蓝藻水华与2种藻食性水生动物的相互作用
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摘要
在营养盐管理难以奏效的富营养化水体,运用生物操纵原理,通过水生生物群落控制管理来防治蓝藻水华被认为是一种安全、有效的替代方法。为此,藻食性动物与蓝藻水华之间的相互作用关系值得进一步关注。本文以铜锈环棱螺(Bellamya aeruginosa)和尼罗罗非鱼(Oreochromis niloticus)为主要研究对象,研究了微囊藻毒素(MC)在这2种藻食性动物体内的积累及清除规律,进行了产毒铜绿微囊藻(Microcystis aeruginosa)喂食暴露对螺肝脏几种主要酶的影响及对螺肝细胞DNA损伤的系列室内试验,采用微宇宙试验考察了环棱螺对藻华水体的短期生态效应。主要结果如下:
1.在藻华池塘中,环棱螺体内MC含量有明显季节变化,螺体内MC含量与浮游植物细胞内毒素呈显著的正相关,而与溶解于水中的MC浓度无显著相关。尽管低温季节水体MC含量很低,但螺肝胰腺内仍保持较高的MC浓度(1.023-1.887μg/gDW)。螺体内不同组织MC含量有显著差异(P<0.01),依次为肝胰腺>消化道>鳃>腹足。室内喂食暴露试验表明,温度对螺肝胰腺MC的积累有显著影响(P<0.01),15℃时螺肝胰腺MC含量显著高于25℃的相同处理组;分别用单一产毒铜绿微囊藻(蓝藻组)和产毒铜绿微囊藻+栅藻(混合组)对环棱螺进行喂食暴露后发现,尽管混合组饵料中MC含量仅为蓝藻组的1/2左右,但25℃条件下,混合组螺肝胰腺MC含量显著高于蓝藻组(P<0.05)。将喂食暴露后的螺体移入不含藻毒素的栅藻悬浮液进行清除试验时发现,螺肝胰腺中的MC在前10天都得到快速清除,而10天后清除速率明显下降。15℃清除试验后期螺肝胰腺MC残留量要显著高于25℃的MC含量(P<0.05)。
2.微囊藻喂食暴露及毒素清除期间,螺肝脏中酸性磷酸酶(ACP)、碱性磷酸酶(ALP)活性均随螺肝脏中MC含量变化发生相应变化,谷胱甘肽硫转移酶(GST)活性在混合组先被诱导后被抑制,在蓝藻组初期变化不明显后期则表现为诱导。MC清除过程中,ACP、ALP和GST活性均随MC含量下降表现出不同程度的降低。15℃水温条件下,微囊藻喂食暴露对螺肝脏抗氧化酶超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和谷胱甘肽过氧化物酶(GPX)的活性影响在混合组均表现为抑制,在蓝藻组均表现为诱导;25℃时SOD和CAT活性在2个处理组中均表现为诱导,而GPX酶活性均表现为抑制。试验表明,几种螺肝脏抗氧化酶活性对MC暴露的响应与MC浓度、环境温度、暴露时间均有一定的关系。
3.利用彗星试验研究了产毒微囊藻短期喂食暴露(24h)对螺肝细胞DNA的损伤效应。结果表明,混合组和蓝藻组均在暴露试验开始后6h螺肝脏中MC含量达到最高,而后MC含量下降;12h重复染毒(调整藻类密度)后,螺肝脏中MC含量又迅速回升。同期螺肝细胞DNA损伤指标彗星尾长(TL)、彗星尾距(TM)和彗星尾部DNA百分含量(Tail DNA%)也随螺肝脏中MC含量发生相应变化。实验期间,混合组和蓝藻组的DNA损伤指标均显著高于对照组,混合组显著高于蓝藻组。
4.采用室外微宇宙试验研究了不同密度环棱螺(高密度组200ind./100L;中密度组100ind./100L;低密度组50ind./100L)对藻华水体的短期(10d)生态效应。结果表明,水体溶氧和pH均随螺投放密度的增加而下降,不同处理组之间的氮盐浓度没有显著差异,而正磷酸浓度存在极显著差异(p<0.01),螺类密度越高正磷酸浓度上升越多。中、高密度组水体内蓝藻水华被完全遏制,蓝藻和藻类总生物量显著下降,绿藻和隐藻比例上升;而藻类多样性指数呈现出随螺类密度增加而下降的趋势。螺的投放对浮游动物的群落结构也产生了显著的影响,试验结束时,中、高密度组轮虫数量显著低于对照组和低密度组,浮游动物多样性指数则与螺投放密度呈显著正相关(r=0.9834)。
5.实验条件下,罗非鱼对铜绿微囊藻的摄食率随鱼体重及水温的升高而增加;25℃时罗非鱼对微囊藻的消化率变化于58.6%至78.1%之间,平均消化率达67.5±6.41%。微囊藻喂食暴露试验表明,蓝藻组和混合组罗非鱼体内的MC含量没有显著差异(P>0.05),两处理组均在喂食暴露第21天后,鱼体内MC含量基本达到稳定。MC在鱼不同组织的分布存在显著差异,鱼肝脏和肠道MC含量较为接近,但均极显著地高于肌肉中的含量(P<0.01)。MC清除试验表明,罗非鱼肠道MC的清除速率最大,肝脏其次,肌肉最小。
The prevention and treatment of cyanobacterial bloom by the principle of biomanipulation controlling aquatic community is considered as a safe and effective alternative method in water body that the management of nutrient would be difficult to prove effective. Therefore, more attention should be paid to the interaction between cyanobacterial bloom and phytoplanktivorous animal. This thesis, with Bellamya aeruginosa and Oreochromis niloticus as the main research object, researched the regulation of accumulation and degradation of microcystin in these two kinds of phytoplanktivorous animals. A series of laboratory tests are conducted to study the influence of toxic Microcystis aeruginosa on the six kinds of enzyme of lives and DNA damage in the hepatocellular of snail after feeding exposure. The microcosm experiment is employed to investigate the short-term ecological effect of Bellamya on the bloom water body. The main results are summarized as follows:
1. In the bloom pond, MC in the body of Bellamya exhibited an obvious seasonal change, and a significant positive correlation was found between the contents of MC in snail and the intracellular microcystin (IMC) of phytoplankton, but there was no significant correlation between tissues and the extracellular Microcystin (EMC). Although the concentration of MC in water body was not high in low temperature season, the value of MC in hepatopancreas remained at 1.023-1.887μg/gDW. The contents of MC in various tissues of animals showed a significant difference (P<0.01), and the amount of MC accumulation was in sequence of hepatopancreas> digestive tract> branchia> abdominal foot. The feeding exposed experiment indicated that the temperature had a significant effect on the accumulation of MC in hepatopancreas (P<0.01): the concentration of MC in hepatopancreas at 15℃was higher than that at 25℃. It was found that, at 25℃, although MC in the mixed diet was about half of BG treatment, the value of MC in hepatopancreas of snails fed by toxic M. aeruginosa + Scenedesmus (mixed) was significantly higher than that fed by toxic M. aeruginosa (BG) (P<0.05). When the degradation experiment was carried out to probe into the dynamics of MC of the tissues in the suspension without MC after feeding exposure, it was found that MC in hepatopancreas had been cleared quickly in the first ten days after removal, but the clearance rate decreased obviously after that. At 15℃the residual amount of MC in hepatopancreas was significantly higher than it was 25℃at the later stage of the degradation experiment (P<0.05).
2. According to the dynamics of MC in liver, the activities of acid phosphatases (ACP) and alkaline phosphatases (ALP) change correspondingly during the period of feeding exposure and clearance. The activities of glutathione-S-transferase (GST) was induced initially and then inhibited in the mixed treatment, the change of initial activity of GST was not obvious in the BG treatment, but increased remarkably at the late stage of the exposure. The activity of these enzymes decreased in different degrees with the concentration of MC lowering during the degradation of MC. The activities of antioxidant enzymes superoxide dismutase (SOD), catalase (CAT) and glutathione peroxidase (GPX) in liver of snail were inhibited in the mixed treatment and induced in BG treatment at 15℃, whereas the SOD and CAT activities were all induced but the GPX activities was inhibited in both treatments at 25℃. The tests indicate that the response of activities of antioxidant enzymes to MC exposure may have a correlation with the concentration of MC, ambient temperature and the duration of exposure.
3. The single cell gel electrophoresis technique (Comet assay) was employed to study the damage effects of short-term feeding toxin Microcystis (24h) on DNA of hepatocyte from snail. The results showed that the MC content in the liver of B. aeruginosa all reached the maximum after 6h exposure in both experimental treatments, and the MCs content decreased afterwards. The MCs content rose again rapidly after 12h exposure (adjusting density of alga). Meanwhile, the comet tail length (TL), tail moment (TM) and tail DNA% in tissues showed a time-dependent and a dose-dependent change in response to the MC content. In the mixed treatment, the parameters of DNA damage were significantly higher than that in BG treatment, both experimental treatments higher than that in the controlled group during the experimental period.
4. The microcosm experiment was conducted to investigate the short-term (10d) ecological effect of different densities Bellamya (Higher-density: 200ind./100L, Middle-density: 100ind./100L, Lower-density: 50ind./100L) on the bloom water body. The results showed that dissolved oxygen (DO) and pH decreased with the increasing density of animal, the concentration of nitrogen had no significant difference among the different treatments, there was a significant difference in the concentration of (PO43+) among the three groups (p<0.01), and the higher the concentration of PO43+ was, the faster the density of orthophosphoric acid increased. The cyanobacterial bloom were completely controlled in the higher- and middle-density treatments, the biomass of cyanophyta and phytoplankton decreased significantly, the proportion of chlorophyta and cryptophyta increased, the index of phytoplankton diversity showed a decreasing trend with the increasing density of animal. The occurrence of Bellamya also produced a prodigious influence on the community structure of zooplankton. The quantity of rotifer in the high- and middle-density treatments was significantly lower than in the controlled and low-density treatments at the later stage of the experiment. A significant positive correlation was found between the index of zooplankton diversity and the density of animal (r=0.9834).
5. The feeding rate of tilapia on M. aeruginosa increased with the increase of body weight and the water temperature under the experimental condition. At 25℃, the digestibility of tilapia varied from 58.6% to 78.1%, and the mean digestibility was 67.5±6.41%. The feeding exposure indicated that there was no significant difference between BG and mixed treatment in the content of MC in fish body (P>0.05), the MC content in both treatments reached stability after 21d exposure basically. There were significant differences in the distribution of MC in different fish tissues. Although the liver and the intestine have the similar MC, they have a significantly higher content than the muscle (P<0.01). The MC degradation experiment indicated that the clearance rate of MC was in the sequence of in intestine >in liver>in muscle.
1.在藻华池塘中,环棱螺体内MC含量有明显季节变化,螺体内MC含量与浮游植物细胞内毒素呈显著的正相关,而与溶解于水中的MC浓度无显著相关。尽管低温季节水体MC含量很低,但螺肝胰腺内仍保持较高的MC浓度(1.023-1.887μg/gDW)。螺体内不同组织MC含量有显著差异(P<0.01),依次为肝胰腺>消化道>鳃>腹足。室内喂食暴露试验表明,温度对螺肝胰腺MC的积累有显著影响(P<0.01),15℃时螺肝胰腺MC含量显著高于25℃的相同处理组;分别用单一产毒铜绿微囊藻(蓝藻组)和产毒铜绿微囊藻+栅藻(混合组)对环棱螺进行喂食暴露后发现,尽管混合组饵料中MC含量仅为蓝藻组的1/2左右,但25℃条件下,混合组螺肝胰腺MC含量显著高于蓝藻组(P<0.05)。将喂食暴露后的螺体移入不含藻毒素的栅藻悬浮液进行清除试验时发现,螺肝胰腺中的MC在前10天都得到快速清除,而10天后清除速率明显下降。15℃清除试验后期螺肝胰腺MC残留量要显著高于25℃的MC含量(P<0.05)。
2.微囊藻喂食暴露及毒素清除期间,螺肝脏中酸性磷酸酶(ACP)、碱性磷酸酶(ALP)活性均随螺肝脏中MC含量变化发生相应变化,谷胱甘肽硫转移酶(GST)活性在混合组先被诱导后被抑制,在蓝藻组初期变化不明显后期则表现为诱导。MC清除过程中,ACP、ALP和GST活性均随MC含量下降表现出不同程度的降低。15℃水温条件下,微囊藻喂食暴露对螺肝脏抗氧化酶超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和谷胱甘肽过氧化物酶(GPX)的活性影响在混合组均表现为抑制,在蓝藻组均表现为诱导;25℃时SOD和CAT活性在2个处理组中均表现为诱导,而GPX酶活性均表现为抑制。试验表明,几种螺肝脏抗氧化酶活性对MC暴露的响应与MC浓度、环境温度、暴露时间均有一定的关系。
3.利用彗星试验研究了产毒微囊藻短期喂食暴露(24h)对螺肝细胞DNA的损伤效应。结果表明,混合组和蓝藻组均在暴露试验开始后6h螺肝脏中MC含量达到最高,而后MC含量下降;12h重复染毒(调整藻类密度)后,螺肝脏中MC含量又迅速回升。同期螺肝细胞DNA损伤指标彗星尾长(TL)、彗星尾距(TM)和彗星尾部DNA百分含量(Tail DNA%)也随螺肝脏中MC含量发生相应变化。实验期间,混合组和蓝藻组的DNA损伤指标均显著高于对照组,混合组显著高于蓝藻组。
4.采用室外微宇宙试验研究了不同密度环棱螺(高密度组200ind./100L;中密度组100ind./100L;低密度组50ind./100L)对藻华水体的短期(10d)生态效应。结果表明,水体溶氧和pH均随螺投放密度的增加而下降,不同处理组之间的氮盐浓度没有显著差异,而正磷酸浓度存在极显著差异(p<0.01),螺类密度越高正磷酸浓度上升越多。中、高密度组水体内蓝藻水华被完全遏制,蓝藻和藻类总生物量显著下降,绿藻和隐藻比例上升;而藻类多样性指数呈现出随螺类密度增加而下降的趋势。螺的投放对浮游动物的群落结构也产生了显著的影响,试验结束时,中、高密度组轮虫数量显著低于对照组和低密度组,浮游动物多样性指数则与螺投放密度呈显著正相关(r=0.9834)。
5.实验条件下,罗非鱼对铜绿微囊藻的摄食率随鱼体重及水温的升高而增加;25℃时罗非鱼对微囊藻的消化率变化于58.6%至78.1%之间,平均消化率达67.5±6.41%。微囊藻喂食暴露试验表明,蓝藻组和混合组罗非鱼体内的MC含量没有显著差异(P>0.05),两处理组均在喂食暴露第21天后,鱼体内MC含量基本达到稳定。MC在鱼不同组织的分布存在显著差异,鱼肝脏和肠道MC含量较为接近,但均极显著地高于肌肉中的含量(P<0.01)。MC清除试验表明,罗非鱼肠道MC的清除速率最大,肝脏其次,肌肉最小。
The prevention and treatment of cyanobacterial bloom by the principle of biomanipulation controlling aquatic community is considered as a safe and effective alternative method in water body that the management of nutrient would be difficult to prove effective. Therefore, more attention should be paid to the interaction between cyanobacterial bloom and phytoplanktivorous animal. This thesis, with Bellamya aeruginosa and Oreochromis niloticus as the main research object, researched the regulation of accumulation and degradation of microcystin in these two kinds of phytoplanktivorous animals. A series of laboratory tests are conducted to study the influence of toxic Microcystis aeruginosa on the six kinds of enzyme of lives and DNA damage in the hepatocellular of snail after feeding exposure. The microcosm experiment is employed to investigate the short-term ecological effect of Bellamya on the bloom water body. The main results are summarized as follows:
1. In the bloom pond, MC in the body of Bellamya exhibited an obvious seasonal change, and a significant positive correlation was found between the contents of MC in snail and the intracellular microcystin (IMC) of phytoplankton, but there was no significant correlation between tissues and the extracellular Microcystin (EMC). Although the concentration of MC in water body was not high in low temperature season, the value of MC in hepatopancreas remained at 1.023-1.887μg/gDW. The contents of MC in various tissues of animals showed a significant difference (P<0.01), and the amount of MC accumulation was in sequence of hepatopancreas> digestive tract> branchia> abdominal foot. The feeding exposed experiment indicated that the temperature had a significant effect on the accumulation of MC in hepatopancreas (P<0.01): the concentration of MC in hepatopancreas at 15℃was higher than that at 25℃. It was found that, at 25℃, although MC in the mixed diet was about half of BG treatment, the value of MC in hepatopancreas of snails fed by toxic M. aeruginosa + Scenedesmus (mixed) was significantly higher than that fed by toxic M. aeruginosa (BG) (P<0.05). When the degradation experiment was carried out to probe into the dynamics of MC of the tissues in the suspension without MC after feeding exposure, it was found that MC in hepatopancreas had been cleared quickly in the first ten days after removal, but the clearance rate decreased obviously after that. At 15℃the residual amount of MC in hepatopancreas was significantly higher than it was 25℃at the later stage of the degradation experiment (P<0.05).
2. According to the dynamics of MC in liver, the activities of acid phosphatases (ACP) and alkaline phosphatases (ALP) change correspondingly during the period of feeding exposure and clearance. The activities of glutathione-S-transferase (GST) was induced initially and then inhibited in the mixed treatment, the change of initial activity of GST was not obvious in the BG treatment, but increased remarkably at the late stage of the exposure. The activity of these enzymes decreased in different degrees with the concentration of MC lowering during the degradation of MC. The activities of antioxidant enzymes superoxide dismutase (SOD), catalase (CAT) and glutathione peroxidase (GPX) in liver of snail were inhibited in the mixed treatment and induced in BG treatment at 15℃, whereas the SOD and CAT activities were all induced but the GPX activities was inhibited in both treatments at 25℃. The tests indicate that the response of activities of antioxidant enzymes to MC exposure may have a correlation with the concentration of MC, ambient temperature and the duration of exposure.
3. The single cell gel electrophoresis technique (Comet assay) was employed to study the damage effects of short-term feeding toxin Microcystis (24h) on DNA of hepatocyte from snail. The results showed that the MC content in the liver of B. aeruginosa all reached the maximum after 6h exposure in both experimental treatments, and the MCs content decreased afterwards. The MCs content rose again rapidly after 12h exposure (adjusting density of alga). Meanwhile, the comet tail length (TL), tail moment (TM) and tail DNA% in tissues showed a time-dependent and a dose-dependent change in response to the MC content. In the mixed treatment, the parameters of DNA damage were significantly higher than that in BG treatment, both experimental treatments higher than that in the controlled group during the experimental period.
4. The microcosm experiment was conducted to investigate the short-term (10d) ecological effect of different densities Bellamya (Higher-density: 200ind./100L, Middle-density: 100ind./100L, Lower-density: 50ind./100L) on the bloom water body. The results showed that dissolved oxygen (DO) and pH decreased with the increasing density of animal, the concentration of nitrogen had no significant difference among the different treatments, there was a significant difference in the concentration of (PO43+) among the three groups (p<0.01), and the higher the concentration of PO43+ was, the faster the density of orthophosphoric acid increased. The cyanobacterial bloom were completely controlled in the higher- and middle-density treatments, the biomass of cyanophyta and phytoplankton decreased significantly, the proportion of chlorophyta and cryptophyta increased, the index of phytoplankton diversity showed a decreasing trend with the increasing density of animal. The occurrence of Bellamya also produced a prodigious influence on the community structure of zooplankton. The quantity of rotifer in the high- and middle-density treatments was significantly lower than in the controlled and low-density treatments at the later stage of the experiment. A significant positive correlation was found between the index of zooplankton diversity and the density of animal (r=0.9834).
5. The feeding rate of tilapia on M. aeruginosa increased with the increase of body weight and the water temperature under the experimental condition. At 25℃, the digestibility of tilapia varied from 58.6% to 78.1%, and the mean digestibility was 67.5±6.41%. The feeding exposure indicated that there was no significant difference between BG and mixed treatment in the content of MC in fish body (P>0.05), the MC content in both treatments reached stability after 21d exposure basically. There were significant differences in the distribution of MC in different fish tissues. Although the liver and the intestine have the similar MC, they have a significantly higher content than the muscle (P<0.01). The MC degradation experiment indicated that the clearance rate of MC was in the sequence of in intestine >in liver>in muscle.
引文
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