Chinese Journal of Animal Nutrition >

2024 , Vol. 36 >Issue 10: 6212 - 6222

DOI: https://doi.org/10.12418/CJAN2024.529

REVIEW

Biological Functions of N-Carbamylglutamate and Its Application in Animal Production

  • YAN Chen , 1, 2 ,
  • PENG Kai 1 ,
  • CAO Junming 1, 3 ,
  • ZHAO Hongxia , 1, *
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  • 1 Collaborative Innovation Center of Aquatic Sciences, Key Laboratory of Animal Nutrition and Feed in South China of Ministry of Agriculture and Rural Affairs, Guangdong Key Laboratory of Animal Breeding and Nutrition, Institute of Animal Science, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China
  • 2 College of Fisheries of Huazhong Agricultural University, Wuhan 430000, China
  • 3 Institute of Animal Health, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China
*professor, E-mail:

Received date: 2024-04-02

  Online published: 2024-10-14

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Abstract

As a kind of green functional additives, N-carbamoylglutamate (NCG) can adjust the intestinal barrier function, improve intestinal morphology, enhance digestion ability; improve antioxidant enzyme activities, enhance intestinal and liver antioxidant capacity; regulate the expression of pro-inflammatory factors and anti-inflammatory factors, alleviate inflammatory response, and enhance the body’s immunity; regulate body metabolism, improve mitochondrial function, and reduce fat deposition; promote the development of maternal gonads and the growth of juveniles, improve the reproductive performance of animals. Based on a review of relevant literature at home and abroad, this review summarized the biological functions of NCG in regulating intestinal barrier, body metabolism, immunity, antioxidant capacity, reproductive performance, and its application the inbreeding of pigs, poultry, ruminants and aquatic animals, so as to provide reference for the research and application of NCG in animal production.

Key words： N-carbamylglutamate; physiological function; intestinal barrier; metabolism; animal production

Cite this article

YAN Chen , PENG Kai , CAO Junming , ZHAO Hongxia . Biological Functions of N-Carbamylglutamate and Its Application in Animal Production[J]. Chinese Journal of Animal Nutrition, 2024 , 36(10) : 6212 -6222 . DOI: 10.12418/CJAN2024.529

精氨酸是产生一氧化氮、多胺的底物,参与蛋白质、肌酸等的合成,在调控机体营养代谢和免疫反应中起重要作用[1-2]。但是,精氨酸成本较高,而直接在饲粮中补充晶体精氨酸可能会产生氨基酸拮抗效应,而且精氨酸易被机体内精氨酸酶降解。N-氨甲酰谷氨酸(NCG)与N-乙甲酰谷氨酸(NAG)的作用类似,可激活氨基甲酰磷酸合成酶-1(CPS-1),促进内源性精氨酸的合成[3]。研究显示,作为精氨酸内源激活剂,NCG可提高生长性能[4],改善肠道功能和免疫力[4⇓-6],提高肝脏和血浆的抗氧化能力[7-8]。本文综述了NCG的生物学功能及其在动物养殖中应用的研究进展,以期为NCG在动物生产中的应用提供参考。

1 NCG的结构和特性

NCG分子式为C6H10N2O5,相对分子质量为190.15,纯品为无色透明晶体。NCG不溶于水,微溶于有机溶剂,活性物质等电点为3.02,其盐类溶于水。NCG能激活线粒体中CPS-1和吡咯啉-5-羧酸合成酶(P5CS),促进谷氨酰胺和脯氨酸合成瓜氨酸,进而促进精氨酸内源合成[9-10]。

2 NCG的生物学功能

2.1 调控肠道屏障

肠道作为动物机体的重要器官,不仅具有消化和吸收营养物质的功能,还能作为免疫系统来抵御各种疾病。肠道屏障是指肠道防止肠腔内的有害物质穿过肠黏膜进入体内其他组织器官和血液循环的结构与功能的总和,由机械屏障、化学屏障、微生物屏障和免疫屏障组成[11]。NCG通过改善肠道结构、提高肠道菌群丰度、增强肠道抗氧化能力和免疫水平、降低炎症反应等保持肠道屏障功能的稳定。

2.1.1 NCG对肠道机械屏障的影响

肠道机械屏障由肠黏膜上皮细胞及紧密连接蛋白组成。NCG对肠道机械屏障的影响表现在以下2个方面:1)增加肠黏膜杯状细胞数[12],改善肠道绒毛高度、隐窝深度及绒毛高度/隐窝深度值,提高肌层厚度和绒毛宽度,增强肠道的吸收能力[13];2)上调肠道中闭合蛋白(occludin)、密封蛋白(claudin)和闭锁小带蛋白(ZO)的表达,降低肠道通透性,保持肠道的屏障功能[14]。研究显示,NCG能增加断奶仔猪肠道绒毛高度及绒毛高度/隐窝深度值,改善空肠形态[15]。

2.1.2 NCG对肠道化学屏障的影响

肠道的化学屏障主要包括覆盖在肠上皮起保护和润滑作用的以黏蛋白为主的黏液和消化液等。研究显示,NCG能提高杂交鳢(Channa maculata♀×Channa argus♂)肠道淀粉酶(AMS)、胰蛋白酶、脂肪酶、Na+-K+-ATP酶活性,进而提高营养物质消化率[16-17]。NCG能升高黄鲶鱼肠道精氨酸脱羧酶、鸟氨酸脱羧酶、二胺氧化酶活性和一氧化氮含量,进而促进营养转化[18]。

2.1.3 NCG对肠道微生物屏障的影响

肠道微生物屏障主要为益生菌与肠黏膜黏附结合形成的菌膜屏障,肠道内的菌群与机体共生,维持肠道内的微生态平衡[19]。研究显示,NCG能改善肠道微生物组成及代谢产物,改善肠道环境:NCG可显著提高杂交鳢肠道厚壁菌门及变形菌门的丰度[13];增加杂交绵羊后肠毛螺菌属的丰度,降低毛梭菌属的丰度,改变后肠短链脂肪酸(SCFA)含量,增加丙酸含量,减少乙酸和己酸含量[20];宫内发育迟缓(IUGR)羔羊补充NCG升高了肠道梭菌、乳酸菌和链球菌的相对丰度,降低葡萄球菌的相对丰度,升高乙酸、丁酸和丙酸含量[14]。

2.1.4 NCG对肠道免疫屏障的影响

肠道免疫屏障主要由淋巴细胞、分泌性免疫球蛋白A(sIgA)等抗菌物质和淋巴组织组成,其中起核心作用的是sIgA。sIgA由淋巴细胞和浆细胞产生,对保护肠黏膜起重要作用,能够结合病原体和毒素,与补体和溶菌酶共同发挥作用[21]。NCG保护肠道免疫屏障包含3种主要途径:1)通过淋巴细胞和促炎因子表达激活肠道免疫系统提高免疫力。研究显示,CD4+作为T淋巴细胞在免疫反应中起重要作用,而sIgA是由固有层内淋巴细胞数量决定的,且sIgA分泌细胞严重依赖于活化的T细胞分泌的一些细胞因子,如白细胞介素-10(IL-10)[22];NCG通过刺激淋巴细胞增殖和细胞因子合成,提高大肠杆菌攻击新生仔猪回肠sIgA、CD4+浓度和γ-干扰素(IFN-γ)、白细胞介素-2(IL-2)、白细胞介素-4(IL-4)和IL-10含量,提高肠道黏膜的免疫功能[23]。2)通过提高抗氧化酶活力、抑制细胞凋亡改善抗氧化能力。研究显示,NCG能提高大鼠空肠中过氧化氢酶(CAT)活性和谷胱甘肽(GSH)含量,降低丙二醛(MDA)含量[5];提高断奶仔猪空肠中总抗氧化能力(T-AOC)、超氧化物歧化酶(SOD)活性,上调空肠中CAT、SOD1、谷胱甘肽过氧化物酶4(Gpx4)、谷氨酸半胱氨酸连接酶催化亚单位(GCLC)、谷氨酸半胱氨酸连接酶修饰亚单位(GCLM)和芳香烃受体(AhR)、细胞色素P450 1A1(CYP1A1)mRNA的表达[15];提高IUGR羔羊空肠中诱导型一氧化氮合酶(iNOS)、内皮型一氧化氮合酶(eNOS)mRNA和蛋白质丰度并通过一氧化氮(NO)依赖途径调节抗氧化酶活性,改善氧化还原状态[24];通过抑制IUGR羔羊死亡受体依赖的凋亡途径和线粒体凋亡途径来抑制内质网应激诱导的细胞凋亡和氧化损伤[25]。3)下调促炎因子及细胞凋亡相关基因表达降低炎症反应。研究显示,胱天蛋白酶(Caspase)家族成员通过激活启动子型Caspase在生物过程中对细胞凋亡的调节起重要的调节作用[26];NCG通过下调炎症相关基因肿瘤坏死因子-α(TNF-α)、白细胞介素-1β(IL-1β)和白细胞介素-8(IL-8)以及Caspase家族和转化生长因子-β1(TGF-β1)的mRNA水平缓解炎症反应[27],减少IUGR羔羊髓样分化因子88(MyD88)、核因子-κB(NF-κB)和Toll样受体4(TLR4)表达[28];NCG调节Treg细胞分泌TGF-β1、IL-10、信号转导和转录激活因子5(STAT5),调节Th17细胞分化,改善Treg细胞功能,减轻脂多糖(LPS)诱导的肠道炎症[29];中性粒细胞(PMN)是机体抵抗病原菌入侵的主要防线,NCG能增加淋巴细胞数量和增强PMN功能,减轻奶牛的炎症和氧化应激[30]。

2.2 调节机体代谢

2.2.1 调节脂质代谢

细胞外信号调节激酶(ERK)活化促进关键脂肪形成调节因子CCAAT-增强子结合蛋白(C/EBP)α、β和δ的表达[31],哺乳动物雷帕霉素靶蛋白(mTOR)能催化蛋白激酶B(AKT)、胰岛素样生长因子-1受体(IGF-1R)、核糖体蛋白S6激酶(S6K)和转录因子EB(TFEB)的磷酸化[32]。NCG抑制ERK1/2-mTOR-S6K1信号通路的磷酸化,下调脂肪细胞分化基因C/EBPα的表达,提高脂肪分解基因甘油三酯脂肪酶(ATGL)的mRNA水平,降低血浆极低密度脂蛋白(VLDL)和肝脏甘油三酯(TG)及游离脂肪酸(NEFA)的积累[27]。NCG通过上调蛋鸡肝脏肉碱棕榈酰转移酶1A(CPT1A)的表达调节腺苷酸活化蛋白激酶(AMPK)信号通路,促进脂肪酸进入线粒体进行β-氧化,下调肝脏脂肪酸合成酶(FASN)的表达,调节过氧化物酶体增殖物激活受体(PPAR)信号通路,促进脂解引起脂肪酸增加,实现对肝脏脂质代谢的调节;此外,NCG 还能增加血清和肝脏中NO的水平,增加蛋黄中多不饱和脂肪酸(PUFA)含量[33]。在生长育肥猪上的研究显示,NCG上调了肌肉中脂质代谢相关基因激素敏感脂肪酶(HSL)、乙酰辅酶A羧化酶(ACC)和过氧化物酶体增殖物激活受体γ辅助活化因子-1α(PGC-1α)的表达水平,提高肌肉C18∶1n9c和单不饱和脂肪酸(MUFA)含量[34]。日本鲈鱼体内和体外前脂肪细胞试验结果均表明,NCG可降低FASN、胆固醇合成相关基因羟甲基戊二酰辅酶A还原酶(HMGCR)、脂蛋白脂酶(LPL)和瘦素的mRNA水平,减少脂肪沉积[35]。在满足50% NRC营养要求的饲料中添加NCG能降低IUGR湖羊胎儿血清中TG、NEFA、胆固醇和β-羟基丁酸水平,改善脂代谢[36]。NCG处理的獭兔通过上调血清中生长激素(GH)和胰岛素样生长因子-1(IGF-1)水平、升高NO合成酶活性、降低肝脏FASN活性、降低肾周和皮下脂肪百分比、血清TG含量,抑制肝脏脂肪生成,减少脂肪沉积[37]。

2.2.2 调节能量代谢

AMPK是细胞和全身能量稳态的关键调节因子[38],PGC-1α可以调节线粒体的生物发生和功能[39]。给IUGR哺乳湖羊羔羊补饲NCG能抑制AMPK-PGC1α-线粒体转录因子A(TFAM)通路,降低PGC-1α和TFAM表达,提高ATP含量以及异柠檬酸脱氢酶、柠檬酸合成酶和α-酮戊二酸脱氢酶复合物活性,促进三羧酸循环,进而改善肝脏能量状态和线粒体功能[40⇓-42]。

2.2.3 调节氨氮代谢

尿素循环有5个关键酶参与,分别为CPS-1、鸟氨酸转氨甲酰酶(OTC)、氨基琥珀酸合成酶(ASS)、氨基琥珀酸裂解酶(ASL)和精氨酸酶(ARG)。研究显示,NCG能提高黄颡鱼(Pelteobagrus fuluidraco)幼鱼肝脏中ASS和ASL活性,增加尿素循环效率,降低血清氨、尿素浓度,减轻氨应激的影响[43]。NCG能增加大鼠尿液中肌酐、乙醇、乳酸盐和肌氨酸的水平,降低乙酰胺、乙酸盐、瓜氨酸、肌酸、甘氨酸、N-乙酰谷氨酸盐、苯乙酰甘氨酸、丙酮和对羟基苯乙酸盐的水平,改变与氮代谢和肠道微生物群代谢相关的尿液代谢组[44]。NCG还能升高荷斯坦牛血浆中NO和精氨酸水平,降低粪氮、尿氮含量,促进饲粮中蛋白质向乳蛋白的转化,提高氮利用率,改善氮代谢[10,45]。

2.2.4 改善氨基酸转运及合成

视黄醇结合蛋白1(RBP1)是将视黄醇从肝脏储存转运至组织的载体蛋白,NCG能上调IUGR哺乳羔羊RBP1表达,促进维生素A对生殖的调节,从而提高蛋鸡的生产性能;同时,NCG还能上调钙转运相关基因兰尼碱受体2(RYR2)和L-型电压依赖型钙通道β2(CACNB2)表达以及下调受体活性修饰蛋白3(RAMP3)及SH3和富含半胱氨酸结构域的蛋白(STAC)表达,改善蛋壳厚度[20]。NCG通过提高氨基酸转运蛋白溶质载体家族38成员2(SNAT2)、溶质载体家族1成员1(EAAC1)、溶质载体家族3成员1(SLC3A1)和溶质载体家族3成员2(SLC3A2)mRNA的表达改善与鸟氨酸循环相关的氨基酸的空肠转运[15]。NCG能够增加IUGR哺乳羔羊回肠中磷酸化哺乳动物雷帕霉素靶蛋白(pmTOR)/mTOR比值,通过调控mTOR信号转导以及溶质载体家族1成员5(SLC1A5)和肽转运蛋白1的表达来影响肠道氨基酸谱[46-47]。

2.3 调节免疫和抗氧化能力

研究表明,NCG能下调花鲈炎症相关因子IL-1β及凋亡因子Caspase-3、Caspase-8的mRNA水平,上调抗炎因子IL-10的mRNA水平,降低炎症和凋亡细胞因子的表达[35]。Wang等[48]发现,NCG能提高镜鲤(Cyprinus carpio)肠道溶菌酶活性,降低促炎因子TNF-α、IL-1β和IL-8的mRNA表达。p53作为细胞代谢波动的调节剂[49],可以通过内源性B淋巴细胞瘤-2(Bcl-2)或Bcl-2关联X蛋白(Bax)和外源性肿瘤坏死因子受体超家族成员6(Fas)或Fas配体(FasL)促进细胞凋亡[50-51]。NCG能提高营养不良母羊IUGR胎儿脐血中免疫球蛋白A(IgA)、IL-1β和IL-10水平,降低Bax、Fas和p53 mRNA表达和Caspase-3活性,增加Bcl-2 mRNA表达,改善胸腺中的细胞凋亡和免疫功能,促进胎儿生长[52]。
研究显示,饲料添加0.03% NCG能显著提高杂交鳢血浆中过氧化氢酶(CAT)和过氧化物酶活性[16],NCG能提高中间球海胆(Strongylocentrotus intermedius)体腔液中SOD和CAT活性[53]。Zhao等[18]的试验结果表明,在黄颡鱼饲料添加0.025% NCG能显著提高血清谷胱甘肽过氧化酶(GSH-Px)、葡萄糖过氧化物酶活性和T-AOC。Cao等[8]研究了NCG对大鼠的作用,结果表明,NCG能提高肝脏和血浆T-AOC,增加GSH含量,降低MDA含量。

2.4 提高繁殖性能

NCG通过提高A型子叶iNOS和溶质载体家族15成员1(SLC15A1)的mRNA表达水平上调转运蛋白、增强营养物质转运和改善妊娠母羊胎盘氨基酸运输,通过减少孕酮受体(PGR)的mRNA表达水平降低母体血清中的17β-雌二醇和孕酮水平,通过减少成纤维细胞生长因子2(FGF2)的mRNA表达水平降低A型子叶毛细血管面积密度(CAD)和毛细血管表面密度(CSD),使胎儿得到更好的发育[54]。NCG能通过抑制IGF-1和促卵泡生成素(FSH)诱导雌二醇和孕酮产生,并刺激颗粒细胞(GC)增殖,减缓卵泡分化,调节卵巢功能[55-56];NCG还能降低血清皮质醇浓度,提高瘦素和甲状腺素(T4)的浓度,改善母羊的母体和胎盘发育[57]。NCG能上调后备母猪绒毛膜尿囊中内皮型一氧化氮合酶(eNOS)、血管内皮生长因子-A(VEGF-A)、胎盘生长因子(PLGF)和血管生成素-2(ANG-2)基因表达,改善母猪妊娠结局[58]。解偶联蛋白-1(UCP-1)可调节线粒体内膜棕色脂肪组织(BAT)氧化还原[59],骨成型蛋白7(BMP7)可调控PR结构域蛋白16(PRDM16)基因表达,激活棕色脂肪形成[60]。NCG能提高精氨酸家族氨基酸和葡萄糖生成氨基酸的浓度,促进骨桥蛋白(OPN)和整联蛋白αv、β3 mRNA的表达,促进内蒙古白绒山羊子宫内膜的发育;同时,NCG还能促进UCP-1和BMP7的mRNA表达,改善胎儿BAT的储存,从而有助于胎儿的早期发育[61]。NCG诱导母猪子宫内膜蛋白质丰度改变,增加整联蛋白、踝蛋白(talin)和eNOS表达,促进胚胎发育[62]。蛋白激酶-Ⅰ(PKG-Ⅰ)能被环鸟苷一磷酸活化,改善促分裂原活化蛋白激酶级联(ERK1/2和p38)的磷酸化,增加内皮细胞的增殖、迁移和分化,促进血管生成[63]。NCG能促进牦牛卵泡发育,增加大卵泡(直径>10 mm)的数量,同时通过增强卵巢类固醇激素合成促进雌激素合成,并调节戊糖磷酸途径和三羧酸循环等葡萄糖代谢相关途径[64];NCG可上调鸡PKG-Ⅰ、Raf1和p-p38的表达,增加卵巢血管生成和细胞增殖[65]。NCG可增加蛋鸡子宫血管数量、子宫黏膜皱襞高度、子宫内膜绒毛长度和子宫黏膜皱襞面积[66]。NCG可上调血清代谢物包括阿魏酸、肉桂酰甘氨酸、3-苯乳酸、和γ-谷氨酰谷氨酸的浓度,改善母猪活产仔猪数量和仔猪出生体重,提高繁殖性能[67]。NCG通过提高精氨酸、谷氨酰胺、谷氨酸和脯氨酸浓度诱导磷酸化蛋白激酶B(p-PKB)、磷酸化核糖体蛋白S6激酶B1(p-S6K1)及磷酸化信号转导和转录激活因子3(p-STAT3)表达,激活PI3K/Akt/mTOR信号通路与信号转导和转录激活因子3(STAT3)磷酸化,提高细胞黏附活性,改善大鼠妊娠结局[68]。高浓度NCG通过抑制小鼠下丘脑神经元细胞GT1-7促性腺激素释放激素(GnRH)和神经元型一氧化氮合酶(nNOS)的mRNA的表达以及下调与GnRH合成和分泌相关基因的mRNA表达来直接和间接调节GnRH,从而影响性腺发育[69]。

3 NCG在动物生产中的应用

3.1 在水产动物生产中的应用

研究表明,NCG能显著提高大菱鲆(Scophthalmus maximus)幼鱼的增重率、特定生长率和蛋白质效率,降低饲料系数和内脏指数[70];而在花鲈(Lateolabrax japonicus)[71]和杂交鳢[16]上的研究结果显示,饲料适量添加NCG能提高蛋白质沉积率,对鱼体增重率、特定生长率无显著影响;在尼罗罗非鱼(Oreochromis niloticus)幼鱼饲料中添加0.2% NCG,幼鱼的增重率和蛋白质效率显著升高,其他添加量的NCG对生长性能无显著影响[72];Li等[73]研究发现,NCG促进杂交鲟(Acipenser schrenckii ♀×A. baerii ♂)的生长,鱼体终末重、增重率和蛋白质效率显著升高,Wang等[48]在镜鲤上得到相同结论。研究表明,饲料中适量添加NCG能显著提高大菱鲆幼鱼肌肉精氨酸含量和全鱼粗蛋白质含量,说明NCG有助于精氨酸内源合成,促进蛋白质合成[74]。在饲料中适量添加NCG能降低花鲈腹脂率,提高脂肪沉积率,说明NCG可能使脂肪向肌肉转移[71]。NCG能显著降低罗非鱼的肝体比,显著提高全鱼脂肪含量,脂肪有由肝脏转移至肌肉的倾向[72]。上述研究结果表明,饲料中适量添加NCG可显著改善鱼体蛋白质含量和脂肪分布,提高鱼类生长性能。

3.2 在猪生产中的应用

Feng等[67]试验结果表明,饲粮中添加0.05% NCG可通过增加血清NCG浓度、改变肠道微生物组和血清代谢物改善活产仔猪数量和出生体重变异系数,说明NCG可以通过改善母猪的活产仔猪数和仔猪出生体重均匀性来提高繁殖性能。对仔猪进行口服给药试验,发现50 mg/kg BW NCG能提高平均日增重和血清精氨酸、谷氨酸、瓜氨酸和鸟氨酸浓度,增加空肠乳糖酶活性和盲肠乳杆菌属数量[6]。Ye等[75]研究了低蛋白质饲粮中添加NCG对肥育猪胴体性状和肉质的影响,发现NCG能增加肥育猪背最长肌面积,减少背膘生长,可生产高亮氨酸含量的功能性猪肉。Wang等[76]在生长猪饲粮中添加0.1% NCG后发现,粗蛋白质、精氨酸、组氨酸、亮氨酸、苯丙氨酸、缬氨酸、丝氨酸和酪氨酸的消化率提高,表明NCG能促进肠道消化。Hu等[15]研究发现,饲粮中添加0.05% NCG能改善断奶仔猪的终末体重和平均增重,提高仔猪生长性能。上述研究结果表明,饲料中添加NCG能改善猪的卵泡功能、肠道形态和消化能力,提高了猪的生长性能和繁殖能力。

3.3 在反刍动物生产中的应用

Sun等[77]对湖羊胎儿脐静脉血浆代谢谱进行分析后发现,氨基酸、碳水化合物和能量、脂质以及氧化应激代谢途径发生了改变,表明NCG能参与调控机体代谢。Zhou等[64]研究了饲粮添加NCG对牦牛卵泡发育的影响,发现NCG能增强卵巢类固醇激素合成,调节葡萄糖代谢。Zhu等[78]对NCG对西藏荷斯坦奶牛瘤胃发酵参数及细菌群落多样性的影响进行了研究,发现NCG可以提高瘤胃氮利用率和总挥发性脂肪酸含量,增加瘤胃微生物多样性,提高厌氧菌Patescibacteria的相对丰度,使荷斯坦奶牛更好地适应低氧环境,Li等[79]的研究结果与之相似。Li等[79]研究发现,NCG能降低热应激奶牛瘤胃微生物群中变形菌门和放线菌门的丰富度和多样性,提高瘤胃微生物蛋白、挥发性脂肪酸含量和丙酸盐摩尔比,降低瘤胃pH、氨氮含量和乙酸/丙酸比值,提高氮的利用率。上述研究结果表明,NCG能改变反刍动物瘤胃微生物组成和代谢产物,改善瘤胃的发酵功能。

3.4 在家禽生产中的应用

Ma等[80]研究发现,饲粮中添加NCG可调节公鸡睾丸细胞色素P450的表达,促进体内促性腺激素释放激素和睾酮的合成,进而促进性腺和曲细精管的发育。维生素A被认为在卵巢类固醇生成、卵母细胞成熟和生殖道黏膜上皮发育中起关键作用[81]。Ma等[82]研究发现,0.08% NCG能调节蛋鸡子宫维生素A代谢,并能调节子宫功能,从而提高产蛋率和蛋品质。NO与脂质代谢存在密切联系,NO通过激活大鼠胆固醇代谢的关键因子肝固醇调节元件结合蛋白-2和增加低密度脂蛋白受体的表达[83],以及通过可逆的蛋白质S-亚硝基化对线粒体脂肪酸代谢发挥调节作用[84]。Liu等[33]研究发现,在蛋鸡饲粮中添加0.08% NCG能提高血清和肝脏中NO含量,调节肝脏脂质代谢,生产富含PUFA的功能性鸡蛋。Ma等[66]研究发现,饲粮中添加1% NCG显著提高了蛋鸡的蛋壳厚度、蛋壳强度、蛋壳百分率、蛋壳比重和蛋壳有效厚度,显著降低了乳突层有效厚度、乳突间隙宽度和乳头结节,表明NCG能够改善蛋壳质量。上述研究结果表明,饲料中添加NCG能促进家禽性腺发育,改善蛋鸡的生产性能、鸡蛋营养和蛋壳质量,提高鸡蛋品质。

4 小结

NCG作为一种功能性饲料添加剂,具有价格低、添加量少等优点,在动物生产上具有广泛应用前景。研究表明,NCG能改善肠道绒毛高度、隐窝深度、肌层厚度和紧密连接蛋白的表达,提高淀粉酶、脂肪酶等消化酶活性,增加肠道菌群中有益菌的丰度,调节肠道屏障功能;提高CAT、SOD等抗氧化酶活性,增强肠道和肝脏抗氧化能力;上调抗炎因子的表达,降低促炎因子和凋亡细胞因子的表达,增强机体免疫力,缓解炎症反应;上调脂解相关基因表达,减少脂肪沉积,促进三羧酸循环,改善肝脏能量状态;促进母体卵巢、胎盘和幼体发育,调节激素分泌,提高动物繁殖性能。目前,NCG在动物上的研究主要集中于畜禽和鱼类,其发挥生物学功能的具体作用机理尚未充分阐明,在这些过程中涉及的酶、激素、转录因子和信号通路还需要进一步地研究并阐明。此外,NCG对不同养殖动物、不同生长阶段、不同环境条件下的适宜添加量还有待于明确,其对动物吸收、代谢、生理功能的作用及机制还需要深入探索。

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