2. 青海畜牧兽医职业技术学院, 湟源 812100
2. Qinghai Vocational and Technical College of Animal Husbandry and Veterinary Medicine, Huangyuan 812100, China
硒是反刍动物必需的微量营养素,具有抗氧化活性、抗炎、抗诱变、抗癌、抗病毒、抗菌、抗真菌、抗寄生虫等功能,硒有助于维持机体正常的生理功能[1]。新的研究表明,硒可以调节免疫、影响细胞内信号传递和基因调控,而且还可以缓解应激[2], 硒兼有营养、解毒等多种生物学功能,是生命的“保护剂”,硒与动物的健康密切相关,在草地放牧系统中发挥着重要作用[3]。世界上许多地区的土壤中硒含量都很低[4-6], 我国许多放牧地区也属于缺硒地区,新疆天山北部放牧地区[7]、青藏高原东北边缘放牧草地[8]、青海湖布哈河上游地区高寒草甸[9]等地区属于严重硒缺乏地区,而硒缺乏会导致反刍动物生长缓慢、免疫水平下降,继而影响反刍动物的健康及生产力。因此,在缺硒地区,反刍动物饲粮中需要添加硒,以满足对硒的需求,而饲粮中添加硒首先要研究硒对反刍动物各方面的影响,因此本文主要通过阐述硒对反刍动物瘤胃发酵、生产性能、免疫功能、抗氧化性能的影响,以期为硒在反刍动物生产及应用中提供参考。
1 硒对反刍动物瘤胃发酵和营养物质消化率的影响 1.1 促进瘤胃发酵硒对反刍动物瘤胃发酵有重要的影响,瘤胃微生物需要微量元素来维持正常的功能[10]。研究表明,羟基蛋氨酸硒能促进奶牛瘤胃发酵[11],绵羊饲喂纳米硒可以提高其瘤胃发酵和饲料利用率[12]。还有研究发现,饲粮中添加0.3 mg/kg酵母硒有助于促进育成湖羊的瘤胃发酵[13]。硒能够促进瘤胃发酵的原因可能是硒刺激了瘤胃微生物或酶活性[14-15];同时添加硒保护了瘤胃纤毛虫种群[16],还有研究发现,添加纳米硒可以增加瘤胃微生物数量,从而提高氨氮利用率[17];饲粮中复合添加硒和维生素E,可以降低瘤胃pH,同时增加了绵羊瘤胃内的挥发性脂肪酸(VFA)含量、原虫总数[18],因此添加硒提高了瘤胃发酵。但也有研究表明,绵羊饲粮补充0.2 mg/kg DM硒酸钠或亚硒酸钠时,对瘤胃微生物功能没有显著影响,瘤胃液硒含量与瘤胃细菌数量呈显著负相关,VFA含量及乙酸、丙酸比值差异不显著,没有显著影响瘤胃微生物的功能[19],这可能与绵羊对无机硒的吸收较低、饲粮中的硒多以不溶性形式存在相关,从而降低了硒的有效性,同时硒的添加效果与硒的添加形式、添加量及添加持续时间、动物生理状态、拮抗作用等因素有关,其原因还不明确,还需要深入研究。
综合上述研究,硒促进反刍动物瘤胃发酵的主要原因可能是添加硒增加了总VFA含量,把乙酸为主的发酵模式转变为丙酸发酵模式,提高了丙酸的含量[14, 18, 20],同时补充硒能显著增加瘤胃微生物数量和活性[15],另外,有机硒有更高的生物利用率,可以加快瘤胃微生物的繁殖,增加总VFA含量[21], 添加硒通过提高瘤胃微生物功能,提高了瘤胃氮利用率,促进了瘤胃发酵,但是具体机理还需要深入研究。
1.2 提高营养物质消化率饲粮补硒可以提高绵羊全消化道中干物质(DM)、有机物(OM)、粗蛋白质(CP)、粗脂肪(EE)、中性洗涤纤维(NDF)、酸性洗涤纤维(ADF)消化率,饲粮中添加纳米硒可提高反刍动物的瘤胃发酵,饲料转化效率也较添加酵母硒显著提高[12],同时添加酵母硒也可以提高羔羊的营养物质消化率[22],研究还发现西门塔尔牛基础饲粮中添加0.3、0.6 mg/kg的蛋氨酸硒,OM、CP消化率显著高于对照组[23]。羟基蛋氨酸硒可以促进泌乳中期奶牛瘤胃发酵,提高营养物质表观消化率[11],同样饲粮中添加酵母硒也可以提高泌乳期奶牛营养物质的消化率[14]。
2 硒对反刍动物生产性能的影响 2.1 提高生长性能硒可以提高反刍动物的生长性能。硒对小牛出生后的发育非常重要,牛奶中硒含量不足可能导致小牛硒缺乏,影响小牛健康生长,母牛补充硒可防止这种情况发生[24]。母牛补充硒可以增加其体重[25],硒还可以提高湖羊的生长性能,添加0.3 mg/kg酵母硒通过提高饲粮中营养物质表观消化率和增加组织中硒的沉积来提高了育成湖羊的生长性能[13]。研究表明,母羊在妊娠后期补充含硒的瘤胃缓释丸,可以提高断奶期羔羊的体重[26]。杜寒杂交羊饲粮添加0.2 mg/kg的酵母硒时,其平均日增重(ADG)最大,随着硒添加量的增加,ADG又开始降低, 结果说明,适量的硒可以提高杜寒杂交羊的生长性能[27]。但是也有研究发现,硒来源及其添加量对犊牛日增重无显著影响[28]。
2.2 促进泌乳补充硒可以促进反刍动物泌乳性能。研究表明,当母羊从怀孕到产期补充高水平的硒时,可以提高其产奶量[29]。荷斯坦奶牛补充0.3 mg/kg DM羟基蛋氨酸硒,可提高其产奶量,但是会降低乳脂率[30]。硒提高反刍动物泌乳量以及乳成分原因可能在于硒的含量影响了乳腺的血管分布,促进了乳腺发育,同时补充硒可以为增加牛奶产量提供更多的营养[31],同时泌乳奶牛的硒含量是影响乳腺硒蛋白活性的重要调节因子[32],硒提高了牛乳腺上皮细胞内和细胞外谷胱甘肽过氧化物酶(GSH-Px)等硒蛋白的体外表达,促进了牛乳腺上皮细胞的生长和活力[33]。此外,硒降低了热应激诱导的氧化应激,从而提高了母牛的泌乳性能。但也有许多研究表明,对不缺硒的奶牛补充不同硒源和剂量的硒,都不能显著影响产奶量或乳的成分[34-36],饲喂各种水平的有机或无机硒并不影响奶牛产奶量和乳成分,原因可能是与反刍动物不同生理状态、硒添加水平及添加方式有关[36-37]。
3 硒对反刍动物免疫功能的影响及作用机制硒在反刍动物体内的代谢与机体免疫关系密切,它可以提高反刍动物免疫功能[38],它是硒蛋白的重要成分,硒蛋白在内分泌、免疫、炎症中发挥着多种作用[39]。硒对反刍动物免疫水平有着重要的影响,饲粮补充适量的硒可以提高反刍动物的免疫功能,而缺乏硒会导致反刍动物免疫水平降低,并产生一系列的病变。
3.1 补充硒提高免疫功能在细胞和分子水平上研究表明,饲粮中适当补充硒,对母羊和羔羊的免疫水平具有很重要的作用[40],牛血清硒含量与其免疫反应和健康之间存在正相关性[41-42]。研究发现,有机和无机硒都可以提高羔羊的体液免疫反应[43],研究表明奶牛饲粮中添加硒其体细胞数低于对照组[44],硒还可以改善氧化应激,减轻奶牛乳腺炎和子宫炎症状[45]。表 1归纳了硒对反刍动物免疫功能的影响,大部分研究表明硒可以提高反刍动物的免疫水平。
反刍动物缺乏硒会影响其免疫功能。研究表明,低水平的硒可能会提高反刍动物传染病和慢性病的发病率和死亡率[40],缺硒甚至还会导致肉牛死亡[53]。而且它对牛免疫系统有巨大的影响[46],缺乏硒与免疫抑制有关[53],缺硒通常表现为胎盘潴留、子宫炎、乳腺炎、流产、生育力下降和感染易感性增加,在小牛、羔羊中,缺硒产生白肌病[54],同时缺硒还与胎膜残留率增加、临床乳腺炎、小牛死亡率和乳体细胞数增加有关,而补充硒之后降低了这些临床疾病的发生率[55]。也有研究表明,血清硒含量与子宫内膜炎发生有关, 患有子宫内膜异位症的奶牛在分娩后往往呈现较低水平的硒[56]。研究还发现,硒缺乏导致幼畜产生克山病,这表明硒与病毒感染之间存在着一定的关系[57]。
3.3 硒提高免疫功能的机制硒存在于25种硒蛋白中,而大多数硒蛋白已被证明具有抗炎作用[48],硒蛋白参与调节氧化应激和抗炎反应过程,低水平的硒会增加死亡风险,导致免疫功能低下,而高水平的硒或补充硒有抗病毒作用[48]。一方面,硒通过多种硒蛋白发挥生物学作用来提高免疫功能;另一方面,硒通过提高中性粒细胞功能来提高免疫水平,研究表明全血硒含量与中性粒细胞黏附呈显著正相关[59],缺硒导致中性粒细胞活动减少,降低了中性粒细胞的功能,而中性粒细胞是细胞内杀死侵入的病原体所必需的,同时GSH-Px是一种抗氧化剂,可以保护中性粒细胞不受有毒活性氧分子的破坏[60]。而在非特异性免疫上,硒可以提高吞噬细胞(包括单核-巨噬细胞和中性粒细胞)的功能,它们依赖于氧化还原的机制清除病原微生物[61]; 在红细胞免疫上硒也有重要的影响,饲粮中添加富硒益生菌能有效提高山羊红细胞数量[62]; 在特异性免疫上,硒蛋白水平还影响免疫细胞发育以及免疫反应[63]; 在体液免疫上,细胞内硒水平可能在B细胞激活过程中发挥了重要作用,因此提高了体液免疫水平[64]。
4 硒对反刍动物抗氧化性能的影响及作用机制硒的抗氧化作用是硒生理生化作用的基础,硒是组织和细胞抗氧化防御必不可少的营养成分,硒主要通过其在GSH-Px中的作用参与抗氧化。饲粮添加硒可以显著提高血液和组织中GSH-Px的活性[65]。
4.1 提高抗氧化性能硒在细胞抗氧化系统中起关键作用。表 2归纳了硒对反刍动物抗氧化性能的影响,研究表明硒可以通过提高GSH-Px活性来提高不同反刍动物的抗氧化性能,而且还可以有效缓解氧化应激。
以上研究表明,硒可以显著提高牦牛、山羊、奶牛、肉牛的抗氧化水平,而缺乏硒会降低反刍动物的抗氧化水平。
4.2 提高抗氧化性能的机制硒是构成GSH-Px的活性中心的元素[72-73],硒主要通过发挥GSH-Px作用来实现抗氧化,而GSH-Px是第1个被证明可以防止细胞膜氧化损伤的硒酶,硒通过抗氧化、消除自由基的生物学作用,显著提高了反刍动物机体的抗氧化系统功能[74],GSH-Px活性与可溶性硒呈正相关[75],研究表明硒主要通过依赖GSH-Px等硒蛋白发挥抗氧化功能[76], 硒蛋白主要是指含有硒代半胱氨酸(Sec)的蛋白,被称为第21种氨基酸,而硒蛋白的主要特征是调节氧化还原状态和保护机体免受氧化损伤[77]。硒可以提高抗氧化性能的机制是GSH-Px可以清除过氧化氢(H2O2), 并将脂质过氧化物(ROOH)还原成相应稳定的醇和水, 从而减轻细胞膜多不饱和脂肪酸过氧化作用, 以及ROOH分解产物引起的细胞损伤[78],其反应式为:
不同硒蛋白在不同的机体内发挥着重要的抗氧化功能,从而提高了反刍动物的抗氧化能力,表 3归纳了硒蛋白在组织细胞浆、红细胞、细胞内、细胞外、血浆、肾脏、肺脏、肠黏膜中起到的抗氧化作用。
综上所述,硒在反刍动物体内广泛存在,在不同的功能领域中发挥着重要作用。硒对反刍动物有着非常重要的作用,它能够通过促进反刍动物瘤胃发酵以及营养物质在瘤胃内的降解,提高反刍动物对饲粮的利用率,改善机体免疫力,防治疫病,提高机体的抗氧化水平,从而促进反刍动物生长。
目前硒在反刍动物应用方面研究较多,但是相关机理研究较少,还需要从以下5个方面着手:1)需要进一步研究硒在不同反刍动物生产中的适宜添加量以及作用效果和作用机制。2)研究硒的添加形式,比如有机硒和螯合形式的硒对反刍动物的作用,以此提高硒的利用率。3)研究不同反刍动物在不同生理阶段的硒需要量及硒对动物健康的影响。4)通过体外筛选和体内验证的方式得出适合于不同反刍动物的硒添加剂和添加水平。5)研究影响反刍动物对硒吸收和利用的因素。
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