广东农业科学
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作者简介:

赵红霞(1976—),女,博士,研究员,研究方向为水产动物营养与饲料,E-mail:zhaohongxia8866@163.com

通信作者:

曹俊明(1962—),男,博士,研究员,研究方向为水产动物营养与饲料,E-mail:junmcao@163.com

中图分类号:S963.73

文献标志码:A

文章编号:1004-874X(2020)11-0135-09

DOI:10.16768/j.issn.1004-874X.2020.11.015

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目录contents

    摘要

    水产饲料添加剂具有提高水产动物摄食率和饲料转化效率、促进生长和发育、调节机体代谢机能或免疫功能、增强抗病力、改善水产品品质等作用。近年来,国内外已有较多关于水产饲料添加剂的研究报道,在传统配合饲料中使用水产饲料添加剂可调控水产动物生长、品质、免疫和抗病抗应激能力,能产生良好的经济效益、社会效益和环境效益,因此水产饲料添加剂已成为科研热点和商业焦点。综述了免疫刺激剂、促生长和抗氧化剂、诱食剂、植物提取物等新型水产饲料添加剂在水产动物中的作用机制、应用效果和适宜添加剂量,并指出其存在的问题和未来的研究方向,以期为促进我国水产饲料添加剂的研究与应用提供参考。

    Abstract

    Aquatic feed additives play important roles in raising the feeding rate of aquatic animal and the feed conversion efficiency, promoting growth and development, regulating body metabolic function or immune function, strengthening disease resistance and improving quality of aquatic products. In recent years, many studies about aquatic feed additives at home and abroad have been reported. In order to produce good economic, social and environmental benefits, aquatic feed additives are used in traditional compound feeds to improve the growth and quality and regulate immunity and disease-resistant response ability of aquatic animals. Therefore, aquatic feed additives have been a hot spot of scientific researches and a focal point of business. The mechanism, application effect and appropriate dose of aquatic additives in aquatic animals, such as immunologic stimulants, growth promoters, antioxidants, feeding attractants and plant extracts were summarized. The existing problems in the previous researches and future research directions of aquatic feed additives were pointed out to provide references for the promotion of the research and application of aquatic feed additives in China.

    关键词

    水产动物饲料添加剂营养免疫抗氧化

  • 水产饲料添加剂是在饲料生产加工、使用过程中添加的少量或微量物质,对强化基础饲料营养价值、提高动物生产性能、保证动物健康、节省饲料成本和改善水产品品质等方面有明显的效果。水产饲料添加剂可分为营养性(氨基酸添加剂、维生素添加剂、微量元素添加剂等)和非营养性饲料添加剂(免疫刺激剂、抗氧化剂、促生长剂、诱食剂等)。 水产饲料添加剂是配合饲料的核心,种类繁多、成分多样、功能广泛,虽然在水产配合饲料中用量少, 但却发挥着重要作用。我国水产动物营养研究与饲料工业经过40 多年的快速发展,取得了巨大成就。 1991 年我国水产饲料产量75 万t,2019 年则达到2200 多万t,而水产养殖产量达到5000 万t,占世界总产量的2/3[1]。然而,随着水产养殖规模化、 集约化程度的提高,养殖过程中面临着大量的如拥挤、环境、代谢等因子的激烈应激,使水产动物抗病力减弱,易暴发大规模病害,影响水产养殖业的健康可持续发展。因此,迫切需要寻求解决水产养殖安全生产的新途径。应用营养调控提高水产动物抗氧化应激能力和免疫力是一种有效途径,使用水产饲料添加剂也是一种有效手段。加强营养与免疫学方面的研究工作将有利于推动水产养殖业的健康可持续发展,因而具有重要的理论意义和实际应用价值。研究表明,水产饲料添加剂在提高饲料转化效率、促进营养吸收、增强抗病能力、减少环境污染等方面都有积极作用。广东省农业科学院动物科学研究所近10 多年来较系统深入地开展了水生经济动物特别是华南地区特色水产动物的抗氧化应激和免疫力营养调控研究,并开展水产动物的营养需要研究、功能性添加剂研发与应用,以及鱼虾环保高效饲料的研发和推广等,在水产动物抗氧化应激和免疫力营养调控等方面取得了创新性成果。本文综述了免疫刺激剂、促生长和抗氧化剂、诱食剂、 植物提取物等水产饲料添加剂的研究进展,以期为促进我国水产饲料添加剂的研究与应用提供参考。

  • 1 水产饲料功能性添加剂的开发应用

  • 1.1 免疫刺激剂:β-1,3-葡聚糖

  • β-葡聚糖广泛分布于真菌、细菌和植物中, 是目前研究最广泛的免疫刺激剂之一。饲料中添加 β-1,3-葡聚糖能促进凡纳滨对虾的生长性能、提高免疫功能和抗环境应激能力[2-3]。研究表明,长期摄食免疫刺激剂的鱼类可能产生免疫抑制和抗病力下降[4],摄食较高剂量的免疫刺激剂可能对鱼虾的存活和健康产生负面影响[5]。广东省农业科学院动物科学研究所开展了 β-1,3-葡聚糖在凡纳滨对虾、斑节对虾、草鱼、黄颡鱼和花鲈中的应用研究,获得了 β-1,3-葡聚糖在不同水产动物中的最适添加量:凡纳滨对虾饲料中添加量为336.5 mg/kg, 斑节对虾和草鱼饲料中添加量为400 mg/kg,黄颡鱼饲料中添加量为600 mg/kg,花鲈饲料中添加量为415.10 mg/kg[6-12]

  • 1.1.1 β-1,3-葡聚糖在凡纳滨对虾中的应用研究

  • β-葡聚糖使用剂量、使用时间与效应关系的研究至关重要,其实际作用效果与养殖水产动物的各个生长阶段、投喂方式和添加时间密切相关。王计刚等[6]研究凡纳滨对虾在停饲 β-1,3-葡聚糖后0、 7、14、21、28、35 d的免疫酶活性变化动态规律, 发现连续摄食 β-1,3-葡聚糖14 d后停饲的凡纳滨对虾具有最高的免疫保护率,停饲28 d相对免疫保护率达到77.01%。长期投喂 β-1,3-葡聚糖对凡纳滨对虾血液代谢物和肌肉免疫相关酶活性产生显著影响,饲料中分别添加250、500 mg/kg β-1,3-葡聚糖14、42、56 d,能够显著提高低盐度养殖凡纳滨对虾营养物质代谢和非特异性免疫功能[7-9]。 β-1,3-葡聚糖的间隔投喂方式显著提高凡纳滨对虾生长、血清代谢和抗亚硝酸氮应激能力,14 d间隔投喂0.1%β-1,3– 葡聚糖可能通过促进能量代谢和脂多糖/ß-1,3-葡聚糖结合蛋白、酚氧化物酶原和超氧化物歧化酶mRNA表达而增强凡纳滨对虾抗亚硝酸氮的应激能力。

  • 1.1.2 β-1,3-葡聚糖在花鲈中的应用研究

  • 饲料中添加 β-1,3-葡聚糖可显著提高花鲈的免疫和抗氧化能力,400 mg/kg和600 mg/kg β-1,3– 葡聚糖均能显著提高血清和肝脏免疫球蛋白M、补体C3 含量,血清超氧化物歧化酶、碱性磷酸酶、酸性磷酸酶活性,以及一氧化氮含量和总抗氧化能力, 显著降低血清和肝脏丙二醛含量。通过开展氯化铵应激试验,添加400、600 mg/kg β-1,3-葡聚糖均显著降低应激96 h的累计死亡率,相对保护率分别为60.0%和58.1%[10]。与未添加对照相比,应激后摄食 β-1,3-葡聚糖的花鲈血清酸性磷酸酶、 碱性磷酸酶和总抗氧化能力显著升高,丙二醛含量显著降低,饲料中添加一定水平的 β-1,3-葡聚糖可显著降低氨氮应激累计死亡率和提高抗氧化能力[11]

  • 1.1.3 β-1,3-葡聚糖在黄颡鱼中的应用研究

  • 饲料中分别添加200、600、1000 mg/kg β-1,3-葡聚糖,无论采取连续投喂还是间隔投喂方式,对黄颡鱼幼鱼生长性能、体成分、血清生化指标、免疫、 抗氧化和抗嗜水气单胞菌感染能力均没有显著影响,但间隔投喂1000 mg/kg β-1,3-葡聚糖显著降低黄颡鱼肝体比,显著升高肾脏白细胞介素-1β、 白细胞介素-6 和肿瘤坏死因子-α 水平。腹腔注射 β-1,3-葡聚糖可以增强黄颡鱼成鱼血清免疫球蛋白M、白细胞介素-6 水平和超氧化物歧化酶活性,降低嗜水气单胞菌感染48 h后的累计死亡率, 提高抗嗜水气单胞菌感染能力[12]

  • 1.2 促生长、抗氧化剂:谷胱甘肽

  • 谷胱甘肽( GSH)是广泛存在于细胞中的一种生物活性肽,其中在动物肝细胞、酵母和小麦胚芽细胞中较为丰富,一般达0.1~10.0 mmol/L[13]。 GSH具有活性巯基(- SH)和 γ -酰胺键,这使得它具有多种生理功能,能参与机体自由基清除、 免疫应答激活、毒素螯合、细胞完整性保护和部分物质吸收等多方面生理活动,在临床医学和营养学上已有大量相关研究[14]。水产动物饲料中添加GSH,能够增强水产动物的抗氧化应激能力,改善其机体健康,并促进生长。

  • 1.2.1 GSH在对虾中的应用研究

  • 通过对虾肝细胞离体培养试验表明,谷胱甘肽对肝细胞增殖有显著促进作用,可降低肝细胞RNA/DNA比值,提高肝细胞中类胰岛素生长因子(IGF-1)含量,超氧化物歧化酶(SOD)、谷胱甘肽过氧化物酶(GSH-PX) 活性,以及SOD、过氧化氢酶(CAT)表达量,并降低丙二醛(MDA)含量,对肝胰腺线粒体损伤有显著保护方面[15]。饲料中添加GSH能显著增加凡纳滨对虾的增重率、成活率、饲料转化效率和血清、 肝胰腺、肌肉中的蛋白浓度,提高虾肝胰腺中谷胱甘肽还原酶(GR)、GSH-PX、血清碱性磷酸酶、 酸性磷酸酶活性、总抗氧化能力(T-AOC)、血淋巴细胞计数等,并降低组织MDA含量[16]。GSH通过激发凡纳滨对虾的非特异免疫功能,提高凡纳滨对虾成活率,改善离子铵对凡纳滨对虾造成的氧化胁迫。以增重率为评价指标,GSH在凡纳滨对虾饲料中的适宜添加量为174.13 mg/kg[17]

  • 1.2.2 GSH在草鱼中的应用研究

  • 初体质量约8.50 g的草鱼,在56 d饲养期中分别投喂添加5 种不同剂量GSH(添加量分别为0、100、200、 300、400 mg/kg)的试验饲料,研究发现,饲料中添加GSH能够提高草鱼特定生长率、存活率和饲料效率,其中200 mg/kg剂量组达到显著水平[18]。 饲料中添加GSH明显提高草鱼肝胰指数,显著升高血清IGF-1 水平、血液白细胞数目,并提高草鱼对嗜水气单胞菌的抵抗能力,其中200 mg/kg GSH组草鱼攻毒后存活率达到最高[19]。以特定生长率为评价指标,GSH在草鱼饲料中的适宜添加量为350 mg/kg[18]

  • 1.2.3 GSH在罗非鱼中的应用研究

  • 饲料中添加一定量的谷胱甘肽能显著提高吉富罗非鱼幼鱼的生长性能,提高全鱼粗蛋白与粗脂肪含量、血清和肝脏IGF-1 水平及非特异性免疫相关酶活性,诱导肝脏生长激素(GH)、IGF-1、SOD、谷胱甘肽S-转移酶(GST)mRNA表达。以增重率为评价指标,吉富罗非鱼幼鱼饲料中GSH的适宜添加量为355.13 mg/kg[20]。饲料添加320 mg/kg GSH显著提高罗非鱼肝脏GST、GR、SOD、CAT活性和抗超氧阴离子(anti-O2-),降低肝脏MDA含量[21]。 在亚硝基氮应激96 h内,饲料添加320 mg/kg GSH显著降低应激的累计死亡率,饲料中添加GSH通过激活吉富罗非鱼的抗氧化性能提高其抗亚硝基氮应激能力[21]

  • 1.2.4 GSH在花鲈中的应用研究

  • 饲料添加GSH对花鲈幼鱼的生长性能和常规营养成分无明显影响,添加400 mg/kg花鲈增重率比对照组提高6.2%, 明显增强血清T-AOC、GSH-PX、GST活性和GSH含量,显著提高肝脏SOD和GSH-PX活性及白细胞介素-6 含量,抗亚硝态氮96 h累计死亡率明显低于对照组[22]。以生长性能、血清和肝脏抗氧化指标、抗亚硝态氮应激能力为评价指标,花鲈幼鱼饲料中GSH适宜添加量为300~400 mg/kg[22]

  • 1.2.5 GSH在黄颡鱼中的应用研究

  • 饲料添加适量GSH能提高黄颡鱼幼鱼生长、免疫及抗氧化性能和上调黄颡鱼igf1 基因的表达,但过量添加GSH会产生负面效果[23]。饲料中添加适量GSH可降低黄颡鱼幼鱼血细胞活性氧浓度,维持血细胞内的钙离子平衡,上调bcl2 基因和下调bax基因表达,抑制血细胞凋亡[23]。此外,饲料中添加适量GSH可降低黄颡鱼氨氮应激后血清皮质醇浓度和累计死亡率,改善应激过程中的抗氧化性能,上调hsp70 基因表达,有效提高其抗氨氮应激能力[23]。以特定生长率为评价指标,GSH在黄颡鱼饲料中的适宜用量为357.69 mg/kg[24]

  • 1.3 新型高效诱食剂

  • 1.3.1 二甲-β-基丙酸噻亭(DMPT)

  • DMPT是从海洋植物中提取的生物活性物质,大量存在于海藻和盐生高等植物体内。DMPT具有多种功能,能够作为有效的甲基供体,对哺乳动物、禽类及水产动物的营养代谢具有促进作用,可提高一些鱼类的游泳能力和抗应激能力,刺激虾类蜕壳和生长,具有类似维生素的功能[25-26]。赵红霞等[27]研制开发了以DMPT为主的复合诱食剂,在凡纳滨对虾饲料中可以替代乌贼膏发挥诱食作用, 饲料中添加400 mg/kg DMPT可以显著提高对虾的增重率,并对凡纳滨对虾血液渗透压具有调节作用。DMT(二甲基噻亭)为DMPT的同系物,饲料中添加200~400 mg/kg DMT对鲫鱼有明显的促生长效果,并有利于鲫鱼体形改善(相对体形变长) 以及提高出肉率,同时提高鲫鱼的免疫学功能、 肝胰脏功能,减轻运输应激对鱼体造成的损伤, 对应激造成的负作用有一定抑制作用。

  • 1.3.2 酵母提取物

  • 陈晓瑛等[28]研究比较了凡纳滨对虾对3 种酵母产品(鲜酵母粉、酵母核苷酸、 啤酒酵母水解物)的诱食效果,并探讨了诱食效果最佳的产品在高低蛋白水平下对凡纳滨对虾生长性能的影响。将基础饲料中3%秘鲁鱼粉等量替换为鲜酵母粉、酵母核苷酸、啤酒酵母水解物,添加氧化钇(Y2O3)、氧化钕(Nd2O3)、氧化镧(La2O3) 和氧化镱(Yb2O3)作为惰性标物,用于测定选食偏好,结果发现对虾对不同饲料的选食偏好依次为酵母核苷酸组>鲜酵母粉组>鱼粉组>啤酒酵母水解物组。在此基础上,采用2×2 双因素设计2 个蛋白水平(36%、45.27%)和2 个酵母核苷酸水平(0、 0.8%)的试验以探讨酵母核苷酸在不同蛋白水平下对凡纳滨对虾的生长的影响。结果表明,低蛋白水平能显著提高对虾存活率、终末体均质量、增重率、 特定生长率、摄食量和蛋白质保留率,降低饵料系数;饲料中添加酵母核苷酸能显著提高对虾存活率, 提高终末体均质量、增重率和特定生长率,降低饵料系数。酵母核苷酸具有较好的诱食效果,饲料添加0.8%酵母核苷酸能促进摄食,提高生长性能, 在低蛋白水平下作用更显著[28]

  • 2 其他新型饲料添加剂的开发应用

  • 2.1 核苷酸

  • 核苷酸是遗传物质DNA、RNA的重要组成成分,在机体内具有许多重要的生理生化功能。研究表明,动物处于快速生长、应激和病理状况下, 内源核苷酸往往不能满足各种代谢旺盛的组织和细胞的需要,因此补充外源核苷酸尤为重要[29]。 对大菱鲆、罗非鱼、虹鳟、大西洋鲑、美国红鱼的研究发现,外源核苷酸能够促进生长性能、肠道功能和免疫相关因子活性,核苷酸缺乏会削弱肝脏、 肠道和免疫系统的功能[29]。饲料中添加5 种核苷酸混合物(5′-腺苷酸∶5′-胞苷酸∶5′-尿苷酸二钠∶5′-肌苷酸二钠∶5′-鸟苷酸二钠=1∶1∶1∶1∶1)能够改善凡纳滨对虾幼虾的生长性能,促进摄食,改善肝胰腺功能和肠道形态,提高凡纳滨对虾体内的非特异性免疫相关因子水平, 降低脂质过氧化物含量,从而提高对虾的抗氧化能力和非特异性免疫功能,核苷酸混合物在凡纳滨对虾幼虾饲料中的适宜添加量为400~600 mg/kg[30]。 研究发现,在凡纳滨对虾低蛋白质(37.92%)饲料基础上添加400 mg/kg核苷酸混合物可明显提高饲料干物质、粗蛋白质、粗脂肪、灰分和总磷表观消化率,粗脂肪和总磷表观消化率显著高于高蛋白(46.45%)对照组,但与低蛋白(37.92%)对照组相比无明显差异;显著提高了饲料C16:0、C17:0、 C20:0、C22:1、C18:3n-3、DHA表观消化率,C14:0、 C16:1 的表观消化率显著高于高蛋白(46.45%)对照组,但与低蛋白(37.92%)对照组无明显差异; 与高、低蛋白质水平饲料相比,在低蛋白质饲料中添加核苷酸混合物具有改善凡纳滨对虾对饲料营养物质表观消化率的效果,特别是对粗脂肪、总磷和脂肪酸的作用明显[31]。而在黄颡鱼的基础饲料中分别添加0.2、0.4、0.6、1.0、1.5 g/kg混合核苷酸, 1.5 g/kg添加组黄颡鱼的增重率、特定生长率、饲料效率、蛋白质沉积率和血清胆固醇水平显著高于对照组,饲料中添加核苷酸能够提高黄颡鱼生长性能,促进脂类代谢和增强抗氧化能力[32]

  • 2.2 低聚木糖

  • 低聚木糖(xylo-oligosaccharides,XOS)是功能性寡糖常用种类,是由2~7个D-木糖经β-1,4糖苷键结合成的直链或支链低度聚合糖,有效成分以木二糖和木三糖为主,具有稳定性好、有效用量少、无残留及来源丰富等特点[32]。XOS克服了抗生素、活菌益生素、酸化素等的不足,对饲料加工和储存环境具有很高的耐受性,能抵抗胃酸的灭活作用,在水产饲料中的应用前景十分广阔。 目前,已有欧洲鲈鱼、异育银鲫、奥尼罗非鱼、 草鱼、宝石鲈、斑点叉尾鮰和凡纳滨对虾饲料添加XOS进行生长性能、消化酶活性和肠道菌群等相关研究[33]。张荣斌等[34]研究表明,饲料中添加200 mg/kg低聚木糖能显著促进对虾生长性能,降低饵料系数,提高消化酶活性,提高对虾总蛋白, 改善中肠肠道形态,提高血清PO、LZM活性和肝胰腺T-AOC活性,降低肝胰腺MDA含量,显著提高anti-O2-,显著增强抗白斑综合征病毒能力,显著提高肠道细菌数量和双歧杆菌数量,降低弧菌数量,影响肠道细菌种类[35]。另有研究表明,低聚木糖能显著改善奥尼罗非鱼肠道形态,增加基层厚度和皱襞面积,显著提高肠道双歧杆菌和乳酸杆菌数量,明显提高抗嗜水气单胞菌感染的能力。

  • 2.3 有机硒

  • 硒是动物的必需微量元素,对维持动物的正常生长、促进发育具有重要作用。硒提高水产动物生产性能可能通过以下3 条途径:(1)硒通过提高水产动物的免疫力和抗氧化能力来提高生产性能[36];(2)硒通过5′-脱碘酶介导甲状腺激素的形成和降解,控制生长激素基因的表达和生长激素的合成,从而促进生长[37];(3)硒是许多酶的组成部分和激活因子,具有分解、合成、稳定酶蛋白四级结构、调整酶活性等多种生物作用,硒亦可能通过提高消化酶活性,增加消化吸收速率,促进动物生长[36]。硒在水产动物中的抗氧化作用被广泛研究,其能够增强抗氧化的能力在水产动物体液水平以及基因水平上均已被证实。Hu等[38]系统开展硒的营养生理及其抗氧化应激调控研究,确定了黄颡鱼幼鱼对硒的适宜需要量为0.20 mg/kg。 适量的硒能提显著高黄颡鱼的抗氧化能力,缺硒和高硒激活ROS信号分子,诱导细胞凋亡对黄颡鱼造成氧化应激,高硒诱导Ca2+ 信号分子产生,在细胞凋亡中发挥重要作用,黄颡鱼通过上调hsp70 基因表达降低高硒对机体产生的氧化应激[38]。研究发现,有机硒的促生长效果优于无机硒,能显著提高全鱼和肌肉组织中硒的沉积及肝脏抗氧化能力, 酵母硒通过提高鱼体血糖含量,降低尿素氮排出, 提高黄颡鱼的抗低温胁迫能力[39]

  • 2.4 抗菌肽

  • 抗菌肽是机体的内源性抗生素,是先天性免疫体系的重要组成部分,可广谱灭杀革兰氏阴性菌、 阳性菌等多种有害菌,对真菌、螺旋体、包膜病毒等微生物也有抗菌活性,甚至对某些病毒和肿瘤细胞有抑制作用[40]。研究表明,抗菌肽在凡纳滨对虾、 罗非鱼、锦鲤、草鱼等具有提高免疫和促进生长的作用,其作为一种安全、环保、高效的饲料添加剂具有广阔的应用前景[41]。李林等[42]采用超声波诱导的方式筛选出生长状况良好的蝇蛆,通过组织匀浆、乙酸铵浸提、热处理、冻融处理、盐析、透析、 蒸馏浓缩等方法提取家蝇抗菌肽,应用分光光度法测定其浓度和提取率,比浊法测定其抗菌活性。饲料中添加0.2%~1.0%家蝇抗菌肽可显著提高黄颡鱼的增重率、特定生长率和饲料效率,并显著提高白细胞吞噬百分比,以副溶血弧菌进行攻毒,随抗菌肽添加量的增加,黄颡鱼死亡率有下降趋势。饲料中添加适量的家蝇抗菌肽对凡纳滨对虾有一定的促生长作用,可提高凡纳滨对虾的免疫能力和抗白斑综合征病毒能力,添加0.2%抗菌肽可显著提高对虾特定生长率、饲料效率和血清过氧化物酶(POD)、 溶菌酶(LZM)等酶活性[43]

  • 2.5 植物提取物

  • 植物提取物指通过物理、化学或生物学方法从植物中提取出来的天然生物活性成分,具有某类功效的天然产品。不同的植物提取物活性成分对水产动物具有不同的作用效果,发挥主要作用的植物成分主要包括多糖、生物碱、甙类、有机酸类、类固醇类、精油、植物色素及单宁等。

  • 2.5.1 缩合单宁

  • 天然植物提取物缩合单宁是由黄烷-3-醇单体组成的多酚类聚合物,广泛存在于自然界,具有显著的抗氧化、免疫刺激、抗菌、抗炎、抗寄生虫等生物活性。与化学合成的抗氧化剂相比,缩合单宁具有无污染、无残留、安全高效等诸多优点,在抵御氧化应激对机体危害、增强机体抗氧化能力、改善肠道健康等方面有巨大的研究价值和开发潜力,是理想的抗生素替代品[44]。缩合单宁可促进鱼类生长,提高免疫力和缓解氧化损伤和[45-46]。Peng等[47]研究表明,饲料中添加缩合单宁不影响花鲈幼鱼末体重、增重率、特定生长率、 成活率、饲料系数、摄食量、蛋白质效率、肥满度、 肝体比和脏体比,但可显著提高鲈鱼肠道菌群多样性,抑制Cetobacterium和Aeromonas菌属,促进Clostridium和Brevinema相对丰度增加。随着缩合单宁添加量升高,血清胆固醇和血清甘油三酯含量显著降低,血清总抗氧化能力、过氧化氢酶和超氧化物歧化酶活性显著升高;肝脏总抗氧化能力、超氧化物歧化酶和谷胱甘肽S转移酶活性显著升高; 肝脏SOD和Nrf2 mRNA的表达量显著上升。饲料中添加缩合单宁可显著降低花鲈对急性硫酸铜应激的累计死亡率,提高应激花鲈血清超氧化物歧化酶、 谷胱甘肽过氧化物酶和肝脏总抗氧化能力、过氧化氢酶活性。

  • 2.5.2 桑叶黄酮

  • 黄酮类化合物是一类化学结构相似的植物次生代谢产物,具有抗癌、抗氧化、调节脂代谢、抗菌抗病毒、抗炎等生理功能[48-49]。黄酮类化合物作为一种天然、绿色、环保抗氧化剂, 可提高鱼类生长性能和抗氧化能力。水飞蓟素(从水飞蓟果实中分离得到的一种黄酮类抗氧化剂)能够显著提高草鱼的增重率和特定生长率,降低饲料系数[50]。在饲料中添加槐属植物黄酮能显著提高罗非鱼的增重率、特定生长率、蛋白质效率和肥满度并降低饲料系数[51],在饲料中添加10~20 mg/kg大豆黄酮能降低大菱鲆幼鱼血清MDA水平,显著提高血清SOD和GSH-PX的活性,提高抗氧化能力[52]。广东省农业科学院动物科学研究所在国内首次开展有关桑叶黄酮提高水产动物生长、抗氧化及其机理的研究。研究表明,在饲料中添加适量桑叶黄酮可改善吉富罗非鱼的生长性能,显著提高其抗氧化能力及抗亚硝酸盐应激能力,改善肌肉品质, 提高抗氧化酶SOD和CAT基因表达水平,初步判断其抗氧化作用可能与Nfr2 信号通路有关[53]。在凡纳滨对虾饲料中添加桑叶黄酮可提高血清和肝胰腺总抗氧化能力,降低血清肝胰腺丙二醛和脂质过氧化物的含量,提高凡纳滨对虾抗氧化能力,添加50 mg/kg桑叶黄酮可促进凡纳滨对虾肠道发育,增加肠道菌群多样性[54]。以血清T-AOC为评价指标,桑叶黄酮在凡纳滨对虾饲料中的适宜添加量为56.18 mg/kg[55]

  • 3 展望

  • 在大力推进水产健康养殖转型升级、重视水产品安全与环保以及加速水产供给侧改革的大背景下,研发和推广应用水产饲料添加剂是水产饲料工业发展的重要内容,也是推动水产养殖业健康、 可持续发展的有效措施[56]。水产饲料添加剂的功能广泛,在动物营养和动物养殖中具有重要的地位和作用,但目前尚有许多关键技术问题需要突破,这也将是今后主要的研究方向:

  • (1)水产动物品种繁多,水产添加剂在不同水产动物中的适宜添加剂量、添加方式和添加时间仍需要继续深入探索;

  • (2)我国地域辽阔,应针对不同养殖环境、养殖模式、养殖季节、养殖阶段开展水产饲料添加剂在不同养殖条件下的应用效果和适宜添加量研究;

  • (3)细化水产饲料添加剂的产品定位, 根据生长性能、健康状态、形体参数、繁殖性能、 产品品质、抗病抗应激能力等不同评判标准,制定不同生产目的的产品使用标准;(4)针对不同养殖环境、不同生长阶段水产动物营养代谢差异, 研究水产动物对重要营养物质消化、吸收、转动和代谢的途径和特点,阐明营养代谢的关键调控元件与信号网络。

  • (责任编辑 崔建勋)

  • 曹俊明,博士,二级研究员,国务院特殊津贴专家,广东省丁颖科技奖获得者。现任广东海洋大学党委书记,华中农业大学兼职教授、博士生导师,华南农业大学、上海海洋大学硕士生导师,兼任中国畜牧兽医学会动物营养学分会理事、中国水产学会营养与饲料专业委员会委员、广东水产学会副理事长暨营养与饲料专业委员会主任委员。 主要从事水产动物抗氧化和免疫力的营养调控、功能性添加剂与新型饲料蛋白源的研究开发。主持或参加国家自然科学基金、广东省自然科学团队项目等各级各类项目60 多项,获得科技成果奖励8 项,其中作为第一完成人获农业农村部中华农业科技奖一等奖1 项、广东省科学技术奖二等奖1 项、广东省农业技术推广奖一等奖2 项。发表科技论文130 余篇,其中SCI收录20 篇,主编和参编专著各1 部;在GenBank登录基因3 个;申请专利10 项,获授权专利4 项;制定广东省地方标准1 项;参与开发新产品40 多个,产品和技术推广到100 多家水产饲料企业,新增产值50 多亿元。培养研究生45 人。

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