大型海藻对有害赤潮微藻克生效应的实验生态学研究
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摘要
随着沿海水域富营养化的加剧,赤潮在世界范围内的沿海频繁爆发,已成为一种严重的海洋灾害。自上世纪90年代以来,我国赤潮的发生呈现出“大范围、高频率、多种类、快节奏”的特点,并有连年增加的态势,这对原本已经非常严峻的近岸水域生态环境和沿岸水产业的可持续发展造成了极大的威胁。采用切实有效的赤潮防治措施是赤潮研究的重要内容之一。许多学者提出了一些有效的方法,包括物理法(絮凝法,如粘土)、化学法(药剂法,如硫酸酮)和生物防治法(微生物,如细菌、病毒)等治理有害藻华。虽然这些方法在短期内可能有效,但实际应用的尚少,但总体来说,物理和化学法都是治标不治本,只能暂时杀死或除掉水体中的赤潮生物(甚至其它生物),并不能为水体中的营养物质提供一个输出渠道,因此也就无法防止赤潮的再次发生。鉴于添加外来物质可能对海洋生态系统产生可知或不可预见的影响,利用海洋环境中的生态因子进行赤潮的防控已经越来越引起人们的重视,在这种情况下,如果能找到一种既能抑制赤潮微藻生长又能吸收水体中营养盐的经济型水生生物,达到治标治本,应是最为理想的赤潮防治途径。其中利用大型海藻与微藻间的相互作用来预防或控制赤潮是一个新的研究方向。
本文研究三种代表性的大型海藻孔石莼(Ulva pertusa) (绿藻)、小珊瑚藻(Corallina pilulifera) (红藻)和鼠尾藻(Sargassum thunbergii) (褐藻)对三种赤潮微藻赤潮异弯藻(Heterosigma akashiwo)、亚历山大藻(Alexandrium tamarense )和中肋骨条藻(Skeletonema costatum)的克生作用,并初步提取、分离和鉴定了小珊瑚藻和鼠尾藻组织内抑制有害赤潮微藻生长的克生物质,探讨了小珊瑚藻和鼠尾藻对有害赤潮微藻克生作用的化学物质本质,为利用沿海区域的各种大型海藻来治理赤潮生物提供了科学依据。主要实验结果如下:
1.三种大型海藻对三种有害赤潮微藻克生效应的比较研究
通过排除营养盐、光照和pH等因素的影响,利用共培养系统研究了大型海藻孔石莼、小珊瑚藻和鼠尾藻的新鲜组织、干粉末、水溶性组织抽提液及培养水过滤液对三种有害赤潮微藻赤潮异弯藻、亚历山大藻及中肋骨条藻生长的作用,共存的实验结果表明,三种大型海藻的新鲜组织和干粉末对三种有害赤潮微藻的生长均有强烈的抑制作用,在较高的浓度下还能完全杀死赤潮异弯藻细胞和中肋骨条藻细胞,证明了三种大型海藻对三种有害赤潮微藻具有克生效应,并且大型海藻对有害赤潮微藻的克生效应具有普遍性。各个浓度孔石莼、小珊瑚藻和鼠尾藻水溶性抽提液对三种有害赤潮微藻的生长皆有抑制作用,抑制作用随浓度的升高而加强,这表明大型海藻组织中具有抑制有害赤潮微藻生长的克生物质,这进一步证明克生物质的存在是三种大型海藻对三种有害赤潮微藻产生克生效应的主要方式之一。三种有害赤潮微藻在一次性三种大藻培养水过滤液添加方式下生长没有受到显著抑制,而在半连续孔石莼及鼠尾藻培养水添加方式下其生长均受到了显著抑制,表明这三种大型海藻新鲜组织能够向环境中释放克生物质来抑制有害赤潮微藻的生长,同时也说明了大型海藻所分泌的克生物质容易降解。三种大型海藻新鲜组织对赤潮异弯藻和中肋骨条藻的克生效应由强到弱依次为:孔石莼>小珊瑚藻>鼠尾藻、对亚历山大藻的克生效应为鼠尾藻>孔石莼>小珊瑚藻;三种大型海藻干粉末对赤潮异弯藻的克生效应由强到弱依次为:鼠尾藻>孔石莼>小珊瑚藻,对中肋骨条藻和亚历山大藻的克生效应为:孔石莼>小珊瑚藻>鼠尾藻,表明大型海藻对有害赤潮微藻的克生效应具种的差异性。
2.小珊瑚藻和鼠尾藻组织提取物对赤潮异弯藻和亚历山大藻的生长抑制作用
以小珊瑚藻和鼠尾藻为大型海藻的代表,以赤潮异弯藻和亚历山大藻为目标微藻,研究了不同浓度的小珊瑚藻和鼠尾藻组织蒸馏水及4种具有不同极性的有机溶剂(极性从高到低:甲醇>丙酮>乙醚>氯仿)提取物对两种有害赤潮微藻的生长抑制作用。两种大型海藻组织甲醇提取物在5种提取物中具有最高的生长抑制活性。在相对较高浓度的此种提取物作用下,两种有害赤潮微藻均完全死亡。较高浓度的小珊瑚藻和鼠尾藻组织蒸馏水提取物对两种有害赤潮微藻的生长具有显著的抑制作用,但是在最高浓度下也不能使这两种有害赤潮微藻完全死亡。另外三种有机溶剂的提取物对两种有害赤潮微藻的生长无明显的影响。结果表明,小珊瑚藻和鼠尾藻组织中含有对赤潮异弯藻和亚历山大藻的生长有抑制作用的活性物质,且这些物质具有相对较高的极性。对两种大型海藻的甲醇提取物进行了液液萃取,然后将其分离为石油醚相、乙酸乙酯相、正丁醇相和蒸馏水相。并用这4相对两种有害赤潮微藻进行克生物质的生物活性检测,结果表明石油醚相和乙酸乙酯相具有较强的杀藻活性,脂肪酸可能为小珊瑚藻和鼠尾藻组织内克生物质的主要成分。
3.小珊瑚藻和鼠尾藻组织内克生物质的鉴定
通过对小珊瑚藻和鼠尾藻的石油醚相和乙酸乙酯相进行气相色谱——质谱(GC/MS)联用分析,并核对标准图谱和有关参考文献,鉴定出了12种脂肪酸,其中不饱和脂肪酸占大部分。对小珊瑚藻的鉴定中得出,其中饱和脂肪酸占20.81%,而不饱和脂肪酸成分占65.45%,对鼠尾藻的鉴定中得出,饱和脂肪酸占18.82%,而不饱和脂肪酸成分占58.62%。纯化合物对赤潮异弯藻和亚历山大藻的活性检测结果表明,9,12,15-十八碳三烯酸、6,9,12,15-十八碳四烯酸、5Z,8Z,11Z,14Z-二十碳四烯酸、5Z,11Z,14Z,17Z-二十碳四烯酸和5Z,8Z,11Z,14Z,17Z-二十碳五烯酸具有很强的杀藻活性。
小珊瑚藻和鼠尾藻组织中含有多种不饱和脂肪酸,其中一些不饱和脂肪酸对赤潮异弯藻和亚历山大藻具有强烈的杀藻效果,而且不饱和程度越高其克生效果越明显。这就表明不饱和脂肪酸可能是小珊瑚藻和鼠尾藻组织内抑制赤潮异弯藻和亚历山大藻生长的克生物质中的重要组成成分之一。
4.温度、盐度、光照、pH值对小珊瑚藻克制赤潮异弯藻及鼠尾藻克制亚历山大藻效果和两种有害赤潮微藻生长的影响
本实验共设置了4个环境因子:温度(T)、盐度(S)、光照(L)和pH值(P)。每两个不同的因子之间分别设计双因子实验,共6个处理组合,即温度和盐度、温度和光照、温度和pH值、光照和pH值及盐度和光照,以检验这4个因子对小珊瑚藻克制赤潮异弯藻和鼠尾藻克制亚历山大藻效果及对两种赤潮微藻生长的影响,得出了赤潮异弯藻和亚历山大藻的生长率及大藻克制有害赤潮微藻生长的抑制率,并且检验这4个因子两两之间是否存在交互作用。每个因子分别设置4个处理水平,即:温度(15、20、25和30°C)、盐度(10、20、30和40)、光照(20、100、200和400μmol m~(-2)s~(-1))和pH值(5.5、7、8.5、10)。实验结果表明,温度、盐度、光照和pH值这四个环境因子均能显著影响赤潮异弯藻和亚历山大藻的生长,并且能显著影响小珊瑚藻克制赤潮异弯藻以及鼠尾藻克制亚历山大藻的效果。小珊瑚藻克制赤潮异弯藻以及鼠尾藻克制亚历山大藻的最高抑制效果不是发生在有害赤潮微藻单养时的最适条件,而是发生在环境因子胁迫时,较低的温度(15°C)、较低的盐度(10)、较高的光照强度(400μmol m~(-2)s~(-1))和高pH值(10)均能增强小珊瑚藻克制赤潮异弯藻以及鼠尾藻克制亚历山大藻的效果。
Severe outbreaks of harmful algal blooms (hereafter, HABs) cause serious problems with regard to effective utilization of water resources such as fisheries and water-supply reservoirs, moreover, about 2000 cases of human poisoning resulting from algal toxin are reported every year. Because of the severe economic and public health problems caused by harmful microalgae, many studies about the blooms of harmful microalgae have been conducted. Some promising methods have been developed including the use of yellow losses to sediment red tide organisms, chemical agents such as copper sulfate, hydrogen peroxide, and some biological controls in the form of viruses or bacteria. Although these methods seem effective in some short-term experiments, they may have potentially dangerous environmental consequences; therefore there are very limited investigations of a direct and specific control of marine harmful algal blooms with few environmental side-effects.
To look for HABs control agents that are efficient and benign to the environment, many scientists show a growing interest for growth inhibition of HAB species by allelopathic substances released by other aquatic organisms. Macroalgae and microalgae were known to have an antagonistic relationship in both natural and experimental aquatic ecosystems.
The present studies deal with the allelopathic effects of three macroalgae Ulva pertusa (Chlorophyta), Corallina pilulifera (Rhodophyta) and Sargassum thunbergii (Phaeophyta) on three red tide microalgae Heterosigama akashiwo, Alexandrium tamarense and Skeletonema costatum, and isolation and characterization of these allelochemicals. The three main objectives of these studies were to provide proof of the allelopathic effects of three macroalgae on these three microalgae, to gain an insight into the chemical interactions between macroalgae and microalgae in coastal areas, and to present evidence for the feasibility of using many macroalgae to control the growth of red tide microalgae. Results show that:
1. Comparative studies on the allelopathic effects of three macroalgae on three microalgae
We conducted laboratory experiments under stable environmental conditions, precluding the nutrient, pH, light conditions and effects of bacteria in order to investigate the allelopathic effects of fresh tissue, dry powder and aqueous extracts of three macroalga U. pertusa, C. pilulifera and S. thunbergii on the growth of microalgae H. akashiwo, A. tamarense and S. costatum were evaluated using coexistence culture systems in which concentrations of three macroalga were varied. The effects of macroalga culture medium filtrate on three microalgae were also investigated. The results coexistence assay showed that the growth of the three microalgae was strongly inhibited by using fresh tissues and dry powder; this indicated that there are the allelopathic effects of three macroalgae on three microalgae. Aqueous extracts of various concentrations of three macroalga can inhibit the growth of three microalgae and indicated that there are allelochemicals in three macroalgae; The effects of the macroalga culture medium filtrate exhibited no apparent inhibitory effect under initial filtrate addition but show apparent inhibitory effect under semi-continuous addition, which suggested that continuous release of small quantities of rapidly degradable allelochemicals from the fresh tissue of macroalga were essential to effectively inhibit the growth of three microalgae. In our study, we also concluded that the allelopathic effects of macroalgae on red tide microalgae show universality and species-specific.
2. Growth inhibition of H. akashiwo and A. tamarense by the extracts of C. pilulifera and S. thunbergii
Growth inhibitory effects of different concentrations of distilled water extracts and 4 organic solvent (methanol, acetone, ether and chloroform) extracts of C. pilulifera and S. thunbergii on the two microalgae H. akashiwo and A. tamarense were investigated. Methanol extracts of the highest growth inhibitory activity on two microalgae, and they killed all the cells of two microalgae at relatively higher concentrations. Growth of the two microalgae was significantly inhibited by the distilled water extracts of two macroalgae at relatively higher concentrations while cells of the two microalgae did not die completely even at the highest concentration. The other 3 organic solvent extracts of C. pilulifera and S. thunbergii had no apparent effect on the two microalgae, which suggested that the growth inhibitory substances from the extracts of two macroalgae had relatively high polarities. The methanol extracts of the two macroalgae were partitioned to petroleum ether phase, ethyl acetate phase, butanol phase and distilled phase by liquid-liquid fractionation. The bioassays of the activity of every fraction were carried out on two microalgae, indicated that petroleum ether phase and ethyl acetate phase had strong algicidal effect on the microalgae, fatty acids are probably main allelochemicals in the tissue of macroalgae.
3. Isolation and characterization of allelochemicals in the tissues of C. pilulifera and S. thunbergii
The petroleum ether phase and ethyl acetate phase of the methanol extracts of C. pilulifera and S. thunbergii were analyzed by GC/MS. The results of GC/MS revealed that there were 12 fatty acids in the 2 phase, and most of them were unfatty acids. The results of bioassays demonstrated that 5 unsaturated fatty acids in the tissue of C. pilulifera and S. thunbergii had strong algicidal effect on H. akashiwo and A. tamarense, and the effective concentrations were below 7.1 mg l~(-1). The results this study demonstrated that unsaturated fatty acids in the tissue of C. pilulifera and S. thunbergii might be one of the components of the allelochemicals in the macroalgae. Taken together, it is speculated that unsaturated fatty acids in the tissue of C. pilulifera and S. thunbergii has a potential to be considered as allelochemicals for the control of several harmful algae. However, it is not clear yet whether this compound can practically be applied to open waters. So further studies on the algicidal should focus on efficacy and effects on the algal control spectrum and mode of action are needed.
4. Influence of temperature, salinity, irradiance and pH on the effectiveness of allelopathic effects of C.pilulifera on H. akashiwo and S.thunbergii on A. tamarense and two microalgae growth
Two-factor combinations of different levels of temperature (10, 15, 25 and 30), salinity (10, 20, 30 and 40), irradiance (20,100,200 and 400μmol m~(-2)s~(-1)) and pH (5.5, 7, 8.5 and 10) on the effectiveness of allelopathic effect of C. pilulifera on H. akashiwo or S. thunbergii on A. tamarense and growth of the two microalgae were investigated. Two-factor combinations of different levels of temperature, salinity, irradiance and pH had significant (p<0.01) influence on the mean growth rates of two microalgae, and on the microalgal growth inhibition rates by C. pilulifera or S. thunbergii. The optical growth conditions for H. akashiwo was: 25°C, 30, 100μmol m~(-2)s~(-1) and pH=8.5, and for A. tamarense was: 20~25°C, 30, 100μmol m~(-2)s~(-1) and pH=7-8.5. The effectiveness of allelopathic effects of C. pilulifera on H. akashiwo or S. thunbergii on A. tamarense was increased by low temperature (15°C), low salinity (20), high irradiance (400μmol m~(-2)s~(-1)) and high pH (10).
本文研究三种代表性的大型海藻孔石莼(Ulva pertusa) (绿藻)、小珊瑚藻(Corallina pilulifera) (红藻)和鼠尾藻(Sargassum thunbergii) (褐藻)对三种赤潮微藻赤潮异弯藻(Heterosigma akashiwo)、亚历山大藻(Alexandrium tamarense )和中肋骨条藻(Skeletonema costatum)的克生作用,并初步提取、分离和鉴定了小珊瑚藻和鼠尾藻组织内抑制有害赤潮微藻生长的克生物质,探讨了小珊瑚藻和鼠尾藻对有害赤潮微藻克生作用的化学物质本质,为利用沿海区域的各种大型海藻来治理赤潮生物提供了科学依据。主要实验结果如下:
1.三种大型海藻对三种有害赤潮微藻克生效应的比较研究
通过排除营养盐、光照和pH等因素的影响,利用共培养系统研究了大型海藻孔石莼、小珊瑚藻和鼠尾藻的新鲜组织、干粉末、水溶性组织抽提液及培养水过滤液对三种有害赤潮微藻赤潮异弯藻、亚历山大藻及中肋骨条藻生长的作用,共存的实验结果表明,三种大型海藻的新鲜组织和干粉末对三种有害赤潮微藻的生长均有强烈的抑制作用,在较高的浓度下还能完全杀死赤潮异弯藻细胞和中肋骨条藻细胞,证明了三种大型海藻对三种有害赤潮微藻具有克生效应,并且大型海藻对有害赤潮微藻的克生效应具有普遍性。各个浓度孔石莼、小珊瑚藻和鼠尾藻水溶性抽提液对三种有害赤潮微藻的生长皆有抑制作用,抑制作用随浓度的升高而加强,这表明大型海藻组织中具有抑制有害赤潮微藻生长的克生物质,这进一步证明克生物质的存在是三种大型海藻对三种有害赤潮微藻产生克生效应的主要方式之一。三种有害赤潮微藻在一次性三种大藻培养水过滤液添加方式下生长没有受到显著抑制,而在半连续孔石莼及鼠尾藻培养水添加方式下其生长均受到了显著抑制,表明这三种大型海藻新鲜组织能够向环境中释放克生物质来抑制有害赤潮微藻的生长,同时也说明了大型海藻所分泌的克生物质容易降解。三种大型海藻新鲜组织对赤潮异弯藻和中肋骨条藻的克生效应由强到弱依次为:孔石莼>小珊瑚藻>鼠尾藻、对亚历山大藻的克生效应为鼠尾藻>孔石莼>小珊瑚藻;三种大型海藻干粉末对赤潮异弯藻的克生效应由强到弱依次为:鼠尾藻>孔石莼>小珊瑚藻,对中肋骨条藻和亚历山大藻的克生效应为:孔石莼>小珊瑚藻>鼠尾藻,表明大型海藻对有害赤潮微藻的克生效应具种的差异性。
2.小珊瑚藻和鼠尾藻组织提取物对赤潮异弯藻和亚历山大藻的生长抑制作用
以小珊瑚藻和鼠尾藻为大型海藻的代表,以赤潮异弯藻和亚历山大藻为目标微藻,研究了不同浓度的小珊瑚藻和鼠尾藻组织蒸馏水及4种具有不同极性的有机溶剂(极性从高到低:甲醇>丙酮>乙醚>氯仿)提取物对两种有害赤潮微藻的生长抑制作用。两种大型海藻组织甲醇提取物在5种提取物中具有最高的生长抑制活性。在相对较高浓度的此种提取物作用下,两种有害赤潮微藻均完全死亡。较高浓度的小珊瑚藻和鼠尾藻组织蒸馏水提取物对两种有害赤潮微藻的生长具有显著的抑制作用,但是在最高浓度下也不能使这两种有害赤潮微藻完全死亡。另外三种有机溶剂的提取物对两种有害赤潮微藻的生长无明显的影响。结果表明,小珊瑚藻和鼠尾藻组织中含有对赤潮异弯藻和亚历山大藻的生长有抑制作用的活性物质,且这些物质具有相对较高的极性。对两种大型海藻的甲醇提取物进行了液液萃取,然后将其分离为石油醚相、乙酸乙酯相、正丁醇相和蒸馏水相。并用这4相对两种有害赤潮微藻进行克生物质的生物活性检测,结果表明石油醚相和乙酸乙酯相具有较强的杀藻活性,脂肪酸可能为小珊瑚藻和鼠尾藻组织内克生物质的主要成分。
3.小珊瑚藻和鼠尾藻组织内克生物质的鉴定
通过对小珊瑚藻和鼠尾藻的石油醚相和乙酸乙酯相进行气相色谱——质谱(GC/MS)联用分析,并核对标准图谱和有关参考文献,鉴定出了12种脂肪酸,其中不饱和脂肪酸占大部分。对小珊瑚藻的鉴定中得出,其中饱和脂肪酸占20.81%,而不饱和脂肪酸成分占65.45%,对鼠尾藻的鉴定中得出,饱和脂肪酸占18.82%,而不饱和脂肪酸成分占58.62%。纯化合物对赤潮异弯藻和亚历山大藻的活性检测结果表明,9,12,15-十八碳三烯酸、6,9,12,15-十八碳四烯酸、5Z,8Z,11Z,14Z-二十碳四烯酸、5Z,11Z,14Z,17Z-二十碳四烯酸和5Z,8Z,11Z,14Z,17Z-二十碳五烯酸具有很强的杀藻活性。
小珊瑚藻和鼠尾藻组织中含有多种不饱和脂肪酸,其中一些不饱和脂肪酸对赤潮异弯藻和亚历山大藻具有强烈的杀藻效果,而且不饱和程度越高其克生效果越明显。这就表明不饱和脂肪酸可能是小珊瑚藻和鼠尾藻组织内抑制赤潮异弯藻和亚历山大藻生长的克生物质中的重要组成成分之一。
4.温度、盐度、光照、pH值对小珊瑚藻克制赤潮异弯藻及鼠尾藻克制亚历山大藻效果和两种有害赤潮微藻生长的影响
本实验共设置了4个环境因子:温度(T)、盐度(S)、光照(L)和pH值(P)。每两个不同的因子之间分别设计双因子实验,共6个处理组合,即温度和盐度、温度和光照、温度和pH值、光照和pH值及盐度和光照,以检验这4个因子对小珊瑚藻克制赤潮异弯藻和鼠尾藻克制亚历山大藻效果及对两种赤潮微藻生长的影响,得出了赤潮异弯藻和亚历山大藻的生长率及大藻克制有害赤潮微藻生长的抑制率,并且检验这4个因子两两之间是否存在交互作用。每个因子分别设置4个处理水平,即:温度(15、20、25和30°C)、盐度(10、20、30和40)、光照(20、100、200和400μmol m~(-2)s~(-1))和pH值(5.5、7、8.5、10)。实验结果表明,温度、盐度、光照和pH值这四个环境因子均能显著影响赤潮异弯藻和亚历山大藻的生长,并且能显著影响小珊瑚藻克制赤潮异弯藻以及鼠尾藻克制亚历山大藻的效果。小珊瑚藻克制赤潮异弯藻以及鼠尾藻克制亚历山大藻的最高抑制效果不是发生在有害赤潮微藻单养时的最适条件,而是发生在环境因子胁迫时,较低的温度(15°C)、较低的盐度(10)、较高的光照强度(400μmol m~(-2)s~(-1))和高pH值(10)均能增强小珊瑚藻克制赤潮异弯藻以及鼠尾藻克制亚历山大藻的效果。
Severe outbreaks of harmful algal blooms (hereafter, HABs) cause serious problems with regard to effective utilization of water resources such as fisheries and water-supply reservoirs, moreover, about 2000 cases of human poisoning resulting from algal toxin are reported every year. Because of the severe economic and public health problems caused by harmful microalgae, many studies about the blooms of harmful microalgae have been conducted. Some promising methods have been developed including the use of yellow losses to sediment red tide organisms, chemical agents such as copper sulfate, hydrogen peroxide, and some biological controls in the form of viruses or bacteria. Although these methods seem effective in some short-term experiments, they may have potentially dangerous environmental consequences; therefore there are very limited investigations of a direct and specific control of marine harmful algal blooms with few environmental side-effects.
To look for HABs control agents that are efficient and benign to the environment, many scientists show a growing interest for growth inhibition of HAB species by allelopathic substances released by other aquatic organisms. Macroalgae and microalgae were known to have an antagonistic relationship in both natural and experimental aquatic ecosystems.
The present studies deal with the allelopathic effects of three macroalgae Ulva pertusa (Chlorophyta), Corallina pilulifera (Rhodophyta) and Sargassum thunbergii (Phaeophyta) on three red tide microalgae Heterosigama akashiwo, Alexandrium tamarense and Skeletonema costatum, and isolation and characterization of these allelochemicals. The three main objectives of these studies were to provide proof of the allelopathic effects of three macroalgae on these three microalgae, to gain an insight into the chemical interactions between macroalgae and microalgae in coastal areas, and to present evidence for the feasibility of using many macroalgae to control the growth of red tide microalgae. Results show that:
1. Comparative studies on the allelopathic effects of three macroalgae on three microalgae
We conducted laboratory experiments under stable environmental conditions, precluding the nutrient, pH, light conditions and effects of bacteria in order to investigate the allelopathic effects of fresh tissue, dry powder and aqueous extracts of three macroalga U. pertusa, C. pilulifera and S. thunbergii on the growth of microalgae H. akashiwo, A. tamarense and S. costatum were evaluated using coexistence culture systems in which concentrations of three macroalga were varied. The effects of macroalga culture medium filtrate on three microalgae were also investigated. The results coexistence assay showed that the growth of the three microalgae was strongly inhibited by using fresh tissues and dry powder; this indicated that there are the allelopathic effects of three macroalgae on three microalgae. Aqueous extracts of various concentrations of three macroalga can inhibit the growth of three microalgae and indicated that there are allelochemicals in three macroalgae; The effects of the macroalga culture medium filtrate exhibited no apparent inhibitory effect under initial filtrate addition but show apparent inhibitory effect under semi-continuous addition, which suggested that continuous release of small quantities of rapidly degradable allelochemicals from the fresh tissue of macroalga were essential to effectively inhibit the growth of three microalgae. In our study, we also concluded that the allelopathic effects of macroalgae on red tide microalgae show universality and species-specific.
2. Growth inhibition of H. akashiwo and A. tamarense by the extracts of C. pilulifera and S. thunbergii
Growth inhibitory effects of different concentrations of distilled water extracts and 4 organic solvent (methanol, acetone, ether and chloroform) extracts of C. pilulifera and S. thunbergii on the two microalgae H. akashiwo and A. tamarense were investigated. Methanol extracts of the highest growth inhibitory activity on two microalgae, and they killed all the cells of two microalgae at relatively higher concentrations. Growth of the two microalgae was significantly inhibited by the distilled water extracts of two macroalgae at relatively higher concentrations while cells of the two microalgae did not die completely even at the highest concentration. The other 3 organic solvent extracts of C. pilulifera and S. thunbergii had no apparent effect on the two microalgae, which suggested that the growth inhibitory substances from the extracts of two macroalgae had relatively high polarities. The methanol extracts of the two macroalgae were partitioned to petroleum ether phase, ethyl acetate phase, butanol phase and distilled phase by liquid-liquid fractionation. The bioassays of the activity of every fraction were carried out on two microalgae, indicated that petroleum ether phase and ethyl acetate phase had strong algicidal effect on the microalgae, fatty acids are probably main allelochemicals in the tissue of macroalgae.
3. Isolation and characterization of allelochemicals in the tissues of C. pilulifera and S. thunbergii
The petroleum ether phase and ethyl acetate phase of the methanol extracts of C. pilulifera and S. thunbergii were analyzed by GC/MS. The results of GC/MS revealed that there were 12 fatty acids in the 2 phase, and most of them were unfatty acids. The results of bioassays demonstrated that 5 unsaturated fatty acids in the tissue of C. pilulifera and S. thunbergii had strong algicidal effect on H. akashiwo and A. tamarense, and the effective concentrations were below 7.1 mg l~(-1). The results this study demonstrated that unsaturated fatty acids in the tissue of C. pilulifera and S. thunbergii might be one of the components of the allelochemicals in the macroalgae. Taken together, it is speculated that unsaturated fatty acids in the tissue of C. pilulifera and S. thunbergii has a potential to be considered as allelochemicals for the control of several harmful algae. However, it is not clear yet whether this compound can practically be applied to open waters. So further studies on the algicidal should focus on efficacy and effects on the algal control spectrum and mode of action are needed.
4. Influence of temperature, salinity, irradiance and pH on the effectiveness of allelopathic effects of C.pilulifera on H. akashiwo and S.thunbergii on A. tamarense and two microalgae growth
Two-factor combinations of different levels of temperature (10, 15, 25 and 30), salinity (10, 20, 30 and 40), irradiance (20,100,200 and 400μmol m~(-2)s~(-1)) and pH (5.5, 7, 8.5 and 10) on the effectiveness of allelopathic effect of C. pilulifera on H. akashiwo or S. thunbergii on A. tamarense and growth of the two microalgae were investigated. Two-factor combinations of different levels of temperature, salinity, irradiance and pH had significant (p<0.01) influence on the mean growth rates of two microalgae, and on the microalgal growth inhibition rates by C. pilulifera or S. thunbergii. The optical growth conditions for H. akashiwo was: 25°C, 30, 100μmol m~(-2)s~(-1) and pH=8.5, and for A. tamarense was: 20~25°C, 30, 100μmol m~(-2)s~(-1) and pH=7-8.5. The effectiveness of allelopathic effects of C. pilulifera on H. akashiwo or S. thunbergii on A. tamarense was increased by low temperature (15°C), low salinity (20), high irradiance (400μmol m~(-2)s~(-1)) and high pH (10).
引文
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