动物营养学报 >

2024 , Vol. 36 >Issue 12: 7493 - 7501

DOI: https://doi.org/10.12418/CJAN2024.637

综述

植物精油的生物学功能及其调控仔猪肠道健康的研究进展

  • 闫志强 , 1, 2 ,
  • 余远迪 1, 2 ,
  • 刘志云 1, 2 ,
  • 付利芝 , 1, 2, *
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  • 1 重庆市畜牧科学院,重庆 402460
  • 2 国家生猪技术创新中心,重庆 402460
*付利芝,研究员,E-mail:

闫志强(1989—),男,河南洛阳人,副研究员,硕士,从事兽用中药和天然药物研究。E-mail:

Copy editor: 菅景颖

收稿日期: 2024-05-24

  网络出版日期: 2024-12-12

基金资助

国家生猪技术创新中心先导项目(NCTIP-XD/B12)

国家重点研发计划(2023YFD1301600)

国家重点研发计划(2021YFD1301101)

收起

Research Progress on Biological Functions of Plant Essential Oils and Their Regulation on Intestinal Health in Piglets

  • YAN Zhiqiang , 1, 2 ,
  • YU Yuandi 1, 2 ,
  • LIU Zhiyun 1, 2 ,
  • FU Lizhi , 1, 2, *
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  • 1 Chongqing Academy of Animal Sciences, Chongqing 402460, China
  • 2 National Center of Technology Innovation for Pigs, Chongqing 402460, China
*professor, E-mail:

Received date: 2024-05-24

  Online published: 2024-12-12

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摘要

植物精油是从植物的花、叶、根、果实或种子中萃取提炼出的芳香族化合物,具有抗炎、抗菌、抗氧化等生物学功能及来源广泛、毒副作用小、不易产生耐药性等特点,作为一种抗生素替代品广泛应用于仔猪生产中。本文从植物精油的分类、理化性质、生物学功能及其对仔猪肠道健康的影响等方面进行综述,以期为植物精油在仔猪生产中的高效应用提供参考。

关键词： 植物精油; 生物学功能; 仔猪; 肠道健康

本文引用格式

闫志强 , 余远迪 , 刘志云 , 付利芝 . 植物精油的生物学功能及其调控仔猪肠道健康的研究进展[J]. 动物营养学报, 2024 , 36(12) : 7493 -7501 . DOI: 10.12418/CJAN2024.637

Abstract

Plant essential oils are aromatic compounds extracted and refined from flowers, leaves, roots, fruits, or seeds of plants. They have many biological functions such as anti-inflammatory, antibacterial, and antioxidant, with a wide range of sources, minimal toxic side effects, and they are not prone to developing drug resistance. As a substitute for antibiotics, they are widely used in piglet production. This article provides a review of the classification, physicochemical properties, biological functions, and effects of plant essential oils on the intestinal health of piglets, in order to provide reference for the efficient application of plant essential oils in piglet production.

Key words： plant essential oils; biological function; piglets; intestinal health

肠道健康是影响仔猪生长发育的关键因素,由于仔猪消化系统和免疫系统发育不健全,其肠道疾病常常发生,致使养殖成本增加和生产效益下降[1]。因此,寻找一种有效维持仔猪肠道健康的方法提高其生产性能,一直是养猪业关注的重点。植物精油作为一种天然的提取物,具有抗菌、抗炎、抗氧化等多种生物学功能,在动物生产中应用具有调节肠道菌群平衡、维持肠道屏障完整性、改善肠道免疫等功能,对提高动物生长性能和促进机体健康有积极作用[2]。本文将对植物精油的分类、理化性质、生物学功能进行综述,并重点介绍植物精油对仔猪肠道健康方面的调控作用,以期为养猪业提供一种安全有效和可行的肠道保健措施,从而提高仔猪的生长性能和养殖效益。

1 植物精油的分类及理化性质

根据提取部位不同,植物精油可分为花精油(如玫瑰精油)、叶精油(如薄荷精油)、茎精油(香茅精油)、根精油(如姜精油)、果实精油(如柠檬精油)等。植物精油通常为无色或淡黄色具有芳香性和挥发性油状液体,密度为0.8~1.0 g/mL,不溶于水、脂溶性强,性质相对稳定,但易受光、热和氧气等外界因素影响,其活性成分为挥发性化合物,易从消化道吸收到血液循环中,并在体内进行有效分布和运输。不同种类的植物精油其活性成分不同,因此具有不同的功效和用途[3]。常用于仔猪生产的植物精油有牛至精油、百里香酚、香芹酚、茴香精油、迷迭香精油、山苍子精油以及复合植物精油等。

2 植物精油的生物学功能及其作用机制

2.1 抗氧化

氧化应激不仅会破坏免疫细胞的结构和功能,导致畜禽免疫力下降,还会损伤生殖细胞,影响精子和卵子的质量和数量,降低受精率。植物精油中含有酚类、萜烯类、类黄酮等多种抗氧化活性成分,且不同成分其抗氧化机制不尽相同,如酚类化合物含有大量的酚羟基,可以电离出氢自由基(H·),H·与游离基反应,从而阻止氧化反应,此外,其还可激活核因子E2相关因子2(Nrf2)信号通路发挥抗氧化作用。Nrf2信号通路激活与氧化应激密切相关,当动物机体受到氧化应激时,激活Nrf2,促进移位至细胞核,活化血红素氧合酶-1(HO-1),提高抗氧化酶[超氧化物歧化酶(SOD)和谷胱甘肽过氧化物酶(GSH-Px)]的活性,进而减少活性氧(ROS)的产生[4]。而萜烯类化合物可通过直接清除多余的ROS、提高抗氧化酶活性实现抗氧化作用[5]。香叶醇通过激活Nrf2/HO-1信号通路,促进Nrf2和HO-1蛋白表达,增强抗氧化酶(SOD和GSH-Px)的活性,进而实现抗氧化作用[6]。Dong等[7]通过自由基清除试验测定百里香精油的抗氧化活性发现,百里香精油有较明显的抗氧化活性,且抗氧化活性与其浓度成正比,在浓度为0.05 mg/mL时抗氧化活性最弱,在浓度为1.0 mg/mL时抗氧化活性最强。

2.2 抗病原微生物

病原微生物与畜禽健康有着密切的关系,其可通过食物、水、空气或直接接触等途径传播,畜禽感染后出现消化道、呼吸道或泌尿道等症状,严重者可导致死亡。研究发现,植物精油可有效缓解或对抗病原微生物的感染,在仔猪生产中有较大的应用潜力[8-9]。

2.2.1 抗菌

植物精油具有抗菌活性强、抗菌谱广及不易产生耐药性细菌等优点[10],且可通过多种途径实现其抗菌效果(表1)。植物精油的抗菌机制主要有以下几种:1)抑制细菌生物膜的形成。细菌生物膜主要由脂质、磷脂、蛋白质等组成,植物精油因其强脂溶性可破坏细菌细胞膜的完整性,导致细胞内容物泄漏,从而杀死细菌。例如,肉桂精油可通过抑制定植扩散、胞外多糖合成和细胞表面特性来抑制金黄色葡萄球菌生物膜的形成,进而实现抗菌活性[11]。2)抑制细菌内能量代谢。植物精油可通过抑制能量代谢过程中相关酶(三磷酸腺苷酶、苹果酸脱氢酶、琥珀酸脱氢酶等)的活性,影响细菌内三羧酸循环,抑制能量的产生。例如,砂仁精油可显著降低金黄色葡萄球菌的Na+-K+-ATP酶和Ca2+-Mg2+-ATP酶活性,影响能量代谢,同时改变细胞内Na+、Ca2+浓度,增加细胞膜的渗透性,进而起到抗菌效果[12]。3)破坏细菌DNA结构。植物精油可影响DNA合成过程中相关酶(如拓扑异构酶)的活性或直接破坏DNA结构,进而影响细菌的遗传信息。例如,山苍子精油和菊花油的有效成分与细菌DNA和RNA聚合酶的活性位点之间具有高亲和力,可有效影响大肠杆菌、金黄色葡萄球菌、铜绿假单胞菌和化脓性链球菌的复制,进而表现出较强的抑菌活性[13-14]。综上可知,植物精油与抗生素的抗菌机制有一定的区别(图1),且不同的植物精油间抗菌谱和抗菌途径不同,临床应用时应先对其进行筛选,以获得抗目标菌效果明显的植物精油。
表1 不同植物精油的抗菌机制

Table 1 Antibacterial mechanisms of different plant essential oils

植物精油
Plant essential oils		 细菌名称
Bacterial name		 作用机制
Mechanism		 参考文献
References
香芹酚Carvacrol 假单胞菌 抑制生物膜的形成 [15⇓-17]
牛至油Oregano oil 肠沙门氏菌、李斯特菌
桂花精油
Acanthus polystachyus essential oil		 粪大肠杆菌、白色念珠菌、铜绿
假单胞菌和金黄色葡萄球菌
香叶醇Geraniol 假单胞菌 抑制铜绿假单胞菌毒力基因的表达 [18]
生姜精油
Ginger essential oil		 大肠杆菌、金黄色葡萄球菌 抑制与细菌能量代谢和三羧酸
循环相关基因的表达		 [19]
艾草精油
Artemisia argyi essential oil		 金黄色葡萄球菌、大肠
杆菌和白色念珠菌		 破坏细菌细胞壁结构 [20-21]
柠檬马鞭草精油
Lemon verbena essential oil		 假单胞菌
图1 植物精油和抗生素的抗菌机制对比

Plant essential oils:植物精油;Antibiotics:抗生素;Promotion:促进;Inhibition:抑制;Bacteria:细菌;Fungus:真菌;Cell wall:细胞壁;Cell membrane:细胞膜;Cytoplasm:细胞质;Plasmid:质粒;Ribosome:核糖体;Out membrance protein:外膜蛋白;Nucleus:细胞核;Mitochondria:线粒体;Conidia:分生孢子;Vesicles:囊泡;Metulae:初生小梗;Stipes:茎节;Foot cells:足细胞;Endoplasmic reticulum:内质网。

Fig.1 Comparison of antibacterial mechanisms between plant essential oils and antibiotics

2.2.2 抗病毒

植物精油对多种病毒(如单纯疱疹病毒、流感病毒、黄热病毒等)表现出显著的抗病毒活性[8]。植物精油因其亲脂性,易溶于病毒的包膜中,引起包膜破裂,影响病毒的复制[22];同时,植物精油还可干扰病毒包膜和刺突蛋白的形成,破坏病毒衣壳或蛋白质包膜[23],进入和穿透宿主细胞与病毒蛋白直接结合,使其失活或阻断病毒DNA和RNA的复制,抑制病毒子代生成,进而避免病毒感染[24]。例如,Zheng等[25]通过鼻内给予流感病毒构建病毒感染小鼠模型,使用香芹酚进行治疗后发现,香芹酚通过调节Toll样受体(TLR)/维甲酸诱导基因Ⅰ样受体样受体(RLR)模式识别并抑制甲型流感病毒复制,进而减轻因过度免疫反应引起的肺组织损伤;Ksouri等[26]采用分子对接技术对新型冠状病毒(SAS-CoV-2)的主要蛋白酶和植物精油进行分析发现,肉桂单宁B1、肉桂单宁B2对SAS-CoV-2与主要蛋白酶的异体部位有显著结合的能力,可能会阻止该病毒的附着,进而起到抗病毒作用。此外,植物精油还可提高机体免疫球蛋白和相关细胞因子含量。例如,Niu等[27]在饲粮中添加植物精油进行饲喂试验发现,植物精油可提高断奶仔猪血清免疫球蛋白A(IgA)、免疫球蛋白G(IgG)、免疫球蛋白M(IgM)含量,降低血清中炎症相关细胞因子[白细胞介素-1β(IL-1β)、白细胞介素-6(IL-6)、肿瘤坏死因子-α(TNF-α)]的含量,且通过包被工艺制备的植物精油效果更好。综上可知,植物精油主要通过破坏病毒结构、影响病毒黏附、侵入等方式直接抑制病毒复制和调控机体免疫应答(体液免疫和细胞免疫)方式间接抑制病毒2种途径发挥抗病毒作用。

2.2.3 抗寄生虫

植物精油含有单萜衍生物,具有较好的抗锥虫、疟原虫、利什曼原虫及肠道原虫活性[28-29]。Bailén等[30]从唇形科和菊科植物中提取23种植物精油进行抗利什曼虫试验发现,百里香酚和香芹酚是抗利什曼虫的主要活性成分。Ezzatkhah等[31]使用生姜精油对弓形虫进行体内和体外研究发现,生姜精油可抑制弓形虫的生长发育(体外),增加弓形虫感染小鼠的存活率,其机制可能是通过增强机体免疫、降低肝脏氧化应激程度及调节炎症反应等方式实现的。

2.3 抗炎

炎症是动物机体对于损伤、感染或刺激的一种非特异性的防御和修复反应,但过度或长期的炎症反应可能会导致组织损伤。植物精油的抗炎活性已得到广泛的研究和证实。在对脂多糖(LPS)诱导的C57BM/6J小鼠肠道炎症模型的研究中发现,山苍子精油可通过降低肝脏和肠道中TNF-α、IL-6和IL-1β的表达水平起到抗炎效果[32]。在LPS诱导的RAW264.7细胞炎症模型的研究发现,山香精油可抑制核因子-κB(NF-κB)的激活,降低炎性细胞中诱导型一氧化氮合酶和环氧合酶的mRNA表达水平,进而使IL-6、IL-1β和TNF-α等促炎细胞因子的含量减少[33]。在右旋糖酐硫酸钠诱导小鼠结肠炎模型的研究中发现,胡椒精油抑制Toll样受体4(TLR4)和丝裂原活化蛋白激酶(MAPK)的活化,调节TLR4/MAPK信号通路,进而抑制促炎细胞因子TNF-α、IL-1β和IL-6的表达[34]。由此可见,植物精油通过多种途径抑制因过度炎症反应引起的机体损伤,一方面可通过直接抑制相关炎性介质的释放(IL-1β和IL-6)发挥抗炎作用;另一方面可通过影响相关炎症通路的激活,实现抗炎效果。因此,植物精油可在生猪养殖中的断奶应激、热应激及运输应激方面有较大的应用潜力。

2.4 提高免疫力

提高免疫力有助于畜禽抵御病原微生物的入侵,减少疾病的发生和更好地应对环境变化和应激情况,提高适应能力和生存率。植物精油在免疫方面的作用主要体现在抗过敏、炎症、感染和肿瘤等方面[35-36]。Yeh等[37]通过蒸馏法从芸香科植物中提取吴茱萸精油,并作用于前列腺癌细胞发现,其活性成分可抑制前列腺癌细胞的生长和调节脾脏细胞因子白细胞介素-2(IL-2)、白细胞介素-10(IL-10)表达,起到治疗效果。刘娇等[38]研究显示,饲粮中添加植物精油可提高肉鸭血清中IgG和IgM的含量,进而提高机体免疫力。Yang等[39]研究发现,饲粮中添加植物精油可提高断奶仔猪血清中补体3和补体4的含量,改善机体的免疫状况。综上所述,植物精油可通过调节机体细胞免疫(细胞因子)和体液免疫(免疫球蛋白)提高机体的免疫力。

3 植物精油对仔猪肠道健康的影响

仔猪肠道在早期阶段发育并不完善,主要体现在以下几方面:1)肠道表面积相对较小,肠道上皮细胞的功能和数量有限,吸收能力较弱;2)胃酸分泌量较少,胃肠道的消化酶活性较低,消化能力不足;3)肠道菌群未完全建立,肠道黏膜较薄弱,易患肠道疾病。因此仔猪阶段需要特殊的饲养管理和营养策略来满足其营养需求,促进肠道健康生长和发育。

3.1 植物精油对仔猪肠道发育的影响

植物精油多为芳香族化合物,这些芳香成分可通过瘦素抵抗、交感神经/副交感神经的活动或下丘脑中神经肽Y(NPY)/刺鼠相关蛋白(AgRP)、可卡因和苯丙胺调节转录本(CART)/原黑素皮质素(POMC)的mRNA表达来发挥食欲调节作用[40]。此外,植物精油还可促进胃肠道消化酶的分泌[41]、改善肠道结构和功能[42](如调节肠上皮细胞的增殖和迁移、改善肠道发育、增加绒毛高度、降低隐窝深度)、促进与生长相关的激素(如生长激素、甲状腺激素等)分泌,以促进仔猪肠道和机体生长[43]。

3.2 植物精油对仔猪肠道非特异性免疫的影响

肠道非特异性免疫是一种通过多种细胞和分子相互作用的防御机制,包括黏膜屏障、肠道微生物群、肠道免疫细胞,其主要通过阻止病原菌的侵入,维持肠道结构和黏膜屏障的完整性,调节肠道微生物群的组成和功能,进而维持肠道健康。研究发现,植物精油可促进仔猪肠道发育,改善肠道形态,促进消化酶、抗氧化酶、溶菌酶等的分泌,增加肠道菌群的多样性,从机械、化学、生物屏障多方面改善仔猪肠道健康,减少仔猪因断奶、环境改变、冷热应激等造成的肠道损伤[44⇓-46]。

3.2.1 植物精油对仔猪肠道机械屏障的影响

肠道机械屏障由肠上皮细胞、黏液层和细胞间的紧密连接组成,可选择性渗透水、电解质、小分子营养物质等,阻止细菌、毒素和其他有害物质进入血液循环[47],能有效保护肠黏膜、维持肠道菌群平衡和调节肠道免疫,在肠道健康中起着重要的保护和调节作用[48]。Liang等[49]在饲粮中添加植物精油饲喂28日龄的断奶仔猪后发现,植物精油可提高仔猪肠道闭合蛋白和闭锁蛋白的mRNA表达水平以降低大肠杆菌引起的腹泻。Yang等[50]的研究显示,山苍子精油可增加仔猪十二指肠或空肠的绒毛高度,进而维持仔猪肠道形态,促进肠道健康。Zhao等[51]研究发现,植物精油可提高仔猪空肠中黏蛋白-2和溶菌酶的蛋白表达,增多空肠黏液和杯状细胞,促进肠道干细胞分化。由此可见,植物精油可通过促进仔猪肠道发育、改善肠道形态及提高紧密连接蛋白表达等方式增强仔猪肠道机械屏障功能。

3.2.2 植物精油对仔猪肠道化学屏障的影响

肠道化学屏障主要由口腔、胃、胰腺等分泌的消化酶与胃肠道黏液层的抗菌肽和免疫球蛋白及胃肠道肠腔腺体分泌的糖蛋白等组成的防御屏障,可有效阻止有害物质和微生物直接接触肠道上皮细胞,从而对肠道健康有着极为重要的保护作用。Shao等[52]研究发现,在断奶仔猪饲粮中添加混合植物精油可降低血液中溶菌酶活性和皮质醇水平,减轻断奶应激。Dieguez等[53]研究发现,饲粮中添加植物精油可提高断奶仔猪肠道中乳糖酶、麦芽糖酶和蔗糖酶活性,促进其对营养物质的消化吸收,并能改善肠道健康。Zhao等[54]报道,饲粮中添加植物精油可显著提高仔猪空肠中胰蛋白酶、脂肪酶和胰凝乳蛋白酶的等消化酶活性。综上可知,植物精油可促进仔猪肠道消化酶和溶菌酶活性,进而增强仔猪肠道化学屏障功能。

3.2.3 植物精油对仔猪肠道生物屏障的影响

肠道微生物群不仅可通过竞争性排除和抑制病原菌的生长进而维持肠道内的微生态平衡,还能激活宿主的免疫反应,促进肠道非特异性免疫。据报道,植物精油可通过调动物节机体肠道菌群多样性和结构,改善肠道健康[55-56]。Arzola-Alvarez等[57]通过对植物精油与2株猪腹泻肠毒素大肠杆菌菌株和猪粪便细菌菌群混合培养发现,其可降低肠杆菌科的总量,但增加乳杆菌科的数量;同时研究发现,断奶仔猪饲粮中添加植物精油可提高肠道组织中双歧杆菌属、粪杆菌属、普雷沃氏菌科NK3B31(Prevotellaceae_NK3B31)和拟杆菌属的丰度,降低克里斯滕森菌科R-7(Christensenellaceae_R-7)和密螺旋体属的丰度,调节肠道菌群多样性,以降低因断奶应激造成的肠道炎症反应和致病菌入侵肠道的机会[58-59]。Li等[60]报道,断奶仔猪饲粮中添加植物精油可促进结肠中芽孢杆菌纲(Bacilli)、乳杆菌目(Lactobacillales)、链球菌科(streptococcaceae)、韦荣球菌科(Veillonellaceae)和巨球型菌属(Megasphaera)等有益细菌的生长,并证实肠道细菌属与肠道微生物群相关代谢物的改变高度相关。此外,Mo等[61]研究发现,微囊化植物精油可显著降低断奶仔猪盲肠和结肠中潜在致病菌和有益菌的相对丰度比,这可能是其降低腹泻指数的重要原因。综上可知,植物精油一方面可提高仔猪肠道有益菌的丰度和数量,促进消化酶的分泌和营养物质的消化吸收;另一方面可抑制有害菌的增殖,减少和清除有害菌产生的毒素,改善仔猪肠道生物屏障,促进仔猪肠道健康。

3.3 植物精油对仔猪肠道特异性免疫的影响

特异性免疫主要有细胞免疫和体液免疫2种形式。细胞免疫通过T淋巴细胞识别抗原片段,进行激活并增殖产生效应细胞攻击和清除抗原。体液免疫则是通过B淋巴细胞产生和分泌特异性抗体与抗原结合,中和抗原或激活补体系统,清除抗原。在断奶仔猪饲粮中添加不同剂量的茶树精油进行饲喂后发现,50 mg/kg的茶树精油可显著提高十二指肠及回肠黏膜中分泌型免疫球蛋白A的含量[62]。饲粮中添加牛至精油可上调断奶仔猪空肠组织中干扰素-ε、干扰素-ω的表达量,改善仔猪的肠道免疫水平[63]。Zhang等[64]研究发现,饲粮中添加茶树精油可提高断奶仔猪血清中抗炎细胞因子IL-10的含量,以调节仔猪的免疫状态。综上可知,植物精油可通过促进免疫球蛋白的分泌,调节肠道的体液免疫,同时促进相关细胞因子的表达,提高仔猪肠道细胞免疫水平。

4 小结与展望

植物精油可通过改善仔猪肠道组织形态、促进消化酶和溶菌酶分泌、调节肠道菌群多样性及增强肠道非特异性与特异性免疫等多种途径改善仔猪肠道健康,可广泛地应用于生猪养殖中,促进生猪绿色高效养殖。但植物精油因其易挥发分解、不溶于水和有特殊气味等性质,局限了其在仔猪生产上的应用。此外,植物精油在猪饲粮中的添加剂量亦不明确。因此,未来应优化植物精油的制备工艺,深入探讨其在猪不同生长阶段饲粮中的最佳添加剂量,为植物精油在生猪养殖中的应用提供参考。

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