动物营养学报 >

2023 , Vol. 35 >Issue 4: 2129 - 2142

DOI: https://doi.org/10.12418/CJAN2023.201

综述

枸杞多糖的生理功能及其在动物生产中的应用

  • 刘杰 , 1, 2, 3 ,
  • 李柏珍 2 ,
  • 伍树松 2 ,
  • 张桂杰 , 1, 3, *
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  • 1 宁夏大学动物科学系,银川 750021
  • 2 湖南农业大学动物科学技术学院,长沙 410128
  • 3 宁夏草牧业工程技术研究中心,银川 750021
*张桂杰,教授,博士生导师,E-mail:

刘 杰(1997—),男,湖南邵阳人,硕士研究生,从事动物营养与饲料科学研究。E-mail:

Copy editor: 菅景颖

收稿日期: 2022-10-07

  网络出版日期: 2023-04-12

基金资助

宁夏回族自治区自然科学基金(2020AAC03079)

宁夏高等学校一流学科建设(草学学科)资助项目(NXYLXK2017A01)

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Physiological Functions of Lycium barbarum Polysaccharide and Its Application in Animal Production

  • LIU Jie , 1, 2, 3 ,
  • LI Baizhen 2 ,
  • WU Shusong 2 ,
  • ZHANG Guijie , 1, 3, *
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  • 1 Department of Animal Science, Ningxia University, Yinchuan 750021, China
  • 2 College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China
  • 3 Ningxia Grassland and Animal Husbandry Engineering Technology Research Center, Yinchuan 750021, China
*professor, E-mail:

Received date: 2022-10-07

  Online published: 2023-04-12

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摘要

随着我国饲料“禁抗”时代的来临,天然的植物提取物逐渐成为一种新型替代抗生素的饲料添加剂。枸杞多糖是从枸杞及其副产物中提取出来的植物多糖类物质,主要由甘露糖、葡萄糖、半乳糖、阿拉伯糖、半乳糖醛酸和鼠李糖等组成,具有抗氧化、抗肿瘤以及提高机体免疫力等多种生理功能,是枸杞及其副产物的主要活性成分。本文综述了枸杞多糖的主要制备工艺、生理功能和作用机制及其在动物生产中应用的研究进展,旨在为枸杞多糖作为新型饲料添加剂在动物生产中的应用提供参考。

关键词： 枸杞多糖; 生理功能; 作用机制; 动物生产

本文引用格式

刘杰 , 李柏珍 , 伍树松 , 张桂杰 . 枸杞多糖的生理功能及其在动物生产中的应用[J]. 动物营养学报, 2023 , 35(4) : 2129 -2142 . DOI: 10.12418/CJAN2023.201

Abstract

With the advent of the era of “Prohibition of Antibiotics” comes to feed in China, the natural plant extracts have gradually become a new type of feed additive to replace antibiotics. Lycium barbarum polysaccharides (LBP), plant polysaccharides extracted from Lycium barbarum and its by-products, mainly consists of mannose, glucose, galactose, arabinose, galacturonic acid and rhamnose and so on. LBP has various physiological functions such as anti-oxidation, anti-tumor and immunity improvement. It is also the main active components of Lycium barbarum and its by-products. This paper reviewed the main preparation technologies, physiological functions, action mechanisms of LBP and its research progress in animal production, aiming to provide theoretical reference for the application of LBP as a new type of feed additive in animal production.

Key words： Lycium barbarum polysaccharide; physiological function; action mechanism; animal production

饲用抗生素能够促进动物健康成长,但长期滥用抗生素会造成畜产品抗生素残留、产生耐药菌等问题,严重的危害人体健康和破坏生态环境。研究和开发新型替抗产品已成为饲料行业发展的必然趋势。近年,越来越多的研究表明,植物提取物具有抗氧化、调节免疫、促进动物生长等功效,已成为新型饲料添加剂的研究热点[1-2]。枸杞(Lycium chinense Miller)是我国传统的药食同源物种,属茄科枸杞属植物,资源丰富,广泛分布在我国半干旱和干旱地区。枸杞营养丰富,并含有多糖、黄酮、花青素等多种活性物质,而多糖是其主要的活性成分。枸杞多糖(Lycium barbarum polysaccharides,LBP),具有提高动物生产性能、改善产品品质、提高抗病能力、改善肠道菌群和提高抗氧化能力等作用(图1)[3-4],是一种无污染、无残留的天然绿色新型饲料添加剂。本文综述了枸杞多糖的主要制备工艺、生理功能和作用机制,以及在动物生产中应用的研究进展,为枸杞多糖在绿色健康养殖应用和推广方面提供参考依据。
图1 枸杞多糖的来源、萃取、功能和应用

Fig.1 Source, extraction, function and application of Lycium barbarum polysaccharide[3⇓-5]

1 枸杞多糖的概述

1.1 枸杞多糖的来源及结构

枸杞,又名天精,是一种茄科落叶灌木的中草药,其根、茎、叶、果都具有丰富的医药价值和营养作用,在我国有着2 000多年的栽种历史[5]。枸杞种植分布广泛,主要集中在干旱和半干旱地区,我国枸杞属品种有9个,其中种7个、变种2个(表1)[6-7],主产区分布在宁夏、新疆、河北等地,种植面积达82 000 hm2,年产浆果约95 000 t,是全球枸杞种植面积和产量最大的国家[8]。枸杞含有丰富的多糖、黄酮、花青素等活性物质,而多糖是其主要的活性成分;枸杞副产物中的枸杞枝叶、残次果及其加工产生的残渣中均含有丰富的枸杞多糖[9-10]。枸杞多糖是一种易溶于水,可以与蛋白质或多肽类结合的酸性复杂多糖,具有抗氧化、抗肿瘤、调节免疫、降低血糖和提高动物产品质量等诸多功效。
表1 我国枸杞属品种及多糖含量

Table 1 Varieties and polysaccharide content of Lycium in China[6-7]

项目
Items		 名称
Name		 主产区
Main production area		 多糖含量
Polysaccharides content/(mg/g)		 备注
Note
种Species
1 枸杞 陕西、山西、河北 12.36±0.05
2 宁夏枸杞 宁夏、新疆、内蒙古、陕西、山西、河北 13.35±0.06
3 黑果枸杞 新疆、西藏 6.86±0.11
4 截萼枸杞 山西、内蒙古、甘肃 12.88±0.06
5 新疆枸杞 新疆 13.30±0.25
6 柱筒枸杞 新疆 9.98±0.09
7 云南枸杞 云南 14.43±0.12
变种Variant
1 北方枸杞 河北、山西、陕西 10.92±0.07 为枸杞变种
2 黄果枸杞 宁夏 9.69±0.42 为宁夏枸杞变种
枸杞多糖在结构上属于高分子杂多糖,主要由糖类、糖醛酸类和蛋白质类组成,其中糖类主要由甘露糖、葡萄糖、半乳糖、阿拉伯糖、半乳糖醛酸和鼠李糖等组成(图2)[10]。枸杞多糖在空间结构上表现为侧链长、分支多和整体松散的状态,因此分子结构式不具有明确性,也没有固定的组成比例(表2)[11]。目前,研究报道中枸杞多糖的主链主要有-[-α-D-Xyl]2(1,2)-[-α-D-Glc]18-(1,4)-[-α-Man]3-(1,2)-[-α-L-Rha]3-(1,4)-(-β-D-GalN)1-(1,6)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,6)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,4)-[-β-D-GalN]1-(1,6)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,4)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,6)-[-β-D-GalN]1-(1,6)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,4)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,4)-[-β-D-GalN]1-(1,4)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,4)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,4)-[-β-D-GalN]1-(1,6)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,6)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,4)-[-β-D-GalN]1-(1,6)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,4)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,6)-[-β-D-GalN]1-(1,6)-、[-α-D-Xyl]2-(1,2)-[-α-D-Glc]18-(1,4)-[-α-D-Man]3-(1,2)-[-α-L-Rha]3-(1,6)-[-β-D-GalN]1-(1,4)-等8种[12],侧链主要有1,4-β-D-GlcpA、β-D-Galp、1,6-β-D-Galp、α-L-Araf、β-L-Araf、1,5-α-L-Araf和β-L-Rhap等7种[13]。
图2 枸杞多糖的主要糖类结构

Fig.2 Structure of main carbohydrates of Lycium barbarum polysaccharide[10-11]

表2 枸杞多糖的单糖组分及结构

Table 2 Lycium barbarum polysaccharide monosaccharide composition and structure

方法
Methods		 分子质量
Molecular
weight/kDa		 单糖组成
Monosaccharide
composition		 摩尔比
Molar ratio		 参考文献
References
柱前衍生化HPLC法
Pre-column derivatization HPLC method		 4.92 甘露糖、鼠李糖、葡萄糖、
D-半乳糖胺、D-木糖		 5.52∶5.11∶28.06∶
1.00∶1.70		 [12,14]
凝胶渗透色谱法
Gel permeation chromatography		 45 葡萄糖、半乳糖、
阿拉伯糖、鼠李糖		 1.4∶49.8∶47.8∶1.2 [15]
DEAE SepharoseTM Fast Flow &
Sephacryl S-300 HR		 80 鼠李糖、阿拉伯糖、
半乳糖、乳糖醛酸		 3.13∶53.55∶39.37∶3.95 [16]
气相色谱法
Gas chromatography		 31.25 木糖、阿拉伯糖、半乳糖、
甘露糖、葡萄糖、鼠李糖		 1.74∶0.57∶4.54∶3.47∶
2.94∶1.00		 [17]

1.2 枸杞多糖的制备工艺

目前报道的枸杞多糖的提取方法众多,主要分为溶剂浸提法、超声波辅助提取法、超临界流体萃取法、微波辅助提取法、超高压提取法、高速剪切技术辅助提取法和酶解法等(表3)。上述提取制备工艺各有差异,溶剂浸提法操作简单、成本低,但提取时间长、易受外界环境影响;超声波辅助提取法和微波辅助提取法提取时间短、效率高,但容易破坏多糖结构,改变其活性物质;超临界流体萃取法和高速剪切技术辅助提取法提取效率高、纯度高,但设备价格昂贵、成本高;酶解法提取速度快、环保高效,但容易受温度和浓度的影响。
表3 枸杞多糖的提取方法

Table 3 Extraction method of Lycium barbarum polysaccharide

提取方法
Extraction methods		 最优工艺
Optimal process		 提取率
Extraction rate/%		 参考文献
References
溶剂浸提法
Solvent leaching method		 料液比1∶30(g/mL),
时间2 h,温度50 ℃		 4.88 [18]
超声波辅助提取法
Ultrasonic-assisted extraction method		 料液比1∶20(g/mL),
时间40 min,温度60 ℃,超声功率180 W		 9.19 [19]
超临界流体萃取法
Supercritical fluid extraction method		 二氧化碳流量25 L/h,时间2 h,
温度45 ℃,压力10 MPa		 18.59 [20]
超高压提取法
UHP extraction method		 料液比1∶15(g/mL),
压力460 MPa,时间17 min		 12.1 [21]
微波辅助提取法
Microwave-assisted extraction method		 料液比1∶26(g/mL),
时间1.8 min,微波功率300 W		 9.57 [22]
提取方法
Extraction methods		 最优工艺
Optimal process		 提取率
Extraction rate/%		 参考文献
References
高速剪切技术辅助提取法
High-speed shear technology-
assisted extraction method		 料液比1∶30(g/mL),温度60 ℃,剪切速率
8 000 r/min, 4 min后60 ℃水浴中加热提取,时间2 h		 24.79 [23]
酶解法
Enzymatic digestion method		 酶浓度0.3%,时间140 min,温度49.6 ℃ 13 [24]
回应面法
Response surface method		 料液比1∶38(g/mL),时间80 min,温度73 ℃,
超声功率185 W		 12.54 [25]

2 枸杞多糖的生理功能及其作用机制

2.1 抗氧化

动物体内发生生化反应时,会导致原子和基团缺乏电子,从而形成化学性质活泼、具有强烈氧化作用的自由基[26]。当动物机体自由基动态平衡遭到破坏后,会过量产生自由基,对动物机体大分子物质造成结构性损坏,是诱发疾病和衰老的原因之一[27]。抗氧化剂具有抑制扩散和消除自由基的功能,可以保护组织细胞免受伤害,维持机体正常功能[28]。枸杞多糖具有较强的抗氧化作用,其抗氧化功能已被诸多研究所证实[29]。在体外抗氧化性试验中,枸杞多糖对ABTS自由基、超氧阴离子自由基( O 2 -·)、羟自由基(·OH)和二苯代苦味酰基自由基(DPPH)有很好的抗氧化效果[30-31]。在体内的抗氧化性试验中,枸杞多糖对超氧化物歧化酶(SOD)的活性有明显提升,可以进一步降低自由基带来的多不饱和脂肪酸过氧化,从而降低线粒体膜等生物膜受到的自由基的损害[32-33]。因此,枸杞多糖可作为抗氧化剂应用于动物生产中。
研究表明,枸杞多糖可以通过转录因子核因子E2相关因子2(Nrf2)/抗氧化反应元件(ARE)通路调控抗氧化功能[34]。Nrf2是氧化应激反应的最主要因子,在非应激状态下与调控因子Kelch样环氧氯丙烷相关蛋白1(Keap1)相结合,以低含量的形式存在于细胞质中,当受到氧化应激刺激时,会迅速与Keap1解离进入到细胞核中,与ARE结合共同调控SOD、血红素氧合酶-1(HO-1)、过氧化氢酶(CAT)、谷胱甘肽过氧化物酶(GSH-Px)和还原型烟酰胺腺嘌呤二核苷酸磷酸(NADPH):醌氧化还原酶1(NQO1)等抗氧化酶的表达,从而增强抗氧化能力[35-36]。枸杞多糖通过磷脂酰肌醇3-激酶(PI3K)/丝氨酸-苏氨酸蛋白激酶(Akt)通路启动Nrf2,使SOD和GSH-Px活性升高,降低活性氧(ROS)和过氧化脂(LPO)的含量,起到抗氧化作用[37]。Varoni等[38]研究发现,在大鼠饲粮中添加300 mg/kg的枸杞多糖可以显著提高血浆中GSH-Px活性,降低MDA含量,提高总抗氧化能力。此外,枸杞多糖还可以通过下调诱导性一氧化氮合成酶(iNOS)mRNA表达,调控其抗氧化功能[39]。一氧化氮(NO)具有神经保护和神经毒性作用,它的形成依赖iNOS,浓度过高时会产生大量的·OH和二氧化氮(NO2)自由基,造成氧化应激损伤[40]。抑制iNOS mRNA转录可以减少NO产生,提升抗氧化能力,减轻氧化应激损伤。枸杞多糖通过抑制NADPH的产生,减少活性氧基团生成,提高SOD和GSH-Px的活性,降低MDA的含量[41]。Zhao等[42]研究发现,枸杞多糖可以促进神经元细胞表面释放SOD,提高GSH-Px活性,降低氧化应激损伤,起到保护神经的作用。

2.2 抗肿瘤

机体在受到刺激后,氧化代谢紊乱造成DNA损伤,信号通路促进血管内皮生长因子加速表达是诱发肿瘤原因之一[43-44]。枸杞多糖可以减少断裂DNA氧化,调节自身免疫系统,阻止肿瘤细胞继续生长,减少肿瘤细胞膜的流动性,诱导肿瘤细胞凋亡[45]。在动物机体内,枸杞多糖可以启动肿瘤坏死因子-α(TNF-α)基因,提高其表达效率,诱导癌细胞凋亡,达到抗肿瘤的作用[46-47]。枸杞多糖在体外诱导小鼠肝癌细胞(H22)存活率和荷瘤小鼠抗肿瘤活性与不同分子质量的关系显示,枸杞多糖对H22有明显的抑制作用,当其分子质量在40~350 kDa区间时可以显著抑制肿瘤生长和降低肿瘤重量[48]。Western印迹试验显示枸杞多糖对半胱氨酸蛋白酶8(Caspase-8)和Bcl-2蛋白的表达有抑制作用,对Bax蛋白的表达有促进作用[49]。
研究发现,枸杞多糖可以通过核因子-κB(NF-κB)信号通路调节炎症反应[50]。NF-κB是一种可以调控炎症反应的核转录因子,正常情况下,NF-κB与细胞质中的抑制蛋白结合成无活性的NF-κB抑制蛋白(IκB),当受到机体刺激时NF-κB被启动,NF-κB启动途径分为典型途径和非典型途径[51-52]。NF-κB典型途径,在受到刺激后,转化生长因子β(TGEβ)迅速启动转化生长因子激酶1(TAK1),而TAK1启动IκB激酶(IKK)复合物,使三聚体中IκB蛋白磷酸化,解离和释放出NF-κB p50蛋白(p50)、NF-κB p65蛋白(p65)和NF-κB c-Rel蛋白(c-Rel)等;NF-κB非典型途径,主要是作用于NF-κB p100蛋白(p100)上,合成NF-κB p52蛋白(p52)和NF-κB RelB蛋白(RelB)[53]。枸杞多糖可以通过NF-κB典型途径调控炎症反应,增加细胞质中的p65、NF-κB亚基和抑制蛋白IκB-α的蛋白表达,减少NF-κB转入细胞核中,抑制促炎反应,达到抗肿瘤效果[54]。Qi等[55]在临床研究中发现,枸杞多糖还可以通过启动蛋白激酶细胞外信号调节激酶(ERK)和p53的表达,抑制人乳腺癌细胞(MCF-7)的凋亡。Mao等[56]在研究结肠癌细胞生长效应和机制时发现,1 000 mg/L的枸杞多糖可以中断细胞周期,显著抑制结肠腺癌细胞(SW-480)的生长。

2.3 免疫调节

免疫系统是机体重要的防御系统,由免疫器官、免疫细胞和免疫分子构成。免疫细胞在机体分布广泛,包括淋巴细胞、巨噬细胞和树突状细胞等,主要参与体内免疫过程的细胞是T淋巴细胞和B淋巴细胞[57]。T淋巴细胞是机体重要免疫细胞之一,而枸杞多糖能够增加T淋巴细胞总量及TH细胞亚群占比,提高淋巴细胞转化率,加快机体免疫恢复和提升[58]。B淋巴细胞在体液免疫中发挥极为重要的作用,枸杞多糖可以提高B淋巴细胞数量,提高抗体水平,从而提升体液免疫的效果[59-60]。单核巨噬细胞是一种具有吞噬杀伤病原体、抗原呈递等免疫功能的细胞,枸杞多糖可以使单核巨噬细胞活跃度增强和吞噬指数上升[61-62]。研究显示,枸杞多糖可作用于各类型的单核巨噬细胞,增加其NO的生成量,提升细胞内部溶菌酶与SOD的活性[63];自然杀伤细胞(NK细胞)是机体内一种非特异的免疫杀伤细胞,枸杞多糖可以提升NK细胞活性、数量和杀伤力[64]。
研究显示,枸杞多糖可以通过NF-κB和促分裂素原活化蛋白激酶(MAPK)信号通路调控机体免疫活性。转录因子NF-κB是免疫的中枢调节因子,其经典途径与非经典途径均对T淋巴细胞和B淋巴细胞发育活化起到重要作用[65]。在经典途径中,p65和p65亚基对CD4+CD25+调节性T细胞发育起到促进作用,可以增加CD8+T细胞的数量,当p65抑制或缺失,会减弱T淋巴细胞免疫活性[66]。在非经典途径中,当T淋巴细胞受体(TCR)对抗原识别或B淋巴细胞受体(BCR)激活后,诱导膜相关鸟苷酸激酶蛋白1(CARMA1)、黏膜相关淋巴组织蛋白1(MALT1)和B细胞淋巴瘤因子10(BCL10)相互作用,随后被蛋白激酶磷酸化和泛素化后形成CARMA1-BCL10-MALT1(CBM)复合物,CBM复合物启动NF-κB信号通路,促进T淋巴细胞存活和增殖[67]。在NK细胞中的细胞因子和趋化因子启动过程中,NF-κB通过促进细胞毒性介质表达来控制NK细胞溶解功能[68]。枸杞多糖通过NF-κB p65核转位促进T淋巴细胞和B淋巴细胞增殖,提高CD4+、CD8+淋巴细胞活性,增加血清中免疫球蛋白M(IgM)和免疫球蛋白G(IgG)含量,从而提高动物机体的免疫力[69-70]。Xu等[54]研究发现,NF-κB和MAPK复合体免疫分子调控脂多糖(LPS)诱导牛乳腺上皮细胞炎症反应中,枸杞多糖可以通过调控NF-κB和MAPK信号启动过氧化物酶体增殖物激活受体γ(PPARγ)来抑制疾病的发生。

2.4 其他功能

枸杞多糖还具有保护生殖功能、治疗骨质疏松、降压降脂和保肝护肝等作用。枸杞多糖对生殖功能的保护主要通过保护生精细胞及调节体内性激素水平,使生精细胞Ⅰ型胶原α1(Col1α1)和Ⅰ型胶原α2(Col1α2)凋亡蛋白的表达显著下降,增加生精干细胞的数量,提高附睾精子质量及生精数量,维持睾丸生精能力和雄性性特征表现[71]。同时,枸杞多糖还可以修复卵巢损伤,提升卵泡刺激素、黄体生成素、睾酮和雌二醇水平[72-73]。在治疗骨质疏松方面,枸杞多糖通过降低破骨细胞的活性,来降低机体骨胶原的代谢速度,防止过度转换,减少钙、磷和镁等矿物质流失程度,对骨质疏松的骨代谢起调节作用[74-75]。在降压降脂和保肝护肝等方面,枸杞多糖可以改善人肝肿瘤细胞(Hep G2)胰岛素水平,通过血清白蛋白和总蛋白含量提升,抑制谷丙转氨酶(GPT)、谷草转氨酶(GOT)和γ-谷氨酰转肽酶(γ-GT)的活性[76-77]。另外,枸杞多糖还可以增加肝脏内基质金属蛋白酶-9(MMP-9)的活性,减少基质金属蛋白酶抑制剂-1(TIMP-1)的含量,增强降解细胞间质的作用,延缓肝纤维化进程[78-79]。
综上所述,枸杞多糖具有抗氧化、抗肿瘤、提高免疫力和保肝护肝等功能,其作用机制如图3所示。
图3 枸杞多糖的作用机制

NADPH:还原型烟酰胺腺嘌呤二核苷酸磷酸 reduced nicotinamide adenine dinucleotide phosphate;NOx2:NADPH氧化酶2 NADPH oxidase 2;NOx4:NADPH氧化酶4 NADPH oxidase 4;NOx5:NADPH氧化酶5 NADPH oxidase 5;ROS:活性氧 reactive oxygen;SOD:超氧化物歧化酶 superoxide dismutase;GSH-Px:谷胱甘肽过氧化物酶 glutathione peroxidase;Nrf2:核因子E2相关因子2 nuclear factor E2 related factor 2;PI3K/Akt:磷脂酰肌醇3-激酶/丝氨酸-苏氨酸蛋白激酶 phosphatidylinositol 3-kinase/serine-threonine protein kinase;IgM:免疫球蛋白M immunoglobulin M;IgG:免疫球蛋白G immunoglobulin G;p65 subunit:p65亚基;NF-κB subunit:核因子-κB亚基;IκB-α protein: IκB-α蛋白;ERK:细胞外调节蛋白激酶 extracelluar regulated protein kinase。

Fig.3 Mechanism of action of Lycium barbarum polysaccharide[34,37,54-55,69-70,78-79]

3 枸杞多糖在动物生产中的应用

3.1 在猪生产中的应用

Yin等[80]研究发现,在断奶仔猪的饲粮中添加4 000 mg/kg枸杞多糖可以明显提高平均日增重,降低腹泻率和料重比,显著增加血清中IgG、IgM、白细胞介素(IL)-10、IL-2和TNF-α的含量;同时,还可以增加十二指肠和空肠的绒毛高度,降低隐窝深度,增加绒毛高度与隐窝深度的比值,改善肠道内容物菌群结构,增加乳杆菌属和粪杆菌属的丰度,降低肠杆菌科和肠球菌科的丰度。Chen等[81]研究也表明,枸杞多糖可以提高血清中SOD、CAT和GSH-Px活性,降低MDA含量;增加回肠和盲肠中的乳酸杆菌、双歧杆菌和双歧杆菌数量,减少大肠杆菌和厚壁菌数量,改善肠道环境,提高仔猪生长性能。黄红卫等[82]在饲粮中分别添加0、5%、10%的枸杞渣,结果显示,5%枸杞渣为最佳添加量,可以显著提高育肥猪的日增重、屠宰率、瘦肉率和眼肌面积,增加系水力,改善肉品质。Yang等[83]研究了枸杞对改善公猪精液品质的影响,结果表明,按每100 kg体重每天在饲粮中添加40 g干枸杞,可以显著提高精子的进行性运动,改善精子活力,增加射精总精子数,提高精子浓度,降低精子头部去除率、尾部缺陷率、颈部缺陷率、中段缺陷率和原生质液滴缺陷,提高精液品质。此外,冯妮等[84]在猪精液中分别加入2和8 mg/mL的枸杞多糖,结果显示,枸杞多糖可以显著提高获能精子存活率和顶体完整率,降低精子坏死率,提高SOD活性,降低MDA含量,保持精子完整形态。
综上所述,在猪生产中应用枸杞多糖能够增强免疫力,调整肠道菌群结构,提高精液品质,改善胴体品质并提高生产性能。

3.2 在家禽生产中的应用

Long等[85]在1日龄爱拔益加肉鸡饲粮中分别添加1 000、2 000、4 000 mg/kg的枸杞多糖,结果显示,2 000 mg/kg枸杞多糖可以显著提高肉鸡的平均日增重和平均日采食量,降低料重比,提高肠道消化酶活性,增加肠道淀粉酶、蛋白酶和脂肪酶活性,促进营养物质消化,提高肉鸡的生长性能;可以增加血清和肝脏中SOD和GSH-Px的活性,降低肝脏中MDA的含量,提高肉鸡的抗氧化性能;还可以显著提高血清中IgG、免疫球蛋白A(IgA)、IL-2、IL-4、IL-6、γ-干扰素(IFN-γ)和TNF-α的含量,提高肉鸡的抗病能力。王建东等[86]研究也表明,在肉鸡饲粮中添加2%枸杞多糖免疫增效剂可以显著增加血清IgG含量,提高肉鸡免疫器官发育,增强体液免疫能力。此外,李松桥等[87]研究发现,在黄羽肉鸡饲粮中加入4%枸杞渣与甘草混合添加剂,能够显著提高肉鸡的胸肌率,改善胴体颜色和肌肉嫩度。张海燕等[88]在蛋鸡饲粮中分别添加0、50、100 mg/kg的枸杞多糖,结果表明,50 mg/kg枸杞多糖能够显著提高平均产蛋重,降低料重比,增强蛋壳厚度,提高蛋鸡的产蛋性能和蛋品质;可以增强胸腺、法氏囊和脾脏重量和活性,调节蛋鸡的免疫功能,显著增加蛋鸡血液中CD3+、CD4+、CD8+的含量,提高蛋鸡的抗病原体能力。Maisto等[89]研究发现,在蛋鸡饲粮中加入5%的辣木叶和10%的枸杞可以提高蛋黄中叶黄素、角黄素和隐黄质的含量,降低蛋黄中胆固醇含量,从而达到改善蛋品质功能。此外,韩占兵等[90]研究也发现,在蛋鸡饲粮中添加5%枸杞渣可以提高蛋鸡产蛋率,降低蛋重和料蛋比,改善蛋黄颜色,提高蛋鸡的生产性能和蛋品质。
综上所述,在家禽生产中应用枸杞多糖可以显著提高家禽的生长性能,提高机体免疫能力,改善产品品质并提高生产性能。

3.3 在反刍动物生产中的应用

郑梦荷等[91]在犊牛饲粮中分别添加0、5、10、15 g/d的枸杞多糖,结果显示,10 g/d枸杞多糖可以显著增加犊牛的平均日增重、平均日采食量,降低料重比,提高生长性能;可以改善犊牛蛋白质、脂肪和糖代谢,增加犊牛血清中血清总蛋白、总胆固醇、甘油三酯和葡萄糖含量;还可以促进激素分泌,增加犊牛血清中类胰岛素一号增长因子、生长激素、三碘甲状腺原氨酸、甲状腺激素和胰岛素水平,改善脂代谢和促进生长。马吉锋等[92]在肉牛基础饲粮中添加500 g/kg枸杞多糖增效剂,结果发现,枸杞增效剂可以提高肉牛血液中免疫球蛋白含量、淋巴细胞和白细胞比例,进一步促进细胞因子分泌。此外,侯鹏霞等[93]的研究发现,在肥育牛饲粮中添加2.95 kg/(d·头)的枸杞枝条发酵饲料可以提高育肥牛的平均日增重,增加血清中IgA、IgG和IgM含量,但过量添加枸杞枝条会降低血清中IL-2、IL-4、IL-13的含量。王建东等[94]在围产期奶牛饲粮中添加3 g/(d·头)的枸杞多糖免疫增效剂,结果显示,饲喂枸杞多糖免疫增效剂后可以显著提高奶牛围产期外周血免疫球蛋白含量,增加血清中IgG、IgA、IgM的含量;可以提高细胞因子IL-2、IL-13、IL-17、IL-19、IFN-α和IFN-γ的含量;还可以提高免疫力,增加白细胞和淋巴细胞的比例。剧浩等[95]在羔羊饲粮中分别添加0、0.1%、0.3%、0.5%的枸杞多糖,结果显示,0.3%枸杞多糖添加量为最佳,可以显著提高羔羊的干物质、粗蛋白质、有机物和氮表观消化率,促进羔羊生长;可以提高羔羊血清中IgA和IgM的含量,增强羔羊免疫力。李庆敏等[96]研究也发现,在育肥滩羊饲粮中添加5%的枸杞渣可以显著提高滩羊的干物质采食量、氮的生物学价值和血清中IgG和IgM的含量,增强机体抗氧化能力和免疫力。此外,Ren等[97]研究了枸杞多糖与海带多糖对绒山羊精液品质的影响,结果表明,4 mg/mL枸杞多糖与1 mg/mL海带多糖可以增强绒山羊精子活力,改善线粒体活性以及膜和顶体的完整性,同时可以提高SOD、CAT和GSH-Px活性,增强对精子的低温保护,提高精液品质。
综上所述,在反刍动物生产中应用枸杞多糖可以提高消化性能,改善精液品质,提高机体免疫力和生产性能。

3.4 在水产动物生产中的应用

谭连杰[98]在卵形鲳鲹饲料中分别添加0、0.05%、0.1%、0.2%、0.4%和0.8%的枸杞多糖,结果表明,0.1%枸杞多糖是最佳添加量,可以显著提高血清中SOD、GSH-Px活性和总抗氧化能力,促进体内蛋白质和脂代谢,改善肌肉品质;可以显著增加血清中补体3(C3)、补体4(C4)含量和溶菌酶活性,提高非特异性免疫力;还可以显著提高血清中高密度脂蛋白含量,显著降低血清中甘油三酯、胆固醇和低密度脂蛋白含量,改善脂代谢。Zhang等[99]研究发现,在罗非鱼饲料中添加2 000 mg/kg枸杞多糖可以显著增强血清中C3含量和IL-1β基因的表达,减少脾脏细胞凋零,提高非特异性免疫力、生长速度以及饲料转化效率。Liu等[100]研究也表明,在四氯化碳诱导后的鲤鱼饲料中添加1%的枸杞多糖,可以显著降低鲤鱼血清中GOT、GPT、乳酸脱氢酶和抗氧化酶的活性,同时可以抑制MDA生产量的增加,增强抗氧化能力。此外,Tang等[101]研究发现,在壬基酚处理后的斑马鱼饲料中添加0.5 mg/L枸杞多糖,可以显著增加血清中的SOD活性,降低MDA含量,减少生精细胞凋亡,上调11β-羟化酶活性,促进雄性激素,改善生殖功能,提高精子质量和数量。
综上所述,在水产动物生产中应用枸杞多糖可以增强抗病能力,改善产品品质,提高精液品质和生长性能。

4 小结

枸杞多糖通过Nrf2、ARE、NF-κB和MAPK等途径发挥抗氧化、抗肿瘤和调节免疫等生理功能,其在动物生产研究中展现出提高生产性能、改善产品品质、提高抗病能力、改善肠道菌群结构和提高精液品质等作用,具有广泛的开发利用价值和应用前景。目前,针对枸杞多糖的研究主要集中在细胞和小鼠上,在动物生产中应用的研究相对有限,特别是在猪、禽和反刍动物上的研究较少,不同物种、不同性别和不同阶段的适宜剂量尚不明确,且其在动物体内的生理功能作用机制也尚不完全清楚。同时,枸杞多糖的制备工艺研究主要集中在枸杞果,利用枸杞茎叶、残次果、枸杞渣等枸杞副产物的制备枸杞多糖的工艺研究较少,不利于在动物生产中推广应用。在今后的研究中应进一步深入探索枸杞多糖的作用机制,加强其在猪、鸡、牛、羊等动物生产应用中的研究,加大在枸杞副产物制备工艺方面的研究,为枸杞多糖作为一种绿色、安全、高效的饲料添加剂应用于动物生产提供新的理论依据。

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