Chinese Journal of Animal Nutrition >

2024 , Vol. 36 >Issue 9: 5497 - 5509

DOI: https://doi.org/10.12418/CJAN2024.469

REVIEW

Research Progress on Effects of Malondialdehyde on Aquatic Feed and Aquatic Animals’ Health

  • PENG Cong , 1, 2 ,
  • TAN Beiping 1, 2 ,
  • LIN Xiangqin 1, 2 ,
  • DENG Junming , 1, 2, *
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  • 1 College of Fisheries, Guangdong Ocean University, Zhanjiang 524088, China
  • 2 Aquatic Animals Precision Nutrition and High-Efficiency Feed Engineering Research Centre of Guangdong Province, Zhanjiang 524088, China
* professor, E-mail:

Received date: 2024-03-25

  Online published: 2024-09-08

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Abstract

Malondialdehyde (MDA) is a significant carbonyl compound that is formed during non-enzymatic oxidation reactions of lipids and sugars, showing high levels of biotoxicity. The main risks associated with MDA in aquaculture include the disruption of essential nutrient structures in feed, such as proteins, lipids, and vitamins, as well as negative effects on animal health. These effects can lead to reduced production performance, compromised intestinal mucosal barrier, liver toxicity, decreased antioxidant capacity, and increased inflammation. This review specifically examined the production of MDA in feed, the mechanisms that influence the structure and function of key nutrients in feed, and the overall health of aquatic animals. The primary goal is to establish a theoretical basis for preventing feed oxidation and ensuring feed safety.

Key words： malonaldehyde; antioxidant function; feed safety; mucosal barrier function; lipid oxidation

Cite this article

PENG Cong , TAN Beiping , LIN Xiangqin , DENG Junming . Research Progress on Effects of Malondialdehyde on Aquatic Feed and Aquatic Animals’ Health[J]. Chinese Journal of Animal Nutrition, 2024 , 36(9) : 5497 -5509 . DOI: 10.12418/CJAN2024.469

近年来,人们对水产品品质的要求越来越高,水产饲料质量在一定程度上决定了水产品品质,是水产养殖业健康可持续发展的关键物质保障。当前,饲料资源日益紧缺,提高饲料资源利用率和保障饲料安全成为水产饲料行业所面临的主要问题[1-2],而有毒有害物质含量是安全优质饲料的评判指标之一[3]。丙二醛(malondialdehyde,MDA)作为脂类和糖类在非酶氧化反应中形成的主要活泼羰基化合物,其破坏蛋白质结构、损伤DNA功能等生物学毒性,以及在饲料生产和水产养殖中的潜在危害逐渐被发现[4-5]。在破坏饲料蛋白质结构方面,极低浓度下MDA就能修饰蛋白质巯基基团、氨基酸残基如蛋氨酸(Met)、赖氨酸(Lys)、半胱氨酸(Cys)、组氨酸(His)和酪氨酸(Tyr)残基等,导致蛋白质变性、失活和交联,并破坏其结构与功能[6-7]。在危害动物机体健康方面,MDA对机体胃肠道损伤、抗氧化能力具有剂量效应[8],故以饲料中MDA含量为标准创建和评估未来饲料安全风险具有可行性。在损伤DNA功能方面,MDA与鸟嘌呤碱基发生反应形成MDA-鸟嘌呤加合物,导致染色体畸变,阻滞转录过程,进而损害DNA功能[9]。目前,系统阐述MDA对水产动物健康影响的报道较少。本文重点阐述MDA来源,破坏饲料营养素结构、危害动物机体健康机制,以期为MDA毒害机理以及绿色可持续水产养殖提供理论参考。

1 MDA概述

1.1 MDA理化性质

MDA是一种具有挥发性且呈无色针状晶体的小分子化合物,化学式为C3H4O2,常带有2个结晶水,另外,MDA具有强亲电子的1,3双不饱和醛结构,对蛋白质、DNA等亲核物质表现出极强的毒性作用[10-11];其纯品在中性条件下以烯醇阴离子形式存在,化学反应性较低,酸性条件下化学反应性较高[12]。此外,MDA含有分子内氢键,质子可以通过该氢键在供体和受体氧原子之间转移(O1-H-O2)[13-14],如图1所示。
图1 MDA烯醇化及质子转移

Fig.1 MDA enolization and proton transfer[14]

1.2 饲料中MDA产生过程

MDA含量是饲料长时间储存过程中过氧化的重要检测指标之一,而MDA产生的主要途径是脂质自氧化[15-16]。脂质氧化从脂质分解成游离脂肪酸(free fatty acids,FFA)和低级脂肪酸(C≤10)开始[17]。一方面,光、热条件能激发光敏剂,产生单线态氧气,进攻多不饱和脂肪酸(polyunsaturated fatty acids,PUFA)双键高密度电荷区域,形成不稳定的六元环;环裂解并将双键向氧转移形成氢过氧化物,最后分解产生MDA等活泼羰基化合物[18⇓-20];另一方面,自由基、过氧化氢等活性物质从PUFA酰基链中获取氢原子并产生脂质自由基(R·),而脂质自由基极易与氧再次反应形成脂质过氧自由基(ROO·),脂质过氧基则通过非特异性氢提作用向附近的PUFA中提取氢原子,接受氢原子后变为极易水解热裂为脂质烷氧自由基(RO·)的脂质氢过氧化物(ROOH),进而氧化生成MDA等活泼羰基化合物[21⇓-23];提供氢原子的不饱和脂肪酸变回脂质自由基进一步发生氧化连锁反应,从而导致脂质的加速腐败,称为自氧化(如图2所示)[24]。饲料脂质氧化过程除了自氧化外,也受到饲料微生物的酶解反应影响。PUFA被分解成游离脂肪酸并进一步反应生成醛、酮、醇等物质,其一般与自氧化同时发生[25]。此外,饲料中脂质过氧化形成MDA的过程与饲料种类(水分、脂质含量)、微生物催化作用和储存时间与条件(光照强度、金属离子催化)等多方面因素密切相关[26]。
图2 脂质自氧化过程产生MDA

RH:不饱和脂肪酸 unsaturated fatty acids;R·:脂质自由基 lipid alkyl radicals;ROO·:脂质过氧自由基 lipid peroxyl radicals;RO·:脂质烷氧自由基 lipid alkoxyl radicals;ROOH:脂质氢过氧化物 lipid hydroperoxide;·OH:羟基自由基 hydroxyl radical。

Fig.2 MDA production by lipid autoxidation process

2 MDA对饲料的影响

蛋白质、脂质、碳水化合物、维生素、矿物质是饲料中的主要营养素。研究发现,脂质过氧化产生的MDA对饲料影响主要体现在改变蛋白质、脂质和维生素结构与功能方面。

2.1 MDA对蛋白质的影响

蛋白质羰基含量反映着饲料蛋白质的氧化程度,是最常见的氧化标记物[27]。Li等[28]用10 mmol/L MDA与10 mg/mL大米蛋白质溶液混合反应后发现,大米蛋白质羰基含量为15.07 nmol/mg,显著高于空白对照组,说明MDA对饲料中蛋白质成分具有氧化作用。宋娟娟等[29]用10 mmol/L MDA处理10 mg/mL豆粕悬浮液后发现,大豆蛋白质也被氧化,试验组豆粕中羰基含量为20 nmol/mg蛋白质,显著高于空白对照组;此外,试验组豆粕中蛋氨酸(Met)、酪氨酸(Tyr)和赖氨酸(Lys)含量分别显著降低至0.55%、1.85%、3.28%,蛋白质体外消化率下降为45%,巯基及总巯基基团含量分别为6 nmol/mg蛋白质、93 nmol/mg蛋白质,显著低于空白对照组,而席夫碱荧光强度逐渐升高至900 nm,说明MDA能将蛋白质中的半胱氨酸(Cys)、组氨酸(His)和Lys残基氧化修饰成席夫碱,从而促进蛋白质羰基化,降低蛋白质消化率。此外,MDA对饲料中蛋白质的结构和功能性质也具有破坏作用。Li等[6]用10 mmol/L MDA氧化10 mg/mL米糠蛋白质溶液后发现,氧化米糠蛋白质羰基含量为10.34 nmol/mg,试验组蛋白质羰基含量以及分子平均粒径均显著高于对照组,蛋白质二级结构中无规卷曲、β-转角、α-螺旋结构含量以及表面疏水性分别为23.82%、8.83%、19.59%、163,均显著下降,说明MDA能破坏蛋白质二级结构,诱导蛋白质共价交联聚集,进而影响其功能。Sun等[30]用10 mmol/L MDA溶液氧化分离核桃蛋白质后发现,核桃蛋白质起泡能力、泡沫稳定性、乳化性和乳化稳定性分别降低至6.37%、5.62%、25.86 m2/g和23.10 min,溶解性降低86%,说明蛋白质理化性质受到破坏。综上所述,一方面,MDA能够修饰蛋白质巯基基团、氨基酸残基(如Cys、His和Lys残基等),造成蛋白质羰基化,并产生席夫碱,影响饲料中蛋白质消化吸收;另一方面,MDA能破坏蛋白质无规则卷曲、β-转角等二级结构,诱导蛋白质发生变性、失活和共价交联聚集,降低其溶解性、起泡能力、泡沫稳定性、乳化性和乳化稳定性等功能活性。

2.2 MDA对脂质的影响

脂质是饲料中主要的能源物质,为动物生长提供必需脂肪酸。不饱和脂肪酸含有双键和烯丙基氢原子,对氧化剂极其敏感,随着不饱和度增加,如花生四烯酸(C20∶4)和二十二碳六烯酸(C22∶6),氧化敏感性也增强[31]。Barrera等[32]认为脂质过氧化的醛类产物(如MDA)能与生物大分子形成共价亲电加成产物来修饰生物大分子(如脂质)。由于MDA极不稳定,易与脂质过氧化产物乙醛以及蛋白质发生反应,产生MDA-乙醛-蛋白质加合物,并伴随着自由基[如超氧阴离子自由基( O 2 -·)]的生成[33]。自由基的产生又能参与前文脂质氧化过程,加速脂质氧化。Linda等[34]也表示脂质过氧化的醛类产物(如MDA)被认为是引发脂质过氧化一级自由基反应的“二级毒性信使”。磷脂是一类含有磷酸基团的极性脂质统称,分为甘油磷脂类与鞘磷脂类。马路凯[35]认为MDA结构中的活性醛基可以与磷脂发生反应生成席夫碱,改变功能性大分子的原有结构。这可能是因为MDA羰基侧链所连基团空间位阻比较小,再加上磷脂中含有亲核性氨基基团,因此二者交联活性较强并产生席夫碱类化合物[36]。综上所述,一方面,MDA与蛋白质和其他脂质过氧化产物发生交联,并伴随自由基产生,间接加速脂质氧化;另一方面,MDA由于其羰基侧链所连基团空间位阻小,与具有亲核氨基基团的磷脂反应,直接改变磷脂结构。

2.3 MDA对维生素的影响

维生素是饲料中一种维持动物正常生理功能不可或缺的微量有机物,其中具有抗氧化功能的维生素主要为维生素C、维生素A、维生素E和维生素P[37]。维生素C具有强还原性,其烯醇式羟基键强度弱易断裂,能与自由基反应生成半脱氢抗坏血酸;维生素A中含有双烯共轭健,能够转移单线态氧中能量,自身变成激发态;二者可作为脂质过氧化自由基高效的淬灭剂[38]。MDA与乙醛及蛋白质反应产生的自由基(如 O 2 -·)可间接与其发生反应,改变维生素C、维生素A结构。维生素E是生育酚、生育三烯酚及具有D-α-生育酚生物活性的衍生物总称,其苯基间位含有羟基活性官能团并具有显著抗氧化作用[39]。维生素P又称芦丁,属生物类黄酮,也具有间位羟基活性官能团[40]。Rosario等[41]研究发现,MDA能与具有间位羟基酚类物质如2,5-二甲基间苯二酚(2,5-dimethylbenzene-1,3-diol)、1,3-二羟基-5-甲基苯(5-methylbenzene-1,3-diol)、橄榄醇(5-pentylbenzene-1,3-diol)和5-烷基间苯二酚(5-alkylbenzene-1,3-diol)发生反应,生成5(或7)-烷基-7(或5)-羟基-4-甲基-4H-色烯-3-羧醛和4-(3甲酰苯)-7-羟基-4H-色烯-3-羧醛等加合物;此外,Rosario等[42]还进一步验证了MDA与具有间位羟基酚类物质反应的可能性。综上所述,MDA能产生自由基间接改变维生素C、维生素A的结构,也能与间位羟基酚类维生素E、维生素P发生反应,产生多种加合产物,但有关MDA对饲料中维生素的影响及反应产物的消化吸收效率和生理功能还需要进一步研究。

3 MDA对水产动物健康的影响

MDA不但能改变饲料主要营养物质的结构功能,而且对水产动物健康也会有一定影响,如降低生产性能、损伤肠道黏膜屏障、诱导肝脏毒性、降低机体抗氧化能力和诱发炎症反应等(图3)。
图3 饲料MDA对水产动物健康的影响

Fig.3 Effects of dietary MDA on aquatic animals’ health

3.1 降低生产性能

以水产动物生长增重评价生产性能具有较好的经济合理性,而水产动物健康快速生长由多种质量维度、空间维度、器官维度等指标客观全面地评估,如特定生长率(specific growth rate,SGR)、增重率(weight gain rate,WGR)、饲料系数(feed conversion ratio,FCR)、肝体比(hepasomatic index,HSI)、脏体比(viserosomatic index,VSI)等[43-44]。MDA对动物机体具有毒副作用,且显著降低水产动物的生产性能。黄雨薇等[45]研究发现,当饲料中MDA含量超过61.04 mg/kg(以饲料所含粗脂肪为基础计)时,草鱼(Ctenopharyngodon idellus)SGR显著低于对照组且呈负相关关系,FCR显著高于对照组且呈正相关关系。陈科全等[46]研究发现,采用MDA含量为123.92 mg/kg(以饲料所含粗脂肪为基础计)饲料饲喂初重为74 g的草鱼72 d后发现,SGR、肥满度和蛋白质沉积率分别为1.51%/d、1.73、31.1%,显著低于对照组;FCR为1.89,显著高于对照组;说明MDA能阻碍草鱼机体对蛋白质的利用,降低生长速度。机体消化吸收能力是水产动物生长的关键因素,而消化酶活性是反映机体消化吸收能力的主要指标[47]。Fan等[8]研究发现,珍珠龙胆石斑鱼(Epinephelus fuscoguttatus♀×E. lanceolatu ♂)消化酶活性与饲料MDA含量呈负线性关系,随着饲料MDA水平的升高,肠道胰蛋白酶、脂肪酶和淀粉酶活性显著降低,鱼体SGR和WGR显著下降,说明MDA能降低消化酶活性,进而阻碍水产动物生长,从而降低生产性能。综上所述,饲料MDA通过降低水产动物体内消化酶活性,阻碍机体对蛋白质利用率,减缓个体生长速度,进而降低水产动物生产性能;另外,大多数水产研究者只围绕质量维度阐述MDA对水产动物生长评价,而多维度多指标评价MDA对水产动物生长和健康的影响尚有待进一步研究。

3.2 损伤肠道黏膜屏障

鱼类肠道不仅具有消化吸收营养物质的功能,而且能参与机体免疫应答、内分泌调节以及新陈代谢过程,是鱼类抵御微生物入侵最复杂的黏膜免疫器官[48-49]。肠道黏膜屏障能够将肠腔环境与内环境分开,防止致病性微生物、毒素、抗原进入机体,是动物体消化系统的第1道屏障,在机体免疫方面发挥着巨大的作用[50-51]。肠道黏膜屏障功能障碍是肠道炎症的先决条件之一[52],而饲料脂质氧化产物MDA对肠道具有显著的生物毒副作用。饲养试验发现,使用含185.04 mg/kg(以饲料所含粗脂肪为基础计)MDA的饲料饲喂草鱼72 d后,与对照组相比,肠道出现假复层柱状上皮结构,且绒毛间隙、杯状细胞显著增加,绒毛密度和高度显著下降,中央乳糜管增大,说明饲料中MDA显著破坏了肠道黏膜结构[46]。体外细胞试验发现,向培养36 h离体草鱼肠道黏膜细胞中加入9.89 μmol/L MDA,6 h后细胞生长抑制率高达52.32%,黏膜细胞活性、总蛋白含量显著降低,细胞集落面积减少,空泡变性严重,线粒体、细胞核出现明显损伤,培养液中MDA含量显著提高[53]。紧密连接蛋白是在肠道机械屏障中构成细胞紧密连接结构的关键蛋白[54]。研究发现,MDA能够下调紧密连接蛋白相关基因表达,损害肠道黏膜紧密连接结构。黄雨薇[55]研究发现,采用含185 mg/kg(以饲料所含粗脂肪为基础计)MDA的氧化饲料饲喂草鱼72 d后,与对照组相比,试验组闭合蛋白(Claudin,如Claudin-3、Claudin-15a)、闭锁蛋白(Occludin)和闭锁小带蛋白(ZO,如ZO-1、ZO-2、ZO-3)基因表达水平显著下调,紧密连接结构出现巨大缝隙,血清二胺氧化酶活性显著高于对照组。综上所述,MDA进入机体一方面会破坏肠道黏膜细胞内部结构,抑制其生长和存活,进而损害肠道黏膜完整性,最终影响机体对营养物质消化吸收,损害机体健康;另一方面会下调Claudin-3、Claudin-15a、Occludin、ZO-1、ZO-2和ZO-3基因表达水平,损伤肠道紧密连接结构,增加肠道黏膜通透性,导致有毒有害物质进入动物机体。

3.3 诱导肝脏毒性

肝脏是机体代谢、解毒以及胆固醇和胆汁酸合成的重要器官。MDA不仅使肝脏细胞膜纤维化,细胞结构受损,而且使线粒体内部结构混乱,嵴数量减少[46,56],其原因可能是MDA对丙酮酸脱氢酶、α-酮戊二酸脱氢酶和苹果酸脱氢酶等线粒体呼吸链关键酶造成不同程度损伤,抑制呼吸链反应,阻碍线粒体功能,产生活性氧,抑制细胞能量代谢,最终损伤肝脏细胞[57⇓-59]。此外,MDA通过降低胆汁酸合成功能,大量胆固醇在肝脏积累,增加肝脏负担,进而产生脂肪肝等病症。肝脏合成胆汁酸过程是以胆固醇为原料,在肝脏中合成初级胆汁酸如胆酸和鹅脱氧胆酸,并排出肝脏[60-61]。胆固醇7α羟化酶(cholesterol 7α-hydroxylase,CYP7A1)是肝脏合成胆汁酸的关键酶[62],3-羟基-3-甲基戊二酰辅酶A还原酶(3-hydroxy-3-methylglutaryl coenzyme A reductase,HMGCR)是以乙酰辅酶A为原料合成胆固醇的关键酶[63]。黄雨薇等[64]研究发现,用富含MDA的饲料饲喂草鱼72 d后发现,肝体比显著上升,血清中高密度脂蛋白胆固醇与低密度脂蛋白胆固醇比值显著降低,肝脏中CYP7A1基因表达显著降低,HMGCR基因表达显著提高,血清和肝脏中总胆固醇含量均显著增加,血清和肠道总胆汁酸含量显著降低,而肝脏总胆汁酸含量也出现不同程度降低,说明MDA使肝脏合成胆汁酸能力降低,大量胆固醇在肝脏中积累,肝脏脂质代谢功能受到干扰。综上所述,MDA通过降低线粒体呼吸链关键酶活性,诱导自由基产生并损伤线粒体内部结构,进而损害肝脏细胞;此外,MDA还通过降低CYP7A1基因和HMGCR基因表达,破坏胆汁酸合成过程,迫使肝脏胆固醇沉积,增加肝脏负荷并诱导肝脏毒害。

3.4 降低机体抗氧化能力,诱发炎症反应

机体中的谷胱甘肽过氧化物酶(glutathione peroxidase,GSH-Px)、超氧化物歧化酶(superoxide dismutase,SOD)和过氧化氢酶(catalase,CAT)等抗氧化酶能清除过量自由基,缓解机体氧化应激[65]。谷氨酸半胱氨酸连接酶催化亚基(glutamate-cysteine ligase catalytic subunit,GCLC)是谷胱甘肽(glutathione,GSH)合成过程中限速酶γ-谷氨酰半胱氨酸合成酶的活性亚基,其能促进GSH/谷胱甘肽S-转移酶(glutathione S-transferases,GSTs)通路的关键活性物质GSH、谷胱甘肽还原酶(glutathione reductase,GR)、GSH-Px以及GSTs等发挥正常的生理功能[66]。研究发现,低水平MDA进入机体后可刺激肠道GCLC基因表达,而高水平MDA抑制肠道GCLC基因表达[67]。林秀秀等[67]用含0、61、124和185 mg/kg(以饲料所含粗脂肪为基础计)MDA的试验饲料饲喂草鱼72 d后发现,与对照组相比,试验组肠道GCLC基因相对表达量分别为1.00、1.39、1.88和1.67,且随着饲料MDA含量升高呈先升高后降低趋势,二者显著正相关。胞液酶GSTpi是具有CAT和脱氯化氢酶生理功能的GSTs同工酶,其常以单体形式与c-Jun氨基末端激酶(c-Jun N-terminal kinases,JNK)形成复合物,抑制JNK活性[68-69]。MDA进入机体后能刺激GSTpi自身形成二聚体,致使复合物解离,释放出JNK,损害机体健康[67]。但由于机体自我修复机制,JNK磷酸化能激活转录因子c-Jun,一方面能促进GSTpi基因转录,从而减少JNK水平;另一方面又能反馈降低JNK活性,缓解机体损伤[70]。用含185 mg/kg(以饲料所含粗脂肪为基础计)MDA的试验饲料饲喂草鱼72 d后发现,与对照组相比,试验组GSTpi基因相对表达量显著上调为1.62,且与饲料MDA含量具有显著正相关[67]。另外,肠道炎症反应与肿瘤坏死因子(tumor necrosis factor,TNF)-α、白细胞介素(interleukin,IL)-1β、IL-6等细胞炎性因子密切相关,而促炎因子受到核因子E2相关因子2(NF-E2-related factor 2,Nrf2)/抗氧化反应元件(antioxidant response element,ARE)、JNK[71]和核转录因子-κB(nuclear factor-kappa B,NF-κB)[72]等信号通路调控。研究发现,Nrf2/ARE信号通路能促进GSH-Px等抗氧化酶的表达[73],而GSH-Px1、GSH-Px2能调控NF-кB信号通路,抑制肠道炎症反应中具有重要作用的NF-κB的合成与释放,下调下游炎性基因表达,缓解肠道炎症反应[74]。此外,研究还显示饲料中添加MDA可提高石斑鱼肠道JNK激酶活性,激活JNK信号通路[75];MDA能够抑制Nrf2/ARE信号通路[76]。以上研究结果说明MDA可能刺激肠道发生炎症反应,但其具体机制有待进一步研究。综上所述,MDA进入机体后一方面上调肠道GCLC基因表达,激活GSH/GSTs信号通路,增加GSH、GR、GSH-Px以及GSTs的生成,暂时提高肠道抗氧化能力,但当MDA含量过高时,会对肠道造成实质性损伤,进而下调GCLC基因表达;另一方面MDA能刺激GSTpi自身形成二聚体,致使复合物解离,释放出JNK,损害机体健康,当MDA损伤作用弱于机体免疫应答时,促进GSTpi基因转录并结合JNK,降低JNK活性,缓解机体损伤。

4 MDA的去除与控制

依据国家标准,石斑鱼、巴沙鱼配合饲料中MDA含量限量分别为10.0[77]、5.0 mg/kg[78](以饲料所含粗脂肪为基础计)。因此,缓解或消除MDA毒副作用是水产养殖业绿色发展亟需解决的问题之一。目前,饲料生产中常用茶多酚[79]、二丁基羟基甲苯[80]、乙氧基喹啉[81]等作为清除过量自由基、预防饲料变质的主要抗氧化剂。此外,玉米黄素[82]和岩藻黄素[83]等天然类胡萝卜素也可通过清除自由基、淬灭单线态氧作用间接降低肉制品存储加工过程中MDA含量和氧化速率。然而,常见抗氧化剂只能延缓脂质、蛋白质氧化速率而无法直接清除MDA,因此需要发掘MDA去除剂。有关MDA清除与控制的相关研究见表1。芦慧勤等[84]将10 mmol/L MDA与10 mmol/L L-Lys混合后注入10倍体积的牦牛肉5 d后发现,L-Lys组牦牛肉过氧化物值和硫代巴比妥酸值均显著低于未添加L-Lys组,而蛋白质羰基含量显著高于未添加L-Lys组。外源添加L-Lys虽然能依靠自身氨基酸残基与MDA反应,从而延缓肌肉MDA含量的增加,但此反应不具备特异性并导致L-Lys被氧化。Rosario等[41]认为含有间位羟基的多酚类物质对MDA具有捕获能力,并在氧化亚麻籽油与小麦粉制备饼干过程加入橄榄醇后发现:橄榄醇组MDA含量为500~750 ng/g,显著低于未加橄榄醇组;加合产物7-羟基-5-戊基-4H-色烯-3-羧醛和7-羟基-4-甲基-5-戊基-4H-色烯-3-羧醛含量分别为1 200~1 500 ng/g、36~48 ng/g,均显著高于未加橄榄醇组[41-42]。由此可见,橄榄醇在制备食物或饲料过程中具有降低MDA含量,减缓MDA形成速率的潜力。但值得注意的是,饲料中橄榄醇使用量以及产生加合产物对水产动物生长和健康是否具有负面影响有待研究。此外,蒋丙婷等[85]研究发现,没食子酸、槲皮素、儿茶素、绿原酸等多酚类物质(含有间位羟基酚)能缓解MDA对花生球蛋白质的氧化作用。综上所述,橄榄醇、没食子酸、槲皮素等含有间位羟基的多酚类物质具有在饲料中应用降低MDA含量的潜力,但仍面临一些挑战。
表1 MDA清除与控制

Table 1 MDA removal and control

作用机制
Action
mechanism		 添加剂
Additive		 添加量
Additive amount		 添加方式
Additive modality		 简要结果
Summary results		 参考文献
References



间位羟基酚捕获MDA
Meso-hydroxyphenol
scavenging of MDA		 橄榄醇 0.16 g/
12.5 g饼干		 氧化亚麻籽油与小
麦粉制备饼干成
型时加入橄榄醇		 减少MDA含量,产生7-羟基-
5-戊基-4H-色烯-3-羧醛和
7-羟基-4-甲基-5-戊基-4H-
色烯-3-羧醛等加合产物		 [42]
槲皮素 10 mg/mL,
等体积		 与MDA溶液和蛋
白质溶液混合		 缓解MDA氧化花生球
蛋白质,其游离氨基
浓度增加83.31%		 [85]






间位羟基酚捕获MDA
Meso-hydroxyphenol
scavenging of MDA		 儿茶素 10 mg/mL,
等体积		 与MDA溶液和蛋
白质溶液混合		 缓解MDA氧化花生球蛋白质,
其羰基浓度降低98.10%,
溶解度增加7.30%		 [85]
绿原酸 10 mg/mL,
等体积		 与MDA溶液和
蛋白质溶液混合		 缓解MDA氧化花生球蛋白质,
其表面疏水性与羰基
浓度显著降低		 [85]
没食子酸 10 mg/mL,
等体积		 与MDA溶液和蛋
白质溶液混合		 缓解MDA氧化花生球蛋白质,
其表面疏水性降低35.57%,
内源荧光强度显著增大		 [85]
氨基酸残基消耗MDA
Amino acid residue
consumption of MDA		 L-
赖氨酸		 10 mmol/L,
肉重10%		 与10 mmol/L MDA
溶液混合注入牦牛肉		 降低牦牛肉过氧化物值和
MDA含量,抑制肌肉脂质和
蛋白质氧化		 [84]



清除自由基、
淬灭单线态氧
Scavenging radicals
and quenching
singlet oxygen		 玉米黄素 配方体系1% 玉米黄素与固化剂、
水混合后加入腌肉配方		 降低腌肉MDA含量与
肌肉氧化速率		 [82]
岩藻黄素 配方体系
0.04%		 以配方成分
形式添加		 清除羟基等自由基;降低火腿
肉香肠MDA含量与氧化速率		 [83]

5 小结与展望

MDA作为脂质主要过氧化产物,其毒副作用在饲料生产中主要体现为改变营养素结构与功能,分为3个方面:1)MDA能够修饰蛋白质巯基基团、氨基酸残基,造成蛋白质羰基化,破坏蛋白质结构,诱导蛋白质变性、失活和共价交联聚集,降低其化学特性,进而影响饲料中蛋白质的消化吸收;2)MDA通过直接与具有亲核氨基基团的磷脂反应,改变磷脂结构,或与蛋白质和脂质过氧化产物乙醛发生交联,产生自由基,间接加速脂质氧化;3)MDA直接与间位羟基酚类维生素E、维生素P发生反应,产生多种加合产物,改变维生素结构与功能,也能产自由基间接改变维生素A、维生素C结构。MDA在水产养殖中的毒副作用,主要体现为:1)降低生产性能,MDA通过降低水产动物体内消化酶活性,阻碍机体对饲料蛋白质的利用,减缓鱼体生长速度,降低水产动物生产性能;2)损伤肠道黏膜屏障,MDA通过破坏肠道黏膜细胞内部结构并抑制其存活,损伤肠道紧密连接结构,破坏肠道黏膜屏障功能,增加肠道黏膜通透性;3)诱导肝脏毒性,MDA通过降低线粒体呼吸链关键酶活性,损伤线粒体内部结构,破坏胆汁酸合成过程,迫使胆固醇在肝脏中积累,增加肝脏负担并产生肝脏毒性;4)降低抗氧化能力,诱发炎症反应。MDA通过引发肠道细胞羰基应激,以及刺激GSTpi自身形成二聚体,致使复合物解离,诱导JNK过渡磷酸化,激活JNK信号通路;MDA还能通过抑制Nrf2/ARE信号通路,下调GCLC基因和GPX2基因表达,降低GSTs的生成,抑制GSH/GSTs信号通路。因此,MDA具有明确的氧化损伤毒副作用,以过氧化值、酸价等评价指标为基础再结合MDA含量检测能更全面准确评估饲料脂质氧化程度及毒害物质含量,可将其作为评价水产动物饲料安全的新型指标;随着对MDA研究的不断深入,水产行业逐渐朝向绿色可持续方向发展,但仍有部分问题值得重视:1)不同鱼类品种及其不同生长阶段对MDA的耐受性研究覆盖面尚不完全,在水产领域确定饲料中MDA限量值有待进一步完善;2)MDA损害肠道黏膜屏障仅存在于物理屏障方面,关于化学、免疫和生物屏障方面有待深入研究;3)目前尚未见MDA对肌肉品质影响的相关报道,肌肉是连接消费者与养殖户的桥梁,因此也有待进一步研究;4)寻找清除饲料中MDA,缓解MDA毒副作用的方法还需要进一步研究。

References

Publishing order | Descend order by publishing year | Descend order by cited within
[1]
ONDIBA R N, OGELLO E O, KEMBENYA E, et al. Future demand and supply of aquafeed ingredients:outlines to commercialize non-conventional protein ingredients to enhance aquaculture production for food security in sub-Saharan Africa[J]. Aquatic Ecosystem Health & Management, 2022, 25(4):75-84.

[2]
武宁宁, 王大为, 崔正国, 等. 我国水产品追溯体系研究进展[J]. 广东海洋大学学报, 2023, 43(5):113-118.

WU N N, WANG D W, CUI Z G, et al. Research progress of Chinese aquatic product traceability system[J]. Journal of Guangdong Ocean University, 2023, 43(5):113-118. (in Chinese)

[3]
KULMAKOVA N, MAGOMADOV T, KOSTOMAKHIN N, et al. Safety and quality of feed for piglets[J]. IOP Conference Series:Earth and Environmental Science, 2021, 935(1):012012.

[4]
ITZIOU A. The role of lysosomal membrane stability,malondialdehyde levels and DNA damage as pollution biomarkers in evaluating biological cleaning products using Mytilus galloprovincialis[J]. Water Practice and Technology, 2022, 17(9):1914-1922.

[5]
ZHU X S, MA Z H, ZHANG X Y, et al. Effect of malondialdehyde-induced oxidation modification on physicochemical changes and gel characteristics of duck myofibrillar proteins[J]. Gels, 2022, 8(10):633.

[6]
LI F, WU X J, WU W. Effects of oxidative modification by malondialdehyde on the in vitro digestion properties of rice bran protein[J]. Journal of Cereal Science, 2021, 97:103158.

[7]
尤翔宇. 过氧自由基和丙二醛氧化对米糠蛋白结构、功能性质和消化性质的影响[D]. 硕士学位论文. 长沙: 中南林业科技大学, 2019.

YOU X Y. Effects of oxidative modification on structure functional properties and digestibility of rice bran protein by peroxyl radicals and malondialdehyde[D]. Master’s Thesis. Changsha: Central South University of Forestry and Technology, 2019. (in Chinese)

[8]
FAN J T, ZHANG Y M, ZHOU H, et al. Dietary malondialdehyde damage to the growth performance and digestive function of hybrid grouper (Epinephelus fuscoguttatus ♀×E. lanceolatu ♂)[J]. Animals, 2023, 13(19):3145.

[9]
YATES S A, DEMPSTER N M, MURPHY M F, et al. Quantitative analysis of malondialdehyde-guanine adducts in genomic DNA samples by liquid chromatography/tandem mass spectrometry[J]. Rapid Communications in Mass Spectrometry, 2017, 31(9):762-770.

DOI PMID

[10]
付鑫浪吉. 饲料组胺和丙二醛对巴沙鱼生长性能、胃肠道健康和肌肉品质的影响[D]. 硕士学位论文. 广东: 广东海洋大学, 2023.

FU X L J. Effects of dietary histamine and malondialdehyde on growth performance,gastrointestinal health and muscle quality of striped catfish (Pangasianodon hypophthalmus)[D]. Master’s Thesis. Guangdong: Guangdong Ocean University, 2023. (in Chinese).

[11]
ZHANG J, YANG Z H, ZHANG S S, et al. Investigation of endogenous malondialdehyde through fluorescent probe MDA-6 during oxidative stress[J]. Analytica Chimica Acta, 2020, 1116:9-15.

DOI PMID

[12]
BAHAVARNIA F, BAGHBAN H N, ESKANDANI M, et al. Microfluidic paper-based colorimetric quantification of malondialdehyde using silver nanoprism toward on-site biomedical analysis:a new platform for the chemical sensing and biosensing of oxidative stress[J]. RSC Advances, 2023, 13(43):30499-30510.

[13]
WU Q Z, YANG S B, LI Q Z. Triel bond formed by malondialdehyde and its influence on the intramolecular H-bond and proton transfer[J]. Molecules, 2022, 27(18):6091.

[14]
TRUJILLO C, SÁNCHEZ-SANZ G, ALKORTA I, et al. Resonance assisted hydrogen bonds in open-chain and cyclic structures of malonaldehyde enol:a theoretical study[J]. Journal of Molecular Structure, 2013, 1048:138-151.

[15]
VANDEMOORTELE A, HEYNDERICKX P M, LELOUP L, et al. Kinetic modeling of malondialdehyde reactivity in oil to simulate actual malondialdehyde formation upon lipid oxidation[J]. Food Research International, 2021, 140:110063.

[16]
MA L, LIU G, LIU X. Malondialdehyde,4-hydroxy-2-hexenal,and 4-hydroxy-2-nonenal in vegetable oils:formation kinetics and application as oxidation indicators[J]. European Journal of Lipid Science and Technology, 2019, 121(7):1900040.

[17]
ZHANG W Y, ZHAO X, WANG C, et al. Investigating roles of oil pre-oxidation on emulsion stability:dispersed-phase and interfacial-film both matter[J]. Food Hydrocolloids, 2024, 151:109873.

[18]
谭冬飞, 王少雷, 张清阳, 等. 光诱导乳脂肪氧化的研究进展[J]. 乳业科学与技术, 2021, 44(1):39-43.

DOI

TAN D F, WANG S L, ZHANG Q Y, et al. A review of recent studies on the light-induced oxidation of milk[J]. Journal of Dairy Science and Technology, 2021, 44(1):39-43. (in Chinese)

DOI

[19]
HOU Y X, FU Q, KUANG Y F, et al. Unsaturated fatty acid-tuned assembly of photosensitizers for enhanced photodynamic therapy via lipid peroxidation[J]. Journal of Controlled Release, 2021, 334:213-223.

DOI PMID

[20]
LEE M M S, LIN D M, CHAU J H C, et al. Adipocyte-targeting type Ⅰ AIE photosensitizer for obesity treatment via photodynamic lipid peroxidation[J]. ACS Nano, 2023, 17(11):11039-11053.

[21]
MACHADO M, RODRIGUEZ A L M, GOMES A M, et al. Vegetable oils oxidation:mechanisms,consequences and protective strategies[J]. Food Reviews International, 2023, 39(7):4180-4197.

[22]
CHEMAT A, SONG M, LI Y, et al. Shade of innovative food processing techniques:potential inducing factors of lipid oxidation[J]. Molecules, 2023, 28(24):8138.

[23]
HUANG Z H, GUAN W L, LYU X, et al. Impacts of acute ammonia-N exposure on the muscle quality of whiteleg shrimp (Penaeus vannamei): novel insights into lipid and protein oxidation[J]. Food Chemistry, 2024, 437(Pt 1):137781.

[24]
PHAN S. A selective review of lipid autoxidation and rancidity,antioxidants,and dithiooxamide[J]. Asian Journal of Medicine and Health, 2022, 20(4):55-79.

[25]
CUCINIELLO R, DI MEO F, FILOSA S, et al. The antioxidant effect of dietary bioactives arises from the interplay between the physiology of the host and the gut microbiota:involvement of short-chain fatty acids[J]. Antioxidants, 2023, 12(5):1073.

[26]
田兴舟, 卢琦. 花青素在乳脂氧化中的作用及其机制[J]. 动物营养学报, 2020, 32(8):3476-3482.

DOI

TIAN X Z, LU Q. Role of anthocyanin in oxidation of milk fat and its mechanism[J]. Chinese Journal of Animal Nutrition, 2020, 32(8):3476-3482. (in Chinese)

[27]
刘岩, 李敏, 金枝, 等. 生物保鲜剂真空浸渍对冰温贮藏罗非鱼蛋白特性的影响[J]. 广东海洋大学学报, 2022, 42(2):88-94.

LIU Y, LI M, JIN Z, et al. Effect of vacuum impregnation assisted biological preservatives on the protein properties of tilapia fillets during ice-temperature storage[J]. Journal of Guangdong Ocean University, 2022, 42(2):88-94. (in Chinese)

[28]
LI F, WU X J, WU W. Effects of malondialdehyde-induced protein oxidation on the structural characteristics of rice protein[J]. International Journal of Food Science & Technology, 2020, 55(2):760-768.

[29]
宋娟娟, 谢婷, 刘文涵, 等. 丙二醛对豆粕蛋白质氧化的影响及茶多酚的缓解作用[J]. 南京农业大学学报, 2023, 46(2):324-332.

SONG J J, XIE T, LIU W H, et al. Effects of malonaldehyde on protein oxidation of soybean meal and the mitigation effect of tea polyphenols[J]. Journal of Nanjing Agricultural University, 2023, 46(2):324-332. (in Chinese)

[30]
SUN L G, WU Q Z, MAO X Y. Effects of oxidation modification by malondialdehyde on the structure and functional properties of walnut protein[J]. FOODS, 2022, 11(16):2432.

[31]
DROMANT M, MAS-BARGUES C, ESCRIVÁ C, et al. Lipid peroxidation as measured by chromatographic determination of malondialdehyde.Human plasma reference values in health and disease[J]. Free Radical Biology and Medicine, 2021, 165:27-28.

[32]
BARRERA G, PIZZIMENTI S, DAGA M, et al. Lipid peroxidation-derived aldehydes,4-hydroxynonenal and malondialdehyde in aging-related disorders[J]. Antioxidants, 2018, 7(8):102.

[33]
ZIMMERMAN M C, CLEMENS D L, DURYEE M J, et al. Direct antioxidant properties of methotrexate: inhibition of malondialdehyde-acetaldehyde-protein adduct formation and superoxide scavenging[J]. Redox Biology, 2017, 13:588-593.

DOI PMID

[34]
ADAMS L K, BENSON E E, STAINES H J, et al. Effects of the lipid peroxidation products 4-hydroxy-2-nonenal and malondialdehyde on the proliferation and morphogenetic development of in vitro plant cells[J]. Journal of Plant Physiology, 1999, 155(3):376-386.

[35]
马路凯. 植物油中丙二醛、4-羟基-2-己烯醛和4-羟基-2-壬烯醛的热响应机制研究[D]. 博士学位论文. 广州: 华南理工大学, 2019.

MA L K. Formation mechanism of malondialdehyde,4-hydroxy-2-hexenal,and 4-hydroxy-2-nonenal in vegetable oils during thermal processmg[D].Ph.D.Thesis. Guangzhou: South China University of Technology, 2019. (in Chinese)

[36]
贾丰. 基于席夫碱反应改性的麦醇溶蛋白凝胶体系的构建、性能及其形成机理研究[D]. 博士学位论文. 广州: 华南理工大学, 2021.

JIA F. Preparation,properties,and formation mechanisms of schiff base reaction-modified gliadin gel-system[D].Ph.D.Thesis. Guangzhou: South China University of Technology, 2021. (in Chinese)

[37]
LIU A, TO VAN P T H, DUMAS A, et al. Vitamin nutrition in shrimp aquaculture:a review focusing on the last decade[J]. Aquaculture, 2024, 578:740004.

[38]
周敏娜. 维生素类清除自由基机理的密度泛函理论计算[D]. 硕士学位论文. 新乡: 河南师范大学, 2011.

ZHOU M N. Density functionai theory calculation on principle vitamin eliminating free radicals[D]. Master’s Thesis. Xinxiang: Henan Normal University, 2011. (in Chinese)

[39]
任颖. α-生育酚及α-生育三烯酚对生长育肥猪抗氧化能力及肉品质的影响[D]. 硕士学位论文. 武汉: 华中农业大学, 2020.

REN Y. The effect of α-tocopherol and α-tocotrienol on antioxidant capacity and meat quality of growing-finishing pigs[D]. Master’s Thesis. Wuhan: Huazhong Agricultural University, 2020. (in Chinese)

[40]
娄永, 马志欢, 邵香敏, 等. 芦丁及其衍生物研究进展[J]. 化学与粘合, 2023, 45(6):565-571.

LOU Y, MA Z H, SHAO X M, et al. Research progress in rutin and its derivatives[J]. Chemistry and Adhesion, 2023, 45(6):565-571. (in Chinese)

[41]
ZAMORA R, ALCON E, HIDALGO F J. Malondialdehyde trapping by food phenolics[J]. Food Chemistry, 2023, 417:135915.

[42]
ZAMORA R, ALCON E, HIDALGO F J. Addition of olivetol to crackers decreases malondialdehyde content and produces malondialdehyde-olivetol adducts[J]. Food Chemistry, 2024, 432:137046.

[43]
张玉, 龚埜, 李文飞, 等. 饲料磷脂水平对乌鳢幼鱼生长性能、抗氧化能力及脂肪沉积的影响[J]. 广东海洋大学学报, 2023, 43(6):9-18.

ZHANG Y, GONG Y, LI W F, et al. Effects of dietary phospholipid level on growth performance,antioxidant capacity and fat deposition of juvenile snakehead (Channa argus)[J]. Journal of Guangdong Ocean University, 2023, 43(6):9-18. (in Chinese)

[44]
DU Z, TURCHINI G M. Are we actually measuring growth?An appeal to use a more comprehensive growth index system for advancing aquaculture research[J]. Reviews in Aquaculture, 2022, 14(2):525-527.

[45]
黄雨薇, 叶元土, 蔡春芳, 等. MDA对草鱼肠道黏膜结构屏障损伤和肝胰脏、肠道胆固醇、胆汁酸合成影响[J]. 水生生物学报, 2016, 40(4):869-878.

HUANG Y W, YE Y T, CAI C F, et al. The damage of intestinal mucosa barrier structure and the effect of cholesterol and bile acid synthesis pathway in the liver and intestine under MDA in grass carp (Ctenopharyngodon idella)[J]. Acta Hydrobiologica Sinica, 2016, 40(4):869-878. (in Chinese)

[46]
陈科全, 叶元土, 蔡春芳, 等. 饲料丙二醛对草鱼生长、肝胰脏及肠道结构和功能的影响[J]. 水生生物学报, 2016, 40(4):779-792.

CHEN K Q, YE Y T, CAI C F, et al. Effects of MDA on the growth performance,structure and function of hepatopancreas and intestine of grass carp (Ctenopharyngodon idellus)[J]. Acta Hydrobiologica Sinica, 2016, 40(4):779-792. (in Chinese)

[47]
FENG L, FENG L, JIANG W D, et al. The beneficial effects of exogenous protease K originated from Parengyodontium album on growth performance of grass carp (Ctenopharyngodon idella) in relation to the enhanced intestinal digestion and absorption capacities[J]. Aquaculture, 2023, 563(Part 1):738929.

[48]
李晋南, 王常安, 张圆圆, 等. 高淀粉水平下精氨酸对松浦镜鲤肠道形态与功能的影响[J]. 广东海洋大学学报, 2022, 42(1):39-46.

LI J N, WANG C A, ZHANG Y Y, et al. Effect of arginine supplementation in high starch diets on intestinal digestive enzyme activities and intestinal morphology of Songpu mirror carp (Cyprinus carpio L.)[J]. Journal of Guangdong Ocean University, 2022, 42(1):39-46. (in Chinese)

[49]
KORTNER T M, AFANASYEV S, KOPPANG E O, et al. A comprehensive transcriptional body map of Atlantic salmon unveils the vital role of the intestine in the immune system and highlights functional specialization within its compartments[J]. Fish & Shellfish Immunology, 2024, 146:109422.

[50]
GUO Y L, FENG L, JIANG W D, et al. Impaired intestinal physical barrier of on-growing grass carp by deficient iron under Aeromonas hydrophila infection[J]. Aquaculture, 2023, 563(Part 1):738915.

[51]
ZHANG H L, RAN C, TEAME T, et al. Research progress on gut health of farmers teleost fish: a viewpoint concerning the intestinal mucosal barrier and the impact of its damage[J]. Reviews in Fish Biology and Fisheries, 2020, 30(4):569-586.

[52]
LI A L, YANG S J, HAN X T, et al. Progress of research on the alleviation of intestinal inflammation by regulating intestinal mucosal function with postbiotics[J]. Food Bioscience, 2024, 57:103437.

[53]
姚仕彬, 叶元土, 蔡春芳, 等. 丙二醛对离体草鱼肠道黏膜细胞的损伤作用[J]. 水生生物学报, 2015, 39(1):133-141.

YAO S B, YE Y T, CAI C F, et al. Damage of MDA on intestinal epithelial cells in vitro of grass carp (Ctenopharyngodon idella)[J]. Acta Hydrobiologica Sinica, 2015, 39(1):133-141. (in Chinese)

[54]
王晓艳, 李宝山, 孙永智, 等. 植物蛋白源诱发鱼类肠炎及修复的研究进展[J]. 广东海洋大学学报, 2023, 43(6):37-46.

WANG X Y, LI B S, SUN Y Z, et al. Research progress of plant protein-induced enteritis of fish and its repair[J]. Journal of Guangdong Ocean University, 2023, 43(6):37-46. (in Chinese)

[55]
黄雨薇. 氧化鱼油、丙二醛对草鱼(Ctenopharyngodon idellus)肝胰脏、肠道胆固醇、胆汁酸合成代谢的影响[D]. 硕士学位论文. 苏州: 苏州大学, 2015.

HUANG Y W. The effect of cholesterol and bile acid metabolism synthesis pathway in the liver and intestine under oxidized fish oil and MDA in Ctenopharyngodon idellus[D]. Master’s Thesis. Suzhou: Soochow University, 2015. (in Chinese)

[56]
蔡建光, 汤华, 唐晖, 等. 丙二醛对大鼠海马神经元结构的破坏和钙离子稳态的影响[J]. 生命科学研究, 2011, 15(4):283-289.

CAI J G, TANG H, TANG H, et al. The effects of malondialdehyde on the cellular structures of hippocampal neurons and its calcium homeostasis in SD rats[J]. Life Science Research, 2011, 15(4):283-289. (in Chinese)

[57]
龙建纲, 王学敏, 高宏翔, 等. 丙二醛对大鼠肝线粒体呼吸功能及相关脱氢酶活性影响[J]. 第二军医大学学报, 2005, 26(10):1131-1135.

LONG J G, WANG X M, GAO H X, et al. Malonaldehyde inhibits respiratory function and enzyme activities in isolated rat liver mitochondria[J]. Academic Journal of Naval Medical University, 2005, 26(10):1131-1135. (in Chinese)

[58]
SHI N B, YAN X, ADELEYE A S, et al. Effects of WS2 nanosheets on N2-fixing cyanobacteria:ROS overproduction,cell membrane damage,and cell metabolic reprogramming[J]. Science of the Total Environment, 2022, 849:157706.

[59]
FUKUSHIMA C T, DANCIL I S, CLARY H, et al. Reactive oxygen species generation by reverse electron transfer at mitochondrial complex Ⅰ under simulated early reperfusion conditions[J]. Redox Biology, 2024, 70:103047.

[60]
KISTER B, VIEHOF A, ROLLE-KAMPCZYK U, et al. A physiologically based model of bile acid metabolism in mice[J]. iScience, 2023, 26(10):107922.

[61]
苏冠宁, 孙赫, 吴高峰, 等. 胆汁酸的肠肝循环及胆汁酸受体在糖脂代谢中的调控作用[J]. 动物营养学报, 2019, 31(8):3511-3521.

DOI

SU G N, SUN H, WU G F, et al. Enterohepatic circulation of bile acids and regulation roles of bile acid receptors in glycolipid metabolism[J]. Chinese Journal of Animal Nutrition, 2019, 31(8):3511-3521. (in Chinese)

[62]
KOUNO T, LIU X, KISSELEVA T, et al. Selective PPARδ agonist seladelpar suppresses bile acid synthesis by reducing hepatocyte CYP7A1 via the fibroblast growth factor 21 signaling pathway[J]. The Journal of Biological Chemistry, 2022, 298(7):102056.

[63]
陈刚, 欧光海, 温震威, 等. 水飞蓟素对杂交石斑鱼幼鱼血清生化指标、脂代谢相关基因表达及肝脏代谢组的影响[J]. 广东海洋大学学报, 2023, 43(4):41-51.

CHEN G, OU G H, WEN Z W, et al. Effects of dietary silymarin supplementation on serum biochemical parameters,lipid metabolism-related gene expression,and liver metabolome in juvenile hybrid grouper (Epinephelus fuscoguttatus ♀×Epinephelus polyphekadion ♂)[J]. Journal of Guangdong Ocean University, 2023, 43(4):41-51. (in Chinese)

[64]
黄雨薇, 叶元土, 蔡春芳, 等. 氧化鱼油对草鱼(Ctenopharyngodon idelluspond)肝胰脏、肠道胆固醇、胆汁酸合成代谢的影响[J]. 基因组学与应用生物学, 2015, 34(8):1636-1646.

HUANG Y W, YE Y T, CAI C F, et al. The effect of cholesterol and bile acid metabolism synthesis pathway related enzymes in the liver and intestine under oxidized fish oil in Ctenopharyngodon idelluspond[J]. Genomics and Applied Biology, 2015, 34(8):1636-1646. (in Chinese)

[65]
汪兰, 夏雨婷, 胡澳, 等. 维生素C对禁食黄颡鱼抗氧化能力和肌肉品质的影响[J]. 广东海洋大学学报, 2024, 44(1):20-26.

WANG L, XIA Y T, HU A, et al. Effects of vitamin C supplementation on antioxidant capacity and muscle quality of yellow catfish (Pelteobagrus fulvidraco) after fasting[J]. Journal of Guangdong Ocean University, 2024, 44(1):20-26. (in Chinese)

[66]
刘慧敏. γ-谷氨酰半胱氨酸合成酶的酶学性质及其应用研究[D]. 硕士学位论文. 南京: 南京师范大学, 2021.

LIU H M. Enzymatic properties of gamma-glutamylcysteine synthetase and its applications[D]. Master’s Thesis. Nanjing: Nanjing Normal University, 2021. (in Chinese)

[67]
林秀秀, 叶元土, 蔡春芳, 等. 丙二醛引起草鱼肠道、肝胰脏谷胱甘肽/谷胱甘肽转移酶通路抗氧化应激[J]. 动物营养学报, 2015, 27(11):3604-3612.

DOI

LIN X X, YE Y T, CAI C F, et al. Malondialdehyde causes glutathione/glutathione transferase pathway oxidative stress in intestine and hepatopancreas of grass carp (Ctenopharyngodon idellus)[J]. Chinese Journal of Animal Nutrition, 2015, 27(11):3604-3612. (in Chinese)

[68]
CASTRO-CALDAS M, MILAGRE I, RODRIGUES E, et al. Glutathione S-transferase pi regulates UV-induced JNK signaling in SH-SY5Y neuroblastoma cells[J]. Neuroscience Letters, 2009, 451(3):241-245.

[69]
JOO Y C, JI H Y, JEA M S, et al. Tat-GSTpi inhibits dopaminergic cells against MPPsup+/sup-induced cellular damage via the reduction of oxidative stress and MAPK activation[J]. Biomedicines, 2023, 11(3):836-836.

[70]
WU Y, FAN Y, XUE B, et al. Human glutathione S-transferase P1-1 interacts with TRAF2 and regulates TRAF2-ASK1 signals[J]. Oncogene, 2006, 25(42):5787-5800.

DOI PMID

[71]
ZHANG L X, WANG J, XU T X, et al. Bicyclol alleviates obesity-induced renal injury by inhibiting JNK and NF-κB-mediated inflammation[J]. International Immunopharmacology, 2024, 129:111609.

[72]
聂一帆, 吴静, 马艳平, 等. 大口黑鲈肿瘤坏死因子-α诱导蛋白2基因克隆及免疫相关基因表达[J]. 广东海洋大学学报, 2022, 42(3):1-10.

NIE Y F, WU J, MA Y P, et al. Molecular cloning of tumor necrosis factor-alpha induced protein 2 gene and expression analysis of immune-related genes associated with it in largemouth bass (Micropterus salmoides)[J]. Journal of Guangdong Ocean University, 2022, 42(3):1-10. (in Chinese)

[73]
彭聪, 黎力之, 张进明, 等. 灵芝多糖的抗氧化作用及其在动物生产中的应用研究进展[J]. 中国畜牧杂志, 2021, 57(9):33-38.

PENG C, LI L Z, ZHANG J M, et al. Research progress on ganoderma lucidum polysaccharide antioxidant and its application in animal production[J]. Chinese Journal of Animal Science, 2021, 57(9):33-38. (in Chinese)

[74]
KOEBERLE S C, GOLLOWITZER A, LAOUKILI J, et al. Distinct and overlapping functions of glutathione peroxidases 1 and 2 in limiting NF-κB-driven inflammation through redox-active mechanisms[J]. Redox Biology, 2020, 28:101388.

[75]
张雨萌. 饲料组胺和丙二醛对珍珠龙胆石斑鱼生长性能、胃肠道健康和肌肉品质的影响[D]. 硕士学位论文. 湛江: 广东海洋大学, 2023.

ZHANG Y M. Effects of dietary histamine and malondialdehyde on growth performance,gastrointestinal health and muscle quality of hybrid grouper (Epinephelus fuscoguttatus ♀×E.lanceolatu♂)[D]. Master’s Thesis. Zhanjiang: Guangdong Ocean University, 2023. (in Chinese)

[76]
赵璐, 孙俊波, 金小琴. 丙二醛通过抑制Nrf2/ARE促进肾小球系膜细胞凋亡[J]. 基础医学与临床, 2015, 35(1):69-73.

ZHAO L, SUN J B, JIN X Q, et al. MDA accelerates the glomerular mesangial cell apoptosis through inhibition of Nrf2/ARE[J]. Basic & Clinical Medicine, 2015, 35(1):69-73. (in Chinese).

[77]
国家市场监督管理总局, 国家标准化管理委员会. 水产配合饲料第6部分:石斑鱼配合饲料:GB/T 22919.6—2024[S]. 北京: 中国标准出版社, 2024.

State Administration for Market Regulation, Standardization Administration of the People’s Republic of China. Aquatic feed-Part 6:formula feed for grouper:GB/T 22919.6—2024[S]. Beijing: Standards Press of China, 2024. (in Chinese)

[78]
国家市场监督管理总局, 国家标准化管理委员会. 水产配合饲料第8部分:巴沙鱼配合饲料:GB/T 22919.8—2024[S]. 北京: 中国标准出版社, 2024.

State Administration for Market Regulation, Standardization Administration of the People’s Republic of China. Aquatic feed-Part 8:formula feed for basa catfish:GB/T 22919.8—2024[S]. Beijing: Standards Press of China, 2024. (in Chinese)

[79]
中华人民共和国农业部. 中华人民共和国农业部公告第2045号[EB/OL].(2013-12-30) [2024-07-21]. http://www.moa.gov.cn/govpublic/XMYS/201401/t20140103_3730193.htm.

Ministry of Agriculture of the PRC. Announcement No. 2045 of the Ministry of Agriculture of the People’s Republic of China[EB/OL].(2013-12-30)[2024-07-21]. http://www.moa.gov.cn/govpublic/XMYS/201401/t20140103_3730193.htm. (in Chinese)

[80]
史弘赫, 刘洋, 王荻, 等. 棉籽浓缩蛋白饲料添加枯草芽孢杆菌和β-葡聚糖对虹鳟幼鱼生长性能、血清生化指标和机体健康的影响[J]. 动物营养学报, 2024, 36(6):3857-3875.

DOI

SHI H H, LIU Y, WANG D, et al. Effects of supplementation of Bacillus subtilis and β-glucan in cottonseed protein concentrate diets on growth performance,serum biochemical indices and organismal health of juvenile rainbow trout (Oncorhynchus mykiss)[J]. Chinese Journal of Animal Nutrition, 2024, 36(6):3857-3875. (in Chinese)

[81]
张卫, 谭北平, 庞奥博, 等. 珍珠龙胆石斑鱼豆粕诱导型肠炎肠道潜在生物标志物筛选[J]. 广东海洋大学学报, 2022, 42(4):1-12.

ZHANG W, TAN B P, PANG A B, et al. Screening of potential biomarkers for soybean meal induced enteritis in pearl gentian grouper (Epinephelus fuscoguttatus♀×Epinephelus lanceolatus♂)[J]. Journal of Guangdong Ocean University, 2022, 42(4):1-12. (in Chinese)

[82]
BURCU O K, EMINE K, MELTEM S. Lipid oxidation in cured meat model systems containing either antioxidant or prooxidant:a comparative study on the determination of malondialdehyde concentration by using conventional,test kit and chromatographic assays[J]. Food Analytical Methods, 2022, 15(10):3283-3296.

[83]
SELLIMI S, KSOUDA G, BENSLIMA A, et al. Enhancing colour and oxidative stabilities of reduced-nitrite turkey meat sausages during refrigerated storage using fucoxanthin purified from the tunisian seaweed Cystoseira barbata[J]. Food and Chemical Toxicology, 2017, 107(Pt B):620-629.

[84]
芦慧勤, 梅议文, 谢琦蓝, 等. L-赖氨酸对丙二醛诱导氧化下牦牛肉品质特性及氧化程度的影响[J]. 食品工业科技, 2023, 44(16):90-98.

LU H Q, MEI Y W, XIE Q L, et al. Effects of L-Lys on the quality characteristics and oxidation degree of yak meat under MDA induced oxidation[J]. Science and Technology of Food Industry, 2023, 44(16):90-98. (in Chinese)

[85]
蒋丙婷, 郭言, 尹可宏, 等. 4种多酚对丙二醛诱导花生球蛋白氧化的改善作用[J]. 中国粮油学报, 2024, 39(6):111-117.

JIANG B T, GUO Y, YIN K H, et al. Ameliorating effect of 4 polyphenols against malondialdehyde-induced oxidation of arachin[J]. Journal of the Chinese Cereals and Oils Association, 2024, 39(6):111-117. (in Chinese)

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