亚硝酸盐对铜绿微囊藻生理特性的影响
摘要
浮游植物占全球初级生产的50%,在淡水水体中主要的藻种是蓝藻门的铜绿微囊藻,水体中氮、磷等营养元素的增加常常引起藻类爆发式的繁殖,形成水华。水华发生时,水体中CO_2浓度下降、pH升高和水体中溶解氧减少,这些条件有利于硝化反应和反硝化反应产生的亚硝酸盐的聚积。另外一些人为的因素也可导致水体中亚硝酸盐浓度的增加。本文研究了亚硝酸盐单独存在时和与硝酸盐、铵盐共存时对铜绿微囊藻生长和生理的影响,探讨了铜绿微囊藻细胞内亚硝酸盐含量的变化对藻生长的影响,对亚硝酸盐影响铜绿微囊藻和四尾栅藻的竞争进行了分析。
在亚硝酸盐单独存在的条件下,浓度小于5mg/L的亚硝酸盐,对铜绿微囊藻的生长没有明显影响,藻能够以亚硝酸盐为氮源。随着培养基中亚硝酸盐浓度逐渐升高到20mg/L,藻的生长受到抑制直至消亡,此时,铜绿微囊藻的光合速率下降,细胞MDA含量增加,细胞内亚硝酸盐含量增加,表明亚硝酸盐通过抑制铜绿微囊藻的光合作用和升高藻体内氧自由基浓度的方式抑制藻的生长。而藻的CAT酶活性、亚硝酸还原酶活性和藻叶绿素a含量增加可能是藻的一种自我保护措施。
硝酸盐存在的条件下(240mg/L),铜绿微囊藻在亚硝酸盐浓度为30mg/L的培养基中依然可以生长。而且,在相同的亚硝酸盐浓度条件下,含硝酸盐培养基中的藻相比只含亚硝酸盐培养基中的藻,有更高的光合速率,更小的MDA含量和藻细胞内亚硝酸盐含量。亚硝酸盐的吸收试验也显示硝酸盐存在的条件下,藻对亚硝酸盐吸收的半饱和常数(K_S)增加,这些结果表明硝酸盐能够抑制藻对亚硝酸盐的吸收,从而减少藻细胞内的亚硝酸盐含量,导致亚硝酸盐对藻抑制的减弱。在硝酸盐和亚硝酸盐共存的培养基中,我们还观察到亚硝酸盐被氧化为硝酸盐,表明此条件下可以诱导铜绿微囊藻生成亚硝酸氧化还原酶。
铵盐存在的培养基中(20mg/L),2mg/L的亚硝酸盐添加,对铜绿微囊藻的生长表现出促进作用,藻的光合速率和细胞内亚硝酸盐含量增加,MDA含量下降。进一步增加亚硝酸盐浓度(4、7、10mg/L),藻的生长受到抑制直至消亡,同时观察到藻的光合速率下降,细胞内亚硝酸盐含量和MDA含量增加。亚硝酸盐的吸收试验显示铵盐存在的条件下,藻对亚硝酸盐吸收的半饱和常数(K_S)减少,表明铵盐能够刺激藻对亚硝酸盐的吸收,增加藻细胞内的亚硝酸盐含量。在一定的范围内,细胞亚硝酸盐含量的增加可以促进藻的生长,但是过高的细胞内亚硝酸盐含量会抑制藻的生长。
硝酸盐浓度为150mg/L和300mg/L的处理组中,相比低CO_2组,高CO_2组藻的生长更好,光合速率增加,细胞内亚硝酸盐含量和MDA含量减少。而硝酸盐浓度为50mg/L的处理组中,高CO_2组和低CO_2组中,藻的生长变化不明显,光合速率、细胞内亚硝酸盐含量和MDA含量都没有明显变化。而且在相同的亚硝酸盐浓度条件下,藻细胞内亚硝酸盐含量的降低,导致培养基中藻生长的增加,这些结果表明细胞内的亚硝酸盐可能是影响藻生长的主要原因。通过对NO_2~--N:NO_3~--N为0:240和30mg/L:240mg/L两个处理组中细胞内亚硝酸盐含量和培养基中硝酸盐浓度变化的30d观察,发现培养基中亚硝酸盐进入藻体和藻排放亚硝酸盐是同时存在的,当培养基中亚硝酸盐浓度过高时,进入藻体的亚硝酸盐大于藻排放的,导致细胞内亚硝酸盐含量增加。相反,则培养基中亚硝酸盐浓度增加。
相比单独培养时,混合培养的条件下,铜绿微囊藻和四尾栅藻的生长都受到抑制,但是四尾栅藻的最大生物量与单独培养时藻的最大生物量的比值大于铜绿微囊藻,而且随着亚硝酸盐浓度的增加这种差距越明显,亚硝酸盐浓度为30mg/L时,铜绿微囊藻完全被抑制。表明两种藻之间存在着竞争,四尾栅藻在竞争中更具优势,亚硝酸盐浓度的升高增强了四尾栅藻的竞争优势。通过两种藻的相互化感作用试验,发现两种藻之间存在着化感作用,而且亚硝酸盐浓度增加化感作用增强。这些现象表明上述的竞争结果是化感作用和亚硝酸盐对藻的毒害作用共同造成的。
Phytoplankton account for approximately 50% of global primary production, and Microcystis aeruginosa blooms are dominant in fresh water. The increase of N and P is the main reason for the explosive growth of phytoplankton. During the water blooms, the concentration of CO_2 and dissolved O_2 decrease and pH increases, this could result in the nitrite accumulation by inhibiting oxidation of nitrite to nitrate or reduction of nitrite to ammonium. On the other hand, human activities and industrialization also lead to an increased accumulation of nitrite in the environment. In this study, the effects of nitrite on the growth and physiology of Microcystis aeruginosa were investigated when nitrite as sole nitrogen source, and under the present of nitrate condition or the ammonium condition; The impacts of intracellular nitrite content change on Microcystis aeruginosa growth were studied, and the competition of Microcystis aeruginosa and Scendesmus quadricauda under different nitrite concentrations was analysed..
When nitrite as sole nitrogen source, Microcystis aeruginosa could take up nitrite for growth, and the growth of alga was not impacted in medium with nitrite concentration below 5mg/L. However, with nitrite concentrations increasing to 20mg/L, the growth of alga was inhibited and photosynthetic activity decreased. Meanwhile, the content of MDA and intracellular nitrite increased. These indicated nitrite inhibited the growth of Microcystis aeruginosa by decreasing photosynthetic activity and increasing the content of oxygen free radical. The increase of chlorophyll a content、CAT activity and nitrite reductase activity might be an adaptive mechanism of alga under high nitrite condition.
When both nitrate and nitrite as nitrogen source, the Microcystis aeruginosa could grow under nitrite concentration of 30 mg/L. Furthermore, compared to medium with only nitrite, the alga had higher photosynthetic activity and less the content of MDA and intracellular nitrite in medium with both nitrate and nitrite under high nitrite concentration condition. The examination of nitrite uptake also showed the half-saturation constant of alga increased when nitrate appearance. These results revealed that nitrate could inhibit the nitrite uptake, which resulted in the decrease of the intracellular nitrite concentration and the inhibition of nitrite on alga growth weakening. In medium containing both nitrate and nitrite, the nitrite could be oxidized to nitrate, suggesting nitrite oxidoreductase has been conduced.
When both ammonium and nitrite as nitrogen source, in medium containing nitrite concentration of 2 mg/L, the growth of alga increased, the photosynthetic activities and intracellular nitrite content also showed increase but MDA content decreased compared the medium of nitrite concentration of 0mg/L. With the nitrite concentrations further increasing to 10mg/L, the growth of alga slowed down and even ceased, the photosynthetic activity decreased but the content of intracellular nitrite and MDA increased. The examination of nitrite uptake showed the half-saturation constant of alga decreased when ammonium appearance, indicating ammonium could promote the uptake of nitrite and increase the intracellular nitrite content. In the range of low intracellular nitrite content, the increase of intracellular nitrite could promote the growth of alga, however, too high intracellular nitrite content could inhibit the growth of alga.
At nitrate concentration of 50mg/L, the growth of alga hadn't significant change in high CO_2 group and low CO_2 group, the intracellular nitrite concentration、the photosynthetic activity and MDA content also remained constant. However, at nitrate concentration of 150mg/L and 300mg/L, the growth of alga tend to be inhibited, the photosynthetic activity of alga decrease but the intracellular nitrite concentration and MDA content were higher in low CO_2 group in contrast with high CO_2 group respectively. In the same way, in the medium with only nitrite, the growth of alga also increased with intracellular nitrite content decreasing. These results showed the increase of intracellular nitrite content was the main reason of the inhibition of growth of alga. By observing the change of intracellular nitrite content and the nitrite concentration in medium containing 0 mg NO_2~--N L~(-1) :240mg NO_3~--N L~(-1) and in medium with 30mg NO_2~--N L~(-1) :240mg NO_3~--N L~(-1), we found there existed nitrite moving in alga and nitrite discharging from alga simultaneously. When the nitrite concentration being high in medium, the nitrite moving in alga was more than nitrite discharging from alga, as a result, the intracellular nitrite content increased. In contrast, the nitrite concentration of medium increased.
Compared to the sole culturing, under the co-culturing condition the growth of M. aeruginosa and Scendesmus quadricauda were inhibited, indicated that there existed competition between M. aeruginosa and S. quadricauda. In co-culturing condition, with nitrite concentration increasing, S. quadricauda became more dominant in competition, and when nitrite concentration of 30mg/L, the growth of M. aeruginosa was inhibited entirely. The examination of mutual allelopathy showed mutual allelopathy was intensified with the increase of nitrite. Therefore, the result of competition of M. aeruginosa and Scendesmus quadricauda was due to under high nitrite concentration (20mg/L、30mg/L) the worse damage of M. aeruginosa and more intense allelopathy would be revealed.
在亚硝酸盐单独存在的条件下,浓度小于5mg/L的亚硝酸盐,对铜绿微囊藻的生长没有明显影响,藻能够以亚硝酸盐为氮源。随着培养基中亚硝酸盐浓度逐渐升高到20mg/L,藻的生长受到抑制直至消亡,此时,铜绿微囊藻的光合速率下降,细胞MDA含量增加,细胞内亚硝酸盐含量增加,表明亚硝酸盐通过抑制铜绿微囊藻的光合作用和升高藻体内氧自由基浓度的方式抑制藻的生长。而藻的CAT酶活性、亚硝酸还原酶活性和藻叶绿素a含量增加可能是藻的一种自我保护措施。
硝酸盐存在的条件下(240mg/L),铜绿微囊藻在亚硝酸盐浓度为30mg/L的培养基中依然可以生长。而且,在相同的亚硝酸盐浓度条件下,含硝酸盐培养基中的藻相比只含亚硝酸盐培养基中的藻,有更高的光合速率,更小的MDA含量和藻细胞内亚硝酸盐含量。亚硝酸盐的吸收试验也显示硝酸盐存在的条件下,藻对亚硝酸盐吸收的半饱和常数(K_S)增加,这些结果表明硝酸盐能够抑制藻对亚硝酸盐的吸收,从而减少藻细胞内的亚硝酸盐含量,导致亚硝酸盐对藻抑制的减弱。在硝酸盐和亚硝酸盐共存的培养基中,我们还观察到亚硝酸盐被氧化为硝酸盐,表明此条件下可以诱导铜绿微囊藻生成亚硝酸氧化还原酶。
铵盐存在的培养基中(20mg/L),2mg/L的亚硝酸盐添加,对铜绿微囊藻的生长表现出促进作用,藻的光合速率和细胞内亚硝酸盐含量增加,MDA含量下降。进一步增加亚硝酸盐浓度(4、7、10mg/L),藻的生长受到抑制直至消亡,同时观察到藻的光合速率下降,细胞内亚硝酸盐含量和MDA含量增加。亚硝酸盐的吸收试验显示铵盐存在的条件下,藻对亚硝酸盐吸收的半饱和常数(K_S)减少,表明铵盐能够刺激藻对亚硝酸盐的吸收,增加藻细胞内的亚硝酸盐含量。在一定的范围内,细胞亚硝酸盐含量的增加可以促进藻的生长,但是过高的细胞内亚硝酸盐含量会抑制藻的生长。
硝酸盐浓度为150mg/L和300mg/L的处理组中,相比低CO_2组,高CO_2组藻的生长更好,光合速率增加,细胞内亚硝酸盐含量和MDA含量减少。而硝酸盐浓度为50mg/L的处理组中,高CO_2组和低CO_2组中,藻的生长变化不明显,光合速率、细胞内亚硝酸盐含量和MDA含量都没有明显变化。而且在相同的亚硝酸盐浓度条件下,藻细胞内亚硝酸盐含量的降低,导致培养基中藻生长的增加,这些结果表明细胞内的亚硝酸盐可能是影响藻生长的主要原因。通过对NO_2~--N:NO_3~--N为0:240和30mg/L:240mg/L两个处理组中细胞内亚硝酸盐含量和培养基中硝酸盐浓度变化的30d观察,发现培养基中亚硝酸盐进入藻体和藻排放亚硝酸盐是同时存在的,当培养基中亚硝酸盐浓度过高时,进入藻体的亚硝酸盐大于藻排放的,导致细胞内亚硝酸盐含量增加。相反,则培养基中亚硝酸盐浓度增加。
相比单独培养时,混合培养的条件下,铜绿微囊藻和四尾栅藻的生长都受到抑制,但是四尾栅藻的最大生物量与单独培养时藻的最大生物量的比值大于铜绿微囊藻,而且随着亚硝酸盐浓度的增加这种差距越明显,亚硝酸盐浓度为30mg/L时,铜绿微囊藻完全被抑制。表明两种藻之间存在着竞争,四尾栅藻在竞争中更具优势,亚硝酸盐浓度的升高增强了四尾栅藻的竞争优势。通过两种藻的相互化感作用试验,发现两种藻之间存在着化感作用,而且亚硝酸盐浓度增加化感作用增强。这些现象表明上述的竞争结果是化感作用和亚硝酸盐对藻的毒害作用共同造成的。
Phytoplankton account for approximately 50% of global primary production, and Microcystis aeruginosa blooms are dominant in fresh water. The increase of N and P is the main reason for the explosive growth of phytoplankton. During the water blooms, the concentration of CO_2 and dissolved O_2 decrease and pH increases, this could result in the nitrite accumulation by inhibiting oxidation of nitrite to nitrate or reduction of nitrite to ammonium. On the other hand, human activities and industrialization also lead to an increased accumulation of nitrite in the environment. In this study, the effects of nitrite on the growth and physiology of Microcystis aeruginosa were investigated when nitrite as sole nitrogen source, and under the present of nitrate condition or the ammonium condition; The impacts of intracellular nitrite content change on Microcystis aeruginosa growth were studied, and the competition of Microcystis aeruginosa and Scendesmus quadricauda under different nitrite concentrations was analysed..
When nitrite as sole nitrogen source, Microcystis aeruginosa could take up nitrite for growth, and the growth of alga was not impacted in medium with nitrite concentration below 5mg/L. However, with nitrite concentrations increasing to 20mg/L, the growth of alga was inhibited and photosynthetic activity decreased. Meanwhile, the content of MDA and intracellular nitrite increased. These indicated nitrite inhibited the growth of Microcystis aeruginosa by decreasing photosynthetic activity and increasing the content of oxygen free radical. The increase of chlorophyll a content、CAT activity and nitrite reductase activity might be an adaptive mechanism of alga under high nitrite condition.
When both nitrate and nitrite as nitrogen source, the Microcystis aeruginosa could grow under nitrite concentration of 30 mg/L. Furthermore, compared to medium with only nitrite, the alga had higher photosynthetic activity and less the content of MDA and intracellular nitrite in medium with both nitrate and nitrite under high nitrite concentration condition. The examination of nitrite uptake also showed the half-saturation constant of alga increased when nitrate appearance. These results revealed that nitrate could inhibit the nitrite uptake, which resulted in the decrease of the intracellular nitrite concentration and the inhibition of nitrite on alga growth weakening. In medium containing both nitrate and nitrite, the nitrite could be oxidized to nitrate, suggesting nitrite oxidoreductase has been conduced.
When both ammonium and nitrite as nitrogen source, in medium containing nitrite concentration of 2 mg/L, the growth of alga increased, the photosynthetic activities and intracellular nitrite content also showed increase but MDA content decreased compared the medium of nitrite concentration of 0mg/L. With the nitrite concentrations further increasing to 10mg/L, the growth of alga slowed down and even ceased, the photosynthetic activity decreased but the content of intracellular nitrite and MDA increased. The examination of nitrite uptake showed the half-saturation constant of alga decreased when ammonium appearance, indicating ammonium could promote the uptake of nitrite and increase the intracellular nitrite content. In the range of low intracellular nitrite content, the increase of intracellular nitrite could promote the growth of alga, however, too high intracellular nitrite content could inhibit the growth of alga.
At nitrate concentration of 50mg/L, the growth of alga hadn't significant change in high CO_2 group and low CO_2 group, the intracellular nitrite concentration、the photosynthetic activity and MDA content also remained constant. However, at nitrate concentration of 150mg/L and 300mg/L, the growth of alga tend to be inhibited, the photosynthetic activity of alga decrease but the intracellular nitrite concentration and MDA content were higher in low CO_2 group in contrast with high CO_2 group respectively. In the same way, in the medium with only nitrite, the growth of alga also increased with intracellular nitrite content decreasing. These results showed the increase of intracellular nitrite content was the main reason of the inhibition of growth of alga. By observing the change of intracellular nitrite content and the nitrite concentration in medium containing 0 mg NO_2~--N L~(-1) :240mg NO_3~--N L~(-1) and in medium with 30mg NO_2~--N L~(-1) :240mg NO_3~--N L~(-1), we found there existed nitrite moving in alga and nitrite discharging from alga simultaneously. When the nitrite concentration being high in medium, the nitrite moving in alga was more than nitrite discharging from alga, as a result, the intracellular nitrite content increased. In contrast, the nitrite concentration of medium increased.
Compared to the sole culturing, under the co-culturing condition the growth of M. aeruginosa and Scendesmus quadricauda were inhibited, indicated that there existed competition between M. aeruginosa and S. quadricauda. In co-culturing condition, with nitrite concentration increasing, S. quadricauda became more dominant in competition, and when nitrite concentration of 30mg/L, the growth of M. aeruginosa was inhibited entirely. The examination of mutual allelopathy showed mutual allelopathy was intensified with the increase of nitrite. Therefore, the result of competition of M. aeruginosa and Scendesmus quadricauda was due to under high nitrite concentration (20mg/L、30mg/L) the worse damage of M. aeruginosa and more intense allelopathy would be revealed.
引文
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