抗生素是指由微生物或高等动植物在生活过程中所产生的一类具有活性或者可以抵抗病原体的次级代谢产物,是一种能干扰其他活细胞生长发育的化学物质。中国科学院研究员应光国课题组研究表明,2013年中国总的抗生素消耗量为16.2万t左右,消耗量约为英国的160倍,其中兽用抗生素占到总体的 52%[1]。畜禽养殖业中的抗生素分为两种:一种为饲用抗生素,通过添加到预混料中达到预防疫病和促生长作用;另一种为治疗用抗生素,是动物出现疫病后通过拌料、注射或者灌服等方式为畜禽治疗用抗生素。饲用抗生素能促进动物生长,防治动物腹泻,但同时也引起药物残留、细菌耐药性等负面问题,对人、动物和生态环境造成严重危害[2]。欧盟2006年下令禁止在饲料中使用抗生素;美国从2014年起禁止在畜禽饲料中使用防御性抗生素;相继,韩国也于2018年禁用在饲料中使用抗生素;农业农村部指出我国药物饲料添加剂将在 2020年全部退出。为保证食品安全,寻找安全、有效的抗生素替代品刻不容缓。
近年来,一些具有抗病促生长作用的替代抗生素的饲料添加剂已开始应用于畜禽养殖业中,并且取得了一定的应用效果。这些抗生素替代品主要有微生态制剂、酸化剂、植物提取物、酶制剂和抗菌肽等。
微生态制剂指的是利用益生菌或益生菌的次级代谢产物制成的制剂。饲用微生态制剂具有绿色、无污染且不易产生耐药性等特点,可作为饲料添加剂长期使用[3]。我国农业农村部公布允许使用的微生物饲料添加剂有34种,主要包括乳酸菌、芽孢杆菌、链球菌、真菌和克鲁维酵母菌等[4]。
1.1.1 益生菌在单胃动物养殖中的应用
在母猪生产中,益生菌可调节母猪机体代谢,还可调节妊娠期母猪肠道微生物和初乳的结构来影响初生仔猪的被动免疫和肠道微生物结构,进而提升仔猪肠道健康潜力[5],提高仔猪生产性能[6-7]。Yu等[8]试验表明,在利用鼠伤寒沙门菌建立的断奶仔猪攻毒模型中,补充鼠李糖乳杆菌可快速清除体内的沙门氏菌,并且缓解由沙门氏菌引起的一系列炎症反应。在肉鸡生产中也得到类似的效果。益生菌可以提高肉鸡生长性能和免疫机能[9-10],改善禽肠道微生物的组成,抑制有害菌的定植[11-12],且包被益生菌的效果更显著[13]。益生菌代谢产生甘露寡糖、果寡糖、菊粉等物质,这些物质被称为益生元,这些物质进入到肠道后可被肠道益生菌利用,促进有益菌的增值。Wang等[14]在断奶仔猪饲料中添加微囊化植物乳杆菌和低聚果糖可有效地提高仔猪的免疫力和结肠中的乳酸菌含量,达到替代抗生素的作用。
不仅如此,益生菌还具有净化畜舍环境的功能。枯草芽孢杆菌可以在动物体内产生分解硫化氢的酶类和氨基化氧化酶等,这些物质可以氧化吲哚,降低动物肠道内氨气、硫化氢的浓度,改善畜舍环境[15]。Smialek等[16]用益生菌混合物(乳酸乳球菌1×109 cfu/g、广布肉毒杆菌 1×109 cfu/g、干酪乳杆菌1×109 cfu/g、植物乳杆菌1×109 cfu/g和酿酒酵母1×107 cfu/g)添加到Ross308肉鸡日粮中,降低了弯曲杆菌对肉鸡胃肠道的侵袭力,净化了家禽饲养环境。
1.1.2 益生菌在反刍动物养殖中的应用
在奶牛生产中,添加一定量的乳酸菌还可以防治奶牛阴道炎,降低发病率[17]。GenÍs等[18]给产犊前3周的奶牛阴道注射2次乳酸菌后,使阴道炎患病率降低了58%,说明乳酸菌在一定程度上可以作为奶牛子宫炎的预防性疗法。细菌代谢产生的中链脂肪酸和短链脂肪酸如乙酸、丙酸对肠道健康也是非常有益的[19]。Kenney等[20]给肉牛补充屎肠球菌和嗜酸乳杆菌(1×109 cfu/d),不仅可以促进生长,还可以改善瘤胃发酵特性,促进乙酸比例提高。
1.1.3 益生菌在水产养殖中的应用
乳酸菌因其特有的生物学价值也被广泛应用到水产养殖中,乳酸菌可以提高鱼的生长性能、免疫应答、抗病力[21],还可以改善水质,防止铝污染引起的水产养殖和食品安全问题[22]。Bogut等[23]用嗜酸乳杆菌、粪肠球菌和干酪乳杆菌饲喂鲤鱼,鲤鱼的生长加快,饲料转化率提高。赵倩等[24]发现产朊假丝酵母、枯草芽孢杆菌和鼠李糖乳杆菌中药发酵制剂均能显著影响鲤的肠道菌群结构,其中鼠李糖乳杆菌中药发酵剂组肠道菌群最丰富。
1.1.4 益生菌在发酵饲料中的应用
我国的优质蛋白质饲料资源如大豆、鱼粉等存在着严重短缺的情况,而一些非常规资源如棉粕、菜粕等非常丰富,但是它们中的一些抗营养因子如棉酚、噁唑烷硫酮、异硫氰酸酯等过量摄入会引起动物中毒,严重限制了非常规饲料的利用[25]。而经过微生物或酶制剂处理的棉、菜粕可有效降低抗营养因子含量,提高动物免疫力。Wang等[26]研究表明,棉粕经过枯草芽孢杆菌和酵母菌混合发酵后可有效降低棉酚含量,提高肉鸡免疫力,并且提高了盲肠乳酸菌的数量。Hu等[27]利用枯草芽孢杆菌、产假丝酵母、粪肠球菌发酵菜粕饲喂肉鸡,结果表明,发酵菜粕有效的降低了异硫氰酸酯和硫代葡萄糖苷的含量,改善了肉鸡肠道形态。
有机酸作为食品防腐剂已有近百年时间了。有机酸主要包括延胡索酸、柠檬酸、乙酸、乳酸、丙酸、苹果酸、山梨酸、戊酮酸等[28]。饲料酸化剂可降低饲料中细菌、霉菌的数量,提高饲料品质和适口性,进入消化道后可降低食糜pH,提高消化酶活性,还可通过杀菌抑菌的作用,维持消化道菌群平衡,从而保证机体健康[29]。有机酸可直接扩散进入细菌细胞内,抑制病原菌体内关键的大分子物质如脂类、蛋白质、DNA和RNA等的代谢活动,破坏其细胞膜的完整性,起到保护机体的作用[30]。
无机酸以其酸性强、成本低的优势,也得到了一些厂家和研究者的青睐。一些无机酸主要包括强酸如硫酸、盐酸,弱酸如磷酸等。有些研究表明,盐酸在促进仔猪生长方面效果甚至优于有机酸[31]。但是过量的氯离子会造成动物中毒,食欲减退[32],在实际生产中要非常注意。
1.2.1 酸化剂在生猪养殖中的应用
已经有大量研究表明了有机酸对畜禽的作用,如有机酸可以提高胃蛋白酶和胰蛋白酶活性[33-34],改善日粮适口性。早期断奶仔猪胃酸分泌能力弱,消化系统不完善,胃酸分泌能力差[35],酸化剂的应用在早期断奶仔猪上效果尤为明显。姚继明等[36]研究发现,日粮中添加复合有机酸化剂(五酸肽)能极显著提高断奶仔猪的日增重和日采食量,降低料肉比。甲酸–柠檬酸复合物不仅可以提高生长育肥猪生长代谢,还可以有效降低了粪便中沙门氏菌的数量[37]。研究表明,柠檬酸、苯甲酸、乳酸、苹果酸均具可以起到防治仔猪腹泻的效果[38]。
1.2.2 酸化剂在禽类养殖中的应用
李万军等[39]研究表明在饲粮中添加益生菌及有机酸复合制剂能够提高大骨鸡生长性能和免疫水平,对肉品质有一定的改善作用。酸化剂在增加采食量的同时,还可以提高饲料粗蛋白、粗脂肪消化率,提高饲料转化效率[40],改善小肠粘膜结构[41-42]。另外,酸化剂一方面可以通过降低胃肠pH值促进维生素A和维生素D的吸收,另一方面与矿物元素形成易被吸收的络合物来促进矿物元素的吸收。Boling等[43]研究表明,在低磷日粮中添加柠檬酸和柠檬酸钠使雏鸡胫骨灰分和日增重分别增加43%和22%,有效的提高了雏鸡植酸磷的利用率,但是对猪的影响很小。酸化剂还可以有效改善肠道微生物区系及肠道屏障功能[44],抑制肠道有害细菌[45]和真菌的繁殖[46],促进动物生长。Ahmed等[47]研究表明肉鸡饲粮中添加有机酸,可以调节肠道大肠杆菌引起的炎症反应,提高生产性能,降低肉鸡死亡率。混合酸化剂(柠檬酸、乳酸、磷酸和延胡索酸)也可显著降低肉鸡盲肠食糜中螺杆菌属细菌的数量,增加乳杆菌属和瘤胃球菌科细菌的数量,并促进有益微生物的生长[48]。
植物体内次级代谢产物含量丰富,主要包括皂苷、多糖、生物碱、挥发油、多酚、黄酮类等多种类型[49]。研究表明,植物提取物可促进畜禽分泌消化酶,降低肠道 pH值,促进营养物质吸收,刺激动物生长[50];可提高机体免疫力,抑制炎症[51];可提高肌肉中肌红蛋白含量,改善肉品质[52];可清理机体中的氧化自由基,消除氧化应激,提高机体的抗氧化能力[53]。近年来,植物提取物行业发展迅速,它是我国中药的主要成分,不仅在我国,乃至在欧美国家已经发展成为一个有广泛市场前景的新兴产业。
1.3.1 饲用植物提取物在生猪养殖中的应用
王珍珊等[54]试验结果显示,添加4.0%苜蓿和紫苏混合植物提取物到育肥猪饲料中,能显著提高日增重,降低料重比,改善胴体品质。植物精油种类非常多,许多都具有较强的杀菌能力。如百里香、肉桂、冬香等精油对猪胸膜肺炎放线杆菌、猪链球菌、支气管败血波氏杆菌、猪放线杆菌、多杀性巴氏杆菌和副猪嗜血杆菌均有较好的抑制效果[55]。给母猪补饲植物多糖、茶多酚、皂苷、类黄酮等植物提取物可有效的提高仔猪初生重和活仔率,降低死胎数,缩短产程时间,同时,改善了仔猪初生重的均匀度[56]。
1.3.2 饲用植物提取物在禽类养殖中的应用
胡贵丽等[57]研究表明,博落回生物碱可刺激口腔内唾液分泌,降低胆碱酯酶活性,提高肉鸡的平均日增重和平均日采食量。植物提取物在促进机体免疫器官发育方面也起到了一定的作用。杜仲叶提取物不仅可以提高机体免疫球蛋白 IgM和IgG的含量,还对肉鸡法氏囊发育具有促进作用[58];姜黄提取物对文昌鸡法氏囊和胸腺发育也有一定的促进作用[59]。植物提取物不仅可以缓解肠道菌群失调引起的腹泻问题,还可以调节肠道物理屏障功能。添加5%牛至精油能够显著增加肉鸡的回肠绒毛高度,提高乳酸菌数量,降低盲肠和回肠内容物中大肠杆菌数量[60];闻爱友等[61]试验表明迷迭香和百里香的复合植物提取物可显著提高肉鸡体内乳酸杆菌和双歧杆菌数量,降低大肠杆菌的数量,与朱宇旌等[62]在断奶仔猪上的研究结果一致。丝兰提取物可提高肉仔鸡空肠绒毛高度和十二指肠绒毛高度[63]。
1.3.3 饲用植物提取物在奶牛养殖中的应用
袁宇等[64]发现肉桂油可抑制黄曲霉的生长,柠檬醛可抑制禾谷镰刀菌的生长,丁香酚可抑制黄曲霉和禾谷镰刀菌生长。关于植物提取物治疗和预防乳房炎的研究也非常多,如蒲公英提取物[65]、皂苷[66]、生物碱[67]都对引起乳房炎的葡萄球菌有明显抑制作用,对提高奶牛的免疫力也有一定的促进作用。
在动物饲料中添加的酶制剂主要分为两类,一类是动物消化道可以分泌的内源性消化酶,另一类是动物本身不能产生的外源性消化酶。作用机制主要包括:(1)补充体内消化酶的不足,刺激内源酶的分泌;(2)破坏植物细胞壁,提高饲料利用率;(3)消除抗营养因子,提高营养价值。在实际生产中,往往根据不同的动物类型和阶段以及饲料原料种类,有选择的进行添加,如木聚糖酶主要用于含有小麦副产物的饲料中,以达到节约成本的目的。
单胃动物不能有效地利用植酸磷,植酸酶可水解植酸,释放有机磷。范秋丽等[68]研究表明,植酸酶可以提高肉鸡生产性能,提高胫骨密度。纤维素、半纤维素、果胶等非淀粉多糖也不能被单胃动物利用,生产中经常通过添加一些纤维素酶、β–葡聚糖酶、木聚糖酶、果胶酶等来提高饲料利用率、降低饲料成本。添加木聚糖酶和β–葡聚糖酶到玉米 DDGS(玉米酒精糟及可溶物)日粮中,可有效提高保育猪DDGS消化率,进一步提高仔猪日增重[69]。李晓洁等[70]研究表明,添加含 β–葡聚糖酶和木聚糖酶的复合非淀粉多糖酶到肉鸡低能玉米–豆粕型日粮中,进一步提升了日粮能量和养分的利用率,显著提高了肉鸡的表观代谢能。在乳仔猪、肉仔鸡和肉仔鸭等幼龄动物中补充一定量的外源性消化酶,如蛋白酶、淀粉酶、脂肪酶等,可有效地提高动物生产性能,降低死淘数[71]。杨贤斌等[72]发现,脂肪酶可有效地提高仔猪的生长性能,促进脂肪代谢,还可有效提高仔猪日粮干物质、总能和氮的表观消化率。
除一些具有消化功能的酶外,某些酶还具有清除体内有害物质,提高机体免疫力的作用。如过氧化氢酶、葡萄糖氧化酶和溶菌酶等酶类,这些酶具有改善动物肠道健康、提高饲料利用率,减轻炎症反应的作用。熊华武[73]研究表明,溶菌酶能够有效地缓解仔猪的腹泻,促进仔猪的生长。过氧化氢酶可明显促进断奶仔猪肠道的发育,改善绒毛、隐窝结构,促进体内各种酶活性,降低机体腹泻率[74]。
抗菌肽广泛分布于自然界的动物、植物和微生物当中,是带正电荷的小分子多肽[75]。抗菌肽通常由10~50个氨基酸组成,具有广谱抗菌活性、热稳定性、抗菌机制独特和不易产生耐药性等特点,且不会导致环境污染问题[76]。不同的抗生素作用机制不同,多数抗生素通过与细菌细胞膜和胞内特异性受体结合,抑制细菌细胞壁或 DNA的合成而发挥作用,容易产生耐药性[77],而抗菌肽是通过与细菌细胞膜的互相作用破坏细菌细胞膜的完整性,使细菌内容物外泄而导致细菌死亡,因而不易产生耐药性[78],具有良好的抗细菌、抗病毒的作用。
天蚕素是最早分离出来的昆虫抗菌肽,也是最早分离得到的抗菌肽。天蚕素具有提高肉鸡生产性能、改善肠道黏膜结构、提高机体免疫力的功效[79]。天蚕素还可以有效缓解大肠埃希菌攻毒引起的断奶仔猪腹泻症状,有效提高日增重[80]。近年来,随着肠道微生物研究的热度增高,抗菌肽在肠道微生物方面的研究越来越多,而且均起到了一定的积极作用。蛙皮素抗菌肽Dermasepin-M可使育肥猪平均日增重提高4.64%,料重比降低 4.71%,十二指肠中大肠杆菌含量降低 11%,乳酸菌含量提高20.59%[81]。Shi等[82]在断奶仔猪饲粮中添加复合抗菌肽,也显著提高了肠道中乳酸杆菌和双歧杆菌的数量,显著降低了大肠杆菌的数量。Liao等[83]研究显示,抗菌肽可以降低对虾肠道有害菌数量,提高有益菌数量,改变肠道微生物区系,从而达到提高生产性能、提高免疫力的效果。体外抑菌试验也证实了抗菌肽La47、Css54对大肠杆菌和金黄色葡萄球菌均能达到与氯霉素和氨苄霉素类似的抑菌效果[84]。
抗菌肽在抵抗病原菌上起到了一定的替代抗生素的效果,具有广谱抗菌活性,但是其生产工艺复杂,产量低,成本高,造成养殖成本大幅增加。
抗生素替代品在动物的生长和健康方面发挥了积极的作用,对生态环境也起到了一定的改善作用,而且无污染、无残留、无副作用。然而,在抗病和促生长方面与抗生素还存在一定距离。而且抗生素替代品本身也存在一些缺陷,如微生态制剂不易存放、易失活,效果不稳定;抗菌肽产品生产成本高,存在转基因安全问题;在使用中为了增强效果,往往采用多种产品复配使用,大大增加了生产成本;抗生素替代类饲料添加剂种类繁多,每种都有其优缺点,很难说清楚哪种添加剂效果最佳;一些厂家以次充好、以假乱真,混淆市场。由此可见,抗生素替代的道路还很长,而且,养殖户需要从饲养管理、饲养环境等多方面双管齐下,减少抗生素的使用。
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Research Progress on Substitutes of Antibiotics Used as Feed Additives