CN108935981A - 围产期奶山羊添加烟酰胺对羔羊糖脂代谢的影响及其机制 - Google Patents
围产期奶山羊添加烟酰胺对羔羊糖脂代谢的影响及其机制 Download PDFInfo
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Abstract
本发明公开了围产期奶山羊添加烟酰胺对羔羊糖脂代谢的影响及其机制,从母体添加烟酰胺入手,研究其对羔羊肠道形态发育及养分转运载体表达、糖代谢和血液生化指标以及脂肪代谢的影响及机制,研究发现围产期奶山羊添加烟酰胺可以提高羔羊肠道对葡萄糖吸收,抑制羔羊肝脏糖异生,促进肝脏脂肪分解,减少肝脏损伤;也可促进羔羊腹脂脂肪沉积和肠道形态发育,促进羔羊生长发育,研究成果为调节羔羊生长发育提供理论依据。
Description
技术领域
本发明属于动物营养领域,具体涉及一种围产期奶山羊添加烟酰胺对子代羔羊糖脂代谢的影响。
背景技术
围产期是奶畜的特殊生理时期,包括妊娠后期和泌乳前期,一般指产前21天至产后21天。孕期和泌乳高峰期奶畜的胎盘、乳腺、脂肪、肌肉、肝脏和胃肠道等组织、器官的代谢会发生巨大改变,以便为母子营养分配和胎儿发育供应葡萄糖和氨基酸,为肝脏和乳腺提供乙酸、丙酸、丁酸等游离脂肪酸(free fatty acid,FFA)用以糖异生和脂肪合成以满足泌乳能量和脂肪需要。胎儿营养和内分泌状态发生改变可能会引起子代代谢、结构、生理的变化,且变化是永久和可遗传的,甚至会引起子代疾病的发生(Metcoff et al.1980)。胎儿和新生儿的环境和营养效应甚至影响成年后的生产性能。分娩前6周,胎儿快速增重,且母体是胎儿的唯一营养来源。所以,妊娠后期保证母体营养对胎儿发育至关重要。有研究表明,母羊分娩前1月添加维生素A促进提高羔羊成活率和生产性能(Eldaim et al.2015);妊娠期母羊添加维生素E降低死胎率(Donnem et al.2015);妊娠后期和泌乳早期对母羊进行补饲,提高羔羊初生重和出生后60天体重(Idris et al.2010)。因此,围产前期和泌乳早期母体营养对于胎儿及出生后早期生长发育至关重要,保证母体围产前期和泌乳早期营养充足可促进子代生长发育,提高生产性能。
烟酸和烟酰胺是维生素B3的两种存在形式,两者在体内可互相转换,烟酰胺是烟酰胺腺嘌呤二核苷酸(NAD+)和烟酰胺腺嘌呤二核苷酸磷酸(NADP+)的主要前体物之一。烟酸主要来源于植物,烟酰胺主要来源于动物,但谷物中烟酸主要以结合型状态存在,利用效率低。反刍动物瘤胃中烟酸的主要来源:(1)饲料中烟酸;(2)瘤胃微生物合成烟酸;(3)色氨酸在肝脏中通过犬尿酸途径转化为烟酸,但转化效率较低,且色氨酸属于必需氨基酸,此部分所占比重较小。
目前关于烟酰胺在糖代谢方面的研究有:围产期奶牛补充烟酸可显著降低肝脏葡萄糖转运载体2(glucose transporter,GLUT2)的表达(Kinoshita et al.2016);以肥胖和糖尿病鼠为模型,添加烟酰胺可调节糖代谢和NAD-sirtuin通路并且可能与改变线粒体功能有关(Yang et al.2013)围产期奶牛添加烟酸改变肝脏糖代谢基因的表达,可能与FoxO1有关(Asako Kinoshita et al.2015)。烟酰胺在脂肪代谢方面的研究有:烟酰胺作为功能性微量营养素对于能量代谢至关重要,烟酰胺长期处理MSC细胞可促进脂肪形成(Shapiroet al.2016)。Liu等(2009)发现烟酰胺可控制牛脂肪前体细胞的脂肪分解和能量平衡;烟酰胺在一定浓度范围内促进肝细胞能量代谢(臧坤等2009;成海建等2013),改善应激奶牛的脂代谢(孙先枝等2015)。烟酸在药理剂量下是一种很好的降脂药物,能降低低密度脂蛋白胆固醇(low-density lipoprotein,LDL)水平,并增加高密度脂蛋白胆固醇(high-density lipoprotein,HDL)水平。限饲荷斯坦奶牛补充烟酸降低血非酯化脂肪酸(NEFA)水平(Pires et al.2007);对围产期奶牛补充12g/d瘤胃保护烟酸,抑制脂肪分解,降低奶能量输出,改善围产期能量平衡,但对采食量没有影响(Yuan et al.2012),补充24g/d烟酸可降低围产后期奶牛血液NEFA水平,抑制脂肪分解(Morey et al.2011);中国杂种育肥牛补充1000mg/kg烟酸增加血清HDL水平,降低LDL、TG、NEFA、总胆固醇(total cholesterol,TC)、糖化血清蛋白水平,提高肉品质(Yang et al.2016),以上可见烟酰胺和烟酸均可调节脂肪代谢。
烟酰胺在肠道发育方面的研究有:烟酰胺可调节肠道黏膜屏障,保护肠道健康,HNF-α和PPARα是调节肠道完整性和功能的主要肠道转录因子,claudin-1、claudin-5和ZO-1都是肠道紧密连接蛋白,烟酸上调这些因子的表达说明烟酸可提高肠道屏障(Wei etal.2015);烟酸缺乏可降低肠道粘膜免疫和肠道生理功能(Feng et al.2016),因此,烟酰胺对于肠道免疫及糖脂代谢方面发挥至关重要的作用。
综上所述,烟酰胺可影响糖代谢、脂肪代谢和促进肠道发育。但目前还缺乏关于母体添加烟酰胺对子代糖脂代谢等影响的研究。羔羊培育是奶山羊养殖过程的重要环节,羔羊阶段打下良好基础对于成年后奶山羊的生长发育以及生产性能十分关键。所以,研究母体烟酰胺对子代羔羊糖脂代谢的影响及其机制,可为羔羊的高效健康生长发育提供理论依据。
发明内容
为了克服现有技术的缺点与不足,本发明从母体添加烟酰胺入手,探究烟酰胺的母体营养干预对羔羊糖脂代谢及肠道形态发育的影响及其机理,为调节羔羊生长发育提供理论依据。
本发明是通过以下技术方案实现的:
本发明的目的之一,提供围产期奶山羊添加烟酰胺在促进羔羊肠道形态发育和提高羔羊肠道对营养物质消化吸收中的应用。
所述应用为提高羔羊十二指肠绒毛高度、十二指肠绒毛高度与隐窝深度比值V/C、羔羊空肠绒毛高度与隐窝深度比值V/C、羔羊回肠绒毛高度以及提高回肠绒毛高度与隐窝深度比值V/C,进而促进羔羊形态发育和提高羔羊对肠道消化吸收能力。
所述应用为促进羔羊空肠葡萄糖转运载体GLUT2和SGLT1的表达,回肠GLUT2的表达,进而促进羔羊肠道葡萄糖的转运和吸收。
本发明的另一个目的,提供围产期奶山羊添加烟酰胺在提高羔羊肝脏功能和减少肝脏损伤中的应用。
本发明的另一个目的,提供围产期奶山羊添加烟酰胺在降低羔羊脂肪肝发生中的应用。
所述应用为可促进羔羊肝脏脂肪分解,抑制甘油三酯合成。
本发明的另一个目的,提供围产期奶山羊添加烟酰胺在促进羔羊腹腔脂肪积累中的应用。
所述应用为促进羔羊腹腔脂肪组织脂肪酸的合成和甘油三酯的合成,使脂肪合成代谢增强,但不减弱脂肪分解代谢。
本发明的另一个目的,提供一种围产期奶山羊饲喂烟酰胺的方法,通过直接口服饲喂烟酰胺或者日粮添加烟酰胺。
优选地,所述方法中每天早上和晚上各饲喂1次,每次2.5g。
本发明的有益效果
本发明首次公开了围产期奶山羊添加烟酰胺可促进羔羊小肠肠道形态发育、提高羔羊肠道营养物质的消化吸收、提高羔羊肝脏功能和减少肝脏损伤、降低羔羊脂肪肝发生和促进羔羊腹脂沉积,并对相关机制进行研究,为通过母体营养调控羔羊生长发育提供理论依据。
附图说明
图1围产期奶山羊添加烟酰胺对羔羊肠道葡萄糖转运载体表达的影响。
图2围产期奶山羊添加烟酰胺对羔羊肝脏糖异生相关酶基因表达的影响。
图3围产期奶山羊添加烟酰胺对羔羊肝脏糖原含量和代谢关键酶基因表达的影响。
图4围产期奶山羊添加烟酰胺对羔羊肝脏SIRT1、PGC1α、FoxO1和SREBP1mRNA水平的影响。
图5围产期奶山羊添加烟酰胺对羔羊肝脏ACC、FAS、SCD mRNA相对表达量的影响。
图6围产期奶山羊添加烟酰胺对羔羊腹脂ACC、FAS、SCD mRNA相对表达量的影响。
图7围产期奶山羊添加烟酰胺对羔羊肝脏甘油三酯代谢关键酶mRNA相对表达量的影响。
图8围产期奶山羊添加烟酰胺对羔羊腹脂甘油三酯代谢关键酶mRNA相对表达量的影响。
图9围产期奶山羊添加烟酰胺对羔羊肝脏脂肪酸氧化和转运关键酶相对表达量的影响。
图10围产期奶山羊添加烟酰胺对羔羊腹脂PGC1α和SREBP1mRNA水平的影响。
具体实施方式
以下通过实施例对本发明特征及其它相关特征作进一步详细说明,以便于同行业技术人员的理解:
试验地点:动物试验在西北农林科技大学畜牧养殖试验基地进行。
试验动物:试验选取2胎次妊娠奶山羊15只,按照体重、采食量配对分为3组,分别为对照组(C组),产后1至28d添加烟酰胺组(P组)、产前21d至产后28d添加烟酰胺组(EP组),烟酰胺添加量为早上和晚上各添加1次,每次2.5g。母羊产羔后,对应组羔羊分别为LC组、LP组和LEP组,半小时内饲喂初乳。分别从各组羔羊中选取5只,人工哺喂对应组母乳至产后28d。先将对应组母乳混匀,然后过滤。先将奶样加热至74℃左右,然后降温至37℃左右哺喂羔羊,哺喂时间分别是05:00、10:00、15:00、20:00。(以下实施例都是以此方法饲喂进行的)
本发明所用引物由西安擎科泽西生物技术有限公司合成。
实施例1围产期奶山羊添加烟酰胺对羔羊肠道形态发育和养分转运载体表达的影响
1、样品采集
1.1采集肠段:羔羊28日龄屠宰,迅速打开腹腔,分离十二指肠、空肠和回肠,分别在肠道中间剪取约3cm的肠段,用生理盐水冲洗干净肠道内容物,放置于4%多聚甲醛溶液进行固定,储存于4℃冰箱以待肠道形态分析。
1.2采集肠道黏膜:羔羊28日龄屠宰,迅速打开腹腔,取出十二指肠、空肠和回肠,用生理盐水冲洗干净肠道内容物后,纵向剪开肠段将其平展,随后用载玻片轻轻刮取肠道黏膜,分装入冻存管,迅速放于液氮中短暂储存,然后转移至-80℃冰箱储存待测。
2、肠道形态测定
将肠段从固定液取出,用水冲洗过夜。按照由低到高的浓度顺序,将肠段组织放置于乙醇中进行脱水,使用二甲苯对肠段进行透明、透蜡,将组织放入放有石蜡的模具中进行包埋、冷却,随后进行切片。石蜡切片经过脱蜡、染色(苏木精-伊红HE染色方法)、透明、中性树胶封片步骤,然后观察切片,选取典型且具有代表性视野,测量十二指肠、空肠和回肠的绒毛高度和隐窝深度,随后计算绒毛高度与隐窝深度的比值。
3、采用RT-PCR法检测各肠道黏膜中相关基因表达水平
选取GAPDH作为内参基因,用Primer Primier 5.0软件设计SGLT1和GLUT2的引物,对组织提取的RNA进行RT-PCR反应,检测各肠道黏膜中相关基因表达水平。
4、结果
4.1围产期奶山羊添加烟酰胺对羔羊小肠形态发育的影响
表1围产期奶山羊添加烟酰胺对羔羊小肠形态的影响
注:同行数据肩标不同小写字母表示差异显著(P<0.05),肩标不同大写字母表示差异极显著(P<0.01),肩标相同或无字母表示差异不显著(P>0.05);n=5;下同。
由表1可知,LP和LEP组十二指肠绒毛高度高于LC组(P<0.05),LEP组回肠绒毛高度高于LC和LP组(P<0.05),空肠绒毛高度各组间无显著差异(P>0.05)。十二指肠、空肠和回肠的隐窝深度各组间差异不显著(P>0.05)。LP组和LEP组十二指肠V/C比值高于LC组(P<0.01),LP组和LEP组空肠V/C比值高于LC组(P<0.05),LEP组回肠V/C比值高于LC和LP组(P<0.01)。
肠道粘膜形态,尤其是绒毛和隐窝结构,是评判小肠消化和吸收能力的重要参照指标,也可反映肠道健康(Albrecht et al.2007)。绒毛高度增加表明养分吸收表面积增大(Caspary et al.1992),绒毛变短、隐窝变深,导致养分吸收少,从而降低生产性能(Xu etal.2003);增大绒毛高度、绒毛高度与隐窝深度比值与促进上皮细胞周转有关(Fan etal.1997),且较长的绒毛高度可有效激活细胞有丝分裂(Samanya et al.2002)。本发明中,母体后期添加烟酰胺提高羔羊十二指肠绒毛高度、空肠的绒毛高度与隐窝深度比值,母体全期添加烟酰胺提高羔羊回肠绒毛高度和绒毛高度与隐窝深度比值。表明母体添加烟酰胺可促进羔羊小肠形态发育,提高羔羊肠道的消化吸收能力。
4.2围产期奶山羊添加烟酰胺对羔羊肠道葡萄糖转运载体表达的影响
如图1所示,羔羊十二指肠GLUT2和SGLT1的相对表达水平各组间无显著差异(P>0.05)。LP组羔羊空肠的GLUT2相对表达水平高于LC组和LEP组(P<0.01),LC组与LEP组间无显著差异(P>0.05)。LP组和LEP组羔羊空肠的SGLT1的相对表达水平高于LC组(P<0.01),LP组和LEP组间无显著差异(P>0.05)。LEP组羔羊回肠的GLUT2相对表达水平高于LC和LP组(P<0.05),LC组与LP组间无显著差异(P>0.05)。LP组和LEP组羔羊回肠SGLT1相对表达水平低于LC组(P<0.01),LP组与LEP组间无显著差异(P>0.05)。
哺乳羔羊采食母乳后,母乳进入皱胃凝结成块,皱胃中凝块裂解,蛋白和脂肪进入十二指肠进行消化,消化终产物多为葡萄糖、半乳糖、乳糖、氨基酸和小肽等,随后小肠将产物吸收利用,为羔羊供应营养促进羔羊生长发育。按照小肠己糖吸收的经典模型,葡萄糖和半乳糖依靠Na依赖的SGLT1转运载体转运通过肠上皮细胞膜,然而从细胞转移至血液循环需要己糖易化扩散转运载体GLUT2。SGLT1和GLUT2在葡萄糖穿过肠细胞基底侧膜进入循环过程中发挥协同作用(Kellett et al.2008)。因此GLUT2和SGLT1在肠道葡萄糖转运、吸收过程中发挥至关重要的作用(et al.2014)。本发明检测十二指肠、空肠和回肠的SGLT1和GLUT2的表达水平,结果显示母体添加烟酰胺对羔羊十二指肠SGLT1和GLUT2的表达未造成显著影响;母体后期添加烟酰胺极显著提高羔羊空肠SGLT1和GLUT2的表达,母体全期添加烟酰胺极显著提高羔羊空肠SGLT1和回肠GLUT2的表达,对空肠GLUT2的表达无显著影响。所以,本试验表明母体添加烟酰胺显著提高羔羊空肠的葡萄糖转运吸收。
实施例2围产期奶山羊添加烟酰胺对羔羊血液生化指标和糖代谢中的影响
1、采集血样
于羔羊14、28日龄,晨饲后3h用真空采血管进行颈静脉采血。部分血液于4℃、转速3500r/min条件下,离心15min获得血浆;一部分血液37℃放置30min后,于转速3500r/min条件下离心15min获得血清,将血清、血浆均放在-80℃备用分析。
2、组织样品采集
羔羊28日龄屠宰,迅速打开腹腔,采集肝脏组织,用生理盐水清洗后置于液氮中储存。
3、测定指标及方法
3.1围产期奶山羊添加烟酰胺对羔羊血浆参数的影响
表2围产期奶山羊添加烟酰胺对羔羊血浆参数的影响
血液生化指标GOT和GPT是反映肝脏健康的重要指标。GOT和GPT可促进天冬氨酸和丙氨酸的α-氨基酸转移至酮戊二酸的α-酮酸上,分别产生草酰乙酸和丙酮酸进入三羧酸循环。本发明,母体全期添加烟酰胺具有降低28日龄羔羊血GOT水平的趋势,烟酰胺处理组28日龄羔羊血GPT水平均显著降低,表明母体添加烟酰胺可以提高羔羊肝脏功能,减少肝脏损伤。
3.2围产期奶山羊添加烟酰胺对羔羊血清激素水平的影响
由表3可知,羔羊血清胰岛素、胰高血糖素、瘦素、IGF1、IGF2、胰岛素敏感指数各组间差异均不显著(P>0.05)。所以,本发明中,奶山羊围产期添加烟酰胺对羔羊血清胰岛素、胰高血糖素水平未造成显著影响。
表3围产期奶山羊添加烟酰胺对羔羊血清激素水平的影响
3.3围产期奶山羊添加烟酰胺对羔羊肝脏糖异生关键酶表达的影响
由图2可知,LP和LEP组PEPCK相对表达水平极显著低于LC组(P<0.01),LEP组PEPCK相对表达水平极显著低于LP组(P<0.01)。羔羊肝脏G6P、FBP、PC的相对表达水平各组间差异均不显著(P>0.05)。
3.4围产期奶山羊添加烟酰胺对羔羊肝糖原含量及肝脏糖原代谢关键酶表达的影响
由图3可知,LEP组羔羊肝脏糖原含量具低于LC和LP组趋势(P<0.10)。LP组和LEP组羔羊肝脏GS和GLUT2相对表达水平显著低于LC组(P<0.05),LP组和LEP组间无显著差异(P>0.05)。羔羊PYGL相对表达水平各组间差异不显著(P>0.05)。
3.5围产期奶山羊添加烟酰胺对羔羊肝脏SIRT1、PGC1α、FoxO1、SRBEP1相对表达水平的影响
由图4可知,LP组和LEP组羔羊肝脏PGC1α、FoxO1相对表达水平显著低于LC组(P<0.05),LP组和LEP组间差异不显著(P>0.05)。SIRT1和SREBP1的相对表达水平各组间差异不显著(P>0.05)。
综上所述,本发明中,奶山羊围产期添加烟酰胺对羔羊血清胰岛素、胰高血糖素水平未造成显著影响。PUN可反映机体蛋白质代谢状态,是蛋白质分解代谢的产物。本发明中,母体添加烟酰胺显著降低羔羊PUN水平,表明羔羊机体蛋白质平衡状况较好,母体添加烟酰胺使羔羊PUN水平降低,羔羊的氨基酸氧化率降低,从而提高了机体对氨基酸的利用。血浆葡萄糖反映机体内葡萄糖生成和组织消耗葡萄糖之间的动态平衡,血糖水平的变化可在一定程度上反映机体所处的生理状态。本发明中,奶山羊围产期添加烟酰胺显著提高14日龄羔羊血浆葡萄糖含量,但对28日龄羔羊血浆葡萄糖含量未造成显著影响,造成这种现象的可能原因是羔羊14日龄时机体稳衡机制尚不完善,易受外界处理影响;也可能是烟酰胺提高组织对葡萄糖利用(Tam et al.2005),长期添加烟酰胺导致血糖含量恢复正常水平。烟酰胺提高血糖水平,降低肌糖原含量,诱导氧化应激的产生(Shi et al.2009)。这与本试验血糖结果不同,可能原因是本试验动物处于正常生理状态,与烟酰胺在病理状态下发挥作用的机制不一致。
本发明中,母体全期添加烟酰胺具有降低羔羊肝糖原含量的趋势。肝糖原含量高低取决于糖原合成和分解之间的平衡,为了明确羔羊肝糖原降低的原因,本发明探究GS和PYGL的定量表达水平。GS定量结果表明,母体全期添加烟酰胺组羔羊肝脏GS的表达显著降低,但对PYGL无显著影响,所以肝糖原降低的可能原因是,母体添加烟酰胺通过抑制羔羊肝脏FoxO1表达,降低GS的基因表达,降低肝糖原合成,导致肝糖原含量降低。
糖异生受PC、PEPCK、FBP、G6P的调控,本试验在转录水平分析糖异生关键酶表达的变化,反映糖异生能力的改变。肝脏糖异生对于维持血糖稳态十分重要,PEPCK是糖异生途径的限速酶,本发明中,母体添加烟酰胺显著降低羔羊肝脏PEPCK的mRNA表达水平,但对FBP、PC和G6P未造成显著影响。肝脏中GLUT2是双向葡萄糖转运载体,可以确保葡萄糖在肝细胞内进出。本试验中,母体添加烟酰胺显著降低羔羊肝脏GLUT2的表达,说明肝脏葡萄糖输出减少。因此,围产期奶山羊添加烟酰胺可能通过抑制PEPCK的表达,抑制羔羊肝脏糖异生,降低肝脏葡萄糖输出。SREBP-1C对于葡萄糖刺激的GLUT2基因表达至关重要,SREBP-1C可激活GLUT2的启动子区,GLUT2受SREBP-1C的调控(Im et al.2005)。但本发明中,母体添加烟酰胺并未影响羔羊肝脏SREBP-1的表达,因此GLUT2的表达发生改变可能是受其他因素影响。本发明中,奶山羊围产全期和围产后期添加烟酰胺均显著降低羔羊肝脏PGC1α和FoxO1的表达。因此,围产期奶山羊添加烟酰胺通过抑制羔羊肝脏PGC1α的表达,抑制FoxO1表达,从而抑制其靶基因PEPCK的表达,抑制羔羊肝脏糖异生,转运出肝脏的葡萄糖量减少,GLUT2表达降低。结合前面PUN结果,PUN水平降低,可用于肝脏糖异生的底物含量减少,可能是导致肝脏糖异生降低的主要原因。肠道吸收葡萄糖增加、肝脏糖异生降低可能是导致28日龄羔羊血糖未发生显著变化的原因。
实施例3围产期奶山羊添加烟酰胺对羔羊脂肪代谢的影响及机制
1、样品采集
羔羊28日龄屠宰,迅速打开腹腔,采集腹腔脂肪组织和肝脏组织,用生理盐水清洗后置于液氮中储存。
2、测定指标及方法
2.1肝脏和腹脂表观指标测定
从-80℃冰箱取出肝脏组织和腹腔脂肪组织,融化后分别加入适量生理盐水和无水乙醇,混合均匀,用匀浆器进行匀浆,将匀浆介质于3000r/min转速下离心20min,取上清分装备用,进行TG、FFA、TC和腹脂甘油含量测定。
2.2肝脏、腹腔脂肪组织酶活测定
从-80℃冰箱取出肝脏组织和腹腔脂肪组织,融化后分别加入适量pH 7.4的PBS和无水乙醇(脂肪组织脂肪含量高,无水乙醇才能萃取),混合均匀,用匀浆器进行匀浆,将匀浆介质于3000r/min转速下离心20min,取上清分装备用,测定酶活。
采用双抗体夹心法测定肝脏和腹腔脂肪组织匀浆中GPAM、AGPAT6、DGAT2、ATGL、HSL。
2.3肝脏和腹脂脂肪代谢关键酶mRNA表达测定
AGPAT6、ACC、FAS、SCD、SREBP1引物序列参考Shi等(Shi et al.2013),ATGL引物序列参考Lin等(Lin et al.2013),DGAT2引物序列参考Bionaz and Loor(2008),GPAM引物序列参考Ma and Corl(2012),HSL引物序列参考Li et al.(2015),用Primer Primier 5.0软件设计CPT1A、ApoB、MTTP的引物,对肝脏和腹脂脂肪代谢关键酶mRNA表达进行测定。
3、结果
3.1围产期奶山羊添加烟酰胺对羔羊肝脏和腹脂FFA、TG、TC水平的影响
表4围产期奶山羊添加烟酰胺对羔羊肝脏和腹脂FFA、TG、TC水平的影响
由表4可知,LP和LEP组肝脏FFA含量显著高于LC组(P<0.05),LP和LEP组间无显著差异(P>0.05)。LEP组羔羊腹腔脂肪组织FFA含量显著高于LC组(P<0.05)。LP和LEP组腹腔脂肪组织TG含量极显著高于LC组(P<0.01),LEP组TG含量极显著高于LP组(P<0.01)。肝脏TG、TC含量和腹腔脂肪组织甘油、TC含量各组间无显著差异(P>0.05)。
3.2围产期奶山羊添加烟酰胺对羔羊脂肪酸从头合成关键酶表达的影响
由图5可知,LEP组羔羊肝脏FAS相对表达水平显著低于LC组(P<0.05),LC和LP组间无显著差异(P>0.05)。LEP组ACC相对表达水平与LC组相比降低了3.52倍,但差异不显著(P>0.05)。SCD相对表达水平各组间差异不显著(P>0.05)。
由图6可知,LEP组羔羊腹脂FAS相对表达水平显著高于LC和LP组(P<0.05),LC和LP组FAS相对表达水平无显著差异(P>0.05)。腹脂ACC、SCD相对表达水平各组间无显著差异(P>0.05)。
3.3围产期奶山羊添加烟酰胺对羔羊甘油三酯代谢关键酶表达的影响
由图7可知,羔羊肝脏GPAM、AGPAT6、DGAT2、ATGL相对表达水平各组间无显著差异(P>0.05)。LP组HSL相对表达水平显著低于LC组(P<0.05),LP和LEP组间差异不显著(P>0.05)。
由图8可知,LEP组羔羊腹脂GPAM相对表达水平极显著高于LC组和LP组(P<0.01)。LP和LEP组AGPAT6相对表达水平极显著高于LC组(P<0.01),LP组和LEP组间AGPAT6相对表达水平无显著差异(P>0.05)。DGAT2相对表达水平各组间无显著差异(P>0.05)。各组羔羊腹脂ATGL、HSL相对表达水平无显著差异(P>0.05)。
3.4围产期奶山羊添加烟酰胺对羔羊肝脏脂肪酸氧化和转运和腹脂转录因子的影响
由图9可知,LEP组羔羊肝脏ApoB相对表达水平显著低于LC组(P<0.05),LC组和LP组间无显著差异(P>0.05)。LEP组羔羊肝脏MTTP相对表达水平极显著高于LC和LP组(P<0.01),LC组和LP组间无显著差异(P>0.05)。CPT1A和ACSL的相对表达量各组间无显著差异(P>0.05)。
由图10可知,LEP组羔羊腹脂PGC1α和SREBP1相对表达水平显著高于LC和LP组(P<0.05),LC和LP组间无显著差异(P>0.05)。
3.4围产期奶山羊添加烟酰胺对羔羊肝脏和腹脂脂肪代谢关键酶酶活的影响
表5围产期奶山羊添加烟酰胺对羔羊肝脏脂肪代谢关键酶酶活的影响
表6围产期奶山羊添加烟酰胺对羔羊腹脂脂肪代谢关键酶酶活的影响
由表5可知,LEP组羔羊肝脏AGPAT6活性显著低于LC和LP组(P<0.05),LC和LP组间无显著差异(P>0.05)。LP组HSL活性显著高于LC组(P<0.05),LEP组和LC组间无显著差异(P>0.05)。GPAM、DGAT2、ATGL酶活性各组间无显著差异(P>0.05)。
由表6可知,羔羊腹脂GPAM、AGPAT6、DGAT2、ATGL和HSL的酶活各组间无显著差异(P>0.05)。
综上结论可知,
1、围产期奶山羊添加烟酰胺对羔羊腹腔脂肪组织脂肪代谢的影响及机制
奶山羊围产期添加烟酰胺组羔羊腹脂TG含量显著增加,表明羔羊脂质沉积增加,脂肪沉积取决于脂肪合成代谢和脂肪分解代谢之间的平衡状态,为了明确羔羊腹脂TG增加的机制,本发明检测脂肪合成相关基因和脂肪分解相关基因表达。
围产全期添加烟酰胺组羔羊腹脂PGC1α的mRNA表达显著升高,PGC1α可调节SREBP1的表达。SREBP1是重要的生脂转录调控因子,可选择性激活脂肪酸从头合成关键基因的表达(Osborne et al.,2000),过表达SREBP1-c可提高脂合成基因的表达,下调脂质氧化基因的表达,增加TG的合成和积累(Li et al.,2014)。本实验中,围产全期添加烟酰胺组羔羊腹脂SREBP1表达呈显著上升现象,这与赵涛涛等(2012)研究一致。参与脂肪酸从头合成和TG合成的酶主要在转录水平被调控(Wang et al.,2015),故本实验主要从转录水平探究母体添加烟酰胺对子代腹脂SREBP1表达及其重要生脂靶基因的调控。ACC、FAS、SCD、GPAM是SREBP1的重要靶基因(Shimano et al.,2001),其中ACC、FAS、SCD是脂肪酸从头合成的3个关键酶,ACC催化乙酰辅酶A形成丙二酰辅酶A,然后乙酰辅酶A和丙二酸单酰辅酶A在FAS催化下形成棕榈酸,最后在SCD催化下延长碳链形成长链脂肪酸。本实验中,围产全期添加烟酰胺组羔羊腹脂FAS基因表达显著升高,说明围产全期添加烟酰胺可促进羔羊腹腔脂肪组织脂肪酸从头合成。
GPAM、AGPAT6、DGAT2是TG合成过程的重要酶。提高GPAM表达可显著促进TG合成(Yuet al.,2017),本发明中,围产全期添加烟酰胺组羔羊腹脂GPAM、AGPAT6的表达极显著提高,围产后期添加烟酰胺组羔羊腹脂AGPAT6的表达极显著上升。由此可知,奶山羊围产期添加烟酰胺可促进羔羊腹脂TG合成,使脂肪合成代谢增强。
ATGL和HSL是TG分解的关键酶。提高ATGL、HSL基因表达可促进脂肪分解(Karbowska et al.,2012;Huang et al.,2017)。本发明中,围产全期及围产后期添加烟酰胺均对羔羊腹脂ATGL、HSL的表达未产生显著差异,表明围产期添加烟酰胺对羔羊脂肪分解代谢未造成显著影响。脂肪合成代谢增强、分解代谢未发生显著变化可能是引起腹脂TG含量增加的原因。
2、围产期奶山羊添加烟酰胺对羔羊肝脏脂肪代谢的影响及机制
奶山羊添加烟酰胺显著增加羔羊肝脏FFA含量,为了探究原因,本试验检测脂肪合成相关基因(SIRT1、PGC1α、SREBP1、ACC、FAS、SCD、GPAM、AGPAT6、DGAT2)基因表达,同时检测脂肪分解基因(ATGL、HSL)、脂肪氧化基因(CPT1A、ACSL)、脂质转运基因(ApoB、MTTP)的基因表达。
本发明中,母体全期添加烟酰胺显著降低羔羊肝脏FAS的表达,对ACC和SCD的表达水平未造成显著影响。本发明中,母体后期添加烟酰胺显著增加羔羊肝脏HSL的表达水平,处理效应对GPAM、AGPAT6、DGAT2和ATGL的表达水平未造成显著影响。由以上定量结果可知,围产期奶山羊添加烟酰胺可通过增加羔羊肝脏HSL的mRNA表达水平促进肝脏脂肪分解,但对DGAT2的mRNA表达未造成显著影响。脂肪代谢关键酶活性结果可知,奶山羊围产全期添加烟酰胺显著降低肝脏AGPAT6的酶活,对GPAM和DGAT2的酶活性未造成显著差异,结合定量结果可知,奶山羊添加烟酰胺通过抑制AGPAT6的活性抑制TG合成,但对mRNA无影响,可能是母体添加烟酰胺对羔羊肝脏AGPAT6的调控主要在蛋白水平,而不是mRNA水平。母体后期添加烟酰胺显著提高HSL的酶活性,结合定量结果可知,母体后期添加烟酰胺通过提高HSL的mRNA水平和酶活性促进肝脏脂肪分解,母体添加烟酰胺对羔羊肝脏HSL的调节体现在mRNA水平和蛋白水平。结合上述结果可知,奶山羊添加烟酰胺可提高肝脏脂肪分解、抑制TG合成。肝脏TG分解增强可能是导致羔羊肝脏FFA增加的原因。
ApoB和MTTP对于脂蛋白的装配、分泌至关重要,MTTP是膜内的脂转运蛋白,可促进ApoB包含脂蛋白的分泌,二者和TG合成受到抑制相关。本发明表明,奶山羊添加烟酰胺可抑制羔羊肝脏TG合成,定量结果可知,母体全期添加烟酰胺显著降低羔羊肝脏ApoB的mRNA表达水平,显著提高MTTP的表达。表明烟酰胺降低羔羊肝脏TG合成、促进TG分解,降低ApoB的分泌。MTTP的mRNA表达水平提高,说明ApoB包含的脂蛋白的分泌增加,VLDL-TG分泌可能会增加,可能会导致血浆TG水平上升。围产期奶山羊添加烟酰胺对羔羊肝脏CPT1A和ACSL的表达未造成显著差异,说明母体添加烟酰胺不影响羔羊肝脏脂肪酸氧化。
烟酰胺对SIRT1的作用众说纷纭,烟酰胺是SIRT1的抑制剂(Li et al.2015;Peledet al.2012;Shan et al.2013),但也有报道称烟酰胺在体外是SIRT1的抑制剂,但在细胞内可能是SIRT1的激活剂(Hwang et al.2017)。但本试验中奶山羊添加烟酰胺对羔羊肝脏SIRT1的相对表达水平未造成显著影响,可能原因是羔羊年龄太小,所需的NAD基础水平明显高于年龄较大动物(Koltai et al.2010),也可能是由于本试验中进入羔羊体内的烟酰胺量显著低于细胞试验中可抑制SIRT1活性的剂量(Fulco et al.2008)。本试验中,奶山羊添加烟酰胺对羔羊肝脏SIRT1和SREBP1的表达均未造成影响,因此母体添加烟酰胺对羔羊肝脏脂肪代谢的调节可能与SIRT1和SREBP1无关。
PGC1α在全身及肝脏糖脂代谢中发挥重要的调控作用。本试验中,奶山羊添加烟酰胺降低羔羊肝脏PGC1α的相对表达水平,表明PGC1α可介导母体添加烟酰胺调控羔羊脂肪代谢的过程。母体添加烟酰胺通过抑制羔羊肝脏PGC1α的表达,抑制其下游靶基因FAS的mRNA表达、AGPAT6的酶活性,提高HSL的mRNA表达和酶活性,降低ApoB的分泌,抑制羔羊肝脏甘油三酯合成,促进脂肪分解。
最后应该说明的是,以上所述仅为本发明的优选实施例而已,并不用于限制本发明,尽管参照前述实施例对本发明进行了详细的说明,对于本领域的技术人员来说,其依然可以对前述实施例所记载的技术方案进行修改,或者对其中部分进行等同替换。凡在本发明的精神和原则之内,所作的任何修改、等同替换、改进等,均应包含在本发明的保护范围之内。上述虽然结合附图对本发明的具体实施方式进行了描述,但并非对本发明保护范围的限制,所属领域技术人员应该明白,在本发明的技术方案的基础上,本领域技术人员不需要付出创造性劳动即可做出的各种修改或变形仍在本发明的保护范围以内。
Claims (10)
1.围产期奶山羊添加烟酰胺在促进羔羊肠道形态发育和提高羔羊肠道消化吸收中的应用。
2.根据权利要求1所述的应用,其特征在于,增加羔羊十二指肠绒毛高度和十二指肠绒毛高度与隐窝深度比值V/C,提高羔羊空肠绒毛高度与隐窝深度比值V/C,增加羔羊回肠绒毛高度以及提高回肠绒毛高度与隐窝深度比值V/C,进而促进羔羊形态发育和提高羔羊肠道消化吸收能力。
3.根据权利要求1所述的应用,其特征在于,促进羔羊空肠葡萄糖转运载体GLUT2和SGLT1的表达,回肠GLUT2的表达,进而促进羔羊肠道葡萄糖的转运和吸收。
4.围产期奶山羊添加烟酰胺在提高羔羊肝脏功能,减少肝脏损伤中的应用。
5.围产期奶山羊添加烟酰胺在降低羔羊脂肪肝发生中的应用。
6.根据权利要求5所述的应用,其特征在于,促进羔羊肝脏脂肪分解,抑制甘油三酯合成。
7.围产期奶山羊添加烟酰胺在促进羔羊腹腔脂肪积累中的应用。
8.根据权利要求7所述的应用,其特征在于,促进羔羊腹腔脂肪组织脂肪酸的合成和甘油三酯的合成,使脂肪合成代谢增强,但不减弱脂肪分解代谢。
9.一种围产期奶山羊饲喂烟酰胺的方法,其特征在于,通过直接口服饲喂烟酰胺或者日粮添加烟酰胺。
10.根据权利要求9所述的方法,其特征在于,所述方法中每天早上和晚上各饲喂1次,每次2.5g。
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