较小的微微型,光合固碳量受UVR的影响较大。而冬季以微微型浮游植物为优势种,且对UVR较敏感。另外,由近岸到远洋,浮游植物优势种群从微型到微微型转变,而UVR对浮游植物固碳量的抑制率也随之升高。近岸富营养海域,浮游植物生物量较高,较大粒径细胞(>20μm)占比例较高,浮游植物固碳量及固碳速率较高,对阳光UVR耐受能力较强;相反,外海寡营养海域,浮游植物生物量较低,较大粒径细胞占比例较低,水体光合固碳量及单位叶绿素固碳速率较低,对UVR也较敏感。河口海域,受陆源输入和潮汐的共同影响,理化环境极为复杂,浮游植物生物量、种群和光合能力随盐度梯度变化较大;表层水体光合固碳量及同化系数,在低盐区(<10)和高盐区(>20)较低,中盐区(10-20)较高、受UVR影响较大。
总之,阳光UVR对浮游植物的影响,因时空变化而不同,也因浮游植物群落的变化而变化,在微微型占优势的水体光合固碳量受UVR影响较大。UVB始终导致负面效应,而UVA在阳光辐射较低的水层或天气下,对浮游植物的固碳量有促进作用。
Atmospheric CO2 uptake by the oceans depends on the“marine biological CO2 pump”driven by marine photosynthesis. Solar ultraviolet radiation (UVR, 280-400 nm) is known to affect the photosynthetic processes and carbon fixation of phytoplankton. Therefore, measurements of marine primary production, without UVR being considered as in traditional methods, can result in significant errors. On the other hand, the depletion of the stratospheric ozone layer has caused an increase of solar UVB (280-315 nm) irradiance reaching the Earth’s surface, which could influence the marine primary productivity. Based on the above significances of UVR-effects, this thesis was designed to investigate UVR effects on carbon fixation of the phytoplankton assemblages from the typical areas of the South China Sea. The relationships of UVR-related physiological impacts with the changes in environments or taxonomical structure were analyzed, and the responses to UVR of phytoplankton were compared from the coasts to pelagic waters. The main results are as follows:
Different features in light transmission of the vessels can lead to the estimation errors on marine photosynthetic carbon fixation as high as 60%. Carbon fixation rates measured with C-14 method using the quartz (UV transmitted), borate glass (partially UV transmitted), polycarbonate (little UV transmitted and little PAR blocked) or common glass vessels (partially UVA transmitted), showed significant differences. Compared with the results from the quartz vessels, the carbon fixation rate could be overestimated by 30% with the common glass vessels, but underestimated by 3% with borate glass or by 30% with polycarbonate vessels, respectively.
Phytoplankton carbon fixation could be enhanced at moderate levels of solar radiation by UVA (315-400 nm), though it brought about inhibition at higher levels. Longer wavelengths of UVB (305-315 nm) were also effective in enhancing the photosynthetic carbon fixation. Under UVR alone treatment (PAR, filtered out), UVR could drive the photosynthetic carbon fixation. Therefore, the carbon fixation by phytoplankton assemblages exhibited higher rates in the presence of UVR on cloudy days, but lower rates on sunny days. The capacity of UVR-utilization was related to the size of phytoplankton cells. Micro-cells (>20μm) were able to use UVR more efficiently than nano-cells (5-20μm), and smaller nano- or pico-cells (<5μm) were unable to use UVR for the photosynthetic carbon fixation. Additionally, more UV-absorbing compounds were detected in the cells with UVR-utilized capacities.
UVR-induced impacts varied among different seasons and spatially. UVR-induced inhibition was lower in summer than that in winter, although solar radiation was high in the former season when the repair rate of UVR-induced damage was faster in the higher temperature. During the summer period, typhoon’s passage influenced the UVR-induced effects on phytoplankton community structure and carbon fixation. The larger cells were dominant in pre-typhoon; while pico-cells were dominant in post-typhoon periods, and UVR decreased the photosynthesis to more extent after typhoon. In winter, the phytoplankton assemblages were dominated by piconano-cells, and more sensitive to UVR. On the other hand, the dominant phytoplankton species changed from micro- to pico-cells from the coastal to off-shore waters, and UVR-induced inhibition on photosynthetic carbon fixation increased with increased distance from the shore or with decreased cell sizes. In the estuaries, physic-chemical environments were very complex because of the influence by the freshwater inputs and the water-exchange with the adjacent open sea; therefore, dramatic changes were found in phytoplankton biomass, taxonomical structure and photosynthetic capacity. Carbon fixation per Chl a or per volume of seawater were lower in the low (<10) or high (>20) salinity zones, but higher in the middle salinity zone (10-20) where more reduction caused by solar UVR was observed on the photosynthetic rates of phytoplankton assemblages.
In summary, solar UVR effects on phytoplankton varied spatially and temporally; and also changed according to the changes in community structure. Solar UVB always led to the negative effects on the phytoplankton photosynthesis; while UVA could promote the carbon fixation at low or moderate levels of sunlight on cloudy days or at deeper depths.
引文
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