2026, Vol. 56
2. 中国海洋大学海洋环境与生态教育部重点实验室，山东 青岛 266100
2, 4-二氨基丁酸(2, 4-diaminobutyric acid, DAB)是一种非蛋白氨基酸，具有两个氨基和一个羧基基团，在中性或碱性条件下呈现正电性，在室温下为无色结晶固体，化学性质较稳定。该化合物最早是在细菌代谢物多粘菌素A(一种多肽类抗生素)的酸性水解物中被发现[1]，之后在其他细菌[2]、植物[3-4]、藻类[5-7]、水生动物[5, 8-9]等生物样品中也有检出。DAB对肝脏[10]和神经系统[11-13]具有毒性作用，可导致小鼠或大鼠出现肝损伤症状和惊厥、抽搐、运动不协调等神经中毒症状[12, 14]；在细胞水平上，也会导致氧化损伤、肿胀和溶解[13, 15]、能量消耗[16]、氨基酸动态失衡[17]，甚至凋亡[18]。
根据DAB的同分异构体β-N -甲氨基-L-丙氨酸(β-N -methylamino-L-alanine, BMAA)提取工艺(见图 1)[19-20]，研究人员将DAB分为游离态、溶解结合态和沉淀结合态三种存在形式，其中游离态和溶解结合态统称为总溶解态。游离态是经三氯乙酸提取后直接溶解在上清液中的毒素分子；溶解结合态是经三氯乙酸提取后分布在上清液中，再经盐酸水解后释放出的毒素分子；沉淀结合态是经三氯乙酸提取后分布在沉淀物中，再经盐酸水解后释放出游离态毒素的蛋白结合态形式。DAB的同分异构体包括BMAA、N -(2-氨乙基)甘氨酸(N -(2-aminoethyl)glycine, AEG)、β-氨基-N -甲基丙氨酸、2, 3-二氨基丁酸、3, 4-二氨基丁酸、3-氨基-2-(氨甲基-)丙酸和2, 3-二氨基-2-甲基丙酸[21]，其中BMAA、AEG是最常见的两种同分异构体(见图 2)。
鉴于DAB在不同营养级的生物样品中被广泛检出，且具有神经毒性，本文对DAB的生物来源、检出情况、致毒机制等进行综述，以期为DAB的毒理学与健康风险研究提供重要参考。
2 神经毒素DAB及其同分异构体的生物来源2008年，研究人员首次在蓝细菌(Calothrix sp.)的单种培养物中检测出DAB[22]，之后在硅藻、甲藻、隐藻等单种培养物中检出DAB。因此，蓝细菌与真核细胞藻类被认为是DAB的生物来源。此外，在多种十字花科(Cruciferae)、豆科(Fabaceae)等植物中也发现该化合物[3-4, 14]。
2.1 DAB毒素的原核生物来源 2.1.1 蓝细菌目前，在微囊藻属(Microcystis )、聚球藻属(Synechococcus )、念珠藻属(Nostoc )、鱼腥藻属(Anabaena )、鞘丝藻属(Lyngbya )、节球藻属(Nodularia )等蓝细菌(也称蓝藻)中检出DAB毒素，且主要以游离态和溶解结合态形式存在[7, 22-28](见表 1)。蓝细菌中DAB毒素的产量具有显著的种间和区域差异性，这种差异可能与蓝细菌的基因组、共生细菌类群和生长环境等多种因素的差异有关。Downing等运用15N(NH4Cl)标记发现，微囊藻在氮限制条件下，DAB产量呈增加趋势[29]；在高磷酸盐水平(8.75 mmol·L-1)、强光照(9 000 Lux)和高温(25、30 ℃)等培养条件下，DAB产量显著增加[27]。这说明蓝细菌DAB的合成可能与其应激响应机制相关。蓝细菌基因组中存在天冬氨酸4-磷酸途径(见图 3)的关键酶编码基因，包括转氨酶等[30]，表明蓝细菌可能通过该代谢途径合成DAB，但目前未有直接的遗传学证据验证这一假设。然而DAB毒素在蓝藻培养体系中的生物来源模糊，已在微囊藻、束毛藻等蓝细菌培养物中发现共生细菌的存在[31-32]，并且以上研究均未排除共生细菌的存在。未来研究需结合分子生物学手段和生态学调查，以阐明DAB的合成机制及其在生态系统中的潜在影响。
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表 1 蓝藻中检出DAB的含量 Table 1 The contents of DAB detected in cyanobacteria |
除蓝细菌外，研究人员在分离自土壤[33-34]、水体[35-36]、大气[37-38]等不同环境介质的细菌样品中检出DAB，甚至在沙漠[39-40]、冰川[41-42]等极端环境中分离的细菌也检出该毒素。细菌中DAB参与肽聚糖交联，并以细胞壁结构成分的形式稳定存在。这些细菌附着或共生于某些动植物，例如艾草、人参等根部[43-44]，夹竹桃和杨树的树皮[45-46]，扁玉螺及高原鼠兔的肠道、奶牛瘤胃等[5, 47-48]。此外，蝙蝠、猕猴、牛等多种生物排泄物分离的细菌[49-51]，以及海洋与淡水环境中细菌形成的生物膜[49-51]均检出DAB毒素。法国泻湖贻贝外壳生物膜中总溶解态DAB的平均浓度为3.3 μg·g-1干重(dry weight, DW)[7]，而加拿大温尼伯湖淡水沉积物生物膜中总溶解态DAB浓度范围为0.5~77.6 ng·g-1 DW[52]。由此来看，产生DAB的细菌类群具有非常高的生物多样性。
关于细菌合成DAB的过程机制研究表明，DAB合成可能与特定转氨酶的催化反应有关，谷氨酸和天冬氨酸-4-半醛可逆反应生成DAB和2-酮戊二酸。不动杆菌(Acinetobacter spp.)、大肠杆菌(Escherichia coli )均含有编码转氨酶的dat(以及ddc基因)基因[53]，在四种假单胞菌(Pseudomonas spp.)[54]、Ⅰ型好氧性甲烷氧化菌(Methylomicrobium alcaliphilum 20Z)[55]中也发现了DAB转氨酶的基因。DAB的这种生物合成途径仍需要进一步研究确认。
2.2 DAB毒素的真核生物来源水环境中真核细胞藻类也是DAB毒素的重要来源。Réveillon等在法国泻湖的硅藻、甲藻、隐藻、绿藻和金藻样品中检出DAB[7-8]；Violi等在澳大利亚淡水环境中分离的5株硅藻中也检出DAB[28]；中国学者在海洋硅藻中检出DAB，其中拟菱形藻的含量最高(406.96 μg·g-1)，远高于国外水环境中藻类样品的检出量[5]。因此，硅藻作为海洋生态系统的重要浮游植物类群和DAB毒素的重要生物来源，潜在威胁海洋生态系统与人类的健康。从毒素的存在形式来看，真核藻类样品中总溶解态DAB含量远高于沉淀结合态毒素的含量(见表 2)。
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表 2 真核藻类DAB的检出量 Table 2 The contents of DAB detected in eukaryotic algae |
研究人员在陆生十字花科、豆科的山黧豆属、猪屎豆属、野豌豆属等高等植物中也检出DAB[4, 14, 24, 57-60]。金边苏铁(Cycas revoluta )的种子[24]、根[27]和德保苏铁(Cycas debaoensis )的叶子[61]以及番茄(Solanum lycopersicum )的花柱代谢物[62]中也检出DAB毒素(见表 3)。
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表 3 高等植物DAB的检出量 Table 3 The contents of DAB detected in plants |
BMAA是DAB的一种同分异构体，最初被发现自苏铁种子，而后发现BMAA的最初来源极有可能是与苏铁珊瑚状根部共生的念珠藻(Nostoc sp.)[63]。之后在不同种类的蓝细菌如微囊藻属(Microcystis )、色球藻属(Chroococcales )、集胞藻属(Synechocystis )和细鞘丝藻属(Leptolyngbya )等样品中普遍检出BMAA[64-65]。由于使用非特异性的氨基酸分析方法使得实验室菌株[65]、野外分离株[66]、野外样品[67]中BMAA检测含量过高，而在HILIC柱使用未衍生化方法中BMAA含量低[68-69]甚至未检出[22]。使得人们对蓝细菌作为BMAA生物来源的结论仍存在争议。Jiang等人首次在5株硅藻[70]以及Violi等人在4株淡水硅藻[56]中检出BMAA，说明海洋硅藻和淡水硅藻可能是BMAA的生物来源。目前仅在实验室培养的海洋链状裸甲藻(Gymnodinium catenatum )[71]以及三角异冒藻(Heterocapsa triquetra )[72]等甲藻样品中检出BMAA，暂未见其他种类的甲藻检出BMAA的报道。Li等人证明了极小海链藻(Thalassiosira minima )中BMAA是由硅藻自身产生的[73]，进一步研究海洋硅藻的分子机制发现，BMAA的产生是由铁限制强烈诱导的，是通过胞内CysK基因催化肽链中半胱氨酸残基和甲胺发生亲核反应原位生成BMAA[74]。BMAA同样存在于细菌中，粉状芽孢杆菌(Bacillus pulvifaciens )产生的一种抗生素galantin Ⅰ经酸水解后可释放S-2, 3-DAP和S-BMAA[75]，该化合物是目前已知的唯一含有BMAA结构的肽类化合物[76]。因此，BMAA的生物来源较为广泛，主要包括硅藻等藻类，同时也涵盖细菌。
AEG已在淡水、海洋、陆生、温泉等多种生境的蓝细菌中被检出，主要包括集胞藻属(Synechocystis sp.)、色球藻属(Chroococcidiopsis sp.)、鱼腥藻属(Anabaena sp.)、念珠藻属(Nostoc sp.)、鞘丝藻属(Lyngbya sp.)以及细鞘丝藻属(leppolyynbya sp.)等[77-78]，这些蓝细菌主要分布于淡水和海洋环境中。AEG在硅藻、甲藻、绿藻等真核生物中也曾被检出，主要包括骨条藻属、亚历山大藻属、原甲藻属、微拟球藻属、衣藻属等[7-8]。目前，关于藻类AEG合成的遗传学证据尚未见报道。迄今为止，针对DAB的其他同分异构体，亦缺乏系统性研究。
3 环境样品中神经毒素DAB及其同分异构体的分布当前，不同国家的研究人员在淡水、空气介质、水生生物等不同样品中检出DAB毒素，表明其在世界范围广泛存在(见图 4)。在环境样品BMAA的调查研究中，科学家通常会同时检测其同分异构体DAB。通常相较于BMAA，藻类样品[7, 8, 23-24, 27-28, 56, 79]、多数水生动物样品[5-8, 23, 80-85]以及南极洲[86]蓝藻藻甸和阿拉伯湾[9]海洋微生物结皮中DAB含量更高，检出率相近或更高。水处理过程中，常规工艺(紫外线、消毒剂氯/氯胺)难以有效去除DAB及其消毒副产物等有害物质[87]，严重威胁饮用水水质安全，危害人体健康。
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( 图中显示检出各个科类生物的DAB最高浓度。The figure shows the highest concentration of DAB detected for each category of organisms. ) 图 4 世界范围内环境生物样品中DAB的检出情况 Fig. 4 Global detection of DAB in environmental and biological samples |
中国淡水环境中浮游植物游离态DAB含量相对较低，0.37~3.82 ng·g-1WW，且未检出BMAA[27]，在太湖水样中也曾检出游离态DAB(1.83~2.09 ng·L-1)而BMAA的浓度范围为0.89~230.8 ng·L-1[88]，说明在水体介质中BMAA的波动性更大，可能受更复杂的环境因素(如藻类群落结构、水文条件等)影响。我国沿海采集的生物样品中DAB毒素的调查结果显示，浮游植物(0.01~12.34 μg·g-1 DW)、浮游动物(0.03~17.56 μg·g-1 DW)、贝类软体动物(0.05~3.82 μg·g-1 WW)、虾和蟹类节肢动物(0.02~1.91 μg·g-1 WW)、鱼类(0.09~0.16 μg·g-1 WW)等不同营养级的生物样品中均含有DAB毒素(见图 5)[5-6, 80-81]，但未表现出明显的生物放大现象，而BMAA在硅藻为主的海洋生态系统的食物网中表现出生物放大现象，这种食物链传递效率的差异可能是BMAA源于其硅藻内源性合成机制[74]直接进入食物链，而DAB可能为细菌外源性来源[73]、易代谢分解而未能形成显著放大。近年来，海洋软体动物(0.04~3.82 μg·g-1 WW)[5-6, 80-81]、节肢动物(0.02~1.91 μg·g-1 WW)[5, 81]中总溶解态DAB毒素水平相对稳定，这提示DAB毒素可能不是由初级生产者浮游植物产生的，且海洋生物可能具有高效的DAB毒素排出机制。
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图 5 中国水环境中采集的生物样品中DAB的检出量 Fig. 5 The contents of DAB detected in biological samples collected from aquatic environments in China |
1901—1904年采集的南极淡水蓝藻藻甸中检出总溶解态DAB[86]，且浓度(0.40~6.56 μg·g-1 DW)(见图 6)和检出率(85.7%)较高[9, 86]，AEG(0.74~6.79 μg·g-1 DW)含量较高而总溶解态BMAA含量较低(0.53 μg·g-1 DW)。后来研究人员在阿拉伯湾的海洋微生物结皮[9](0.19~31.08 μg·g-1 DW)、卡塔尔沙漠的蓝藻土壤结皮(2.8~4.4 μg·g-1 DW)[89]样品中也检出不同浓度的DAB，且随采样深度(0~105 cm)增加逐渐递减至0.5 μg·g-1 DW以下[90]。AEG含量(0.003 4~6.03 μg·g-1 DW；0.7~4.4 μg·g-1 DW)高而BMAA含量(1.4~9.1 ng·g-1 DW；未检出)低，这可能是因为蓝藻种类特异性代谢或极端环境因子调控作用不同。在淡水环境中，也能够检测到DAB(0.01~21.1 μg·L-1)、BMAA(0.01~25.3 μg·L-1)以及AEG(0.01~19 μg·L-1)[91-94]。
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图 6 国外水环境采集的生物样品中DAB的检出量 Fig. 6 The contents of DAB detected in biological samples collected from aquatic environments in other countries and regions |
淡水环境中浮游植物(澳大利亚[95]、加拿大[96])样品的DAB含量(212 ng·g-1~7.61 mg·g-1)明显高于半咸水环境中浮游植物(瑞典波罗的海[97]、法国地中海泻湖[7])样品的DAB含量(平均浓度0.69 μg·g-1 DW)。AEG和BMAA的含量分布也呈现类似规律：淡水环境中AEG(2.02 ng·g-1~23.9 mg·g-1)和BMAA含量(47.26 μg·g-1~2.63 ng·g-1)高于半咸水环境中AEG(ND)和BMAA含量(平均浓度为0.49 μg·g-1 DW)。然而BMAA的含量较低，这可能与其合成物种种类少、降解效率高等多因素有关。该现象与中国水环境中DAB毒素的检出情况截然相反，中国水环境中较高含量的DAB毒素出现在海洋环境样品中，而澳洲和北美洲地区淡水环境样品中DAB的浓度更高。
法国地中海泻湖环境样品中总溶解态DAB和BMAA随着食物链浮游植物-浮游动物-贻贝(0.69、0.92、7.2 μg·g-1 DW；0.49、0.63、4.0 μg·g-1 DW)呈现生物放大现象(见图 6)[7]。但在法国的泻湖和双壳类养殖区采集的贝类样品中DAB含量无显著季节性差异，且较为稳定[7-8, 23]，然而藻类存在显著的季节性动态变化，这表明高密度生长的浮游植物并未使贝类体内的DAB含量显著增加，说明食物链传递对贝类体内DAB含量的贡献不明显。这提示浮游植物可能不是DAB生产者，而细菌可能是其主要的生产者。
美国内布拉斯加州水库的调查结果显示，游离态和沉淀结合态DAB和BMAA在水生植物、鱼类等不同营养级生物中未见明显的生物放大作用[94]。南佛罗里达地区生长的龙虾含有游离态BMAA(0.01~11 μg·g-1 WW)、游离态和沉淀结合态DAB毒素(0.51~2.34 μg·g-1 WW)、以及微量的AEG和β-氨基-N -甲基丙氨酸毒素[98-99]，且BMAA主要富集在卵中，DAB主要富集在卵和尾巴部位[98]。美国海域海豚的脑组织中检出总BMAA(20~328 μg·g-1 WW)和DAB(98~742 μg·g-1 WW)[100]，但西班牙加利西亚生长的海豚的肝脏、肾脏或肌肉样品中未检出BMAA和DAB[101]，这可能与不同海域BMAA和DAB的生物来源差异有关，也可能与毒素在海豚体内的富集部位有关。
藻类膳食补充剂通常是由螺旋藻或其他蓝绿藻制成的，是研究人员关注的焦点之一。最早研究人员在德国市售的一种水华束丝藻的膳食补充剂中检出DAB(0.08 μg·g-1)，未检出BMAA[24]，之后加拿大、北美洲、西班牙市售的藻类膳食补充剂中也检出DAB、BMAA、AEG，其中DAB的检出率≥78%，且含量最高[61, 102-104]。其中，加拿大市售的藻类膳食补充剂检出的BMAA(0.13~2.52 μg·g-1)和DAB含量(0.49~107.06 μg·g-1)最高[104]，而北美洲、西班牙地区市售的藻类膳食补充剂的BMAA未检出，且DAB含量较低(0.003 5~2.40 μg·g-1)[24, 61, 102-103]。此外，研究人员在鲨鱼软骨粉中也检出BMAA(86~265 μg·g-1)和DAB(53~207 μg·g-1)[105]。
4 神经毒素DAB的致毒机制目前有关DAB神经毒性致毒机制的解释主要包括三种作用途径：触发兴奋性毒性、干扰γ-氨基丁酸(γ-aminobutyric acid, GABA)能神经系统、引发肝源性神经损伤(见图 7)[13, 18, 106]。
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图 7 DAB神经毒性的经典致毒机制示意图 Fig. 7 Classical mechanisms of DAB neurotoxicity |
谷氨酸作为中枢神经系统核心兴奋性神经递质，通过激活配体门控离子通道型受体(如N-甲基-D-天冬氨酸(N-methyl-D-aspartate, NMDA)/离子型谷氨酸受体AMPA)介导神经元去极化。受体过度激活引发细胞内Ca2+超载，导致兴奋性毒性，表现为细胞长期的去极化、细胞内Ca2+信号的激活以及细胞凋亡酶的激活[107-110]。DAB能够作为谷氨酸受体(如NMDA、离子型谷氨酸受体(iGluRs))的激动剂引起兴奋性毒性[13, 107]。
DAB在碳酸氢盐存在下形成的β-氨基甲酸酯加合物可激活NMDA受体从而引起神经兴奋性毒性。具体表现为显著的去极化效应和乳酸脱氢酶释放增加[13]，表明DAB可严重破坏细胞膜的完整性[111]。DAB通过调控动作电位相关离子(如Na+、K+)通道，诱导细胞去极化。研究表明，DAB能够导致小鼠艾氏腹水癌细胞内Na+及K+含量下降、Cl-及水含量上升[17]，这种离子组成的改变直接影响细胞膜的电化学性质，一方面引起膜电位去极化甚至无法恢复[112]，另一方面显著降低膜输入电阻[107]。科学家提出了DAB诱导细胞溶解的双重作用机制假说，一种是DAB在胞内蓄积引起持续去极化破坏了渗透平衡[18, 113]，另一种是DAB通过激活机械力敏感型Cl-通道降低膜输入阻力[107, 114]。
DAB进入细胞内是通过钠依赖性的系统A中性氨基酸转运体[107, 115]，该转运过程与L-丙氨酸和G-蛋氨酸存在竞争性抑制关系[18, 107]。DAB对细胞内外游离氨基酸浓度有显著影响，能够抑制原代皮质神经元细胞摄取L-胱氨酸[111]，并导致胶质母细胞外的氨基酸如谷氨酸、天冬氨酸等阴离子氨基酸和丙氨酸、甘氨酸等中性氨基酸的增加[16]。此外，钠离子依赖性的DAB摄入细胞过程消耗大量能量，腺嘌呤核苷三磷酸(Adenosine triphosphate, ATP)是通过Na+/K+-ATP酶获取的[16]。
4.2 DAB干扰GABA能神经系统给药DAB后，研究人员发现出现惊厥反应的大鼠脑组织中GABA浓度升高[10, 106]，DAB通过多种途径干扰GABA能神经系统功能。在神经突触体中，DAB除通过系统A中性氨基酸转运体外，还可经由GABA系统转运[116]。生理状态下，GABA通过激活GABAA受体-氯离子通道复合体，并引起超极化，进而产生抗惊厥效应[117]。DAB作为GABA的结构类似物，能够竞争性结合神经元表面GABA的高亲和力摄取位点[116]，从而干扰正常的抑制性神经传递并引起惊厥。DAB能有效抑制3H-GABA在Na+依赖性神经末梢的摄取过程[118]，DAB和GABA在短期(0~10 min)暴露过程中表现出竞争性抑制伴随细胞内DAB积累，而长期(50~100 min)暴露表现出非竞争性抑制[116]。GABA摄取的抑制可能来自胞外竞争性和胞内非竞争性抑制的混合作用[119]，并呈现非线性特征[120]。然而，由于发现对非钠依赖性GABA通道的抑制是非立体特异性的，这表明DAB的神经毒性机制并非通过抑制GABA摄取，而是主要源于与突触后GABA受体的直接相互作用[119]。
4.3 DAB诱导慢性氨中毒引发神经毒性通过对暴露大鼠的多组织分析发现，其肝脏、血液及脑组织中的尿素浓度均呈现显著性升高。研究发现，DAB通过抑制大鼠肝脏中鸟氨酸氨甲酰基转移酶的活性，从而抑制尿素循环引起肝损伤[10]。尿素循环的抑制导致血氨浓度显著增加，引发慢性氨中毒。而脑中谷氨酰胺浓度的长期微量增加间接地导致继发性脑神经毒性，这一过程是DAB通过阳离子氨基酸转运系统y+穿过血脑屏障[121]引发的。
对DAB与其同分异构体BMAA、AEG的毒性效应开展了比较研究[15, 122-123]，发现这三种化合物的致毒机制有相似或相同之处。其中，BMAA的神经毒性是通过激活谷氨酸受体(如代谢型谷氨酸受体mGluRs、NMDA和离子型谷氨酸受体AMPA)结合和错误嵌入肽链来介导的[21]，而AEG的神经毒性是通过诱导氧化应激和激活谷氨酸受体(代谢型谷氨酸受体5)来介导的[111]。DAB与BMAA的毒性效应存在显著的物种和细胞类型差异性，表现出不同的敏感性。在细胞水平，DAB对小鼠巨噬细胞RAW264.7、小胶质细胞BV-2[15]以及皮质神经元细胞[111]的毒性更强，导致细胞活力降低和死亡率增加，还诱导水蛭Retzius神经元细胞更高的去极化水平[112, 122]。BMAA对N2a细胞、小鼠神经元NSC-34细胞[15, 124]的毒性更强，除了细胞活力降低以外，还诱导NSC-34细胞的膜透性增加、线粒体脱氢酶活性降低[124]，对星形胶质细胞摄取体外L-胱氨酸的抑制作用更强[111]。在生物个体水平，DAB对斑马鱼幼鱼(孵化后6 h)的致死率更高。有一项研究表明同等浓度下BMAA和DAB毒性相近，造成人神经母细胞瘤细胞SH-SY5Y的死亡率无显著差异[125]。DAB与AEG的毒性比较的研究中，DAB对斑马鱼幼鱼(孵化后6 h)的毒性更强，致死率更高[122]，然而AEG对小鼠皮质神经元细胞的毒性更强，其半数致死浓度EC50更低，还对星形胶质细胞摄取体外L-胱氨酸的抑制程度更强[111]。
5 展望本文系统梳理了不同生物样品中神经毒素DAB的检出情况，对其神经毒性致毒机制进行了归纳总结。从现有的文献来看，全球分布的淡水或海洋环境中的细菌、蓝细菌、微藻、大型藻、水生动物及陆生高等植物等多种生物样品中检出DAB毒素，由此引起的健康风险应予以关注。但目前对DAB的真正生产者及其生物合成机制的认识非常有限，且缺少DAB的水生生物的环境质量基准，对其生态环境风险管控提出了挑战。建议今后从以下几个方面开展研究：
(1) 当前有关DAB毒素的真实生产者尚存在争议，真核浮游植物和蓝细菌可能不是DAB毒素的初始生产者，尚未排除这些生物的共生或寄生细菌产生DAB的可能。尽管已有多项研究在放射菌中检出DAB，但人们对其合成机制尚不清楚。开展DAB生物合成机制的探索，将有助于理解其生物学意义。
(2) 深入研究DAB在不同动物、植物生物体中的吸收、分布、代谢及排泄规律，揭示其富集和代谢途径，以进一步解释DAB的毒性机制。
(3) 基于DAB的毒性数据和环境赋存，建立DAB的水生生物的环境质量基准，并开展环境风险评估，提高对该毒素的动态监测与风险预警能力。
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2. Key Laboratory of Marine Environment and Ecology, Ministry of Education, Ocean University of China, Qingdao 266100, China