贻贝对麻痹性贝类毒素的蓄积代谢研究进展

范礼强, 郑关超, 吴海燕, 郭萌萌, 卢 霞, 李 玉, 谭志军, 3

贻贝对麻痹性贝类毒素的蓄积代谢研究进展

范礼强1, 2, 郑关超2, 吴海燕2, 郭萌萌2, 卢 霞1, 李 玉1, 谭志军2, 3

(1. 江苏海洋大学测绘与海洋信息学院, 江苏 连云港 222005; 2. 农业农村部水产品质量安全检测与评价重点实验室, 中国水产科学研究院黄海水产研究所, 山东 青岛 266071; 3. 青岛海洋科学与技术试点国家实验室, 山东 青岛 266237)

麻痹性贝类毒素(paralytic shellfish toxins, PSTs)能够导致贝类、鱼类等海洋生物染毒或死亡, 危害海洋生态安全及人类健康。其中, 贻贝具有强蓄积性特点, 在国外可作为PSTs的指示性生物。近年来, 我国因食用贻贝造成多起PSTs中毒事件成为安全关注重点, 因此急需弄清其风险形成机理。本文通过综述我国近海贻贝等贝类中PSTs的风险程度、贻贝对不同产毒藻来源的PSTs的蓄积作用, 分析环境因子对于蓄积代谢的影响, 深入挖掘贻贝蓄积代谢特异性。进而聚焦贻贝代谢阶段的表达调控过程、影响因素、转化作用及方式等。未来应侧重于摸清我国目标区域贻贝等海洋贝类中PSTs基础风险规律和变化过程, 并据此建立区域性PSTs风险防控技术, 提高我国贻贝中PSTs风险的主动防控能力, 对于保障消费者健康和海洋生态安全具有积极意义。

贻贝; 蓄积代谢; 免疫; 风险评价

近年来, 有害赤潮在世界范围内呈现出频率增加、区域扩大、危害日益严重的发展趋势, 导致贝类中贝类毒素的安全风险愈加严峻[1]。根据所引起的中毒症状, 可将贝类毒素分为: 麻痹性贝类毒素(paralytic shellfish toxins, PSTs)、腹泻性贝类毒素(diarrhetic shellfish poisoning, DSP)、神经性贝类毒素(neurotoxic shellfish poisoning, NSP)、记忆缺失性贝毒(amnesic shellfish poisoning, ASP)和西加鱼毒素(ciguatera fish poisoning, CFP)等[1-2], 其中PSTs是最具代表性的一类, 分布最广、毒性最强[2]。PSTs主要由海洋环境中的亚历山大藻()、裸甲藻()、盾甲藻()等甲藻及生活在淡水中的蓝绿藻产生[3]。双壳贝类滤食有毒藻类, 毒素在贝类体内蓄积或通过食物链作用传递至人类, 引起人类食物中毒[1]。全球每年因PSTs导致的中毒事件约2 000起, 占贝类毒素中毒事件的64%[4]。其中, 双壳贝类(包括贻贝、扇贝、蛤和牡蛎)是最主要的食物链传播来源, 也是美国、加拿大、欧盟等国家及组织开展PSTs管控重点[3], 欧盟限量标准为800 µg STXeq·kg–1[5]。我国是全球第一贝类养殖大国, 同时也是PSTs有害赤潮高发国家之一。

近年来, 我国沿海地区多发、赤潮, 主要分布在渤海(秦皇岛)、东海(浙江、福建及南宁)部分近岸海区[1, 6]。其中, 秦皇岛海域自2016年起春季连续暴发赤潮, 贻贝中毒素蓄积量最高为52 072.8 µg STXeq·kg–1, 超过欧盟限量标准(800 µg STXeq·kg–1)的65倍[7], 可见危害之严重性。同年福建近海暴发赤潮, 其中贻贝中PSTs蓄积含量最高[8-9]。此外Isabel等[10]的研究也表明, 贻贝较其他贝类具有更高的蓄积能力, 它也是造成我国PSTs食物中毒事件的主要物种[11-12]。因此, 评估贻贝对PSTs的蓄积代谢特异性, 对于我国质量安全监管具有重要的现实意义。

贻贝是我国四大经济贝类之一, 紫贻贝、翡翠贻贝和厚壳贻贝是我国沿海常见种类[13], 最适生长温度为5~22 ℃, 盐度为15~40, 是环境适应能力较强的广温广盐类生物。贻贝属于滤食性生物, 并且对不良环境具有强耐受力, 因此国内外学者已经将其用于海洋重金属[14-17]、全氟化合物[16]、农药[18-19]等污染物的监测研究中, 作为生物指示物评估海洋生态系统的污染状况[15]。近十年来, 研究人员在实验室内不断模拟双壳贝类蓄积和清除PSTs过程, 发现贻贝相较其他贝类, 具有蓄积速率快、蓄积量大、清除快等特点[20-21]。这也使得贻贝成为研究PSTs等海洋污染物初始组分的重要指示生物。因此亟需深入探究贻贝在PSTs污染监测过程中的蓄积清除及代谢转化机理, 对于准确评价其安全风险并构建有效防控措施具有重要作用。

1 我国近海贝类中麻痹性贝类毒素风险

我国近海PSTs污染由来已久, 21世纪初北黄海、福建沿海、长江口邻近海域、广东沿海PSTs污染问题较为突出, 中毒事件时有发生[1]。近年来, 渤海和福建沿海PSTs污染较为严重[22]。为准确探究我国近海贝类中PSTs风险程度, 表1总结了2005— 2019年我国近海贝类中PSTs调查结果。其中, 渤海2016年河北秦皇岛贻贝产品PSTs含量超标, 导致10余人中毒且2人死亡。污染调查中发现PSTs最高含量达到52 072.8 µg STXeq·kg–1, 是欧盟限量标准 (800 µg STXeq·kg–1)的65倍, 贻贝污染极为严重。之后2017年、2019 年连续监测, 发现贻贝等贝类中PSTs检出率和残留量都有所下降[23-24], 但为防止PSTs污染的突然爆发, 应针对贝类中PSTs的风险构建有效防控措施, 以便准确认知我国贝类中PSTs的残留风险。黄海海域2007—2008年贝类污染调查中虾夷扇贝的PSTs检出率达到了79.2%, PSTs残留量为262.0~8 339.6 µg STXeq·kg–1[23, 25]。2016年牡蛎等贝类中PSTs最高含量为562.9 µg STXeq·kg–1, 接近欧盟限量标准, 亟待引起关注。东海海域2016—2017年贝类中PSTs检出率和最高残留量分别为74.2% 和23 293.0 µg STXeq·kg–1[26], 直接导致2017年6月福建漳州发生数十人贻贝PSTs中毒[27], 贝类中PSTs风险程度极高。在2017年贻贝污染后续调查中, 发现贝类中PSTs残留量降低至71.0 µg STXeq·kg–1, 存在地域性反复风险。南海海域2005—2006年贝类中PSTs检出率为14.3%, 最高残留量达3 700.0 µg STXeq·kg–1, 是限量标准的4.6倍, 污染严重[28-29]。之后的南海贝类中PSTs污染调查中检出率逐年增加, 2010—2011年是32.9%, 2015年更是达到了86.0%[30], 但南海贝类中PSTs残留量却在逐年下降。综上可见, 随着我国环境污染得到有效控制, 有害赤潮由大面积爆发转为小范围板块化暴发趋势。由于有害赤潮后产生的丰富孢囊沉降至表层沉积物, 受环境影响成为有害赤潮的“种子库”, 显著提高目标海区的PSTs暴发风险。

表1 不同海域贝类中麻痹性贝类毒素分布情况

注: -表示无数据

从2005—2019年的全国近海贝类养殖区PSTs污染调查数据发现: 贝类毒素污染存在季节性差异, 与产毒藻的季节性生长有较大关系[31-32], 尤其是受海水温度影响显著[33]。我国海域南北跨度大, 调查发现南海贝类中PSTs含量多在冬季或早春达到最高。东海纬度高于南海, 贝类中PSTs检出率和超标率也相应推迟到夏季6、7、8月份达到最高[9, 34], 而黄渤海PSTs污染多在4、5月份暴发[33, 35]。贻贝作为我国沿海重要养殖经济贝类, 四大海域贻贝的PSTs检出率和残留量都较高[36], 这也与贻贝具有蓄积速率快、蓄积量大的研究结果相一致。为保障我国贝类产业健康发展和消费者生命安全, 亟需深入探究贻贝中PSTs的蓄积清除及代谢转化规律, 以便更好的对贝类毒素安全风险进行有效管控。

2 贻贝对麻痹性贝类毒素的蓄积、清除特性

贻贝能够快速蓄积大量PSTs, 具有蓄积速率快、蓄积量大、蓄积率高的特点。如表2中所示, 贻贝蓄积速率最高为1 736.0 µg STXeq∙kg–1∙d–1, 是其他四种主要经济贝类的3~15倍[37-42]。邱江兵等[42]和Katsushi等[40]PSTs暴露实验中贻贝的整体蓄积率分别达到13.6%和14.6%, 高于牡蛎的8.3%、菲律宾蛤仔的2.5%和文蛤的12.4%; 贻贝能够蓄积大量PSTs, 暴露实验中贻贝体内PSTs含量分别达到了8 680.0 µg STXeq·kg–1和10 960.0 µg STXeq·kg–1, 约是欧盟限量标准(800 µg STXeq·kg–1)10.6倍和13.7倍。Shumway 等[43]研究表明贻贝神经元轴突对PSTs毒素STX不敏感, 这可能是贻贝PSTs蓄积速率快、蓄积量大的主要原因之一。贻贝组织间蓄积速率和蓄积量存在较大差异[44], 内脏团是贻贝蓄积PSTs的靶器官, 对PSTs蓄积速率和蓄积量都远高于鳃、闭壳肌、性腺等其他组织[42], 内脏中毒素含量有时甚至可以达到整体的90%以上[45]。

表2 不同贝类对麻痹性贝类毒素的蓄积规律

贻贝在暴露实验中能够快速将PSTs清除到体外, 具有清除速度快、清除率高等特点[17, 42]。根据Bricelja等[47]研究, 双壳贝类清除毒素速度大致可以分为: 快、中速清除贝类(6.0%/d~17.0%/d)和慢速清除贝类(0.3%/d~ 4%/d)。贻贝的最高清除速度(1 500.0 µg STXeq∙kg–1∙d–1)和清除速率(17.4%/d) (见表3)分别是其他四种经济贝类的8~15倍和1.3~5.4倍, 属于快速清除贝类。贻贝大约5 d时间, 即可将8 680.0 µg STXeq·kg–1迅速排除至限量标准以下[42]。Katsushi等[40]研究发现贻贝对PSTs清除速度达1 326.6 µg STXeq∙kg–1∙d–1(清除速率为12.1%/d), 远高于其他贝类。Ana等[48]的贻贝PSTs清除实验研究发现, 清除实验中的前24 h贻贝体内PSTs毒性便下降了约66.9%。以上研究结果都证明贻贝在不同环境条件下都具有较强的毒素清除能力。

2.1 环境因子对贻贝蓄积清除PSTs的影响

海洋环境因子如温度、盐度、pH等对贻贝的过滤、摄食等生长生理活动有重要影响, 从而直接影响贻贝对藻毒素的蓄积和清除。温度是环境因子中首要影响因素, 高温同时降低毒素蓄积量并减缓毒素清除速度。Ana等[48]研究了温度对贻贝蓄积、清除能力的影响, 结果表明19 ℃条件下贻贝5天蓄积了1 493.8 µg STXeq·kg–1毒素, 而当温度升高至24 ℃时贻贝5天仅蓄积661.9 µg STXeq·kg–1毒素, 蓄积量降低了约49.0%, 且毒素清除效率也显著降低, 残留时间延长[20]。这是由于高温环境下贻贝对产毒藻的摄食明显减少[49-51]。研究表明, 24 ℃时, 贻贝的过滤率降低至19 ℃时的1/6。此外高温诱发贻贝应激反应, 其参与代谢过程的关键调节酶如糖酵酶丙酮酸激酶(PK)等活性受到抑制[49], 从而使得贻贝清除率下降[52-53]。

表3 不同贝类对麻痹性贝类毒素的清除规律

不同盐度海水中贻贝毒素蓄积速率、蓄积含量存在较大差异。研究发现盐度25.7时贻贝摄食最为活跃, 毒素蓄积含量也相应较高[54-56], 但当盐度降低至5.0时, 贻贝过滤海水速率显著降低, 毒素蓄积能力减弱[57-59]。这是由于低盐度刺激贻贝产生应激反应, 贻贝通过闭合外套膜以降低外界环境对生理活动的影响[21]。因此, 在低盐度环境下, 贻贝过滤率、摄食率降低, 对藻毒素的蓄积能力降低。另外, 研究表明pH降低至7.5时贻贝生长、代谢等生理活动都会显著减慢。Ana等[48]研究了气候变化驱动因素(升温和酸化)对贻贝中PSTs蓄积代谢影响。结果表明上述气候改变可导致残留毒素毒性降低但毒素残留时间延长[60]。在pH降低至7.3时贻贝甚至可能会直接死亡[61-63]。

除温度、盐度、pH会影响贻贝对PSTs的蓄积清除外, 贻贝的种内/种间竞争等也会对贻贝的蓄积、清除产生影响, 竞争会增加它们对食物的需求, 通过鳃过滤更多的水, 因此PSTs蓄积量显著增加[62-64]。

2.2 贻贝对不同产毒藻的蓄积能力评估

研究表明, 产毒藻的毒性强弱、密度以及投喂方式都能够影响贻贝蓄积毒素能力。我国东部沿海主要优势产毒藻为、。由于藻毒素来源不同, 贻贝的蓄积能力具有显著差异[45]。黄德强等[65]研究发现贻贝对于的摄食率显著低于无毒饵料藻。且Lee等[66]研究证明贻贝在毒性大于26 pg STXeq·个–1时摄食率也降低为原来的1/2。在我国近海PSTs调查中[22, 67], 贻贝在赤潮中的蓄积量比在其他产毒藻中的要高。因此, 贻贝的蓄积能力受产毒藻种类、藻密度等因素影响。

此外, PSTs产毒藻的不同密度也会影响贻贝蓄积PSTs。贻贝的摄食率一般随浮游植物密度的增加而升高, 这是由于在较高食物密度下, 过滤同样多海水能够获取更多食物[68]。Negri等[69]对比密度为2×105个/mL和2×104个/mL的蓝绿藻对淡水贻贝毒素蓄积影响时发现: 暴露于密度为2×105个/mL产毒藻中的贻贝体内PSTs毒性水平在2~3天就达到800 µg STXeq·kg–1, 而暴露于2×104个/mL产毒藻中的贻贝5周后只有少量毒素被检测到, 差异显著。赵俊梅等[68]的暴露实验也反映了类似结果, 当产毒藻密度增加时贻贝摄食率升高, PSTs蓄积量增加。

PSTs产毒藻的不同投喂方式也会对贻贝蓄积PSTs产生影响, 连续投喂时贻贝蓄积量要高于定期投喂时的[42, 70]。造成这种现象的原因可能是贻贝长时间处于较低密度的毒藻中, 毒素对其摄食不产生抑制, 贻贝能够不断从水中过滤藻细胞, 毒素更容易在体内累积。而定期投喂高密度的毒藻会使贻贝对毒藻产生生理抑制[69], 并且会较快速把体内PSTs清除进水体中。

3 贻贝对麻痹性贝类毒素的代谢转化

3.1 贻贝对藻毒素代谢转化作用

全球目前已鉴别的PSTs约60余种, 主要包括四大类: 氨基甲酸脂类毒素, 包括石房蛤毒素(STX)、新石房蛤毒素(NEO)和膝沟藻毒素(GTX1-4); N-磺酰氨甲酰基类毒素, 包括GTX5(B1)、GTX6(B2)和C1-C4; 脱甲酰基类毒素, 包括dcSTX、dcNEO、dcGTX1-4; 脱氧脱氨甲酰基类毒素, 包括doSTX、doGTX2和doGTX3等[42, 71]。除了常见的赤潮藻种毒素, 更多的组分仅在贝类体内发现, 为贝类的代谢产物[72-73]。如图1所示, 贝类中常见的PSTs组分间转化形式。

图1 贝类中常见PSTs组分间转化

注: 实线表示还原反应; 点虚线表示水解反应; 横虚线表示异构化

Kim等[74]进行了中PSTs毒素组分与贻贝中PSTs组分对比研究,主要毒素为C2、GTX4, 贻贝暴露初期体内PSTs组分与藻细胞中类似, 随着中毒时间的增加贻贝中主要毒素变为C1和GTX1, 分别是初始C2和GTX4的2.1和2.4倍, Wiese等[75]和Gracia等[39]的研究中也得出相似结论。但与扇贝、牡蛎等相比, 贻贝体内PSTs的代谢转化率相对较低[76]。Choi等[44]用只含有C2毒素的(ATDP)对贻贝开展暴露实验, 净化阶段在贻贝内检测到了GTX2/3等毒素, 证实了贻贝能够将低毒性的N-磺酰氨甲酰基类毒素转化成高毒性的氨基甲酸酯类毒素。于姬等[25]在体外实验研究中同样发现在酸性加热条件下N-磺酰氨甲酰基类毒素(C1-4、GTX5和GTX6)易转化为PSTs组分中毒性最强的氨基甲酸酯类毒素(GTX1-4、STX和NEO)。Paulo等[77]对链状裸甲藻()展开研究,主要毒素为C1-C4、GTX5、GTX6, 而贻贝体内转化成了dcGTX2&3、dcSTX和dcNEO。还原反应是贝类常见的PSTs转化反应, 且还原反应后贝类中PSTs毒性相对更高, 如GTX2&3还原生成STX、C1&2还原生成GTX5、dcGTX2&3还原生成dcSTX[70]。邱江兵等[42]实验中表明GTX6的N1位羟基在贻贝消化腺中能够发生还原反应生成GTX5。

近些年, 在PSTs检测中发现贝类中独有的PSTs代谢物, 如几种新的STX变体(M1-M4)[78-80], 在微藻中没有发现过。根据这些化合物的结构和活性关系, 推测可能是贻贝自体解毒的中间产物[81]。为了解这些新代谢物的来源和生物转化途径, Ding等[82]在实验室条件下用两种(ATHK株和TIO108) 喂养贻贝, 结果表明11-羟基-C2毒素(M1)和11-二羟基-C2(M3)由C2转化而来, 11-羟基-C4毒素(M7)和11-二羟基-C4(M9)由C4转化而来。此外, M2、M4和M6可能是GTX2/3的代谢产物, M8和M10可能是GTX1/4转化而来。这些研究都表明, 贻贝不仅可以对藻中已存在的PSTs进行相互转化, 还可以产生新的PSTs代谢产物, 且新型代谢物与贝类的解毒过程有关。

3.2 代谢转化过程中的影响因素

酶类在PSTs的代谢转化过程中发挥着重要的催化作用, 如表4所示, 主要包括氨甲酰水解酶、磺基转移酶、氨基甲酰酯酶、N-氧化酶和谷胱甘肽还原酶[42, 78, 83-85]。Lin等[83]从贝类消化腺中分离纯化了一种氨甲酰水解酶, 实现R4基团中氨甲酰基(或N-磺酰氨甲酰基)的水解。Yoshida等[78]研究结果表明磺基转移酶可将3’-磷酸腺苷-5’-磷酸硫酸(PAPS)中的硫酸基团转移到STX和GTX2&3的N-21位氨甲基基团, 分别生成GTX5和C1&2。卞中园等[84]表明氨基甲酰酯酶能够催化氨基甲酸酯类毒素(STX、NEO、GTX1-4) C11 位上的氨基甲酰基发生水解反应, 生成脱氨甲酰基类毒素(dcSTX、dcNEO、dcGTX1-4)。邹迎麟等[85]在PSTs生物合成中检测到N-氧化酶可以将GTX2&3转化为GTX1&4。邱江兵等[42]进行了消化腺体外转化实验, 发现贻贝消化腺在谷胱甘肽还原酶的催化下, 能够促使11α-表聚体毒素如GTX1、GTX2、C1向11β-表聚体毒素如GTX3、GTX4、C2转化。

表4 贻贝参与PSTs转化过程调节酶

贝类中PSTs的代谢转化还受到pH、温度和还原剂(谷胱甘肽、半胱氨酸)等因素的影响。如在较高温度和pH值条件下会加速PSTs的差向异构化[86]。双壳贝类组织中毒素的生物转化除了化学和酶促作用的结果, 也可能是由于存在于消化道中细菌转化的结果。Kotaki等[87]研究表明弧菌、假单胞菌能够将CTX1/2/3和NEO毒素转化为STX, 而且有氧条件比无氧条件转化快。

3.3 贻贝对PSTs的适应性机制

研究表明, 贻贝体内的代谢靶器官内脏团会启动抗氧化应激防御系统, 用于减轻PSTs带来的损伤[45, 88]。PSTs进入贻贝体内后, 将产生大量活性氧(ROS), 引起生物体氧化应激, 导致脂质过氧化、蛋白质变性、DNA 损伤等。内脏团和鳃组织内的抗氧化防御系统中的抗氧化酶, 如谷胱甘肽过氧化物酶(glutathione peroxidase, GSH-PX)、超氧化物歧化酶(superoxide dismutase, SOD)、酸性磷酸酶(acid phosphatase, ACP)等会被激活, 以消除ROS, 减少损害[41-42]。Qiu等[89]研究A.tamarense (ATHK)对贻贝抗氧化系统的影响, 发现在毒素蓄积和清除期间, ROS在贻贝内脏团中会迅速产生并消失, SOD和GSH-PX活性增强, 有效清除贻贝体内超氧阴离子自由基和过氧化氢(H2O2)。GSH-PX 还在贻贝体内起到脱毒作用, 其能将脂类过氧化物还原为相应的醇, 并可以代替过氧化氢酶(Catalase, CAT)将游离的H2O2原成水, 同时催化谷胱甘肽(Glutathione, GSH)转变为氧化型[90]。粒细胞分泌的ACP在抗氧化响应过程中活力增强, 鱼烟酰胺腺嘌呤二核甘酸磷酸酶(nicotinamide adenine dinucleotide phosphate, NADPH)是一种还原型辅酶, 含量降低, 部分NADPH可能参与了毒素的生物转化。

血细胞在贻贝天然免疫过程中发挥重要作用, 通过吞噬或包裹活的病原体来保护组织, 并通过炎症过程修复由损伤、中毒和感染引起的组织损伤。通过超微结构观察可以发现PSTs在暴露初期会造成贻贝鳃组织柱状上皮细胞中线粒体和溶酶体增多且聚集, 后期上皮细胞肿胀破裂, 黏液细胞大量释放粘液颗粒, 细胞核萎缩变形, 严重时细胞坏死裂解[41]。Galimany等[86]研究贻贝的免疫反应, 发现血细胞向肠内的渗出, 推测是为了分离肠内有毒藻细胞, 从而将组织损害降到最低。Pousse等[91]研究发现血细胞能将有毒细胞包裹在消化道内, 形成吞噬泡, 吞噬泡与初级溶酶体融合形成次级溶酶体, 次级溶酶体内的ACP就会将其降解清除到体外, 从而减少藻细胞与其他组织的接触。Bianchi等[92]对贻贝进行免疫功能实验研究, 发现PSTs暴露3d后贻贝血细胞吞噬功能显著增强, 以减轻PSTs对组织的损害。但PSTs对贻贝生理活动的影响大多是暂时的, 且在长期暴露于PSTs产毒藻后, 贻贝表现出了良好的生存适应和免疫能力。

4 结论与展望

现有结果表明, PSTs已成为危及我国近海生态及贝类质量安全的关键因子, 其中尤以贻贝中PSTs风险最为严峻, 我国近海中不断检出且多次诱发中毒事件, 亟需进行重点防控。国内外研究表明, PSTs可以被贻贝快速蓄积并极易超过现有安全限量标准, 随后在贻贝体内发生分布、代谢及转化等过程, 而环境因素如温度、pH、盐度、种群密度及产毒藻种类等能够影响贻贝的过滤率、摄食率, 从而改变贻贝对PSTs的蓄积率、蓄积量以及清除率。目前已经从藻和贝中分离鉴定了60多种PSTs及其代谢产物, 它们的残留能力、毒性大小和靶器官等决定了贝类中PSTs的风险表征及危害程度。因此, 后续应侧重于我国近海重点贝类增殖区, 摸清目标区域贻贝等海洋贝类中PSTs基础风险规律和变化过程, 阐明贻贝体内PSTs风险形成的内源过程及调控机理, 比较并评估不同区域贻贝中PSTs风险大小, 并据此建立区域性PSTs风险防控技术, 提高我国贻贝中PSTs风险的主动防控能力, 保障消费者健康安全和产业可持续发展。

[1] 于仁成, 罗璇. 我国近海有毒藻和藻毒素的研究现状与展望[J]. 海洋科学集刊, 2016, 51(1): 155-166.Yu Rencheng, Luo Xuan. Status and research perspectives on toxic algae and phycotoxins in the coastal waters of China[J]. Studia Marina Sinica, 2016, 51(1): 155-166.

[2] 李兆永, 陈军辉, 王帅, 等. 麻痹性贝毒素电喷雾质谱负离子模式分析特征[J]. 质谱学报, 2014, 35(1): 8-15.Li Zhaoyong, Chen Junhui, Wang Shuai, et al. Ms characteristic of paralytic shellfish poisoning toxins detected by electrospray ionization mass spectrometry in negative ion mode[J]. Journal of Chinese Mass Spectrometry Society, 2014, 35(1): 8-15.

[3] Visciano P, Schirone M, Berti M, et al. Marine biotoxins: occurrence, toxicity, regulatory limits and reference methods[J]. Front Microbiol, 2016, 7(10): 1-10.

[4] Wang D, Zhang S, Zhang Y, et al. Paralytic shell­fish toxin biosynthesis in cyanobacteria and dinoflagellates: a molecular overview[J]. Journal of Proteomics, 2016, 135: 132-140.

[5] 王艳, 黄琳, 顾海峰, 等. 中国渤海海域甲藻孢囊的种类多样性和生态地理分布[J]. 植物学报, 2012, 47(2): 125-132. Wang Yan, Huang Lin, Gu Haifeng, et al. Species diversity and eco-geographical distribution of dinofla­gellate cysts in the Bohai sea, China[J]. Chinese Bulletin of Botany, 2012, 47(2): 125-132.

[6] 翟毓秀, 郭萌萌, 江艳华, 等. 贝类产品质量安全风险分析[J]. 中国渔业质量与标准, 2020, 10(4): 1-25.Zhai Yuxiu, Guo Mengmeng, Jiang Yanhua, et al. Analysis on the quality and safety risks of shellfish products[J]. China Fishery Quality and Standards, 2020, 10(4): 1-25.

[7] 张亚亚, 闫国旺, 吴海燕, 等. 基于SPE与SPATT的水体中麻痹性贝类毒素检测方法构建与应用[J]. 海洋与湖沼, 2020, 51(2): 298-306. Zhang Yaya, Yan Guowang, Wu Haiyan, et al. Establishment and application of detection methods to paralytic shellfish poisoning in water based on SPE and SPATT methods[J]. Oceanologia et Limnologic Sinica, 2020, 51(2): 298-306.

[8] Hong N H, Jia L L, Shou E L, et al. Simultaneous determination of twelve paralytic shellfish poisoning toxins in bivalve molluscs by UPLC-MS/MS and its applications to a food poisoning incident[J]. Toxicon, 2019, 6(11): 1-16.

[9] 黄宏南, 张锦宏, 李海杰, 等. 漳州市2018—2019年沿海海域贝类样品毒素监测结果分析[J]. 海峡预防医学杂志, 2020, 26(3): 12-15.Huang Hongnan, Zhang Jinhong, Li Haijie, et al. Analysis on toxin monitoring result of shellfish samples in coastal waters in Zhangzhou city, 2018—2019[J]. Strait Journal of Preventive Medicine, 2020, 26(3): 12-15.

[10] Isabel L D C, Ana P, Ines R, et al. Paralytic shellfish poisoning due to ingestion of contaminated mussels: a 2018 case report in Caparica (Portugal)[J]. Toxi­con: X, 2019, 4(2): 10-17.

[11] Costa P, Robertson A, Quilliam M. Toxin profile of(dinophyceae) from the Portuguese coast, as determined by liquid chromatography tandem mass spectrometry[J]. Marine Drugs, 2015, 13(4): 2046-2062.

[12] Regueiro J, Negreira N, Carreira C A, et al. Dietary exposure and neurotoxicity of the environmental free and bound toxin β-N-methylamino-l-alanine[J]. Food Research International, 2017, 100(3): 1-13.

[13] 张亚亚. 秦皇岛贻贝增殖区麻痹性贝类毒素的源头解析与监控预警技术研究[D]. 连云港: 江苏海洋大学, 2020.Zhang Yaya. Source analysis and forecasting technology of paralytic shellfish toxins in mussels culture area of Qinhuangdao[D]. Lianyungang: Jiangsu Ocean University, 2020.

[14] Degger N, Chiu J, Beverly, et al. Heavy metal contamination along the China coastline: a comprehensive study using artificial mussels and native mussels[J]. Journal of Environmental Management, 2016, 180: 238-246.

[15] Ikkere L E, Perkons I, Sire J, et al. Occurrence of polybrominated diphenyl ethers perfluorinated compounds and nonsteroidal anti-inflammatory drugs in freshwater mussels from Latvia[J]. Chemosphere, 2018, 213: 507-516.

[16] Riani E, Cordova M R, Arifin Z. Heavy metal pollution and its relation to the malformation of green mussels cultured in muara kamal waters, Jakarta Bay, Indonesia[J]. Marine Pollution Bulletin, 2018, 133: 664-670.

[17] Yap C K, Cheng W H, Karami A, et al. Health risk assessments of heavy metal exposure via consumption of marine mussels collected from anthropogenic sites[J]. Science of the Total Environment, 2016, 553: 285-296.

[18] Chmist J, Szoszkiewicz K, Drożdżyński D. Behavioural responses offreshwater mus­sels to pesticide contamination[J]. Archives of Enviro­nmental Contamination and Toxicology, 2019, 77(3): 432-442.

[19] Milun V, Grgas D, Dragičević T L. Assessment of PCB and chlorinated pesticide accumulation in mussels at Kaštela bay (eastern Adriatic)[J]. Science of The Total Environment, 2016, 562: 115-127.

[20] Freitas R, Coppola F, Costa S, et al. The influence of temperature on the effects induced by triclosan and diclofenac in mussels[J]. Science of The Total Environment, 2019, 663: 992-999.

[21] Riisgard H U, Bottiger L, Pleissner D. Effect of salinity on growth of mussels,, with special reference to Great Belt (Denmark)[J]. Open Journal of Marine Science, 2012, 2(4): 167-176.

[22] 梁玉波, 李冬梅, 姚敬元, 等. 中国近海藻毒素及有毒微藻产毒原因种调查研究进展[J]. 海洋与湖沼, 2019, 50(3): 511-524.Liang Yubo, Li Dongmei, Yao Jinyuan, et al. Progresses in investigation and research on phycotoxins and toxic microalgaes in the coastal waters of China[J]. Oceanologia et Limnologic Sinica, 2019, 50(3): 511- 524.

[23] 陈建华, 于仁成, 孔凡洲, 等. 北黄海海域虾夷扇贝体内脂溶性藻毒素分析[J]. 海洋与湖沼, 2014, 45(4): 855-863.Chen Jianhua, Yu Rencheng, Kong Fanzhou, et al. Detection of lipophilic phycotoxin in Patinopecten yessoensis in the northern Yellow Sea[J]. Oceanologia et Limnologic Sinica, 2014, 45(4): 855-863.

[24] 时文博, 高丽娜, 韩现芹, 等. 天津市售主要经济贝类中麻痹性贝类毒素污染现状及特征分析[J]. 河北渔业, 2020, 10(2): 29-33. Shi Wenbo, Gao Lina, Han Xianqin, et al. Analysis of pollution status and composition characteristics of PSP in main economic shellfish sold in Tianjin[J]. Hebei Fisheries, 2020, 10(2): 29-33.

[25] 于姬. 北黄海虾夷扇贝体内麻痹性贝毒研究[D]. 大连: 大连海事大学, 2009. Yu Ji. Study of paralytic shellfish poisoning in Patino­pecten yessoensis from the northern Yellow Sea[D]. Dalian: Dalian Maritime University, 2009.

[26] 彭志兰, 罗海军, 王维洁, 等. 舟山海域麻痹性贝类毒素污染情况及其2种检测方法比较[J]. 食品安全质量检测学报, 2017, 8(4): 1436-1440. Peng ZhiLan, Luo Haijun, Wang Weijie, et al. Investigation of paralytic shellfish poison in waters of Zhoushan and comparison of 2 detection methods[J]. Journal of Food Safety and Quality, 2017, 8(4): 1436- 1440.

[27] 张添林, 陈锦钟, 洪舒萍, 等. 食物中毒一起麻痹性贝类毒素引起的食源性疾病暴发事件调查[J]. 中国食品卫生杂志, 2018, 30(4): 445-448. Zhang Tianlin, Chen Jingzhong, Hong Shuping, et al. Investigation of outbreaks of foodborne diseases caused by paralytic shellfish poisoning[J]. Chinese Journal of Food Hygiene, 2018, 30(4): 445-448.

[28] 江涛, 刘智勇, 李燕俊. 麻痹性贝类毒素污染状况及检测方法的比对研究[J]. 中国热带医学, 2010, 10(11): 1426-1428.Jiang Tao, Liu Zhiyong, Li Yanjun. Comparative study on contamination status and detection methods of paralytic shellfish toxins[J]. China Tropicalmedicine, 2010, 10(11): 1426-1428.

[29] 张纯超. 大亚湾贝类毒素特征研究[D]. 青岛: 中国海洋大学, 2008. Zhang Chunchao. The characteristics of paralytic shellfish toxin and diarrhetic shellfish toxin in Daya bay[D]. Qingdao: Ocean University of China, 2008.

[30] 江天久, 包财, 雷芳, 等. 广东东部沿海麻痹性贝类毒素成分特征分析[J]. 中国水产科学, 2010, 17(1): 119-127. Jiang Tianjiu, Bao Cai, Lei Fang, et al. Analysis on the profiles of paralytic shellfish poisoning in shellfish collected from eastern coast of Guangdong[J]. Journal of Fishery Sciences of China, 2010, 17(1): 119-127.

[31] 过锋, 崔毅, 陈碧鹃, 等. 胶州湾贝类体内有毒有害物质污染状况分析[J]. 渔业科学进展, 2011, 32(6): 115-120. Guo Feng, Cui Yi, Chen Bijuan, et al. Analysis of toxic and harmful substances in shellfish of Jiaozhou Bay[J]. Progress in Fishery Science, 2011, 32(6): 115- 120.

[32] 林祥田, 张明生, 王志坚, 等. 连云港海州湾麻痹性贝类毒素中毒分析[J]. 中国食品卫生杂志, 2005, 17(3): 243-246. Lin Xiangtian, Zhang Mingsheng, Wang Zhijian, et al. Analysis of poisoning characteristics of paralytic shellfish poison in Haizhou bay of Lianyungang, Jiangsu province[J]. Chinese Journal of Food Hygiene, 2005, 17(3): 243-246.

[33] 马丹. 渤海湾天津海域贝类质量调查及活性研究[D]. 天津: 天津大学, 2015. Ma Dan. Research on seashells quality and activity in Tianjin nearshore waters of the Bohai gulf in China[D]. Tianjin: Tianjin University, 2015.

[34] 钱蓓蕾, 徐捷, 王媛, 等. 上海市售贝类产品中麻痹性贝类毒素污染状况调查及其评价[J]. 食品安全质量检测学报, 2012, 3(2): 89-92.Qian Beilei, Xu Jie, Wang Yuan, et al. Investigation and evaluation of paralytic shellfish poison in shellfish collected from aquatic product wholesale market in Shanghai[J]. Journal of Food Safety and Quality, 2012, 3(2): 89-92.

[35] 马元庆, 唐学玺, 刘义豪, 等. 山东半岛近海贝类污染状况调查与评价[J]. 海洋环境科学, 2009, 28(5): 562-565.Ma Yuanqing, Tang Xuexi, Liu Yihao, et al. Investigation and assessment on pollution situation of seashells in Shandong peninsula coast[J]. Marine Environmental Science, 2009, 28(5): 562-565.

[36] 夏远征, 王双双, 辛丘岩, 等. 大连海域贝类麻痹性贝毒的污染状况调查与分析[J]. 食品与机械, 2010, 26(2): 54-56. Xia Yuanzheng, Wang Shuangshuang, Xin Qiuyan, et al. Pollution survey of paralytic shellfish poison (PSP) from aquaculture zones of Changhai’s sea area in Dalian[J]. Journal of Food and Machinery, 2010, 26(2): 54-56.

[37] 刘晓丽. 壳聚糖及其衍生物脱除牡蛎中麻痹性贝类毒素的研究[D]. 湛江: 广东海洋大学, 2011. Liu Xiaoli. Studies on chitosan and its derivatives adsorption for paralytic shellfish poisoning (PSP) in ostrea rivularis could[D]. Zhanjiang: Guangdong Ocean University, 2011.

[38] 沈和定, 付金花, 冉福. 麻痹性贝毒在文蛤体内的累积及净化技术研究[J]. 海洋科学, 2011, 35(7): 45-50. Shen Heding, Fu Jinhua, Ran Fu. Accumulation and detoxification of paralytic shellfish poison(PSP) in hard clam Meretrix[J]. Marine Sciences, 2011, 35(7): 45-50.

[39] Gracia V, Tobke J L, Montoya N G, et al. Experimental exposure of the mussel(d’ Orbigny 1842) to the dinoflagellatefrom Argentine Patagonia[J]. Ecotoxicology, 2020, 29(2): 226-235.

[40] Sekiguchi K, Sato S, Kaga S, et al. Accumulation of paralytic shellfish poisoning toxins in bivalves and an ascidian fed oncells[J]. Fisheries Science, 2001, 67(2): 301-305.

[41] 邴晓菲. 作为饵料组成亚历山大藻对扇贝质量安全形成的影响[D]. 上海: 上海海洋大学, 2017. Bing Xiaofei. Composite effects of Alexandrium spp. as nutritional composition quality and safety of scallop Chlamys farreri[D]. Shanghai: Shanghai Ocean University, 2017.

[42] 邱江兵. 双壳贝类对麻痹性贝毒的代谢转化及其生理生化响应[D]. 青岛: 中国海洋大学, 2014. Qiu Jiangbing. Metabolic transformation of paralytic shellfish toxins by bivalve molluscs and their physiological and biochemical responses[D]. Qingdao: Ocean University of China, 2014.

[43] Shumway S, Cucci T L. The effects of the toxic dinoflagellateon feeding and behavior of bivalve molluscs[J]. Aquatic Toxicol, 1987, 2(10): 9-27.

[44] Choi M C, Hsieh D P H, Lam P K S, et al. Field depuration and biotransformation of paralytic shellfish toxins in scallopand green-lipped mussel[J]. Marine Biology, 2003, 143(5): 927-934.

[45] 颜天, 傅萌, 李钧, 等. 麻痹性贝毒PSP在紫贻贝体内的累积、转化与排出[J]. 海洋与湖沼, 2001, 32(4): 420-427.Yan Tian, Fu meng, Li Jun, et al. Accumulation, trans­formation and elimination of PSP in Mytilus edulis[J]. Oceanologia et Limnologic Sinica, 2001, 32(4): 420-427.

[46] 陈光. 塔玛亚历山大藻所产麻痹性贝毒(PSP)在菲律宾蛤仔体内的累积与排出[D]. 青岛: 国家海洋局第一海洋研究所, 2008.Chen Guang. Accumulation and elimination of paralytic shellfish poisoning (PSP) from Alexandrium tamatense in Venerupis philippinaram[D]. Qingdao: The First Institute of Oceanography, State Oceanic Administration, 2008.

[47] Bricelj V M, Connell L, Konoki K, et al. Sodium channel mutation leading to saxitoxin resistance in clams increases risk of PSP[J]. Nature, 2005, 434(7): 763-767.

[48] Braga A C, Camacho C, Marques A, et al. Combined effects of warming and acidification on accumulation and elimination dynamics of paralytic shellfish toxins in mussels[J]. Environmental Research, 2018, 164: 647-654.

[49] Anestis A, Lazou A, Pörtner H O, et al. Beha­vioral, metabolic, and molecular stress responses of marine bivalveduring long-term acclimation at increasing ambient temperature[J]. Integrative and Comparative Physiology, 2007, 293(2): 911- 921.

[50] Dallas L J, Bean T P, Turner A, et al. Exposure to tritiated water at an elevated temperature: genotoxic and transcriptomic effects in marine mussels[J]. Journal of Environmental Radioactivity, 2016, 164: 325-336.

[51] Filgueira R, Guyondet T, Comeau L A, et al. Bivalve aquaculture-environment interactions in the context of climate change[J]. Global Change Biology, 2016, 22(12): 3901-3913.

[52] Lischka S, Büdenbender J, Boxhammer T, et al. Impact of ocean acidification and elevated temperatures on early juveniles of the polar shelledmortality shell degradation an shell growth[J]. Biogeosciences, 2011, 8(4): 919-932.

[53] Schulte E H. Influence of algal concentration and temperature on the filtration rate of[J]. Marine Biology, 1975, 30(4): 331-341.

[54] Almada P, Villela C. The effects of reduced salinity on the shell growth of small[J]. Marine Biological Association of the United Kingdom, 1984, 64(1): 171-182.

[55] Gruffydd L D, Huxley R, Crisp D J. The reduction in growth ofin fluctuating salinity regimes measured using laser diffraction patterns and the exaggeration of this effect by using tap water as the diluting medium[J]. Marine Biological Association of the United Kingdom, 1984, 64(2): 401-409.

[56] Maar M, Saurel C, Landes A, et al. Growth potential of blue mussels () exposed to different salinities evaluated by a dynamic energy budget model[J]. Journal of Marine Systems, 2015, 148: 48-55.

[57] Riisgard H U, Lüskow F, Pleissner D, et al. Effect of salinity on filtration rates of musselswith special emphasis on dwarfed mussels from the low-saline central Baltic sea[J]. Helgoland Marine Research, 2013, 67(3): 591-598.

[58] Riisgard H U, Lundgreen K, Larsen P S. Potential for production of mini-mussels in Great Belt (Denmark) evaluated on basis of actual and modeled growth of young mussels[J]. Aquaculture International, 2014, 22(2): 859-885.

[59] Riisgard H U, Mulot M, Merino L, et al. Effect of salinity changing rates on filtration activity of mussels from two sites within the Baltic hybrid zone: the brackish Great Belt (Denmark) and the low saline central Baltic sea[J]. Open Journal of Marine Science, 2014, 4(2): 101-109.

[60] Che Y, Ding L, Qiu J, et al. Conversion and stability of new metabolites of paralytic shellfish toxins under different temperature and pH conditions[J]. Journal of Agricultural and Food Chemistry, 2020, 68(5): 1427- 1435.

[61] Hu M, Li L, Sui Y, et al. Effect of pH and temperature on antioxidant responses of the thick shell mussel[J]. Fish & Shellfish Immunology, 2015, 46(2): 573-583.

[62] Parker L, Ross P, O'Connor W, et al. Predicting the response of molluscs to the impact of ocean acidification[J]. Biology, 2013, 2(2): 651-692.

[63] Basile M, Christos O, Andreas P, et al. Effects of long-term moderate hypercapnia on acid base balance and growth rate in marine mussels[J]. Marine Ecology Progress Series, 2005, 293: 109-118.

[64] Capelle J, Stralen M, Wijsman J, et al. Population dynamics of subtidal blue musselsand the impact of cultivation[J]. Aquaculture Environment Interactions, 2017, 9: 155-168.

[65] 黄德强, 暨卫东, 高亚辉, 等. 翡翠贻贝对塔玛亚历山大藻的摄食及毒素积累的初步研究[J]. 台湾海峡, 2003, 22(4): 426-430. Huang Deqiang, Ji Weidong, Gao Yahui, et al. Primary study on grazing rate of Perna viridis on Alexandrium tamarense and its PSP accumulation[J]. Journal Oceanography in Taiwan Strait, 2003, 22(4): 426-430.

[66] Lee J S, Jeon J K, Myung S H, et al. Paralytic shellfish toxins in the musseland dino­flagellatefrom Jinhae bay, Korea[J]. Korean Journal of Fisheries and Aquatic Sciences, 1992, 25(2): 144-150.

[67] 刘仁沿, 刘磊, 梁玉波, 等. 我国近海有毒微藻及其毒素的分布危害和风险评估[J]. 海洋环境科学, 2016, 35(5): 787-800. Liu Renyan, Liu Lei, Liang Yubo, et al. The distribution, impacts and risks of toxic microalgae and phycotoxins in China[J]. Marine Environmental Science, 2016, 35(5): 787-800.

[68] 赵俊梅, 方建光, 包振民, 等. 3种滤食性贝类对塔玛亚历山大藻的摄食研究[J]. 海洋水产研究, 2004, 25(4): 17-22. Zhao Junmei, Fang Jianguang, Bao Zhenmin, et al. Filter-feeding of three bivalves on Alexandrium tamarense[J]. Marine Fisheries Research, 2004, 25(4): 17-22.

[69] Negri A P, Jones G J. Bioaccumulation of paralytic shellfish poisoning (PSP) toxins from the cyanobacteriumby the freshwater mussel[J]. Toxicon, 1995, 33(5): 667-678.

[70] 田华. 麻痹性贝毒的累积、转化、排出过程及预警诊断指标研究[D]. 青岛: 中国海洋大学, 2009. Tian Hua. Study on accumulation, transformation and depuration and diagnoses target of paralytic shellfish poisoning toxins[D]. Qingdao: Ocean University of China, 2009.

[71] Dean K J, Hatfield R G, Lee V, et al. Multiple new paralytic shellfish toxin vectors in offshore north sea benthos, a deep secret exposed[J]. Marine Drugs, 2020, 18(8): 400-412.

[72] Tse S, Lee F, Mak D, et al. Production of paralytic shellfish toxins (PSTs) in toxicis intertwined with photosynthesis and energy production[J]. Toxins, 2020, 12(8): 477-487.

[73] Turner A D, Tarnovius S, Hatfield R G, et al. Application of six detection methods for analysis of paralytic shellfish toxins in shellfish from four regions within Latin America[J]. Marine Drugs, 2020, 18(12): 6-16.

[74] Kim H, Shin I. Comparison of paralytic shellfish toxin profiles ofand blue mussel () in Korea[J]. Food Science and Biotechnology. 2015, 24(2): 751-756.

[75] Wiese M, D Agostino P M, Mihali T K, et al. Neurotoxic alkaloids: saxitoxin and its analogs[J]. Marine Drugs, 2010, 8(7): 2185-2211.

[76] Botelho M J, Marques F, Freitas R, et al. Paralytic shellfish toxin profiles in mussel, cockle and razor shell under post-bloom natural conditions: evidence of higher biotransformation in razor shells and cockles[J]. Marine Environmental Research, 2020, 154: 104839.

[77] Vale P. Metabolites of saxitoxin analogues in bivalves contaminated by[J]. Toxicon, 2010, 55: 162-165.

[78] Cho Y, Ogawa N, Takahashi M, et al. Purification and characterization of paralytic shellfish toxin- transforming enzyme, sulfocarbamoylase Ⅰ, from the Japanese bivalve[J]. Biochimica et Biophysica Acta (BBA) - Proteins and Proteomics, 2008, 1784(9): 1277-1285.

[79] O’Neill K, Musgrave I F, Humpage A. Low dose extended exposure to saxitoxin and its potential neurodevelopmental effects: a review[J]. Environmental Toxicology and Pharmacology, 2016, 48: 7-16.

[80] Goya A B, Tarnovius S, Hatfield R G, et al. Paralytic shellfish toxins and associated toxin profiles in bivalve mollusc shellfish from Argentina[J]. Harmful Algae, 2020, 99: 101910.

[81] Botelho M J, Vale C, Ferreira J G. Seasonal and multi-annual trends of bivalve toxicity by PSTs in Portuguese marine waters[J]. Science of the Total Environment, 2019, 664: 1095-1106.

[82] Ding L, Qiu J, Li A. Proposed biotransformation pathways for new metabolites of paralytic shellfish toxins based on field and experimental mussel samples[J]. Journal of Agricultural and Food Chemistry, 2017, 65(27): 5494-5502.

[83] Lin H, Cho Y, Yashiro H, et al. Purification and characterization of paralytic shellfish toxin transforming enzyme from[J]. Toxicon, 2004, 44(6): 657-668.

[84] 卞中园. 麻痹性贝毒在牡蛎体内蓄积、分布、转化以及羧甲基壳聚糖的脱[D]. 湛江: 广东海洋大学, 2013. Bian Zhongyuan. Study on the accumulation, distribution, transformation for paralytic shellfish poisoning and the depuration of CM-chitosan in oyster[D]. Zhanjiang: Guangdong Ocean University, 2013.

[85] 邹迎麟, 朱明远, Hall S. 两种亚历山大藻产毒过程和毒素特征研究[J]. 黄渤海海洋, 2001, 19(3): 65-70. Zou Yinglin, Zhu Mingyuan, Hall S. Production and characteristics of paralytic shellfish toxins in two species of Alexandrium[J]. Journal of Oceanography of Huanghai&Bohai Seas, 2001, 19(3): 65-70.

[86] Galimany E, Sunila I, Hégaret H, et al. Pathology and immune response of the blue mussel () after an exposure to the harmful dinoflagellate[J]. Harmful Algae, 2008, 7(5): 630-638.

[87] Kotaki Y, Tajiri M, Oshima Y, et al. Identification of a calcareous red alga as the primary source of paralytic shellfish toxins in coral reef crabs and gastropods[J]. Bulletin of the Japanese Society of Scientific Fisheries, 1983, 49(2): 283-286.

[88] Liu Y, Li L, Zheng L, et al. Antioxidant responses of triangle sail musselexposed to harmful algaeand high pH[J]. Chemosphere, 2020, 243: 125-241.

[89] Qiu J, Ma F, Fan H, et al. Effects of feeding, a paralytic shellfish toxin producer, on antioxidant enzymes in scallops () and mussels ()[J]. Aquaculture, 2013, 396(3): 76-81.

[90] Epp O, Ladenstein R, Wendel A, et al. The refined structure of the selenoenzyme glutathione pero­xidase at 0.2nm resolution[J]. European Journal of Bio­chemistr, 1983, 133(1): 51-69.

[91] Pousse E, Sainte M J, Alunno B M, et al. Sources of paralytic shellfish toxin accumulation varia­bility in the Pacific oyster[J]. Toxicon, 2018, 144: 14-22.

[92] Bianchi V A, Langeloh H, Tillmann U. Separate and combined effects of neurotoxic and lytic compounds ofstrains onfeeding activity and hemocyte function[J]. Fish & Shellfish Immunology, 2019, 84: 414-422.

Research progress on the accumulation and metabolism of paralytic shellfish toxin in mussels

FAN Li-qiang1, 2, ZHENG Guan-chao2, WU Hai-yan2, GUO Meng-meng2, LU Xia1, LI Yu1, TAN Zhi-jun2, 3

(1. School of Geomatics and Marine Information, Jiangsu Ocean University, Lianyungang 222005, China; 2. Key Laboratory of Testing and Evaluation for Aquatic Product Safety and Quality, Ministry of Agriculture and Rural Affairs, P. R. China, Yellow Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Qingdao 266071, China; 3. Pilot National Laboratory for Marine Science and Technology (Qingdao), Qingdao 266237, China)

Poisoning with paralysis shellfish toxins (PSTs) can cause death in shellfish, other fish, and other marine organisms as well as hamper the marine ecology and human health. Mussels are highly accumulative and serve as indicators of PSTs in foreign countries. In recent years, several cases of PST poisoning by mussel consumption in China have become the focus of safety attention. Therefore, understanding the mechanism of risk formation is essential and urgent. This study evaluates the risk degree of PSTs in offshore mussels and other shellfishes in China and the accumulation effect of mussels on PSTs from different toxigenic alga sources, analyzes the influence of environmental factors on the accumulation and metabolism, and explores in depth the specificity of mussel accumulation and metabolism. Furthermore, focusing on the expression regulation process of the mussel metabolic stage, our study clarifies the influencing factors, transformation functions, and modes. In future studies, we aim to focus on understanding the basic risk rule and change process of PSTs in mussels as well as other marine shellfish in certain target areas of China and establish regional risk prevention and control technology for active prevention and control of PST risk in mussels of China. This will contribute positively and significantly toward protecting consumer health and the marine ecology.

mussels; accumulation and metabolism; immune; risk assessment

Nov. 2, 2020

S917

A

1000-3096(2021)04-0201-12

10.11759/hykx20201102003

2020-11-02;

2020-12-25

国家重点研发计划项目(2017YFC1600701); 国家自然科学基金面上项目(31772075, 32072329); 2019江苏省研究生科研创新项目(KYCX19_2282)

[National Key R&D Program of China, No.2017YFC1600701; The National Natural Science Foundation of China, No.31772075, 32072329; 2019 Jiangsu Graduate Scientific Research Innovation Program, No.KYCX19_ 2282]

范礼强(1996—), 男, 在读研究生, 研究方向为水产品质量与安全, E-mail: liqiangfan0@gmail.com; 谭志军(1978—),通信作者, 研究员, 主要从事贝类毒素安全检测及评价研究, E-mail: tanzj@ysfri.ac.cn

(本文编辑: 赵卫红)