2. 青海省高原牦牛研究开发中心, 西宁 810016;
3. 青海省高原放牧家畜动物营养与饲料科学重点实验室, 西宁 810016;
4. 青海畜牧兽医职业技术学院, 湟源 812100
2. Qinghai Plateau Yak Research and Development Center, Xining 810016, China;
3. Key Laboratory of Grazing Animal Nutrition and Feed Science of Qinghai Province, Xining 810016, China;
4. Qinghai Vocational and Technical College of Animal Husbandry and Veterinary Medicine, Huangyuan 812100, China
锌存在于器官、组织、骨骼、液体和细胞中,它对反刍动物机体营养生理功能有重要的影响。锌不仅参与和影响体内的三大营养物质和维生素的代谢过程, 还直接或间接参与氨基酸、核酸等物质的代谢过程,而且有超过300种酶的结构和功能依赖锌[1-2]。因此,本文主要通过阐述锌对反刍动物免疫、瘤胃发酵、生产性能、繁殖性能的影响,探讨不同添加形式锌的添加效果和反刍动物锌需要量,以期为锌在反刍动物生产及应用中提供参考。
1 锌对反刍动物免疫功能的影响锌对反刍动物免疫系统正常发育、维持免疫功能有着重要的作用。锌在细胞免疫中的作用对动物抵抗病原体的感染十分重要[3]。有研究表明,给奶牛饲喂低锌饲粮时,牛乳中的体细胞数比较高[4],说明锌具有提高反刍动物体液免疫和细胞免疫的作用。此外,锌还可以提高血清总蛋白和球蛋白含量[5]。Nagalakshmi等[6]通过羊羔试验研究表明,补充15 mg/kg的锌(基础饲粮中锌含量为29.28 mg/kg),羊羔机体可以获得较高的免疫反应,同时补充锌蛋白与补充硫酸锌(ZnSO4)相比,具有更高的抗氧化酶活性和免疫反应。在水牛上的研究结果发现,补充80或140 mg/kg ZnSO4,血清碱性磷酸酶活性和球蛋白含量均升高[7]。给较小的荷斯坦公牛饲喂150 mg/kg ZnSO4,血浆总蛋白、尿素氮、白蛋白含量均显著高于对照组[8]。急性缺锌会导致反刍动物先天和适应性免疫功能下降,而慢性缺锌则会增加炎症的发生率,缺锌还会影响细胞介导的免疫功能,导致单核细胞-巨噬细胞的活化,同时锌具有重要的抗氧化功能[9-10],而缺锌可以引发反刍动物的免疫缺陷, 增加疫病感染的几率。以上研究结果表明锌可以提高反刍动物机体免疫水平和抗氧化功能。
2 锌对反刍动物瘤胃环境的影响锌对反刍动物瘤胃环境有重要的影响,对于反刍动物而言, 锌的营养意义不仅针对动物机体, 而且对瘤胃内环境同样有重要的作用。锌参与了瘤胃微生物的代谢, 它通过对微生物的促生长作用影响瘤胃的正常发酵[11]。Arelovich等[12]研究发现,锌通过延缓尿素中氨的释放,增加反刍动物挥发性脂肪酸中丙酸的比例,同时在饲粮锌含量为250 mg/kg时补充锌,具有降低尿素毒性的可能性,提高反刍动物发酵的能量效率。Černík等[13]研究表明,给山羊羔补充氧化锌(ZnO),其瘤胃乳头较对照组发育良好。Arelovich等[14]在研究中发现,饲粮中添加100 mg/kg的锌可以通过影响瘤胃的某些发酵模式,不同程度地影响动物对补充蛋白质以及低质量粗饲料的利用。添加纳米氧化锌,体外发酵(6~12 h)能够有效地促进瘤胃微生物对饲粮有机物的发酵,增加微生物蛋白产量,提高瘤胃发酵的能量利用效率[15]。Eryavuz等[16]研究发现,在含纤维素的培养液中加入锌发酵48 h时,对照组的纤维素溶解菌和总细菌数量较锌补充组低。但是也有不同的研究发现,羊羔饲粮中高含量的锌会降低挥发性脂肪酸的产生,降低乙酸/丙酸[17],其原因还需要进一步研究。
以上研究表明,饲粮补充锌可通过增加瘤胃中高效酸丙酸的含量、促进瘤胃乳头的发育、提高瘤胃发酵的能量效率以及影响瘤胃发酵模式来促进瘤胃发酵,提高反刍动物对营养物质的利用率,其原因还需要深入研究。
3 锌对反刍动物生产性能的影响 3.1 增重表 1总结了不同形式和不同水平的锌对反刍动物增重的影响。多数研究结论认为饲粮中添加不同锌源和不同锌水平均能提高犊牛、肉牛、羔羊、山羊、绵羊、母羊的平均日增重,从而提高反刍动物的生产性能,但是也有一些研究认为添加锌并不能提高平均日增重,这可能与锌不同添加形式、添加量及添加持续时间、试验动物、动物生理状态、拮抗作用等因素有关。
锌对反刍动物泌乳性能有着重要的影响。郑梦莉等[25]研究发现,饲粮中添加蛋氨酸锌(Zn-Met)和甘氨酸螯合锌均可提高妊娠母羊的生长性能并提高羊奶品质。Salama等[27]研究发现,山羊补充1 g/d Zn-Met,补充组乳清蛋白和非蛋白氮含量显著高于对照组。Smith等[28]研究发现,对于高产奶牛,特别是有蹄问题的奶牛,饲粮中添加Zn-Met可以提供蹄生长所需的营养,并通过帮助保持蹄健康和干物质采食量来提高牛奶产量。Kellogg等[29]研究发现,添加Zn-Met可以提高奶牛泌乳性能,体细胞数减少33.3%, 改善了乳房健康。Cope等[30]研究认为,给奶牛添加NRC(2001)推荐的锌(螯合形式的锌),增加了乳产量。锌氨基酸络合物与ZnSO4的混合补锌,相对于单纯补锌,可以增加奶牛的产奶量,随着锌氨基酸络合物与ZnSO4的比值的增加,产奶量增加的更加明显[31]。
4 锌对反刍动物繁殖性能的影响锌是对反刍动物生殖影响最大的微量矿物元素之一,对反刍动物繁殖性能有着重要的影响。与其他生殖器官相比,生殖细胞中锌的含量要高出数倍,这表明生殖细胞优先积累锌,这可能对生殖细胞的发育、生长和生存很重要[32]。
4.1 对公畜繁殖性能的影响锌对雄性反刍动物精子的发生及初级、次级雄性器官的发育有重要的影响[33]。有研究表明,荷斯坦公牛饲粮中添加锌可提高精子的抗氧化能力,血清中睾酮含量随着饲粮锌添加量的增加而显著增加[34]。任有蛇等[35]研究发现,晋中公绵羊饲粮添加50~100 mg/kg的纳米锌可提高睾丸和附睾中铜锌超氧化物歧化酶(Cu-Zn SOD)mRNA和蛋白表达量。Rahman等[36]研究结果显示,雄鹿补充锌之后精液体积和精子的运动性明显增加,添加100 mg/(头·d)ZnSO4可以改善雄鹿的精液性状和精液血浆抗氧化能力。Arangasamy等[37]研究发现,锌可提高补饲山羊的精子活力,降低精子氧化应激。还有研究发现,缺锌导致山羊睾丸萎缩,性欲下降[38]。Kumar等[39]研究发现,无论是丙酸锌还是ZnSO4均能提高公牛精液的数量和质量。缺锌首先会损害血管紧张素转换酶的活性,进而导致睾酮含量的减少,同时会抑制精子的生成[40],这可能是锌影响雄性反刍动物繁殖功能的重要因素。
4.2 对母畜繁殖性能的影响锌除了对雄性反刍动物繁殖性能有重要影响,对雌性家畜生殖过程的每个阶段也起着重要的作用。王长宏[41]研究表明,奶牛添加Zn-Met,其正常分娩率为80%,较对照组提高了6.67%。有研究发现,锌在肉牛卵母细胞成熟过程中显著影响卵丘细胞内谷胱甘肽含量和DNA完整性,促进胚胎的发育[42]。Abdel-Halim等[43]研究发现,纳米氧化锌影响牛胚胎的发育。
5 不同添加形式的锌添加效果比较不同形式的锌源有着不同的生物效价,因此有必要探讨反刍动物适宜的添加锌源。有研究发现,添加锌源的化学形态对公牛平均干物质采食量、生长速率和饲料转化率均无显著影响,但是与无机锌相比,有机锌化合物可以改善公牛牛蹄健康[44]。Wang等[45]发现,不同锌源对奶牛采食量、乳成分、体细胞数无显著影响,但是螯合能力强的锌比螯合能力适度的锌以及ZnSO4在提高瘤胃发酵、体液免疫反应以及提高泌乳奶牛产奶量方面更有效。Spears[46]对反刍动物锌的来源进行了研究,2种来源的锌有效性无显著差异,Zn-Met喂养的羔羊对锌的表观吸收和ZnO也相似,但Zn-Met中锌的保留率较高,因为该组羔羊尿中锌的排泄倾向较低。Hassan等[47]研究发现,与添加Zn-Met相比,添加ZnSO4显著降低了抗氧化活性相关指标(谷胱甘肽、谷胱甘肽过氧化物酶、超氧化物歧化酶)。有研究发现,纳米氧化锌与传统锌源相比,不仅具有较高的生物学效价,而且具有促进生长、抗菌、免疫调节等多种作用[48]。Kessler等[49]研究发现,红荷斯坦杂交公牛添加不同形式的锌源对平均干物质采食量、生长速率和饲料转化率均无显著影响,Montano等[50]也研究发现,不同锌源对荷斯坦牛生长性能无显著影响。表 2为不同添加形式的锌比较结果,大部分研究认为有机锌生物学效价高于无机形式的锌。
有些研究认为,有机锌的生物学效价高于无机锌,有机形式的锌具有更高的吸收率,粪中残留的锌显著减少,稳定性较好,抗干扰性强。但是到目前为止有机锌的作用机理依然是不甚清楚。尽管人们从不同角度研究了有机锌对家畜的促生长作用,但是各研究结果之间依然存在差异[57],还需要进一步研究。
6 反刍动物锌需要量 6.1 绵羊和山羊Chhabra等[58]研究发现,山羊饲喂锌含量为15 mg/kg的小麦秸秆171~200 d,山羊出现了锌缺乏的临床症状。Jia等[59]研究发现,非哺乳期山羊的锌需要量和牛、羊对锌的需要量一致,为20 mg/kg,绒山羊在产绒期间饲喂锌含量为22 mg/kg的饲粮,提高了绒山羊的生长速率。NRC(1985)营养需要量中,绵羊的锌需要量为20~30 mg/kg, 最大耐受量为750 mg/kg[60]。Masters等[61]研究发现,缺乏锌的母羊在每只出生的羔羊体内沉积了大约63 mg的锌, 这表明发育中的胎儿积累的锌相当于母羊饲粮总摄入量的35%。王洪荣等[62]使用综合法确定绒山羊产绒期常规饲粮锌的需要量为50 mg/kg。金亚倩等[63]研究发现35~50 kg杜泊×晋中绵羊F1代公羔锌净生长需要量为19.57~20.45 mg/kg BW。Ji等[64]通过屠宰试验发现,公羊羔生长所需锌为23.2~23.4 mg/kg。Zhang等[65]通过比较屠宰试验发现,绵羊生长所需的锌为34.21~35.80 mg/kg。从以上研究可以得出羊的锌需要量为20~50 mg/kg。
6.2 奶牛从表 3中可以看出,体重为650 kg、产奶量为40 kg/d的奶牛其锌需要量为40~63 mg/kg DM。
Volden[68]研究发现,生长肉牛锌需要量为30 mg/kg。Freer等[69]研究发现,小的肉牛锌需要量为30 mg/kg。Ott等[17]发现生长牛饲喂900 mg/kg锌时出现了锌中毒,表现为肝铜及日增重下降,而当饲粮锌含量提高至1 700~2 100 mg/kg时,出现异嗜症状。Graham等[70]研究认为,当小牛饲粮过度添加锌(1.5~2.0 g/d),累积剂量为30~66 g时会导致死亡。Jenkins等[71]研究认为,当小牛添加锌含量超过700 mg/kg时,其生产性能会下降。Miller等[72]研究认为,当饲喂2 000 mg/kg锌时奶牛产奶量和采食量均有所下降,饲喂1 000 mg/kg锌时奶牛无不良影响。李万栋[73]在通过体外试验发现,适合于牦牛瘤胃发酵的锌的添加形式为羟基蛋氨酸锌,添加量为30 mg/kg时最有利于瘤胃发酵,最适宜的添加量为20~40 mg/kg。
7 小结综上所述,锌对反刍动物有着非常重要的作用,锌广泛参与多种生物学过程,在酶催化、基因调控和大分子稳定性方面发挥着尤为重要的作用[74]。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,从而促进反刍动物生长、提高繁殖性能、改善机体免疫力、防治疫病等。目前对于锌的应用方面研究较多,但是相关机理研究较少,还需要从以下3个方面着手:1)需要进一步研究锌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)研究锌的添加形式,比如有机锌和螯合形式的锌对反刍动物的作用,以此提高锌的利用率。3)研究添加锌对动物健康的影响。
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