Chinese Journal of Animal Nutrition >

2024 , Vol. 36 >Issue 8: 4847 - 4860

DOI: https://doi.org/10.12418/CJAN2024.415

REVIEW

Research Progress of Fermented Feed in Aquaculture

  • LI Zhengzhong , 1 ,
  • LIU Bo , 1, * ,
  • SHAO Peng 2, 3 ,
  • ZHU Aimin 3
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  • 1 Key Laboratory of Freshwater Fisheries and Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs, Freshwater Fisheries Research Center, Chinese Academy of Fishery Sciences, Wuxi 214081, China
  • 2 Yancheng Shangshui Environmental Biotechnology Engineering Co., Ltd., Yancheng 224400, China
  • 3 Yancheng Academy of Fishery Sciences, Yancheng 224051, China
*professor, E-mail:

Received date: 2024-02-06

  Online published: 2024-08-12

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Abstract

With the rapid development of aquaculture industry and the shortage of fish meal resources, the proportion of plant protein source in feed is gradually increasing, but the anti-nutritional factors in plant protein source lead to low feed utilization, and it also have adverse effects on the health of aquatic animals. In recent years, a large number of studies have shown that fermented feed can improve the growth and palatability of feeds, help to improve the utilization rates of nutrients such as protein, amino acids, small peptides, carbohydrates, etc. by aquatic animals. In addition, biofermentation technology can reduce the antinutritional factors in plant protein sources, increase the nutrient contents of feed, and improve the health level and immunity of aquatic animals. This paper mainly discussed the mechanism of fermented feed, the main strains of fermented bacteria, and the research progress, remaining shortcomings and prospects of fermented feed in aquaculture in order to provide reference for the application of fermented feed in aquaculture.

Key words： fermented feed; fermented strains; aquaculture; research progress

Cite this article

LI Zhengzhong , LIU Bo , SHAO Peng , ZHU Aimin . Research Progress of Fermented Feed in Aquaculture[J]. Chinese Journal of Animal Nutrition, 2024 , 36(8) : 4847 -4860 . DOI: 10.12418/CJAN2024.415

水产养殖有着悠久的历史,始于2 000多年前的中国。目前我国的水产养殖产量占世界水产养殖产量的近68%。根据统计年鉴,2022年我国淡水养殖的产量已达到3 406.38万t,为人类提供了大量的优质动物蛋白质,改善了人们的食物结构[1]。随着集约化水产养殖的发展,养殖水环境污染和养殖动物疾病爆发等问题日益突出,严重的甚至导致养殖动物死亡,造成经济上的重大损失[2]。对此,研究者们研究出了如抗菌肽[3]、中草药提取物[4]、益生菌[5]、后生元[6]等添加剂,用以提高水产动物机体的免疫力与抗病力,以减少病害发生。其中,益生菌类产品因其高安全性、低成本、应用广泛等优点而受到广泛青睐。水产养殖中的益生菌是一类对养殖动物和养殖水体有益的微生物的总称[7]。微生物发酵饲料是在人为控制下利用有益微生物对饲料底物进行发酵的过程,能够改变饲料的营养特性、增强适口性、提高饲料的利用率和改善养殖水环境。
近年来,世界鱼粉资源短缺,价格持续上涨,植物蛋白质源在水产饲料中的占比逐渐增加。植物蛋白质源含有的纤维素和抗营养因子(ANFs)会降低水产动物的消化吸收能力,甚至引发肠炎问题[8],而通过生物发酵技术降低抗营养因子含量被证明是高效且安全的一种方式。益生菌在发酵过程中可对底物进行“预消化”,通过自身代谢活动减少底物中单宁、植酸、游离棉酚和硫甙等抗营养因子的含量,同时增加可溶性蛋白、酸溶性蛋白和小肽等营养物质的含量[9⇓⇓-12]。另外,研究表明,微生物在发酵过程中可以分解膳食纤维,产生多种小分子糖类,有利于肠道微生物的生长[13-14],并且微生物在水产动物肠道生长过程中可产生短链脂肪酸(SCFAs),能够上调抗炎因子的表达水平,增强肠道化学屏障功能[15-16]。发酵饲料的应用能为水产养殖业高质量发展提供坚实的保障。本文就发酵饲料的作用机理、发酵菌种的选择、在水产养殖中的应用以及应用前景进行综述,以期为发酵饲料在水产养殖中的应用提供参考。

1 发酵饲料作用机理

1.1 发酵饲料对发酵底物的作用机制

发酵过程中,益生菌通过降解抗营养因子、释放营养物质途径实现对发酵底物的“预消化”。研究表明,解淀粉芽孢杆菌JY-24通过液体发酵可显著提高辣木叶(Moringa oleifera Lam.)中蛋白质、总酚、黄酮、小肽含量(分别提高了24.53%、82.81%、58.58%和38.63%),同时使单宁含量降低8.4%[17];复合菌发酵日本沼虾基础饲料可显著增加饲料中芽孢杆菌、乳杆菌的数量,增加饲料中消化酶活性以及可溶蛋白、小肽营养物质含量,改善饲料风味[18];芽孢杆菌发酵豆粕可使蛋白质含量提高6.1%,β-伴大豆球蛋白(β-conglycinin)含量降低58%[19];使用鼠李糖乳杆菌发酵麸皮时,发酵麸皮中总蛋白质、总碳水化合物、总酚类和总黄酮类含量明显高于未发酵麦麸,同时降低了植酸含量(从4.20%减少到0.07%)[20];Vo等[21]对花生种子进行不同处理:20 kg花生种子在25~27 ℃、90%相对湿度条件下,每6 h用33~37 ℃水浸泡1次进行发芽(发芽法);20 kg花生种子在90 ℃下蒸煮 2 h,然后磨碎至约50 μm榨油(物理萃取法);取1/2榨油后的花生粕用嗜酸乳杆菌(Lactobacillus acidophilus)在37 ℃下发酵72 h(发酵法),结果显示,发酵法和发芽法降解花生粕单宁酸和生物碱能力优于物理萃取法,且发酵法对单宁酸的降低率明显高于发芽法。

1.2 发酵饲料对肠道菌群的作用机制

水产动物在摄食发酵饲料后,饲料中富含的益生菌在肠道内与致病菌争夺肠道内的生态位产生竞争关系,从而降低致病菌的丰度[22]。除此之外,厌氧生长的益生菌(如酵母菌[23]、芽孢杆菌[24]等)在肠道内生长会消耗氧气、代谢产生乳酸等,在水产动物肠道内制造酸性低氧环境,可抑制有害微生物生长,并为厌氧有益菌创造良好的生存环境。同时,益生菌在发酵过程产生代谢产物[25⇓⇓-28]和抗菌物质(如有机酸、过氧化氢和细菌素[29]),这些代谢产物能够抑制嗜水气单胞菌(Aeromonas hydrophila)[30]和弧菌[31]等有害菌的生长繁殖,调节水产动物肠道内的微生态平衡。

1.3 发酵饲料对细菌代谢产物的作用机制

发酵过程中微生物能代谢产生消化酶(淀粉酶、脂肪酶、蛋白酶、纤维素酶)和营养因子(B族维生素、小肽等)[32-33],辅助水产动物消化吸收饲料中的营养物质。此外,饲料在发酵过程中,大分子蛋白质可被微生物通过代谢活动降解为多肽类物质,部分多肽类物质具有较高的抗氧化活性[34],能够增强水产动物的免疫力与健康,促进生长。

2 发酵饲料使用的主要菌种

发酵饲料的核心之一就是发酵菌种,各种类型的微生物,包括细菌、酵母和真菌已在饲料发酵过程中广泛使用。

2.1 乳杆菌

乳杆菌在水产养殖中得到了广泛的关注,包括植物乳杆菌(Lactobacillus plantarum)、嗜酸乳杆菌、干酪乳杆菌(Lactobacillus casei)、副干酪乳杆菌(Lactobacillus paracasei)、保加利亚乳杆菌(Lactobasillus bulgaricus)、鼠李糖乳杆菌(Lactobacillus rhamnosus)等[35]。乳杆菌在发酵过程中不仅可以降低发酵底物的pH,还能够降低胰蛋白酶抑制因子等抗营养因子含量,产生消化酶、超氧化物歧化酶、乳酸和柠檬酸等,促进蛋白质水解,释放游离氨基酸[36⇓-38]。厉桓[39]研究发现,植物乳杆菌与黄芪粉1∶1固态发酵7 d可降解黄芪大分子蛋白质,增加半乳糖苷酶含量。董奇奇等[40]研究发现,在乳上清液中添加1%~5%鼠李糖乳杆菌发酵20 h,可提高Caco-2细胞核因子E2相关因子2(Nrf2)/醌氧化还原酶1(NQO1)信号通路表达,缓解脂多糖(LPS)诱导的氧化应激,降低炎症反应和肠上皮细胞间通透性,改善肠道屏障。

2.2 芽孢杆菌

在水产养殖中,芽孢杆菌是研究最多的与宿主相关的细菌,包括枯草芽孢杆菌(Bacillus subtilis)、地衣芽孢杆菌(Bacillus licheniformis)、解淀粉芽孢杆菌(Bacillus amyloliquefaciens)、凝结芽孢杆菌(Bacillus coagulans)。这主要是由于芽孢杆菌独特的能力,在水产动物肠道中生长可分泌多种消化酶,并保持肠道处于厌氧环境,有效抑制致病菌的生长,提高水产动物免疫力和对致病菌的抵抗力[41⇓⇓-44]。此外,芽孢杆菌在发酵过程中能降低饲料中的抗营养因子,产生B族维生素(尤其是维生素B12和生物素),提高了饲料的营养价值[45]。另有研究表明,芽孢杆菌还可改善水质,降低水体中致病菌的丰度[46]。

2.3 酵母菌

酿酒酵母(Saccharomyces cerevisiae)、产朊假丝酵母(Candida utilis)、热带假丝酵母(Candida tropicalis)、红酵母属(Rhodotorula)、毕赤酵母属(Pichia)等酵母菌使用历史悠久,作为最早用于发酵的益生菌,其功能性产物包括β-葡聚糖、甘露寡糖和几丁质,另外还可代谢产生酶、低聚糖、氨基酸、肽、有机酸、维生素和其他可溶性因子[47-48]。姜燕等[49]研究表明,使用酿酒酵母发酵刺参饲料可产生乙醇,使饲料具有独特的风味,增加饲料的诱食效果,同时增加饲料中可消化蛋白质含量,改善饲料品质。

2.4 肠球菌

肠球菌是人类和一些动物肠道微生物区系中的一员。研究表明,屎肠球菌可改善罗非鱼(Oreochromis niloticu)的生长性能,增强免疫反应,增加血清补体3(C3)含量和髓过氧化物酶活性[50];提高欧洲鳗鲡(Anguilla anguilla)[51]和褐牙鲆(Paralichthys olivaceus)[52]的免疫能力并抑制由爱德华氏菌和链球菌引起的机体感染。杨成等[53]研究表明,屎肠球菌以25%接种量发酵玉米粉24 h可显著降低发酵底物pH,增加乙酸含量。Shi等[54]研究表明,先用枯草芽孢杆菌再用粪肠球菌进行发酵,可显著降解豆粕中的抗营养因子含量,增加小肽和游离氨基酸的含量。

2.5 木霉菌

木霉菌可产生多种具有生物活性的酶系,如纤维素酶、几丁质酶、植酸酶、木聚糖酶等,可降解结构复杂的多糖和木质素,使与木质素铰链在一起的纤维素和半纤维素游离出来,并进一步降解为多糖和单糖。木霉菌是纤维素酶的主要生产菌,在生产生物燃料、制造纸浆、酿酒等产业中被广泛应用,主要研究对象是里氏木霉(Trichoderma reesei)[55]。随着世界鱼粉资源短缺,植物蛋白质源被大量引入到饲料中,而植物蛋白质的细胞壁含有纤维素、木质素和半纤维素等,影响了水产动物对饲料的消化利用率,且摄食过多纤维素、木质素和半纤维素等会降低水产动物的摄食效率,引发机体炎症反应,降低生长性能[56]。纤维素酶能够破坏植物细胞壁,释放营养物质,提高草鱼(Ctenopharyngodon idella)肠道消化酶活性,增加肠道绒毛高度和杯状细胞数量,保护肠道健康[57]。孙亚楠等[58]研究发现,在菊芋渣表面喷洒0.1%里氏木霉发酵2个月可增加粗蛋白质和粗灰分含量,改善菊芋渣的营养组成。刑胜利等[59]对里氏木霉的发酵条件进行了优化,在发酵5 d后菌体菌丝大量增长,纤维素酶活性不断提高,可高效降解玉米秸秆中的半纤维素和纤维素,产生葡萄糖、阿拉伯糖等。

2.6 基因工程菌

基因工程菌是指将目的基因片段导入细菌体内使其表达,产生所需要的功能的细菌,其核心技术是DNA的重组技术。针对菌种益生性能,可以根据自身需求运用基因工程技术筛选出符合需求的菌种,例如,何锡杲[60]通过基因工程技术将植酸酶基因导入毕赤酵母中得到高产植酸酶菌种,产酶量比优化前提高15%;Zhao等[61]通过CRISPR/Cas9技术得到1株高产淀粉酶的芽孢杆菌,淀粉酶产量提高22%。

3 发酵饲料的发酵方式

发酵方式主要包括单菌发酵、复合菌发酵和菌酶协同发酵,在生产中,多数研究集中在复合菌发酵以及菌酶协同发酵。复合菌发酵即是用2种及以上的有益微生物按照不同比例对底物进行发酵,微生物间可产生协同效应。范阳等[62]根据发酵后豆渣的pH、益生菌数量、还原糖含量、氨态氮浓度等指标,筛选出最适的混菌比例、接种量、水分、发酵温度和发酵时间,确定了混菌发酵豆渣的最优工艺。菌酶协同发酵即在益生菌基础上添加酶制剂协同发酵,酶协同发酵的效果优于单纯使用菌种发酵。研究表明,利用复合菌(枯草芽孢杆菌和植物乳杆菌)与中性蛋白酶协同发酵豆粕,该发酵豆粕替代鱼粉比例小于20%时对大口黑鲈(Micropterus salmoides)的生长没有产生显著影响[63];复合菌(枯草芽孢杆菌,酿酒酵母和植物乳杆菌)与0.1%植酸酶协同发酵可改善菜籽粕的品质[64];枯草芽孢杆菌BJ-1与木瓜蛋白酶等协同发酵可增加棉籽粕的粗蛋白质和氨基酸含量,同时降低粗纤维和游离棉酚含量,提高棉籽粕的品质[65]。

4 发酵饲料在水产养殖中的应用

4.1 发酵饲料对养殖动物生长性能的影响

与传统饲料相比,水产动物更易吸收发酵饲料中的营养物质。Das等[66]研究表明,通过混合菌株(酿酒酵母和枯草芽孢杆菌)固态发酵麻油饼可增加其粗蛋白质含量,降低粗纤维以及单宁、皂苷等抗营养因子含量,发酵麻油饼替代饲料中20%的豆粕可显著提高露斯塔野鳞(Labeo rohita)的生长性能、饲料转化率和营养物质的消化率;在饲料中添加20~50 g/kg嗜酸乳杆菌和酿酒酵母固体发酵浒苔(Enteromorpha prolifera)能显著提高红罗非鱼(Oreochromis mossambicus×Oreochromis niloticus)的增重百分比和特定生长率,降低饲料系数,提高了肝脏和肠道蛋白酶以及淀粉酶活性,增强消化性能[67];在基础饲料中添加5%~10%发酵饲料能显著提高斑点叉尾鮰(Silurus punctatus)的终末体重、特定生长率和蛋白质效率,改善鱼体肌肉营养组成[68];在配合饲料基础上额外添加日投喂量5%~20%的发酵饲料可显著提高中华绒螯蟹(Eriocheir sinensis)的存活率、增重率和特定生长率[69]。
值得注意的是,发酵饲料在基础饲料中比例过大可能无积极效果,甚至影响动物生长与健康。例如,Das等[66]用发酵油饼替代40%豆粕,露斯塔野鳞的生长性能、营养物质消化率和免疫参数与对照组无显著差异;蔡润佳等[70]的研究结果证实,全部饲喂发酵饲料组的蛋白质利用效率要低于基础饲料组;陈晓瑛等[63]研究发现,饲粮中发酵豆粕替代超过20%鱼粉后会显著降低大口黑鲈的终末体重、特定生长率和摄食率。分析原因可能是因为发酵饲料一般含有较多水分,导致摄食的营养物质不足。由此可见,要酌情分析发酵饲料和基础饲料间的比例,发酵饲料占基础饲料的比例应在30%以下。

4.2 发酵饲料对养殖动物抗氧化和免疫性能的影响

水产动物摄食发酵饲料后,饲料中被释放的活性物质可调控水产动物的免疫与健康。研究表明,摄食发酵饲料能提高鲤鱼(Cyprinus carpio)幼鱼血浆中皮质醇和C3含量,且对机体的抗氧化能力没有负面影响[71]。使用发酵辣木叶替代饲料中40%~60%的鱼粉可显著提高异育银鲫(Carassius auratus gibelio)血清中免疫酶[超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、溶菌酶(LZM)]活性和C3含量,降低丙二醛(MDA)含量,抑制脾脏Toll样受体2(TLR2)和肿瘤坏死因子-α(TNF-α)信号通路表达,提高对嗜水气单胞菌的耐受性[72]。饲料中添加20%~30%的微生物发酵剂能显著提升罗氏沼虾(Macrobrachium rosenbergii)肝胰腺中SOD活性[73]。
另外,一些发酵植物和动物成分作为饲料添加剂也可提高水产动物的免疫力。例如,在饲料中添加1%~3%发酵柠檬皮[74-75]、3 mg/kg植物乳杆菌发酵的生物硒[76]、0.2%~0.8%发酵开菲尔[77]、2.5%发酵蚯蚓[78]可分别提高南美白对虾、日本囊对虾(Marsupenaeus japonicas)、斑马鱼等水产动物的非特异性免疫能力、抗氧化能力及抗病力。

4.3 发酵饲料对养殖动物肠道健康的影响

保护水产动物肠道健康是发酵饲料的重要功能。一方面,动物在摄食发酵饲料后,有益菌在肠道定植与致病菌争夺生态位,并分泌抗菌肽、细菌素等抑制致病菌繁殖,达到改善动物肠道菌群结构的目的[79-80]。研究发现,饲喂发酵饲料能提高凡纳滨对虾肠道厚壁菌门、拟杆菌门、芽孢杆菌属和乳杆菌属的丰度,降低弧菌属的丰度[81-82]。葛玲瑞等[83]研究表明,草金鱼(Carassius auratus red var)摄食经过酵母菌和芽孢杆菌发酵后的饲料,肠道双歧杆菌属(Bifidobacterium)的丰度显著增加;陈李婷[84]研究证明,发酵饲料可提高田螺(Cipangopaludina chinensis Gray)肠道中厚壁菌门的丰度和肠道物种多样性。另一方面,益生菌在肠道定植后分泌的代谢产物如SCFAs、乳酸等,被证明可修复宿主肠道黏膜,构建肠道屏障,提高宿主对病原菌的抵抗力[85]。韩瑞玲等[86]研究表明,摄食发酵饲料可增加凡纳滨对虾肠道绒毛高度,使肠上皮细胞绒毛排列更为整齐、紧密。晏明瑶等[87]研究发现,克氏原螯虾(Procambarus clarkii)幼体摄食发酵饲料后肠道皱襞高度和横截面积显著增加。

4.4 发酵饲料对养殖水体的影响

发酵饲料在改善养殖水体方面也有突出的表现。近年来,水产养殖密度不断增大,随之而来的水环境问题日益突出,究其原因是传统饲料中的蛋白质供给量远远超过水产动物本身的需求量,养殖动物未吸收利用的蛋白质随排泄废物一起排出体外,水体中氮、磷元素的不断累积,导致水体环境恶化。
多数养殖户在水质恶化时进行换水或泼洒化学药物,但只能缓解一时,且换水及化学药物易引起水产动物应激反应,而微生物发酵饲料中的益生菌可代谢产生氨基转移酶等,对水体中的污染物起到分解作用,大幅度降低水质污染[88]。王天神[89]研究表明,投喂发酵饲料的试验组养殖水体pH和亚硝酸盐含量均低于对照组;王凤军等[90]研究发现,在养殖第7天发酵饲料组养殖水体的氨氮和亚硝酸盐含量显著低于未发酵组。
表1列举了在水产养殖中常用的发酵菌种制备的发酵饲料的品质改善情况,以及发酵饲料对养殖动物的影响。通过表1可知,益生菌通过分解抗营养因子,增加可消化蛋白质、消化酶活性等,改善水产动物的摄食,提高饲料利用率。
表1 发酵对饲料品质的影响及发酵饲料对养殖动物的影响

Table 1 Effects of fermentation on feed quality and effects of fermented feed on animals

编号
No.		 发酵菌种
Fermented
strains		 发酵底物
Fermented
substrate		 添加量
Addition
level		 养殖动物
Farmed
animals		 发酵对饲料品质的影响
Effects of fermentation on
feed quality		 发酵饲料对养殖动物的影响
Effects of fermented
feed on animals		 参考文献
References
1 复合菌种:枯草芽孢
杆菌+酿酒酵母+
植物乳植杆菌		 日本沼虾
基础饲料		 替代25%的基
础饲料		 日本沼虾 乳酸菌、芽孢杆菌、酵母菌数量↑;
霉菌数量↓;可溶性蛋白
质、小肽含量↑		 FBW、WGR↑;FCR↓;血清SOD活性↑,
MDA含量↓;肠道气单胞菌属丰度↓,
芽孢杆菌属和双歧杆菌属丰度↑		 李正中等[18]
2 鼠李糖乳杆菌 小麦麸皮 替代50%~
100%的麦麸		 罗非鱼 总蛋白、总碳水化合物、总酚类、
总黄酮类含量↑;DPPH、ABTS
活性↑;植酸含量↓		 FBW、WGR、SGR↑;肌肉粗蛋白质含量↑,
粗脂肪含量↓;水体总氨、二氧化氮氧化物
含量↓;血清血红蛋白含量、血细胞比容及
红细胞、白细胞、血小板计数↑		 Flefil等[20]
3 复合菌种:枯草芽孢
杆菌+酿酒酵母+
植物乳植杆菌		 菜籽粕 替代30%的
基础饲料		 黄颡鱼 — FBW、WGR↑;FCR↓;肝体比↑;
肝脏ZO-1、ZO-2 mRNA表达量↓		 贾冰玉等[64]
4 枯草芽孢杆菌 斑点叉尾鮰
基础饲料		 替代5%~10%
的基础饲料		 斑点叉尾鮰 — WGR、SGR↑;FCR↓;肌肉蛋白质含量↑,
灰分含量↓;血清AST、ALT活性↓;肝脏
GSH含量、SOD活性↓;头肾SOD活性↓		 陈启亮
等[68]
5 复合菌种:芽孢杆
菌+乳酸球菌		 蚯蚓 饲料中
添加2.5%		 尖齿胡鲶 粗蛋白质、粗灰分、组氨酸、
苯丙氨酸、亮氨酸、缬氨酸、
脯氨酸、半胱氨酸含量↑		 呼吸爆发活性、血细胞比容、吞噬
活性、吞噬指数、血清SOD活性和
白细胞分化↑		 Nugraha等[78]
6 复合菌种:植物乳植
杆菌+酿酒酵母+
沙福芽孢杆菌		 南美白对虾
基础饲料		 全部饲喂
发酵饲料		 南美白对虾 酸溶蛋白、赖氨酸、
蛋氨酸含量↑		 WGR、体长↑;FCR↓;肝脏消化
酶活性、T-AOC、SOD活性↑;
肠道菌群多样性↑		 Zhang等[82]
7 复合菌种:鼠李糖
乳酪杆菌+枯草芽
孢杆菌+酿酒酵母+
索氏鲸杆菌		 颗粒饲料 替代5%的
基础饲料		 南美白对虾 粗蛋白质、酸溶蛋白、粗脂肪含
量↑;乳酸菌、芽孢杆菌、
酵母菌数量↑		 WGR、SGR↑;FCR↓;血清ACP、
LZM活性↑,MDA含量↓;肠道
变形菌门丰度↑		 韩瑞玲等[86]
8 复合菌种:芽孢杆
菌+酵母菌+植物
乳植杆菌		 克氏原螯虾
配合饲料		 全部饲喂
发酵饲料		 克氏原螯虾 饲料溶失率↓ WGR、SGR↑;FCR↓;肌肉粗蛋
白质、粗脂肪、粗灰分、丝氨酸含量↑;
肝脏胰蛋白酶、脂肪酶、SOD、ACP、
AKP活性↑,MDA含量↓		 姚海行等[92]
9 复合菌种:酿
酒酵母+干酪
乳杆菌		 桑树、莪术
和苧麻
组成的发酵
草药提取物		 基础饲料中添加
300 mg/kg		 尼罗罗非鱼 — WGR、SGR↑;肠道绒毛长度、
淋巴细胞数↑		 Fujaya等[93]
10 真菌菌丝 小麦谷物 发酵小麦谷物替代
未发酵小麦谷物		 虹鳟 — SGR、平均体重↑;肌肉水分含
量↓,脂肪含量↑		 Pascua等[94]
11 酵母菌 葵花籽粕 替代25%
葵花籽粕		 尼罗罗非鱼 粗蛋白质、粗脂肪、粗灰分含量↑;
粗纤维含量↓;总水解氨基酸、
必需氨基酸、非必需氨基酸含量↑;
酚类化合物、植酸、皂苷含量↓		 FBW、WG、SGR↑;
血清ALT、AST活性↓		 Hassaan等[95]
12
酿酒酵母		 麦麸 饲料中添加20% 尼罗罗非鱼 粗蛋白质、总氨基酸含量↑;
纤维素、植酸含量↓		 FBW、WGR、
SGR↑;FCR↓		 Mohammady
等[96]
13 枯草芽孢杆菌 微囊藻副产品 饲料中添加
0.5%或者5%		 南美白对虾 芽孢杆菌数量,总碳水
化合物、黄酮化合物含量,
DPPH活性↑		 WGR、SGR↑;
副溶血弧菌抗性↑		 Lee等[97]
14 黑曲霉 菜籽粕 替代20%鱼粉 真鲷 — FBW、WGR、SGR↑;肌肉
脂肪含量↑;血液血细胞比容↑,
葡萄糖含量↓;肝脏LZM、过氧化物
酶活性↑;对低盐胁迫耐受性↑		 Dossou等[98]
15 菌酶协同发酵:
植物乳杆菌+芽孢
杆菌+酿酒酵母+纤
维素酶+木聚糖酶+
β-葡聚糖酶+β-
甘露聚糖酶		 茶渣 饲料中
添加2%		 大口黑鲈幼鱼 — WGR、FBW、FL↑;肠道消化酶、AKP、
LZM、ACP、GSH-Px、CAT活性以及
T-AOC↑,MDA含量↓;IL-10、
TNF-β、IL-1β、IL-15、TNF-α、
IL-8 mRNA表达量↑		 Hu等[99]
16 复合菌种:酿酒酵
母+索氏鲸杆菌+
鼠李糖乳酪杆菌		 鲤鱼
基础饲料		 替代5%~15%的
基础饲料		 鲤鱼 pH↓;小肽含量↑ 肝脏IL-10、TNF-α mRNA表达量↑;肠道
ZO-1 mRNA表达量及肝素和防御素含量↑;
肠道镰刀菌、固着菌和类杆菌丰度↑		 Zhang等[100]

DPPH:1,1-二苯基-2-苦基肼 1,1-diphenyl-2-picrylhydrazyl;ABTS;2,2'-联氮-双-3-乙基苯并噻唑啉-6-磺酸 2, 2'-azino-bis(3-ethylbenzothiazoline-6-sulfonic acid);FBW:末体重 final body weight;FL:末体长 final body length;WG:增重 weight gain;WGR:增重率 weight gain rate;FCR:饲料系数 feed conversion rate;SGR:特定生长率 special growth rate;ZO-1:闭锁小带蛋白-1 zonula occludens-1;ZO-2:闭锁小带蛋白-2 zonula occludens-2;SOD:超氧化物歧化酶 superoxide dismutase;MDA:丙二醛 malondialdehyde;AST:谷草转氨酶 aspartate aminotransferase;ALT:谷丙转氨酶 alanine aminotransferase;GSH:谷胱甘肽 glutathione;T-AOC:总抗氧化能力 total antioxidant capacity;LZM:溶菌酶 lysozyme;ACP:酸性磷酸酶 acid phosphatase;AKP:碱性磷酸酶 alkaline phosphatase;GSH-Px:谷胱甘肽过氧化物酶 glutathione peroxidase;CAT:过氧化氢酶 catalase;IL-10:白细胞介素-10 interleukin-10;IL-1β:白细胞介素-1β interleukin-1β;IL-15:白细胞介素-15 interleukin-15;IL-8:白细胞介素-8 interleukin-8;TNF-α:肿瘤坏死因子-α tumor necrosis factor-α;TNF-β:肿瘤坏死因子-β tumor necrosis factor-β。

5 小结与展望

发酵饲料在水产养殖中的益生作用愈发突出,发酵饲料产生的营养、生物和化学屏障不仅可以提高水产养殖的产量,还可以改善水产品的品质,实现双赢的效果。但是当前发酵饲料还处于发展阶段,技术条件不是很成熟,还存在较多问题。首先,发酵饲料缺乏统一的生产标准以及评价标准,难以突出发酵饲料的产品特点;其次,发酵菌种主要从动物肠道或者自然界如土壤中分离纯化得到,对菌种的研究不够深入,在使用前需要对其进行安全性评估[91]。另外,发酵饲料缺乏广泛的宣传及技术支持,养殖户对发酵饲料的认识不足并且缺乏技术指导,导致发酵饲料得不到广泛应用。因此,发酵饲料仍然有很大的发展空间,需要科研人员结合实际水产养殖的需求以及要求,选择正确的菌种,为养殖户提供持续的技术支持,推动发酵饲料在水产养殖中的应用。

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