微塑料对海洋桡足类摄食的影响与微塑料的类型、尺寸以及桡足类的种类有关[1, 25]。微塑料对猛水蚤的摄食率和滤水率具有消极作用,且具有剂量-效应关系,即随着微塑料浓度的增大,摄食率和滤水率下降幅度越大。原因可能是由于微塑料对猛水蚤消化系统的机械阻塞,导致摄入微塑料后无法消化。微塑料会聚团或堆积在动物消化道内,“饱腹感”使其摄食率降低,从而摄入营养和能量不足,导致生殖能力等降低[13, 26]。另外,微塑料会附着在桡足类的触角、附肢、背甲、尾叉等部位,影响其活动能力,导致其有“不适感”,从而减少摄食率。除了被动滤食,海洋桡足类能够感知化学物质,当存在藻和微塑料时,桡足类表现为择食性,桡足类会避开摄食与微塑料大小、形状相似的藻[27]。Cole等[28]研究了尼龙对飞马哲水蚤摄食的影响,研究表明以不同藻作为饵料,尼龙纤维和尼龙微粒对飞马哲水蚤的摄食影响不同,飞马哲水蚤表现为选择性摄食,当饵料为圆海链藻、锥状克里普藻时,尼龙纤维显著抑制摄食率,而尼龙微粒对上述两种藻的摄食率没有明显影响。我们的结果显示尼龙6微粒对猛水蚤的摄食率有抑制作用,与Cole等[28]结果不同的原因主要是因为我们所用动物类型、微塑料类型及浓度不同,我们的微塑料浓度高于Cole等[28]所用的浓度。本实验所用的微塑料浓度12.5~50 mg·L-1大约为2.4×107~6.3×109 items·L-1,与饵料浓度1.5 ×108cells·L-1相近;微塑料浓度为100~200 mg·L-1大约为1.9×108~2.5×1010 items·L-1,高于所用的饵料浓度。随着微塑料浓度的增大,尽管桡足类对微塑料有化感作用,但低选择性滤食仍然导致桡足类对微塑料的摄食,并且微塑料浓度越大对摄食率和滤水率的有害作用越大。微塑料对海洋生物不利影响所表现出的剂量-效应关系在Rehse等[22]中已有报道。本实验中的PA 6对猛水蚤的24 h·EC50与Frydkjær等[29]的不规则微塑料聚乙烯(polyethylene,PE)对枝角类大型蚤Daphnia magna 48 h·EC50 (65 mg·L-1)比较相近。Rehse等[22]研究表明1 μm PE对大型蚤96 h·EC50为57.4 mg·L-1。目前已有一些微塑料对浮游动物EC50的报道[22, 29],但微塑料对浮游动物的急性致死效应尚未见报道,因此需要开展微塑料长期暴露对浮游动物的亚致死实验。
本文在摄食率实验测定时发现,在不添加桡足类的对照组中,添加微塑料与无添加微塑料的对照相比,微塑料尼龙6的暴露使三角褐指藻的生长速率降低。采用的尼龙6的浓度与Zhang等[30]的浓度相近,虽然所用的微塑料和藻的类型不同,但对藻的生长抑制效应相近。微塑料对藻可能会产生抑制细胞生长、破坏细胞膜结构、降低叶绿素含量和光合效率、引起细胞氧化损伤等毒性效应[30-31]。Zhang等[30]研究了微塑料对中肋骨条藻的生态影响,结果表明微塑料聚氯乙烯(polyvinyl chloride,PVC)抑制藻的生长和光合作用,电镜扫描结果表明,微塑料的吸附和聚团是造成抑制作用的原因。微塑料对微藻的致毒机理比较复杂,已提出的可能的致毒机理主要有机械损伤、遮蔽效应、氧化损伤、吸附团聚,并且这些致毒效应往往同时存在,致使藻细胞生长代谢异常甚至死亡[19]。微塑料影响藻细胞生长的同时,藻细胞也会产生应激反应以减小、修复微塑料对其造成的损伤[32]。微塑料对藻生理生化、营养的影响[32]推测也会对桡足类的摄食产生间接的影响,尚需进一步证实。
3.2 微塑料对桡足类排泄的影响摄食率的降低直接导致猛水蚤的排粪率随着微塑料浓度的升高而降低。微塑料尼龙6对猛水蚤总排粪率的24 h·EC50(221.2 mg·L-1)高于对正常排粪率的24 h·EC50(84.1 mg·L-1),这与总粪便颗粒中含有异常粪便颗粒有关。由于尼龙6对猛水蚤总排粪率的24 h·EC50仅涉及微塑料对排粪颗粒数量的抑制,无法代表对颗粒大小、密度的影响,因此不能全面衡量微塑料对猛水蚤排泄的消极影响。同时因为微塑料的摄入,猛水蚤排出体积小、短小椭球状的粪便颗粒,且随着微塑料浓度的升高,异常粪便在总粪便颗粒里所占百分比不断升高。微塑料的摄入使粪便颗粒的组成和密度发生改变,导致密度低于正常粪便的密度,而不同密度的微塑料也表现为不同的沉降速率[27]。本实验发现暴露微塑料的粪便不容易沉降,与正常粪便相比,其沉降速率显著降低。同时因为猛水蚤粪便中微塑料含量增大,而微塑料黏度较差,导致其粪便颗粒更易散开,而导致异常粪便的出现[24]。Coppock等[27]指出粪便颗粒的体积与微塑料的形状有关,而且粪便沉降速率与微塑料的类型有关。
3.3 微塑料对桡足类生殖的影响微塑料会降低海洋桡足类的生殖能力和存活率[25]。本实验中,微塑料延迟猛水蚤雌体的首次抱卵,微塑料浓度越高,抱卵率越低。这主要是由于微塑料的摄入导致桡足类摄入的能量和营养不足[33-34],所以延迟抱卵,抱卵率降低。Sussarellu等[26]研究表明微塑料聚苯乙烯对太平洋牡蛎的生殖功能具有伤害作用,卵母细胞的数量、直径显著下降。本实验所用的微塑料浓度暴露未导致桡足类死亡,但推测可能已经对猛水蚤的卵母细胞结构产生影响,并影响其孵化率。本实验数据未能得到微塑料尼龙6对猛水蚤抱卵率的24 h·EC50,随时间变化,48~144 h的EC50各不相同,因此,微塑料对猛水蚤生殖影响的作用机理及时间-效应还需进一步研究。
3.4 其他影响除了摄食、排粪、生殖,微塑料还会对桡足类其他方面产生影响,如代谢以及行为等。Deng等[35]研究表明,5和7 μm的微塑料微球均会引起鱼类肝部的局部感染及脂质积累,代谢组学分析显示,塑料微球会引起鱼类肝部代谢产物的变化并扰乱肝脏部分机制和能量的代谢。Cole等[13]研究表明微塑料聚苯乙烯(Polystyrene,PS)对桡足类Calanus helgolandicus的耗氧率没有明显影响。微塑料对桡足类代谢的影响与所用微塑料类型不同以及微塑料浓度有关。除了上述影响,微塑料对动物的行为还会有影响。丛艺等[36]研究10 μm聚苯乙烯微粒(PS)对沙蚕掘穴行为影响和细胞超微结构的改变,结果表明PS造成沙蚕掘穴行为延长、体壁表皮细胞凋亡和肌肉细胞线粒体水肿。目前,针对PS对海洋生物的研究较多,PS主要用于光学玻璃及仪器、灯罩、包装材料,其发泡塑料可作绝热材料、快餐盒等。与PS不同,尼龙6具有无毒、质轻、优良的机械强度、耐磨性及较好的耐腐蚀性,广泛应用于代替铜的机械、化工等工业中制造轴承、齿轮、泵叶,另外还用于头盔、输液管、医用缝线、渔网等。尼龙的分布也非常广泛,Alam等[37]对Ciwalengke河的微塑料进行了调查研究,结果表明表层海水平均微塑料颗粒为(5.85±3.28)个/L,主要微塑料类型为聚酯纤维(Polyester)和尼龙纤维。因此研究尼龙6对海洋桡足类的毒理效应对于研究微塑料污染对海洋食物网的影响具有重要意义。
4 结论(1) 微塑料尼龙6的短期暴露使猛水蚤摄食率、滤水率和排粪率降低,而且降低幅度与微塑料浓度有关,浓度越大,降低幅度越大。微塑料尼龙6对猛水蚤摄食率、滤水率、排粪率24 h·EC50分别为67.7、62.2、84.1 mg·L-1。另外,尼龙6使猛水蚤的粪便颗粒小型化,由长椭球体变为短小椭球体,可能与其粘度或物理结构的改变有关。微塑料尼龙6暴露导致粪便颗粒的沉降速率显著降低。
(2) 微塑料尼龙6暴露使猛水蚤抱卵延迟,相同培养时间的抱卵率降低。可能是微塑料的存在导致猛水蚤摄食率的降低,进而营养和能量摄入不足,进一步导致对生殖活动的影响。
本实验所用的微塑料浓度高于现场环境,因此用本结果推到现实海洋环境中时,还需考虑现场微塑料种类、浓度、作用时间及其不同的生态效应。
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来源:中国海洋大学