动物营养学报 >

2023 , Vol. 35 >Issue 8: 4850 - 4859

DOI: https://doi.org/10.12418/CJAN2023.450

综述

植物提取物促动物生长作用机制的研究进展

  • 张泽茹 , 1 ,
  • 赵玉全 1 ,
  • 董可 2 ,
  • 沈留红 , 1, *
展开
  • 1 四川农业大学动物医学院,动物疫病与人类健康四川省重点实验室,奶牛疾病研究中心,成都 611130
  • 2 四川育强本草生物技术有限公司,成都 611130
*沈留红,副教授,硕士生导师,E-mail:

张泽茹(2000—),女,山西临汾人,硕士研究生,从事奶牛疾病研究。E-mail:

Office editor: 陈鑫

收稿日期: 2023-02-09

  网络出版日期: 2023-08-10

基金资助

四川省科技厅科技计划项目(2019YJ0650)

四川农业大学学科建设双支计划(2221998011)

四川农业大学本科科研兴趣培养计划(KY20231908)

收起

Research Progress on Mechanism of Action of Plant Extracts to Promote Animal Growth

  • ZHANG Zeru , 1 ,
  • ZHAO Yuquan 1 ,
  • DONG Ke 2 ,
  • SHEN Liuhong , 1, *
Expand
  • 1 Dairy Cows Disease Research Center, Key Laboratory of Animal Disease and Human Health, College of Veterinary Medicine, Sichuan Agricultural University, Chengdu 611130, China
  • 2 Sichuan Yuqiang Herbal Biotechnology Co., Ltd., Chengdu 611130, China
*associate professor, E-mail:

Received date: 2023-02-09

  Online published: 2023-08-10

Fold

摘要

植物提取物(PE)是提取于植物中且具有抗菌、抗氧化和免疫调节等1种或多种生物学功能的天然成分物质,可通过改善饲料适口性,增加动物采食量;促进消化液和消化酶分泌,调节肠道菌群并改善肠道生理形态和功能状态,从而促进营养物质消化吸收;促进生长激素(GH)、胰岛素样生长因子-1(IGF-1)和甲状腺激素(THs)分泌,调节机体生长和代谢;提供维生素D并促进成骨,维持骨组织生长,从而起到促进动物生长的作用。近年来,PE作为天然促生长饲料添加剂已成为国内外研究热点,而目前对PE的研究主要集中在对动物机体某一方面或某一功能的单一影响上。因此,本文针对PE促动物生长作用的机制进行系统综述,为PE的开发利用提供参考。

关键词： 天然植物提取物; 促进动物生长; 作用机制; 饲料添加剂

本文引用格式

张泽茹 , 赵玉全 , 董可 , 沈留红 . 植物提取物促动物生长作用机制的研究进展[J]. 动物营养学报, 2023 , 35(8) : 4850 -4859 . DOI: 10.12418/CJAN2023.450

Abstract

Plant extracts (PE) are natural substances extracted from plants and have one or more biological functions such as antimicrobial, antioxidant and immunomodulation, which can increase animal feed intake by improving feed palatability; promote the secretion of digestive fluid and digestive enzymes, regulate intestinal flora and improve intestinal physiological morphology and functional status, thereby promoting the digestion and absorption of nutrients; promote growth hormone (GH), insulin-like growth factor-1 (IGF-1), and thyroid hormone (THs) secretion, regulate body growth and metabolism; provide vitamin D and promote osteogenesis, maintaining bone tissue growth, thus promoting animal growth. In recent years, PE as a natural growth-promoting feed additive has become a research focus all over the world, and the current research on PE has largely focused on a single impact on a certain aspect or function of the animal bodies. Therefore, this article systematically reviews the mechanism of PE to promote animal growth, so as to provide reference for the development and utilization of PE and further reveal the molecular mechanism.

Key words： natural plant extracts; promote animal growth; mechanism of action; feed additive

动物生长性能是畜牧业生产中十分重要的经济指标,长期以来,抗生素一直被用作预防动物疾病、促进动物生长的饲料添加剂,但抗生素长期甚至超剂量添加已给人类、动物和生态环境造成不可逆转的负向影响[1-2],目前很多国家均已限制或禁止饲料中添加抗生素类添加剂。而植物提取物(plant extracts,PE)是通过物理或化学等方法,从自然植物中提取得到的单一或混合且保留植物原有生物活性及自然因子的物质[1],主要包括黄酮、多酚、多糖、生物碱、有机酸、挥发油和皂苷等,这些生物活性成分大多是植物生长发育过程中的次生性代谢产物[3],具有促生长、抗氧化、抗菌、抗病毒和增强免疫等生物学功能[4-5]。PE通过其活性成分间的单一或协同作用,增加动物采食量,促进营养物质消化吸收,刺激与生长相关激素的分泌以及促进骨组织生长,从而达到促进动物生长的作用,其以天然、安全、副作用小和不易产生耐药性等特性,已成为替抗促生长饲料添加剂的首选原料[3,5]。本文综述了PE促动物生长作用的潜在机制,以期为天然促生长饲料添加剂的开发利用提供理论参考。

1 PE改善饲料适口性,增加动物采食量

采食量是调控动物生长性能的关键因素,而适口性是决定动物对饲料的接受程度、影响采食量的重要因素[6]。PE中的醛类、酚类和芳香烃类等芳香类物质可作为调味剂改善饲料适口性,通过刺激动物嗅觉和味觉等外周感受器,促进动物采食[7-8]。
PE的气味分子进入动物鼻腔到达嗅觉上皮,与嗅觉感受器神经元(olfactory sensory neurons,OSNs)中的嗅觉受体(odorant receptors,ORs)相结合,激活膜上特定的G蛋白偶联受体(G protein-coupled receptors,GPCRs),从而引起环磷酸腺苷(cyclic adenosine monophosphate,cAMP)介导的信号转导通路,导致膜上离子通道开放,OSNs去极化[9]。随后,这些轴突化学信号沿嗅束从嗅球传递至大脑皮层嗅觉区域,而不同气味会触发不同OSNs,引起机体做出不同反应[9⇓-11]。PE经口腔作用于味觉感受器——味蕾,并由专门味觉细胞感知,激活相应特异性受体蛋白并引发信号转导[12-13],一旦动作电位被激活,味觉信息通过神经元传递并到达大脑皮层味觉区域,经处理后的信息协助大脑识别味道并调节动物采食。气味和味道组合作为PE的“提示因子”,可调节动物采食情绪、认知和行为,为判断是否进一步采食提供初步依据[13-14]。Yu等[15]研究发现,八角茴香油可显著升高蛋鸡平均日采食量,可能是因八角油的天然芳香气味和茴香味刺激了蛋鸡的食欲;杜蓥蓥等[16]研究发现,大蒜素和鞣花酸、牛至香酚和柠檬酸PE复合物分别可显著或极显著提高仔猪平均日采食量和平均日增重,可能是由于2种PE复合物能掩盖原饲料中的不适气味以改善其适口性,且大蒜素和柠檬酸的独特气味能促进消化酶分泌,保证饲料中营养物质的消化吸收。综上表明,PE能够改善饲料适口性[17],通过气味刺激促进动物产生食欲,经大脑处理后做出加强唾液分泌和采食等反馈行为[8]。动物有了采食经验后,通过嗅觉和味觉记忆识别[18-19],当再次接触食用过的同类型饲料时就会产生条件反射,引起迷走神经兴奋,乙酰胆碱释放,进一步刺激胃腺分泌胃液及G细胞释放促胃液素,加快胃肠蠕动并提高机体对食物的消化吸收能力,增加动物采食量。

2 PE促进营养物质消化吸收

营养物质提供动物生长、发育和活动所需能量,是维持机体组成和机能必不可少的要素,也是生命活动的物质基础。PE具有促进消化液和消化酶的分泌、调节肠道菌群、改善肠道生理形态和功能状态的作用,有利于营养物质的消化吸收并维持机体生长。

2.1 促进消化液分泌,增强消化酶活性

消化液和消化酶可分解大分子营养物质并加速其消化吸收,参与肠道内众多重要生化过程和物质循环,已有许多研究结果证实PE具有促进消化液和消化酶分泌功能。PE通过促进唾液、肠道黏液和胆汁分泌以及提高内源酶分泌量和活性,改变消化道中饲料形态,降低食糜黏度,进而提高营养物质消化率以保证机体获得充足养分[20-21]。Tiihone等[22]研究发现,饲粮中添加百里香酚和肉桂醛精油混合物可增加丁酸盐浓度及胆囊收缩素、YY肽(PYY)和胰高血糖素样肽2水平,延长食糜在胃中停留时间并刺激胃液和胰液分泌,从而改善营养物质消化吸收。刘福鑫等[20]研究发现,香芹酚可增加肉兔唾液和胆汁分泌,并提高回肠胰蛋白酶、盲肠胰蛋白酶和脂肪酶活性,从而促进肉兔消化。此外,PE在肠道微生物分解转化作用下产生的有机酸(organic acids,OAs)可作为酸化剂降低胃环境pH[23],减慢饲料转运速度以最大限度消化饲料、吸收养分,并对动物胃肠道黏膜产生营养作用。同时,OAs阴离子还可与钙、磷、镁、锌等矿物质络合,促进其吸收和保留,增加其生物利用价值[23⇓-25]。

2.2 调节肠道菌群

肠道具有机体最主要且复杂的微生态系统[26],其包含的细菌多样,对机体生理功能的完善和免疫功能的成熟发挥重要作用[27]。PE可通过抑制肠道有害菌和生物膜形成、促进有益菌生长,减少不利因素导致的肠道微生态失衡,维持肠道正常功能发挥并提高机体生长性能。

2.2.1 抑菌作用

PE可直接抑杀肠道有害菌或促进有益菌生长而间接抑制有害菌繁殖[28-29]。其通过破坏和降解细菌细胞壁中果胶和纤维素,抑制细胞壁肽聚糖合成[30],同时干扰细胞膜中不同脂肪酸、磷脂双层和多糖分子排列,影响膜流动性和通透性,最终导致细菌因重要分子、离子、核酸、氨基酸等渗漏而死亡。细胞膜的破坏会进一步引起细胞内环境pH和无机离子失衡,扰乱物质转运、能量产生和新陈代谢等生命活动。此外,PE能破坏各种细菌酶活性,诱导细胞中ATP泄漏和损失,抑制DNA合成甚至损伤DNA,影响重要毒力基因转录和表达,降低细菌致病力和耐药性,减少菌体生长繁殖[31-32]。如山苍子精油能破坏大肠杆菌O157∶H7细胞膜,影响其呼吸代谢和细胞能量代谢,抑制DNA拓扑异构酶活性和菌体主要毒力基因stx1、stx2、ehxA、eae转录水平[33]。此外,PE中OAs成分以未解离的形式扩散进入细菌内,较高的胞浆pH导致OAs解离并释放出氢离子(H+)和羧酸根(RCOO-)[34],为纠正降低的pH,细菌被迫消耗能量,激活质子泵将H+排出膜外,同时RCOO-干扰DNA、RNA合成并破坏细菌新陈代谢等生理过程,抑制细菌的复制和生长[35-36]。

2.2.2 益生作用

PE中高水平膳食纤维、寡糖和多酚等对益生菌有益的成分[37],大都可顺利通过胃和小肠而不被消化酶降解利用,但大肠中的有益菌可将其发酵转化成短链脂肪酸、乳酸、葡萄糖等,并进行合成代谢,使有益菌得到充足养分而增殖。而需氧的有害菌却无法利用,同时增殖后的有益菌会通过产生OAs等各种途径抑制有害菌,从而调节肠道微生态系统[38⇓-40]。如PE中O-糖基化多酚可被一些微生物水解为苷元和葡萄糖,作为有益菌生长的能量和碳源,刺激益生菌生长[41]。

2.2.3 干扰细菌群体感应(quorum sensing,QS)系统

病原菌吸附于肠道表面后会聚集成群落,并通过QS系统形成生物膜结构,严重影响肠道正常功能发挥。QS是一种细胞间通信机制,通过感知群落成员间分泌的化学信号监测细胞密度,从而在细菌间传递信息并调节与毒力、竞争、致病性和抗性相关的基因表达以适应环境变化,避免受到抗菌剂或免疫系统攻击等影响[42-43]。而PE可作为QS抑制剂,抑制生物膜发育过程的相关因素(如运动性、黏附性、细胞间聚集和通讯等),干扰QS系统的调控,阻碍生物膜形成[44]。此外,PE能根除短期内预先形成的亚微囊生物膜,降低初始生物膜种群量,破坏膜的完整性并导致微菌落分散,降低细胞密度和厚度,破坏多重耐药细菌产生的生物膜结构[45],起到抑制和灭活生物膜的作用,其效果甚至优于标准酸性和碱性化学消毒剂[46]。

2.3 改善小肠生理形态和功能状态

小肠强大的消化吸收养分功能不仅得益于巨大的吸收面积,也与肠上皮细胞(intestinal epithelial cells,IECs)的作用密切相关。IECs含有各种分化的细胞类型,能够吸收养分,分泌黏液、激素和抗菌因子以保护肠道[46⇓-48],而肠绒毛高度(villi height,VH)、隐窝深度(crypt depth,CD)和绒隐比(V/C)是反映IECs增殖、肠道生理形态及功能状态的重要指标[49],VH升高,CD变浅,则IECs增殖率和成熟度越高,吸收养分能力和分泌功能也相对增强[50]。
目前,PE对VH、CD和V/C的正向调节作用已得到广泛研究和证实[51-52]。其可从3个方面刺激IECs的迁移和增殖,改善肠黏膜发育:1)介导IECs多胺-钙离子(Ca2+)信号通路。多胺由肠道菌群代谢产生,为细胞生长、增殖和分化所必需的氨基化合物[53],而细胞内游离Ca2+浓度([Ca2+]cyt)是调控细胞迁移关键因素,其增加依赖于内质网内Ca2+释放或细胞外Ca2+内流[54-55]。PE可增加多胺含量,上调IECs中K+通道蛋白(Kv1.1)表达,促进膜超极化,进而增强细胞外Ca2+流入的驱动力[56-57]。此外,典型的瞬时受体电位1(TRPC1)、小窝蛋白1(caveolin 1,CAV1)、磷脂酶C-γ1(PLC-γ1)、RhoA和Rac1均是对细胞迁移起促进作用的关键因子,且部分因子可相互作用形成蛋白复合物来发挥作用,如TRPC1/RhoA、TRPC1/CAV1、RAC1/PLCγ-1。PE可促进这些调控因子及其复合物的表达,增强Ca2+内流,促进细胞迁移和增殖[54-55,58]。2)PE介导多胺对IECs细胞周期的调控作用。PE可上调细胞周期蛋白(Cyclin)与细胞周期蛋白依赖性激酶(cyclin-dependent kinases,Cdks)2表达,调控G1期到S期的转变以及S期的推进[59],同时下调细胞周期抑制因子p21和p53表达,减少细胞凋亡或生长停滞[60],降低G0/G1期百分比,增加G2/M+S期百分比,从而促进IECs生长和分化[61-62]。3)PE逆转二氟甲基鸟氨酸(difluoromethylornithine,DFMO)对多胺合成和IECs增殖的抑制作用。DFMO作为多胺合成抑制剂,通过降低鸟氨酸脱羧酶(ornithine decarboxylase,ODC)(催化多胺合成中的第1个限速步骤)活性,以减少多胺的生物合成,对IECs增殖起抑制作用[63],PE可逆转DFMO,恢复多胺缺乏的IECs迁移[61-62]。综上所述,PE通过增加多胺含量,介导IECs多胺-Ca2+信号通路并刺激Ca2+调控蛋白及其复合物表达,影响细胞周期分布,逆转DFMO的抑制作用,从而刺激IECs迁移和增殖,促进黏膜生长发育和养分吸收。

3 PE促进生长相关激素分泌

动物生长发育通过外部充足的营养供应和内部激素的共同作用而实现。PE可促进生长激素(growth hormone,GH)、胰岛素样生长因子-1(insulin-like growth factor-1,IGF-1)和甲状腺激素(thyroid hormones,THs)等分泌,调节机体生长和代谢,影响内分泌系统,促进机体生长。

3.1 促进GH和IGF-1分泌

GH具有促进组织生长、刺激细胞增殖、调节新陈代谢等作用[64],其主要通过IGF-1来介导活性发挥并调节骨纵向生长[65-66]。PE能诱导腺垂体中GH分泌[64],GH与生长激素结合蛋白(growth hormone binding protein,GHBP)结合后经血液循环运送至肝脏,启动肝细胞内双面神激酶2-信号传导及转录激活蛋白5(Janus kinase 2-signal transducer and activator of transcription 5,JAK2-STAT5)信号转导机制并促进IGF-1产生[67-68],同时PE可上调IGF结合蛋白(IGF binding protein,IGFBP)3表达[67,69],增加运送至骨组织IGF-1含量,其与生长板细胞膜表面受体结合后可促进软骨细胞增殖、分化,介导软骨内骨形成[68]。PE还可诱导生长板中骨形态发生蛋白-2(bone morphogenetic protein-2,BMP-2)表达,刺激软骨发育[66-67]。Lee等[70]研究发现,黄芪提取物可显著增加大鼠GH的mRNA水平并促进IGF-1产生;Kim等[71]研究发现,黄芪、糙苏和刺五加混合物可增加生长板高度及IGF-1和IGFBP-3水平,刺激软骨细胞增殖,促进骨纵向生长。综上所述,PE可促进GH和IGF-1产生,上调肝脏IGF-1和IGFBP-3以及生长板中局部IGF-1和BMP-2表达,刺激JAK2-STAT5磷酸化以增加循坏IGF-1水平,进而促进软骨细胞增殖,增加生长板高度并诱导骨纵向生长。

3.2 促进THs分泌

THs介导机体代谢和生长发育过程。甲状腺分泌并释放的主要激素是甲状腺素(T4),而主要发挥生物活性的是代谢更快的三碘甲腺原氨酸(T3),T3一部分由甲状腺直接分泌,但大部分来源于T4的转化[65-66]。PE可促进组织水平的T4特异性脱碘转换为T3,随后T3进入细胞核并与甲状腺激素受体(TH receptors,TRs)结合调节广泛的基因[65-66],参与对机体生长的调控,但有无其他作用途径和机制还有待进一步研究。

4 促进骨组织生长

骨组织具有支持和保护身体、储存矿物质、参与机体运动的功能。骨组织的正常发育与机体生长密切相关,PE能促进骨矿化并维持骨骼的完整性,预防骨质流失。

4.1 提供维生素D(vitamin D,VD)

PE中的VD成分可通过其活性代谢产物1,25-双羟维生素D[1,25(OH)2D]调节肠道及肾相关基因转录,进而刺激肠道对钙磷的吸收和肾对钙的重吸收,促进骨矿化[72⇓-74]。在肠道,上皮细胞膜的钙通道瞬时受体电位(transient receptor potential channel subfamily, TRP)V6和钙结合蛋白calbindin(calcium-binding protein calbindin)D9k可介导钙转运,钠磷共转运蛋白可摄取磷酸盐[75];在肾脏,钙通过TRPV5进入远端小管细胞内,与calbindin-D9k和calbindin-D28k结合并在细胞质中转移,随后被钠/钙交换器1(sodium/calcium exchanger 1,NCX1)和质膜钙1b(plasma membrane calcium pump 1b)挤压至膜表面,从而促进钙的重吸收[75]。综上所述,PE通过上调TRPV5、calbindin-D9k、calbindin-D28k、钠/磷共转运蛋白和NCX1表达,促进钙磷吸收,维持胶原基质中足够的钙磷产物,从而促进骨矿化[76-77]。

4.2 促进成骨

骨骼需发育和塑造以适应生长,此过程主要涉及成骨细胞和破骨细胞这2种细胞类型,成骨细胞是骨形成的主要功能细胞,负责骨基质的合成、分泌和矿化[78]。
PE具有成骨活性,其对成骨细胞的促进作用表现于3个方面:1)刺激骨髓间充质干细胞(bone mesenchymal stem cells,BMSCs)增殖,促进其成骨分化。多种骨成分细胞都来源于BMSCs,BMSCs的分化由一系列转录因子控制,不同转录因子决定不同细胞谱系[79]。PE可增强BMSCs自我更新能力,激活骨形态发生蛋白、一氧化氮(nitric oxide,NO)、丝裂原活化蛋白激酶(MAPKs)和典型Wnt/β-catenin通路等成骨相关信号通路[80-81],上调成骨细胞分化的转录因子RUNX2、Osterix和β-catenin以及成骨标志基因碱性磷酸酶(alkaline phosphatase,ALP)、核结合因子-α1(CBF-α1)、骨钙素(osteocalcin,OC)、骨涎蛋白(bone sialoprotein,BSP)和DNA结合抑制剂1(inhibitor of DNA binding and differentiation 1,Id1)等表达,促进BMSCs成骨分化[80,82]。2)抑制原代成骨细胞向脂肪细胞转分化。成骨细胞和脂肪细胞均由BMSCs分化而来,且成骨分化和成脂分化存在相互制约的平衡关系[83]。经PE处理的BMSCs,脂肪生成相关基因、过氧化物酶体增殖物激活受体γ2(peroxisome proliferator activated receptor γ2,PPARγ2)和转录因子CCAAT增强子结合蛋白-β(C/EBP-β)等抑制成骨且驱动脂肪细胞分化的关键因子表达下调[84-85],而PPARγ2和C/EBP-β也受到MAPKs通路和成骨标志基因的抑制,从而减少脂肪生成的转分化并导致BMSCs进一步分化为成骨细胞。3)抑制破骨细胞的生成和活性。由BMSCs和成骨细胞产生的核因子-κB受体活化因子配体(receptor activator of nuclear factor-κB ligand,RANKL),能与破骨细胞前体上NF-κB受体激活子(receptor activator of nuclear factor-κB,RANK)结合,诱导破骨细胞分化和激活,而骨保护素(osteoclastogenesis inhibitory factor,OPG)能阻断RANKL作用,抑制破骨细胞前体细胞分化和融合[86]。PE可通过Wnt/β-catenin通路,在蛋白和mRNA水平上减少RANKL表达并增加OPG分泌,抑制破骨形成[87]。同时,破骨细胞的表型标志物抗酒石酸酸性磷酸酶(tartrate resistant acid phosphatase,TRAP)阳性多核细胞数明显减少,且实际骨吸收面积减小[80]。除了抑制破骨细胞的分化外,PE还能诱导破骨细胞前体细胞G2/M细胞周期暂停,导致细胞凋亡。综上所述,PE通过刺激BMSCs增殖,促进成骨细胞分化,抑制破骨细胞的生成和活性,从而促进骨骼发育,维持正常生长。

5 前景和展望

PE活性成分多样,具有促生长、抗菌、抗氧化、提高免疫力等功效,且不同PE间往往能发挥协同作用。目前PE已作为一种新型饲料添加剂在世界范围内引起了广泛关注,其作为当下一种发展迅速且有效的替抗物质,有望缓解或解决长期困扰畜牧领域发展的抗生素残留、环境污染及食品安全等问题。因此,科学利用PE作饲料添加剂,能更好地促进现代畜牧业的可持续发展。但与此同时,也应看到PE在应用中尚存的问题,并对其毒理学和安全性进行全面系统的研究和评价。

6 小结

PE具有来源广泛、安全高效、纯天然等优点,是一种极具潜力的促生长剂。其促生长作用机制包括:改善饲料适口性,增加动物采食量;促进消化液和消化酶分泌,调节肠道菌群并改善肠道生理形态和功能状态,从而促进营养物质消化吸收;促进GH、IGF-1和THs分泌,调节机体生长和代谢;提供VD并刺激成骨分化,从而促进骨组织生长,防止骨质流失。

参考文献

原文顺序 | 文献年度倒序 | 文中引用次数倒序
[1]
LILLEHOJ H, LIU Y H, CALSAMIGLIA S, et al. Phytochemicals as antibiotic alternatives to promote growth and enhance host health[J]. Veterinary Research, 2018, 49(1):76.

DOI PMID

[2]
PHILLIPS I. Withdrawal of growth-promoting antibiotics in Europe and its effects in relation to human health[J]. International Journal of Antimicrobial Agents, 2007, 30(2):101-107.

PMID

[3]
RIGHI F, PITINO R, MANUELIAN C L, et al. Plant feed additives as natural alternatives to the use of synthetic antioxidant vitamins on poultry performances,health,and oxidative status:a review of the literature in the last 20 years[J]. Antioxidants(Basel), 2021, 10(5):659.

[4]
NAI J J, ZHANG C, SHAO H L, et al. Extraction,structure,pharmacological activities and drug carrier applications of Angelica sinensis polysaccharide[J]. International Journal of Biological Macromolecules, 2021, 183:2337-2353.

DOI

[5]
CHENG G Y, HAO H H, XIE S Y, et al. Antibiotic alternatives:the substitution of antibiotics in animal husbandry?[J]. Frontiers in Microbiology, 2014, 5:217.

[6]
TEPPER B J, BARBAROSSA I T. Taste,nutrition,and health[J]. Nutrients, 2020, 12(1):155.

DOI

[7]
张定华, 谭占坤, 杜敏, 等. 植物提取物的作用机制及在畜禽生产中的应用[J]. 饲料研究, 2021, 44(13):153-156.

ZHANG D H, TAN Z K, DU M, et al. Mechanism of plant extract and its application in livestock and poultry production[J]. Feed Research, 2021, 44(13):153-156. (in Chinese)

[8]
王旭东, 邵丽霞, 唐仁龙, 等. 调味剂对动物嗅觉、味觉、肠道功能的调控研究进展[J]. 中国畜牧杂志, 2018, 54(2):15-20.

WANG X D, SHAO L X, TANG R L, et al. Research progress on transduction pathway of palatability enhancers and their application[J]. Chinese Journal of Animal Science, 2018, 54(2):15-20. (in Chinese)

[9]
BUCK L B. Unraveling the sense of smell (Nobel lecture)[J]. Angewandte Chemie International Edition, 2005, 44(38):6128-6140.

DOI

[10]
SOUDRY Y, LEMOGNE C, MALINVAUD D, et al. Olfactory system and emotion:common substrates[J]. European Annals of Otorhinolaryngology,Head and Neck Diseases, 2011, 128(1):18-23.

DOI

[11]
KAUPP U B. Olfactory signalling in vertebrates and insects:differences and commonalities[J]. Nature Reviews Neuroscience, 2010, 11(3):188-200.

DOI

[12]
WILSON C E, LASHER R S, YANG R B, et al. Taste bud connectome:implications for taste information processing[J]. Journal of Neuroscience, 2022, 42(5):804-816.

DOI

[13]
TARUNO A, NOMURA K, KUSAKIZAKO T, et al. Taste transduction and channel synapses in taste buds[J]. Pflügers Archiv:European Journal of Physiology, 2021, 473(1):3-13.

DOI

[14]
轩俊丽, 张莉, 杜立新. 嗅觉与味觉对动物采食作用研究进展[J]. 中国畜牧兽医, 2015, 42(12):3231-3238.

XUAN J L, ZHANG L, DU L X. Research progress on effects of olfactory and taste on animal food intake[J]. China Animal Husbandry&Veterinary Medicine, 2015, 42(12):3231-3238. (in Chinese)

[15]
YU C Y, WEI J D, YANG C W, et al. Effects of star anise (Illicium verum Hook.f.) essential oil on laying performance and antioxidant status of laying hens[J]. Poultry Science, 2018, 97(11):3957-3966.

DOI

[16]
杜蓥蓥, 王小辉, 蔡旭滨, 等. 2种植物提取物复合物对断奶仔猪生长性能以及肠道形态结构、免疫功能和胰岛素样生长因子-Ⅰ含量的影响[J]. 动物营养学报, 2022, 34(2):830-838.

DOI

DU Y Y, WANG X H, CAI X B, et al. Effects of two plant extract compounds on growth performance and intestinal morphology,immune function and insulin-like growth factor-Ⅰ content of weaned piglets[J]. Chinese Journal of Animal Nutrition, 2022, 34(2):830-838. (in Chinese)

[17]
CLOUARD C, VAL-LAILLET D. Impact of sensory feed additives on feed intake,feed preferences,and growth of female piglets during the early postweaning period[J]. Journal of Animal Science, 2014, 92(5):2133-2140.

DOI

[18]
MIRANDA M I. Taste and odor recognition memory:the emotional flavor of life[J]. Reviews in the Neurosciences, 2012, 23(5/6):481-499.

[19]
LLEDO P M, GHEUSI G, VINCENT J D. Information processing in the mammalian olfactory system[J]. Physiological Reviews, 2005, 85(1):281-317.

DOI

[20]
刘福鑫, 孔繁根, 王国洲, 等. 香芹酚对肉兔生长性能、养分表观消化率、肠道消化酶活性、免疫器官发育及抗氧化能力的影响[J]. 动物营养学报, 2022, 34(3):1875-1884.

DOI

LIU F X, KONG F G, WANG G Z, et al. Effects of carvacrol on growth performance,nutrient apparent digestibilities,intestinal digestive enzyme activities,immune organ development and antioxidant capacity of meat rabbits[J]. Chinese Journal of Animal Nutrition, 2022, 34(3):1875-1884. (in Chinese)

[21]
OSO A O, SUGANTHI R U, REDDY G B M, et al. Effect of dietary supplementation with phytogenic blend on growth performance,apparent ileal digestibility of nutrients,intestinal morphology,and cecal microflora of broiler chickens[J]. Poultry Science, 2019, 98(10):4755-4766.

DOI

[22]
TIIHONEN K, KETTUNEN H, BENTO M H L, et al. The effect of feeding essential oils on broiler performance and gut microbiota[J]. British Poultry Science, 2010, 51(3):381-392.

DOI PMID

[23]
TUGNOLI B, GIOVAGNONI G, PIVA A, et al. From acidifiers to intestinal health enhancers:how organic acids can improve growth efficiency of pigs[J]. Animals(Basel), 2020, 10(1):134.

[24]
NGUYEN D H, SEOK W J, KIM I H. Organic acids mixture as a dietary additive for pigs-a review[J]. Animals(Basel), 2020, 10(6):952.

[25]
RODJAN P, SOISUWAN K, THONGPRAJUKAEW K, et al. Effect of organic acids or probiotics alone or in combination on growth performance,nutrient digestibility,enzyme activities,intestinal morphology and gut microflora in broiler chickens[J]. Journal of Animal Physiology and Animal Nutrition, 2018, 102(2):e931-e940.

DOI

[26]
HEINTZ-BUSCHART A, WILMES P. Human gut microbiome:function matters[J]. Trends in Microbiology, 2018, 26(7):563-574.

DOI

[27]
王梦璇, 董丽, 袁浩洋, 等. 植物提取物在断奶仔猪饲粮中应用的最新研究进展[J]. 中国饲料, 2021(13):1-6.

WANG M X, DONG L, YUAN H Y, et al. Recent progress in the application of plant extracts in the diet of weaned piglets[J]. China Feed, 2021(13):1-6. (in Chinese)

[28]
LIU Q, MENG X, LI Y, et al. Antibacterial and antifungal activities of spices[J]. International Journal of Molecular Sciences, 2017, 18(6):1283.

DOI

[29]
KUMAR SINGH A, CABRAL C, KUMAR R, et al. Beneficial effects of dietary polyphenols on gut microbiota and strategies to improve delivery efficiency[J]. Nutrients, 2019, 11(9):2216.

DOI

[30]
TEETHAISONG Y, AUTARKOOL N, SIRICHAIWETCHAKOON K, et al. Synergistic activity and mechanism of action of stephania suberosa forman extract and ampicillin combination against ampicillin-resistant staphylococcus aureus[J]. Journal of Biomedical Science, 2014, 21(1):90.

DOI

[31]
SAKKAS H, PAPADOPOULOU C. Antimicrobial activity of basil,oregano,and thyme essential oils[J]. Journal of Microbiology and Biotechnology, 2017, 27(3):429-438.

DOI

[32]
TARIQ S, WANI S, RASOOL W, et al. A comprehensive review of the antibacterial,antifungal and antiviral potential of essential oils and their chemical constituents against drug-resistant microbial pathogens[J]. Microbial Pathogenesis, 2019, 134:103580.

DOI

[33]
高永生, 金斐, 朱丽云, 等. 植物精油及其活性成分的抗菌机理[J]. 中国食品学报, 2022, 22(1):376-388.

GAO Y S, JIN F, ZHU L Y, et al. Antimicrobial mechanism of plant essential oils and its active ingredients[J]. Journal of Chinese Institute of Food Science and Technology, 2022, 22(1):376-388. (in Chinese)

[34]
FERRONATO G, PRANDINI A. Dietary supplementation of inorganic,organic,and fatty acids in pig:a review[J]. Animals(Basel), 2020, 10(10):1740.

[35]
ZHANG X P, YANG Y N, ZHANG F, et al. Traditional Chinese medicines differentially modulate the gut microbiota based on their nature (Yao-Xing)[J]. Phytomedicine, 2021, 85:153496.

DOI

[36]
SUIRYANRAYNA M V A N, RAMANA J V. A review of the effects of dietary organic acids fed to swine[J]. Journal of Animal Science and Biotechnology, 2015, 6:45.

DOI PMID

[37]
ZHU W, ZHOU S X, LIU J H, et al. Prebiotic,immuno-stimulating and gut microbiota-modulating effects of lycium barbarum polysaccharide[J]. Biomedicine & Pharmacotherapy, 2020, 121:109591.

DOI

[38]
CAO Y, MA Z F, ZHANG H X, et al. Phytochemical properties and nutrigenomic implications of yacon as a potential source of prebiotic:current evidence and future directions[J]. Foods, 2018, 7(4):59.

DOI

[39]
KOLIDA S, GIBSON G R. Prebiotic capacity of inulin-type fructans[J]. The Journal of Nutrition, 2007, 137(Suppl.11):2503S-2506S.

DOI

[40]
MILUTINOVIĆ M, DIMITRIJEVIĆ-BRANKOVIĆ S, RAJILIĆ-STOJANOVIĆ M. Plant extracts rich in polyphenols as potent modulators in the growth of probiotic and pathogenic intestinal microorganisms[J]. Frontiers in Nutrition, 2021, 8:688843.

DOI

[41]
PENA R T, BLASCO L, AMBROA A, et al. Relationship between quorum sensing and secretion systems[J]. Frontiers in Microbiology, 2019, 10:1100.

DOI PMID

[42]
AZIMI S, KLEMENTIEV A D, WHITELEY M, et al. Bacterial quorum sensing during infection[J]. Annual Review of Microbiology, 2020, 74:201-219.

DOI PMID

[43]
ALAM K, FARRAJ D A A, MAH-E-FATIMA S, et al. Anti-biofilm activity of plant derived extracts against infectious pathogen-Pseudomonas aeruginosa PAO1[J]. Journal of Infection and Public Health, 2020, 13(11):1734-1741.

DOI

[44]
MAHESHWARI M, ABUL QAIS F, ALTHUBIANI A S, et al. Bioactive extracts of Carum copticum and thymol inhibit biofilm development by multidrug-resistant extended spectrum β-lactamase producing enteric bacteria[J]. Biofouling, 2019, 35(9):1026-1039.

DOI

[45]
SAKARIKOU C, KOSTOGLOU D, SIMÕES M, et al. Exploitation of plant extracts and phytochemicals against resistant Salmonella spp.in biofilms[J]. Food Research International, 2020, 128:108806.

DOI

[46]
NOAH T K, DONAHUE B, SHROYER N F. Intestinal development and differentiation[J]. Experimental Cell Research, 2011, 317(19):2702-2710.

DOI PMID

[47]
ALLAIRE J M, CROWLEY S M, LAW H T, et al. The intestinal epithelium:central coordinator of mucosal immunity[J]. Trends in Immunology, 2018, 39(9):677-696.

DOI

[48]
陈祥宇, 耿浩川, 侯建库, 等. 复合植物提取物对蛋鸡生产性能、肠道黏膜形态及盲肠菌群结构的影响[J]. 动物营养学报, 2021, 33(4):2033-2043.

DOI

CHEN X Y, GENG H C, HOU J K, et al. Effects of compound plant extracts on performance,intestinal mucosal morphology and cecal microflora structure of laying hens[J]. Chinese Journal of Animal Nutrition, 2021, 33(4):2033-2043. (in Chinese)

[49]
CHENG H, CHEN J F, TANG S G, et al. Effects of essential oil/palygorskite composite on performance,egg quality,plasma biochemistry,oxidation status,immune response and intestinal morphology of laying hens[J]. Poultry Science, 2022, 101(4):101632.

DOI

[50]
LONG S F, WANG Q Q, HE T F, et al. Maternal dietary Forsythia suspensa extract supplementation induces changes in offspring antioxidant status,inflammatory responses,intestinal development,and microbial community of sows[J]. Frontiers in Veterinary Science, 2022, 9:926822.

DOI

[51]
FARAHAT M, IBRAHIM D, KISHAWY A T Y, et al. Effect of cereal type and plant extract addition on the growth performance,intestinal morphology,caecal microflora,and gut barriers gene expression of broiler chickens[J]. Animal, 2021, 15(3):100056.

DOI

[52]
SUGIYAMA Y, NARA M, SAKANAKA M, et al. Comprehensive analysis of polyamine transport and biosynthesis in the dominant human gut bacteria:potential presence of novel polyamine metabolism and transport genes[J]. The International Journal of Biochemistry & Cell Biology, 2017, 93:52-61.

DOI

[53]
SONG H P, HOU X Q, LI R Y, et al. Atractylenolide I stimulates intestinal epithelial repair through polyamine-mediated Ca2+ signaling pathway[J]. Phytomedicine, 2017, 28:27-35.

DOI

[54]
RAO J N, LIU S V, ZOU T T, et al. Rac1 promotes intestinal epithelial restitution by increasing Ca2+ influx through interaction with phospholipase C-(gamma)1 after wounding[J]. American Journal of Physiology.Cell Physiology, 2008, 295(6):C1499-C1509.

[55]
ZENG D, HU C, LI R L, et al. Polysaccharide extracts of Astragalus membranaceus and Atractylodes macrocephala promote intestinal epithelial cell migration by activating the polyamine-mediated K+ channel[J]. Chinese Journal of Natural Medicines, 2018, 16(9):674-682.

DOI

[56]
SHI Y X, ZHU H B, LI R L, et al. Effect of polysaccharides from Sijunzi decoction on Ca2+ related regulators during intestinal mucosal restitution[J]. Phytomedicine, 2019, 58:152880.

DOI

[57]
ZHANG D, ZHU Y P, LI Z J, et al. The role of Astragalus polysaccharides in promoting IEC-6 cell migration from polyamine-mediated Ca2+ regulation[J]. International Journal of Biological Macromolecules, 2022, 207:179-192.

DOI

[58]
FAGUNDES R, TEIXEIRA L K. Cyclin E/CDK2:DNA replication,replication stress and genomic instability[J]. Frontiers in Cell and Developmental Biology, 2021, 9:774845.

DOI

[59]
KARIMIAN A, AHMADI Y, YOUSEFI B. Multiple functions of p21 in cell cycle,apoptosis and transcriptional regulation after DNA damage[J]. DNA Repair, 2016, 42:63-71.

DOI

[60]
涂小华, 杨光勇, 杨欣, 等. 四君子汤含药血清对IEC-6细胞增殖及RhoA、p21、Cdk2表达的影响[J]. 中成药, 2022, 44(7):2125-2131.

TU X H, YANG G Y, YANG X, et al. Effects of Sijunzi decoction-medicated serum on IEC-6 cell proliferation and RhoA,p21 and Cdk2 expressions[J]. Chinese Traditional Patent Medicine, 2022, 44(7):2125-2131. (in Chinese)

[61]
涂小华, 杨光勇, 杨欣, 等. 四君子汤含药血清对IEC-6细胞增殖及多胺调控NPM及p53表达的影响[J]. 中国药理学通报, 2021, 37(12):1762-1769.

TU X H, YANG G Y, YANG X, et al. Effects of Sijunzi decoction-containing serum on IEC-6 cell proliferation and expression of NPM and p53 regulated by polyamines[J]. Chinese Pharmacological Bulletin, 2021, 37(12):1762-1769. (in Chinese)

[62]
SONG H P, LI R L, CHEN X, et al. Atractylodes macrocephala Koidz promotes intestinal epithelial restitution via the polyamine-voltage-gated K+ channel pathway[J]. Journal of Ethnopharmacology, 2014, 152(1):163-172.

DOI

[63]
BOLAMPERTI S, GUIDOBONO F, RUBINACCI A, et al. The role of growth hormone in mesenchymal stem cell commitment[J]. International Journal of Molecular Sciences, 2019, 20(21):5264.

DOI

[64]
TEDESCHI L, VASSALLE C, IERVASI G, et al. Main factors involved in thyroid hormone action[J]. Molecules, 2021, 26(23):7337.

DOI

[65]
李焰, 杨小燕, 林跃鑫. 银杏叶提取物对肉鸡屠宰性能、血清生化指标和甲状腺激素浓度的影响[J]. 中国畜牧杂志, 2007, 43(23):24-26.

LI Y, YANG X Y, LIN Y X. Effects of ginkgo biloba extract on slaughtering performance and serum biochemical indexes of broiler chickens and the effect of thyroid hormone concentrations[J]. Chinese Journal of Animal Science, 2007, 43(23):24-26. (in Chinese)

[66]
徐之勇, 陈小盘, 汪文阁, 等. 银杏叶提取物对雏鸭生产性能及血清中生长激素、甲状腺素含量的影响[J]. 黑龙江畜牧兽医, 2015, 487(19):206-208.

XU Z Y, CHEN X P, WANG W G, et al. Effects of Ginkgo biloba extract on production performance and serum growth hormone and thyroxine content in ducklings[J]. Heilongjiang Animal Science and Veterinary Medicine, 2015, 487(19):206-208.. (in Chinese)

[67]
LEE D H, LEE S H, CHO N, et al. Effects of eleutherococcus extract mixture on endochondral bone formation in rats[J]. International Journal of Molecular Sciences, 2019, 20(5):1253.

DOI

[68]
LEE D H, LEE S H, LEE Y H, et al. Astragalus extract mixture HT042 increases longitudinal bone growth rate by upregulating circulatory IGF-1 in rats[J]. Evidence-Based Complementary and Alternative Medicine, 2017, 2017:6935802.

[69]
LEE D H, KIM B H, LEE S H, et al. Effects of astragalus extract mixture HT042 on circulating IGF-1 level and growth hormone axis in rats[J]. Children(Basel), 2021, 8(11):975.

[70]
KIM M Y, KIM J Y, LIM D, et al. Skeletal growth and IGF levels in rats after HT042 treatment[J]. Phytotherapy Research, 2012, 26(12):1771-1778.

DOI

[71]
VON ROSENBERG S, WEHR U, BACHMANN H. Effect of vitamin D-containing plant extracts on osteoporotic bone[J]. The Journal of Steroid Biochemistry and Molecular Biology, 2007, 103(3/4/5):596-600.

DOI

[72]
DRORI A, SHABAT Y, BEN YA’ACOV A, et al. Extracts from Lentinula edodes (shiitake) edible mushrooms enriched with vitamin D exert an anti-inflammatory hepatoprotective effect[J]. Journal of Medicinal Food, 2016, 19(4):383-389.

DOI

[73]
BOUILLON R, MARCOCCI C, CARMELIET G, et al. Skeletal and extraskeletal actions of vitamin D:current evidence and outstanding questions[J]. Endocrine Reviews, 2019, 40(4):1109-1151.

DOI

[74]
DUSSO A S, BROWN A J, SLATOPOLSKY E. Vitamin D[J]. American Journal of Physiology-Renal Physiology, 2005, 289(1):F8-F28.

DOI PMID

[75]
GOLTZMAN D. Functions of vitamin D in bone[J]. Histochemistry and Cell Biology, 2018, 149(4):305-312.

DOI PMID

[76]
FENG X, LYU Y, WU Z H, et al. Fructus ligustri lucidi ethanol extract improves bone mineral density and properties through modulating calcium absorption-related gene expression in kidney and duodenum of growing rats[J]. Calcified Tissue International, 2014, 94(4):433-441.

DOI

[77]
PONZETTI M, RUCCI N. Osteoblast differentiation and signaling:established concepts and emerging topics[J]. International Journal of Molecular Sciences, 2021, 22(13):6651.

DOI

[78]
KOMORI T. Regulation of osteoblast differentiation by transcription factors[J]. Journal of Cellular Biochemistry, 2006, 99(5):1233-1239.

PMID

[79]
KIM S W, KIM H G, LEE B E, et al. Effects of mushroom,Pleurotus eryngii,extracts on bone metabolism[J]. Clinical Nutrition, 2006, 25(1):166-170.

DOI

[80]
WANG Z, WANG D, YANG D, et al. The effect of icariin on bone metabolism and its potential clinical application[J]. Osteoporosis International, 2018, 29(3):535-544.

DOI PMID

[81]
CHEN C H, HO M L, CHANG J K, et al. Green tea catechin enhances osteogenesis in a bone marrow mesenchymal stem cell line[J]. Osteoporosis International, 2005, 16(12):2039-2045.

DOI

[82]
TAKADA I, KOUZMENKO A P, KATO S. Wnt and PPARgamma signaling in osteoblastogenesis and adipogenesis[J]. Nature Reviews Rheumatology, 2009, 5(8):442-447.

DOI PMID

[83]
KO C H, SIU W S, WONG H L, et al. Pro-bone and antifat effects of green tea and its polyphenol,epigallocatechin,in rat mesenchymal stem cells in vitro[J]. Journal of Agricultural and Food Chemistry, 2011, 59(18):9870-9876.

DOI

[84]
LI X D, WANG J S, CHANG B, et al. Panax notoginseng saponins promotes proliferation and osteogenic differentiation of rat bone marrow stromal cells[J]. Journal of Ethnopharmacology, 2011, 134(2):268-274.

DOI

[85]
LI X D, CHANG B, CHEN B, et al. Panax notoginseng saponins potentiate osteogenesis of bone marrow stromal cells by modulating gap junction intercellular communication activities[J]. Cellular Physiology and Biochemistry, 2010, 26(6):1081-1092.

DOI

[86]
CHEN B, LI X D, LIU D X, et al. Canonical Wnt signaling is required for Panax notoginseng saponin-mediated attenuation of the RANKL/OPG ratio in bone marrow stromal cells during osteogenic differentiation[J]. Phytomedicine, 2012, 19(11):1029-1034.

DOI

Options
文章导航

/

〈 〉

    联系我们

  • 地址：北京市海淀区圆明园西路2号
    　　　中国农业大学动科动医大楼153室
  • 邮编：100193
  • 电话：010－62817823
  • 网址：http://www.chinajan.com
  • E-mail：cjan@chinajan.com
京公网安备 11010802041926号    京ICP备17050989号-2