动物营养学报 >

2023 , Vol. 35 >Issue 12: 7625 - 7632

DOI: https://doi.org/10.12418/CJAN2023.692

综述

低聚木糖的生理功能及其在畜牧生产中应用的研究进展

  • 韩佳佳 ,
  • 韩锐 ,
  • 付彤 ,
  • 苏传友 , *
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  • 河南农业大学动物科技学院,河南省家畜营养调控与生态养殖国际联合实验室,郑州 450046
* 苏传友,讲师,硕士生导师,E-mail:

韩佳佳(1998—),女,河南洛阳人,硕士研究生,从事动物营养与饲料资源开发利用的研究。E-mail:

Copy editor: 武海龙

收稿日期: 2023-06-12

  网络出版日期: 2023-12-11

基金资助

国家现代农业产业技术体系项目(CARS36)

收起

Research Progress on Physiological Function of Xylo-Oligosaccharide and Its Application in Livestock Production Production

  • HAN Jiajia ,
  • HAN Rui ,
  • FU Tong ,
  • SU Chuanyou , *
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  • International Joint Laboratory of Nutrition Regulation and Ecological Raising of Domestic Animal, College of Animal Science and Technology, Henan Agricultural University, Zhengzhou 450046, China
* lecturer, E-mail:

Received date: 2023-06-12

  Online published: 2023-12-11

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摘要

低聚木糖(XOS)是具有益生元活性的功能性低聚糖,主要来源于木质纤维素材料,如稻壳、玉米芯、秸秆、麦麸等农副产品。XOS在免疫调节、降血脂和调节肠道菌群等方面发挥着重要作用,已成为国内外备受关注的功能性低聚糖之一。本文旨在对XOS的结构、理化性质和生理功能进行系统概述,并讨论目前XOS在畜牧生产中的应用进展,以期为XOS在畜牧生产中的开发利用提供理论依据。

关键词： 低聚木糖; 理化性质; 生理功能; 畜牧生产

本文引用格式

韩佳佳 , 韩锐 , 付彤 , 苏传友 . 低聚木糖的生理功能及其在畜牧生产中应用的研究进展[J]. 动物营养学报, 2023 , 35(12) : 7625 -7632 . DOI: 10.12418/CJAN2023.692

Abstract

Xylo-oligosaccharide (XOS) is functional oligosaccharides with prebiotic activity, mainly derived from lignocellulosic materials, such as rice husk, corn cob, straw, wheat bran and other agricultural products. XOS plays an important role in immune regulation, lowering blood lipids and regulating intestinal flora, and has become one of the functional oligosaccharides that have attracted much attention at home and abroad. In this paper, the structure, physical and chemical properties and physiological functions of XOS are systematically summarized, and the application progress of XOS in animal husbandry production is discussed, in order to provide theoretical basis for the development and utilization of XOS in animal husbandry production.

Key words： xylo-oligosaccharide; physical and chemical properties; physiological function; animal husbandry production

低聚木糖(xylo-oligosaccharide,XOS)是一种难以被肠道消化吸收,具有特殊生理功能的低聚糖,也是一种广泛使用的益生元。国际益生菌和益生元协会(ISAPP)在其益生元定义的最新更新中将XOS确定为新型益生元低聚糖[1]。它可以促进体内有益菌的代谢和增殖。体外和体内研究表明,XOS经特定有益菌发酵后可产生益生元效应,选择性刺激双歧杆菌、乳酸杆菌等有益菌的生长和活性,具有显著的健康益处[2]。XOS作为近年来兴起的一种“超强双歧因子”,被广泛应用于食品、饮料和饲料等领域,已成为国内外备受关注的功能性低聚糖之一。因此,本文介绍了XOS的结构、理化性质和生理功能及其在畜牧生产中的应用,旨在为XOS的进一步发展和应用提供依据。

1 XOS的来源、基本结构及理化性质

1.1 XOS的基本结构及来源

XOS是由2~9个木糖分子通过β-1,4糖苷键连接而形成的低聚糖[3],分子质量一般为200~300 ku,主要包括木二糖、木三糖、木四糖,而木五糖、木六糖、木七糖含量相对较少。与α-链相反,β-1,4链使XOS耐水解,α-链则容易被消化酶消化[4]。据GB/T 35545—2017规定,XOS按照产品形态可分为糖浆(L型)和糖粉(P型);按XOS含量可分为XOS-95型(XOS占比超过95%,下同)、XOS-70型、XOS-35型和XOS-20型。
木聚糖是最常见的多糖半纤维素,而大多数XOS是由富含木聚糖的木质纤维素生物质降解制备的[5-6],其主要来源是玉米芯、秸秆、米糠、麦麸、稻壳、棉籽壳和其他农业废弃物。目前,提取XOS主要有3种方法:自水解、酸水解和酶水解[7⇓-9]。其中,自水解对获得高提取率和高纯度产品的设备要求很高,酸水解会因使用有害试剂而造成环境污染。酶水解因其高便利性、高效率、无污染而成为生产XOS最有前途的方法。然而,从稻壳、玉米芯、秸秆等谷物副产物中提取的XOS含有大量杂质,因此需要通过柱色谱、膜分离和树脂分离等方法进一步分离纯化。

1.2 XOS的理化性质

固体XOS为乳白色或淡黄色粉末,具有优异的物理和化学性能,例如高耐热性和耐酸性[10]。XOS在pH为2.5~8.0时都很稳定,在此pH范围内经100 ℃加热1 h几乎不分解,储存于-10 ℃时也不易冻结。同时,XOS很难被唾液、胃液、胰液和小肠酶液等分解[11],可直接进入大肠。XOS的甜度为蔗糖的40%~50%[12],且与蔗糖的甜味相似。XOS的黏度低于其他低聚糖,可以降低水分活度,提高水溶液中保水的能力[13]。

2 XOS的生理功能

2.1 调节肠道菌群

XOS不能被动物胃肠道消化和吸收,但它可以被肠道微生物代谢[14]。XOS刺激肠道内双歧杆菌等有益菌的生长和活性,进而代谢XOS以释放发酵副产物,如短链脂肪酸(SCFAs)[15]。SCFAs降低肠道的pH,形成肠道酸性环境,为细菌提供适宜的生存环境,有利于肠上皮细胞增殖,从而有利于维持肠道健康,提高肠道免疫力。
在选择性增殖有益菌的同时,XOS还可以有效抑制有害菌的生长繁殖。Moura等[16]发现,XOS可被多种双歧杆菌、乳酸杆菌和芽孢杆菌等有效利用,但不能被大肠杆菌、肠球菌和产气荚膜梭菌等有害细菌降解。其机制可能是双歧杆菌和其他有益细菌可以分泌阿拉伯酶和D-木糖苷酶将XOS水解成单糖,而有害细菌不能产生这些酶[17-18]。此外,XOS本身的结构特性也可以抑制有害细菌的生长[19⇓-21]。Lin等[22]研究发现,人类摄入XOS显著增加了粪便中双歧杆菌属、乳酸杆菌属的数量,并降低了产气荚膜梭菌的数量,从而有效降低了有害菌引起的肠道疾病的发生率。

2.2 调节免疫功能

XOS可以影响免疫系统[4]。Nabarlatz等[23]报道,杏仁壳XOS显示出直接的免疫调节活性。而有益菌代谢XOS产生的SCFAs在调节免疫因子和增强免疫力方面也发挥着重要作用。一方面,SCFAs和G蛋白偶联受体(GPR)之间存在很强的关系,SCFAs可以激活肠上皮细胞上的GPR41和GPR43,以便通过趋化因子和细胞因子快速分泌刺激免疫反应,这些途径介导小鼠的保护性免疫和组织炎症[24]。另一方面,SCFAs调节由T细胞和B细胞介导的适应性免疫,促进肠黏膜和全身抗体反应,从而提高抗病能力[25-26]。此外,Hansen等[27]认为,无论大鼠的微生物群如何,XOS都可以改善肠道屏障功能。然而,XOS调节免疫反应的功能是与调节微生物群有关,还是通过直接结合细胞受体尚不清楚。

2.3 调节脂质代谢

许多研究表明,XOS可以有效降低肥胖者的血脂水平。例如,Sheu等[28]发现连续摄入8 g/d木糖4周后,2型糖尿病患者的脂肪含量显著减少。XOS可以显著降低糖尿病大鼠的血浆胆固醇含量[29]。研究发现,XOS的主要成分之一膳食木二糖可以显著降低肥胖小鼠的血液总胆固醇(TC)和低密度脂蛋白胆固醇(LDL-C)含量[30]。
低聚糖的摄入会降低总生长素和酰化生长素含量。相反,酰化生长素含量的减少会减少食物摄入量,以改善肥胖和控制新陈代谢。研究发现,XOS对胆固醇含量的降低与SCFAs生成的增加有关,SCFAs可有效降低血浆TC含量[31]。尽管人们对XOS降低胆固醇含量的作用机制尚不完全清楚,但一些证据表明,XOS降低血液TC含量可能是通过重塑肠道微生物群组成和上调胆汁酸相关酶的表达介导的[30,32]。

2.4 调节机体抗氧化能力

功能性低聚糖可以通过减少活性氧(ROS)和丙二醛(MDA)的产生来缓解氧化应激,还可以通过促进机体产生抗氧化酶来提高机体的抗氧化能力。因此,功能性低聚糖可以作为抗氧化剂添加到畜禽饲料中,以缓解氧化应激,减少炎症的发生并保持机体健康。
研究表明,XOS表现出抗氧化和自由基清除能力[15],因此在生物医学领域具有潜在的应用前景[21,33]。在一定剂量范围内,XOS的自由基清除能力被证明是剂量依赖性的[34],这种潜力可能归因于酚类化合物的有效释放和氢原子从酚类化合物到自由基的转移[35]。Jagtap等[36]利用1,2-二苯基-2-苦肼(DPPH)自由基试验发现,在XOS浓度为6 mg/mL时,DPPH自由基的清除率为74%,此后随着XOS浓度增加,其抗氧化活性不再有明显变化。

2.5 其他有益作用

大量动物试验证明了XOS在预防龋齿、调节血糖、抗肿瘤、促进矿物质吸收等方面的有益作用[37⇓⇓-40]。XOS不能被口腔中的变形链球菌和其他细菌分解。摄入5%的XOS可以显著降低肥胖小鼠的血糖水平[15]。谷物中的XOS也可以有效改善哺乳动物的血糖水平[41]。XOS有助于SCFAs的产生,SCFAs具有显著的抗肿瘤作用,并且SCFAs的产生降低了肠道的pH,抑制了病原菌和有害菌的生长,从而降低了肿瘤的发病率[42]。当XOS和钙同时摄入时,钙的吸收率提高[43]。Kobayashi等[44]发现,缺铁性贫血小鼠摄入酸性XOS能够抑制肝脏中铁含量的下降,并且可以通过促进铁的吸收、抑制铁的排泄或提高铁的生物利用度来预防缺铁性贫血。同时,XOS在预防心血管疾病、动脉粥样硬化和肠道疾病等方面也发挥着重要作用[45]。

3 XOS在畜牧生产中的应用进展

3.1 在猪生产中的应用进展

仔猪断奶引起的腹泻是一种高度普遍的胃肠道疾病,会对仔猪的免疫系统产生负面影响并引发肠道功能障碍[46-47]。XOS作为益生元物质对维持宿主的肠道健康有积极作用。XOS可以通过改善肠道形态、增加紧密连接蛋白表达以及调节促炎细胞因子与抗炎细胞因子的比例来促进断奶仔猪的肠道健康[48]。Tang等[49]研究发现,XOS能降低促炎细胞因子(白细胞介素-6和白细胞介素-8)含量。研究表明,XOS优先刺激肠道内有益菌(如双歧杆菌和其他乳酸杆菌)的生长和活性,增强免疫功能,促进肠黏膜生长,同时增加仔猪肠道微生物群的多样性[50]。Su等[51]发现,在饲粮中添加0.25 g/kg的XOS可显著降低断奶仔猪腹泻的发生率。Yin等[52]研究了XOS对断奶仔猪肠道菌群的影响,发现XOS影响了乳酸杆菌、链球菌和土酸杆菌在属水平上的相对丰度,干扰素-γ含量的减少和闭锁小带蛋白-1表达的上调则证明了XOS能改善仔猪的炎症状况并增强肠道屏障。Liu等[53]研究发现,添加XOS能显著提高断奶仔猪的平均日增重,而且在整个试验期间,添加XOS组仔猪的平均日增重和饲料效率均高于未添加XOS组。
XOS在生长育肥猪方面的应用较少,可能是由于其对生长性能无显著影响[54-55]。不同剂量的XOS膳食补充剂不会影响猪的生长性能,而是通过增加肌肉粗蛋白质含量来提高猪肉的营养价值。然而,研究报道饲粮中添加200 mg/kg XOS可显著提高保育猪的生长性能[56-57]。Hou等[58]也发现,饲粮中添加0.04% XOS可显著提高仔猪的最终体重和平均日增重。这可能与猪的生长阶段有关,也可能与XOS的来源和剂量有关。

3.2 在反刍动物生产中的应用进展

目前,XOS的研究主要集中于幼龄反刍动物。研究显示,XOS能够降低犊牛、羔羊的腹泻率,提高其免疫力和生长性能。张军华等[59]将0.02% XOS加入羔羊奶中,发现XOS可显著降低羔羊腹泻的发生率,提高其免疫力。王喜明等[60]研究结果显示,XOS能使血清中尿素氮含量下降,对犊牛的腹泻有一定的抑制作用,同时可以降低白蛋白与球蛋白的比值,从而提高机体的免疫能力。
XOS促进动物生长的作用主要是通过选择性促进双歧杆菌等肠道有益菌群在消化道中定植,从而间接达到促生长效应等。王喜明等[61]发现,在XOS组中,乳酸杆菌的数量增加,而大肠杆菌数量呈下降趋势;试验末期,添加5 g/(d·头)XOS的犊牛平均日增重显著高于对照组。XOS在动物生产方面的研究结果表明,在饲粮中添加XOS能够提高营养物质的吸收率,从而有益于动物生产[62]。此外,XOS能提高奶牛采食量,这可能与其能有效提高动物饲料利用率有关。赵磊[63]研究发现,XOS能够对娟姗牛营养物质表观消化率、干物质采食量及产奶性能产生积极影响。同时,XOS可以降低娟姗牛瘤胃甲烷排放,缓解奶牛温室气体排放量。

3.3 在家禽生产中的应用进展

XOS可以改善肉鸡的生长性能,增强内分泌代谢和免疫功能[64]。Yang等[65]发现,XOS可提高肉鸡的生长性能,并在不同水平上改变肠道中微生物群的比例,特别是拟杆菌门和厚壁菌门中的微生物群。Ribeiro等[66]研究了XOS对肉仔鸡生长性能的影响,发现XOS可提高小麦饲粮的营养价值,并且动物生长性能的提高伴随着上消化道微生物群的转移。张伟彬等[67]发现,在饲粮中添加300 mg/kg XOS可以显著提高42日龄肉鸡体重,降低1~42日龄料重比。李玥[68]发现,XOS不仅可以提高肉鸡的生长性能,调节肠道微生物菌群多样性,还能提高对营养物质的吸收利用,改善肉鸡的屠宰性能。
研究表明,饲粮中添加XOS可通过调节蛋鸡营养物质消化率和回肠形态来提高饲料转化率,这可能与细菌多样性的增加和微生物组成的改变有关[62]。Ding等[69]发现,在蛋鸡饲粮中添加XOS对其肠道健康有益,可以提高蛋鸡盲肠中双歧杆菌数量和丁酸含量[70]。Morgan等[71]发现,补充XOS可提高蛋鸡对饲粮中木聚糖的利用率,改善营养利用率、生产性能和胃肠道健康。

4 小结与展望

XOS作为功能性低聚糖,具有低热量、高稳定性和增殖益生菌等特点,在无抗养殖时代拥有巨大的市场潜力。近几十年来,大量研究表明,XOS可以应用于许多食品和饲料产品,并发挥其营养价值。XOS还被证明可以减少与人类健康相关疾病的发生,提高动物的生长和对疾病的抵抗力。XOS的这些功能为其在人类消费和动物生产中的潜在应用开辟了新的视角。然而,XOS的制备和应用还面临着许多挑战。由于缺乏标准化的制备方法,不同批次不同聚合度的XOS质量缺乏一致性。目前,市场上销售的XOS产品以混合物为主,而非单一物质。未来,需要开发出一种具有低成本和高纯度XOS的制备技术。此外,还需要新的研究来进一步阐明XOS的具体分子机制,这些研究可能会提高XOS在预防和治疗人类疾病和动物生产方面的应用价值。

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