大型海藻对不同CO_2浓度的光合生理响应及其生态效应
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摘要
大气CO_2升高引起海水无机碳体系的变化，这势必会对海洋中利用无机碳进行光合作用的海藻等初级生产者产生一定的作用，从而影响海洋初级生产力和海-气界面CO_2交换的变化规律。因此研究不同CO_2浓度对大型海藻光合生理的影响以及大型海藻固定CO_2的生态效应具有重要意义。本文在综述了有关大型海藻对不同CO_2浓度响应的基础上，力图探讨CO_2浓度与光强以及无机氮的协同作用对三种常见大型海藻龙须菜（红藻）、孔石莼（绿藻）和海带（褐藻）生理特性的影响以及大型海藻对海水无机碳体系的调节作用，以期为不同环境条件下大型海藻对不同CO_2浓度的响应以及利用海水养殖或“种植能源”吸收和固定CO_2提供一定的理论依据。主要研究工作包括以下方面：
1.静止和充气培养条件下不同光强对龙须菜光合生理效应的影响。充气显著地促进了龙须菜的生长，而光强过低或者过高均会抑制其生长。在相同的光强下，与充气培养相比，静止培养可抑制藻体的光合作用速率，增大呼吸作用速率；而充气与否对藻体的有效光化学效率（Yield）和光合色素影响不显著。在静止和充气培养条件下，中光组藻体的光合作用速率最大，呼吸作用速率最小；Yield和光合色素随着光强的增大而下降。这些结果说明，低光对生长的影响主要是由于光合作用速率下降引起的，而高光对生长的影响的主要原因是呼吸作用速率的增加。
2. CO_2和光强对三种大型海藻光合生理效应的影响。在正常CO_2浓度水平下，光强对三种大型海藻生长影响显著，在低光和高光下，三种海藻生长均受到抑制。在低光下，高浓度CO_2对三种海藻生长的影响不显著；在中光下，高浓度CO_2对龙须菜的生长有显著的促进作用，而对孔石莼和海带的生长作用不明显；在高光下，高浓度CO_2可促进龙须菜和孔石莼的生长，抑制海带的生长。在低光下，高浓度CO_2可使三种海藻的光合作用速率降低，光合作用速率/呼吸作用速率（P/R）升高，其中孔石莼的P/R显著高于正常CO_2浓度下的；在中光和高光下，高浓度CO_2培养下的龙须菜和孔石莼的光合作用和光合色素要低于正常CO_2浓度培养下的藻体，而高浓度CO_2对海带的光合作用要高于正常CO_2浓度培养下的。在高光下，藻体的呼吸作用速率升高、P/R降低，Chl a和Car含量最低，并且Yield值显著低于其他的光强处理组。
3. CO_2和无机氮浓度变化对三种大型海藻光合生理效应的影响。在正常CO_2浓度下，无机N加富对三种大型海藻的生长、光合作用和光合色素均有显著的促进作用。在N限制的条件下，高浓度CO_2可促进海藻的生长。在对照N海水条件下，相对于孔石莼和海带，高浓度CO_2对龙须菜的生长具有显著的促进作用。在N加富的条件下，高浓度CO_2对龙须菜的生长作用影响不显著，而高浓度CO_2增强了海带和孔石莼的呼吸作用，并且藻体的P/R值显著低于正常CO_2浓度水平下的，这在一定程度上损耗了藻体光合作用同化积累的干物质，从而抑制孔石莼和海带的生长。不管是在正常CO_2浓度下还是CO_2浓度升高的条件下，海带的光合作用速率和光合效率显著低于龙须菜和孔石莼。NO_3~-和NH_4~+加富处理对藻体的生长和光合生理特性没有显著的影响。
4.溶解无机氮加富对海带养殖水体无机碳体系的影响。无机碳体系各组分的变化趋势与无机氮添加浓度和无机氮形态有关。当NO_3~--N和NH_4~+-N浓度范围分别在4.73～52.78μmol L~(-1)和2.56～34.66μmol L~(-1)时，DIC、HCO_3~-和pCO_2均随着营养盐浓度的增加呈下降趋势，其中以NO_3~--3和NH_4~+-3组变化最为明显，均达到最低值，分别为2054和2112μmol L~(-1)、1776和1869μmol L~(-1)、86和114μatm；而当NO_3~--N和NH_4~+-N浓度范围分别为52.78～427.29μmol L~(-1)和34.66～268.33μmol L~(-1)时，DIC、HCO_3~-和pCO_2随着营养盐浓度的增加，其下降幅度逐渐减弱，但实验结束时DIC、HCO_3~-和pCO_2仍低于对照组。NO_3~--N加富组对海带的固碳能力的影响显著高于NH_4~+-N加富组。
5.不同贝藻养殖方式对桑沟湾海域无机碳体系的影响。随着培养时间的延长，贝类组的DIC、HCO_3~-和pCO_2逐渐升高，pH和CO_32-逐渐降低，在42h时，pCO_2比自然海水高出了5.5倍，pH下降了0.8。而在贝藻混养组中，随着藻密度的增加，DIC、HCO_3~-和pCO_2逐渐降低，pH和CO_32-逐渐升高。在本实验中，当贝藻混养比例为1:0.96（湿重）时，养殖水体中pCO_2最低。因此，适当的混养龙须菜和栉孔扇贝可以有效降低水域中养殖贝类释放出的CO_2浓度。
Increased atmospheric CO_2concentration can be predicted to result in a change ofthe components of inorganic carbon system. Aquatic plants use CO_2for photosynthesisand some species use HCO_3~-. The CO_2and HCO_3~-concentration in seawater mustinfluence the photosynthetic carbon fixation of aquatic plants and air-sea CO_2flux.Thus, it’s important to study on the photosynthetic physiological responses ofmacroalgae to different CO_2concentration and their ecological effects. This paperreviewed the responses of macroalgae, trying to explore the ecophysiologicalcharacteristics in three macroalgae Gracilaria lemaneformis (a red macroalga), Ulvapertusa (a green macroalga) and Saccharina japonica (a brown macroalga) and theirecological effects of the fixation carbon, in order to provide theoretical basis forresponse of macroalgae to different CO_2concentrations and controlling the CO_2concentration by the macroalgae cultivation. The main results were as follows:
1. Effect of light intensity on the photosynthetical responses of G. lemaneiformisin non-aerated and aerated cultures. At the same light intensity, non-aerated culturecould decrease the photosynthetic rate and increase respiration rate of algae, comparedto non-aerated culture. But aerated culture doesn’t affect obviously on. Yield andpigments contents. In the moderate light intensity group, there was the maximumphotosynthetic rate and the minimum respiration rate for algae, as well as Yield andpigments contents were progressively decreased with increasing light intensity. Theresults suggested that reduced growth was mainly due to the reduced photosynthesisrate in case of low light intensity, or could be ascribed to the increased respiration ratein case of high light intensity. Aerated culture could relieve the negative effect on thegrowth rate of alga for low light or high light.
2. Effects of carbon dioxide and light intensity on the photosynthetical responsesof three macroalgae. Under the normal CO_2concentration condition, light intensity hada notably significant effect on the growth of macoalgae, and the low light or high light inhibited growth of thalli. In the low light, elevated CO_2concentration had nosignificant influence on the growth of three algae, and decreased the photosyntheticrate of three algae, but increased the P/R, where the growth of U. pertusa wassignificant lower than in the normal CO_2level; in the mediate light, increased CO_2concentration could promote the relative growth rate of G. lemaneformis, have nosignificant effect on U. pertusa, and decrease the growth of S. japonica; in the highlight, elevated CO_2concentration increased the growth of G. lemaneformis and U.pertusa, but inhibited the growth of S. japonica. Compared to the normal CO_2level,moderate light and high light decreased photosynthetic rate and pigments contents of G.lemaneformis and U. pertusa, but increased photosynthetic rate of S. japonica whenthe thalli were cultured under enriched concentration CO_2condition. In the high light,the pigments contents and Yield of the thalli were significantly decreased.
3. Effects of different CO_2and DIN concentrations on the photosyntheticresponses of three macroalgae. The results showed that N supply could significantlyincrease the relative growth rate, photosynthetic rate and pigments contents ofmacroalgae under the normal CO_2concentration condition. When N was limited,elevated CO_2concentration could significantly increase the relative growth rate ofmacroalgae. Under the control N level condition, increased CO_2concentration couldpromote the growth of G. lemaneformis more effectively than U. pertusa and S.japonica. When both CO_2and N were enriched, the growth of G. lemaneformis didn’tincrease significantly, and increased respiration rate of U. pertusa and S. japonica andlow P/R resulted their decreased growth rate. In the any of CO_2concentration, thephotosynthetic rate and photosynthetic efficiency of S. japonica were significantlylower than G. lemaneformis and U. pertusa. There was no significant different betweenenriched NO_3~-and NH_4~+supply on growth and photosynthesis of macroalgae.
4. Effects of DIN enrichment on inorganic carbon system of S. japonica culturewater. The results indicated that inorganic carbon system of culturing S. japonica wascorrelated with the concentrations and forms of DIN. The concentrations of DIC, HCO_3~-and pCO_2generally decreased with the increasing of nitrogen concentrationunder the concentration of4.73~-52.78μmol L~(-1)(NO_3~--N) and2.56-34.66μmol L~(-1)(NH_4~+-N) and were most remarkably affected by the NO_3~--3and NH_4~+-3groups,corresponding with the lowest values2054and2112μmol L~(-1),1776and1869μmolL~(-1),86and114μatm, respectively. However, when the concentration of nitrogen wasbetween52.78-427.29μmol L~(-1)(NO_3~--N) and34.66-268.33μmol L~(-1)(NH_4~+-N), thefalling trends of DIC, HCO_3~-and pCO_2were weakened with nitrogen increasing, butthe concentrations of DIC, HCO_3~-and pCO_2were also lower than the control group.Influence of NO_3~--N addition group on inorganic carbon system of seawater was largerthan NH_4~+-N addition group. And the ability of carbon fixation in NO_3~--N additiongroup was significantly higher than in NH_4~+-N addition group.
5. Carbon dioxide fixation by the seaweed G. lemaneiformis in integratedmulti-trophic aquaculture with the scallop Chlamys farreri in Sanggou Bay, China. Incontrol groups, continuous dissolution of CO_2produced by scallops into the seawaternot only caused an ongoing increase of partial pressure of CO_2(pCO_2),5.5timeshigher than that of natural seawater, but also acidified seawater by0.8units after42hof culture. However, in all seaweed-scallop groups, the higher the algal density, themore CO_2was absorbed; pCO_2was lowest in the maximum density of macrolagae.The results showed that a ratio of bivalve to seaweed was1:0.96(FW), the seawatercould produce the lowest pCO_2. Overall, the integrated culture of seaweed and scallopcould provide an efficient and environmentally friendly means to reduce CO_2emissions from bivalve mariculture.
1.静止和充气培养条件下不同光强对龙须菜光合生理效应的影响。充气显著地促进了龙须菜的生长，而光强过低或者过高均会抑制其生长。在相同的光强下，与充气培养相比，静止培养可抑制藻体的光合作用速率，增大呼吸作用速率；而充气与否对藻体的有效光化学效率（Yield）和光合色素影响不显著。在静止和充气培养条件下，中光组藻体的光合作用速率最大，呼吸作用速率最小；Yield和光合色素随着光强的增大而下降。这些结果说明，低光对生长的影响主要是由于光合作用速率下降引起的，而高光对生长的影响的主要原因是呼吸作用速率的增加。
2. CO_2和光强对三种大型海藻光合生理效应的影响。在正常CO_2浓度水平下，光强对三种大型海藻生长影响显著，在低光和高光下，三种海藻生长均受到抑制。在低光下，高浓度CO_2对三种海藻生长的影响不显著；在中光下，高浓度CO_2对龙须菜的生长有显著的促进作用，而对孔石莼和海带的生长作用不明显；在高光下，高浓度CO_2可促进龙须菜和孔石莼的生长，抑制海带的生长。在低光下，高浓度CO_2可使三种海藻的光合作用速率降低，光合作用速率/呼吸作用速率（P/R）升高，其中孔石莼的P/R显著高于正常CO_2浓度下的；在中光和高光下，高浓度CO_2培养下的龙须菜和孔石莼的光合作用和光合色素要低于正常CO_2浓度培养下的藻体，而高浓度CO_2对海带的光合作用要高于正常CO_2浓度培养下的。在高光下，藻体的呼吸作用速率升高、P/R降低，Chl a和Car含量最低，并且Yield值显著低于其他的光强处理组。
3. CO_2和无机氮浓度变化对三种大型海藻光合生理效应的影响。在正常CO_2浓度下，无机N加富对三种大型海藻的生长、光合作用和光合色素均有显著的促进作用。在N限制的条件下，高浓度CO_2可促进海藻的生长。在对照N海水条件下，相对于孔石莼和海带，高浓度CO_2对龙须菜的生长具有显著的促进作用。在N加富的条件下，高浓度CO_2对龙须菜的生长作用影响不显著，而高浓度CO_2增强了海带和孔石莼的呼吸作用，并且藻体的P/R值显著低于正常CO_2浓度水平下的，这在一定程度上损耗了藻体光合作用同化积累的干物质，从而抑制孔石莼和海带的生长。不管是在正常CO_2浓度下还是CO_2浓度升高的条件下，海带的光合作用速率和光合效率显著低于龙须菜和孔石莼。NO_3~-和NH_4~+加富处理对藻体的生长和光合生理特性没有显著的影响。
4.溶解无机氮加富对海带养殖水体无机碳体系的影响。无机碳体系各组分的变化趋势与无机氮添加浓度和无机氮形态有关。当NO_3~--N和NH_4~+-N浓度范围分别在4.73～52.78μmol L~(-1)和2.56～34.66μmol L~(-1)时，DIC、HCO_3~-和pCO_2均随着营养盐浓度的增加呈下降趋势，其中以NO_3~--3和NH_4~+-3组变化最为明显，均达到最低值，分别为2054和2112μmol L~(-1)、1776和1869μmol L~(-1)、86和114μatm；而当NO_3~--N和NH_4~+-N浓度范围分别为52.78～427.29μmol L~(-1)和34.66～268.33μmol L~(-1)时，DIC、HCO_3~-和pCO_2随着营养盐浓度的增加，其下降幅度逐渐减弱，但实验结束时DIC、HCO_3~-和pCO_2仍低于对照组。NO_3~--N加富组对海带的固碳能力的影响显著高于NH_4~+-N加富组。
5.不同贝藻养殖方式对桑沟湾海域无机碳体系的影响。随着培养时间的延长，贝类组的DIC、HCO_3~-和pCO_2逐渐升高，pH和CO_32-逐渐降低，在42h时，pCO_2比自然海水高出了5.5倍，pH下降了0.8。而在贝藻混养组中，随着藻密度的增加，DIC、HCO_3~-和pCO_2逐渐降低，pH和CO_32-逐渐升高。在本实验中，当贝藻混养比例为1:0.96（湿重）时，养殖水体中pCO_2最低。因此，适当的混养龙须菜和栉孔扇贝可以有效降低水域中养殖贝类释放出的CO_2浓度。
Increased atmospheric CO_2concentration can be predicted to result in a change ofthe components of inorganic carbon system. Aquatic plants use CO_2for photosynthesisand some species use HCO_3~-. The CO_2and HCO_3~-concentration in seawater mustinfluence the photosynthetic carbon fixation of aquatic plants and air-sea CO_2flux.Thus, it’s important to study on the photosynthetic physiological responses ofmacroalgae to different CO_2concentration and their ecological effects. This paperreviewed the responses of macroalgae, trying to explore the ecophysiologicalcharacteristics in three macroalgae Gracilaria lemaneformis (a red macroalga), Ulvapertusa (a green macroalga) and Saccharina japonica (a brown macroalga) and theirecological effects of the fixation carbon, in order to provide theoretical basis forresponse of macroalgae to different CO_2concentrations and controlling the CO_2concentration by the macroalgae cultivation. The main results were as follows:
1. Effect of light intensity on the photosynthetical responses of G. lemaneiformisin non-aerated and aerated cultures. At the same light intensity, non-aerated culturecould decrease the photosynthetic rate and increase respiration rate of algae, comparedto non-aerated culture. But aerated culture doesn’t affect obviously on. Yield andpigments contents. In the moderate light intensity group, there was the maximumphotosynthetic rate and the minimum respiration rate for algae, as well as Yield andpigments contents were progressively decreased with increasing light intensity. Theresults suggested that reduced growth was mainly due to the reduced photosynthesisrate in case of low light intensity, or could be ascribed to the increased respiration ratein case of high light intensity. Aerated culture could relieve the negative effect on thegrowth rate of alga for low light or high light.
2. Effects of carbon dioxide and light intensity on the photosynthetical responsesof three macroalgae. Under the normal CO_2concentration condition, light intensity hada notably significant effect on the growth of macoalgae, and the low light or high light inhibited growth of thalli. In the low light, elevated CO_2concentration had nosignificant influence on the growth of three algae, and decreased the photosyntheticrate of three algae, but increased the P/R, where the growth of U. pertusa wassignificant lower than in the normal CO_2level; in the mediate light, increased CO_2concentration could promote the relative growth rate of G. lemaneformis, have nosignificant effect on U. pertusa, and decrease the growth of S. japonica; in the highlight, elevated CO_2concentration increased the growth of G. lemaneformis and U.pertusa, but inhibited the growth of S. japonica. Compared to the normal CO_2level,moderate light and high light decreased photosynthetic rate and pigments contents of G.lemaneformis and U. pertusa, but increased photosynthetic rate of S. japonica whenthe thalli were cultured under enriched concentration CO_2condition. In the high light,the pigments contents and Yield of the thalli were significantly decreased.
3. Effects of different CO_2and DIN concentrations on the photosyntheticresponses of three macroalgae. The results showed that N supply could significantlyincrease the relative growth rate, photosynthetic rate and pigments contents ofmacroalgae under the normal CO_2concentration condition. When N was limited,elevated CO_2concentration could significantly increase the relative growth rate ofmacroalgae. Under the control N level condition, increased CO_2concentration couldpromote the growth of G. lemaneformis more effectively than U. pertusa and S.japonica. When both CO_2and N were enriched, the growth of G. lemaneformis didn’tincrease significantly, and increased respiration rate of U. pertusa and S. japonica andlow P/R resulted their decreased growth rate. In the any of CO_2concentration, thephotosynthetic rate and photosynthetic efficiency of S. japonica were significantlylower than G. lemaneformis and U. pertusa. There was no significant different betweenenriched NO_3~-and NH_4~+supply on growth and photosynthesis of macroalgae.
4. Effects of DIN enrichment on inorganic carbon system of S. japonica culturewater. The results indicated that inorganic carbon system of culturing S. japonica wascorrelated with the concentrations and forms of DIN. The concentrations of DIC, HCO_3~-and pCO_2generally decreased with the increasing of nitrogen concentrationunder the concentration of4.73~-52.78μmol L~(-1)(NO_3~--N) and2.56-34.66μmol L~(-1)(NH_4~+-N) and were most remarkably affected by the NO_3~--3and NH_4~+-3groups,corresponding with the lowest values2054and2112μmol L~(-1),1776and1869μmolL~(-1),86and114μatm, respectively. However, when the concentration of nitrogen wasbetween52.78-427.29μmol L~(-1)(NO_3~--N) and34.66-268.33μmol L~(-1)(NH_4~+-N), thefalling trends of DIC, HCO_3~-and pCO_2were weakened with nitrogen increasing, butthe concentrations of DIC, HCO_3~-and pCO_2were also lower than the control group.Influence of NO_3~--N addition group on inorganic carbon system of seawater was largerthan NH_4~+-N addition group. And the ability of carbon fixation in NO_3~--N additiongroup was significantly higher than in NH_4~+-N addition group.
5. Carbon dioxide fixation by the seaweed G. lemaneiformis in integratedmulti-trophic aquaculture with the scallop Chlamys farreri in Sanggou Bay, China. Incontrol groups, continuous dissolution of CO_2produced by scallops into the seawaternot only caused an ongoing increase of partial pressure of CO_2(pCO_2),5.5timeshigher than that of natural seawater, but also acidified seawater by0.8units after42hof culture. However, in all seaweed-scallop groups, the higher the algal density, themore CO_2was absorbed; pCO_2was lowest in the maximum density of macrolagae.The results showed that a ratio of bivalve to seaweed was1:0.96(FW), the seawatercould produce the lowest pCO_2. Overall, the integrated culture of seaweed and scallopcould provide an efficient and environmentally friendly means to reduce CO_2emissions from bivalve mariculture.
引文
方建光,匡世焕,孙慧玲,孙耀,周诗赉,宋云利,崔毅,赵俊,杨琴芳,李锋,张爱君,王兴章,汤庭耀.桑沟湾栉孔扇贝养殖容量的研究[J].海洋水产研究,1996a,2:18～32.
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