研究论文

不同谷物来源淀粉对断奶马驹胃肠道葡萄糖转运载体及生长发育相关基因表达的影响

  • 李晓斌 ,
  • 黄新新 ,
  • 臧长江 ,
  • 陈开旭 ,
  • 马晨 ,
  • 张文杰 ,
  • 鲁豪
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  • 新疆农业大学动物科学学院,新疆肉乳用草食动物营养重点实验室,新疆马繁育与运动生理重点实验室,乌鲁木齐 830052

李晓斌(1988—)男,甘肃天水人,副教授,博士,研究方向为动物营养与饲料科学。E-mail:

收稿日期: 2022-11-16

  网络出版日期: 2023-06-08

基金资助

国家自然科学基金项目(31860649)

Effects of Different Grain Sources of Starch on Expression of Glucose Transport Transporters and Growth and Development Related Genes in Gastrointestinal Tract of Weaned Foals

  • LI Xiaobin ,
  • HUANG Xinxin ,
  • ZANG Changjiang ,
  • CHEN Kaixu ,
  • MA Chen ,
  • ZHANG Wenjie ,
  • LU Hao
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  • Xinjiang Key Laboratory of Horse Breeding and Exercise Physiology, Xinjiang Key Laboratory of Herbivore Nutrition for Meat & Milk Production, College of Animal Science, Xinjiang Agricultural University, Urumqi 830052, China

LI Xiaobin, associate professor, E-mail:

Received date: 2022-11-16

  Online published: 2023-06-08

摘要

本试验旨在研究不同谷物来源淀粉对断奶马驹胃肠道葡萄糖转运载体及生长发育相关基因表达的影响,为断奶期马属动物胃肠道发育及科学饲粮配制提供参考依据。选择出生日期相近、5月龄断奶的健康哈萨克马公马驹18匹,随机分为3组,根据精料补充料中谷物不同分别记为玉米组、燕麦组和大麦组,每组6匹。在相同的粗饲料条件下,以2 g/kg BW谷物淀粉为基础,计算马驹精料补充料的补喂量,进行为期60 d的饲养试验。试验结束当天屠宰所有试验马驹,分别取胃、小肠、盲肠及结肠黏膜样品,并测定葡萄糖转运载体和生长发育相关基因相对表达量。结果表明:1)各组之间胃黏膜钠依赖性葡萄糖转运载体1(SGLT1)基因相对表达量无显著差异(P>0.05),燕麦组胃黏膜易化葡萄糖转运载体1(GLUT1)基因相对表达量显著高于玉米组和大麦组(P<0.05)。2)玉米组小肠黏膜SGLT1基因相对表达量显著高于燕麦组(P<0.05),极显著高于大麦组(P<0.01);大麦组小肠黏膜GLUT1基因相对表达量显著高于玉米组和燕麦组(P<0.05)。3)各组之间小肠黏膜表皮生长因子(EGF)、胰岛素样生长因子-1(IGF-1)和胰岛素样生长因子-1受体(IGF-1R)基因相对表达量无显著差异(P>0.05)。4)大麦组盲肠黏膜EGFIGF-1R基因相对表达量极显著高于玉米组和燕麦组(P<0.01)。燕麦组盲肠黏膜EGF基因相对表达量显著高于玉米组(P<0.05)。5)大麦组结肠黏膜EGF基因相对表达量极显著高于燕麦组(P<0.01),玉米组结肠黏膜EGF基因相对表达量显著高于燕麦组(P<0.05)。玉米组结肠黏膜IGF-1R基因相对表达量极显著高于燕麦组和大麦组(P<0.01)。综上所述,给断奶马驹饲喂不同谷物来源淀粉对马驹胃肠道中葡萄糖转运载体及生长发育相关基因表达的影响不同。

本文引用格式

李晓斌 , 黄新新 , 臧长江 , 陈开旭 , 马晨 , 张文杰 , 鲁豪 . 不同谷物来源淀粉对断奶马驹胃肠道葡萄糖转运载体及生长发育相关基因表达的影响[J]. 动物营养学报, 2023 , 35(6) : 3932 -3942 . DOI: 10.12418/CJAN2023.365

Abstract

The aim of this experiment was to investigate the effects of different grain sources of starch on expression of glucose transport transporters and growth and development related genes in gastrointestinal tract of weaned foals, in order to provide a reference for development of the gastrointestinal tract and scientific diet formulation for equine animals during the weaning period. Eighteen healthy Kazakh male foals with similar birth dates and weaned at 5 months of age were selected for the experiment, the foals were randomly divided into three groups, according to the different grains used in the concentrate supplement named: corn group, oats group and barley group, each group contained 6 foals. The amount of concentrate supplement fed to foals was determined on the basis of 2 g/kg BW grain starch under the same roughage conditions, and conducted a 60-day feeding experiment. All foals were slaughtered at the end of the experiment, and the mucosa samples were taken from the stomach, small intestine, cecum and colon, to determine the relative expression levels of glucose transport transporters and growth and development related genes. The results showed as follows: 1) there was no significant difference in gene relative expression level of sodium dependent glucose transporters 1 (SGLT1) in stomach mucosa among all groups (P>0.05), and the gene relative expression level of facilitated glucose transporters 1 (GLUT1) in stomach mucosa of oats group was significantly higher than that of corn group and barley group (P<0.05). 2) The gene relative expression level of SGLT1 in small intestine mucosa of corn group was significantly higher than that of oats group(P<0.05), and significantly higher than that of barley group (P<0.01); the gene relative expression level of GLUT1 in small intestine mucosa of barley group was significantly higher than that of corn group and oats group (P<0.05). 3) There were no significant differences in gene relative expression levels of epidermal growth factor (EGF), insulin-like growth factor-1 (IGF-1) and insulin-like growth factor-1 receptor (IGF-1R) in small intestine mucosa among all groups (P>0.05). 4) The gene relative expression levels of EGF and IGF-1R in cecum mucosa of barley group were significantly higher than those of corn group and oats group (P<0.01). The gene relative expression level of EGF in cecum mucosa of oats group was significantly higher than that of corn group (P<0.05). 5) The gene relative expression level of EGF in colon mucosa of barley group was significantly higher than that of oats group (P<0.01), and the gene relative expression level of EGF in colon mucosa of corn group was significantly higher than that of oats group (P<0.05). The gene relative expression level of IGF-1R in colon mucosa of corn group was significantly higher than that of oats group and barley group (P<0.01). In conclusion, it can be seen that feeding different grain sources of starch have different effects on glucose transport transporters and growth and development related gene expression in the gastrointestinal tract of weaned foals.

谷物中的淀粉含量一般可以达到60%~75%,是马属动物饲粮中重要的非结构性碳水化合物来源[1]。单胃草食家畜胃肠道淀粉主要通过酶解产生葡萄糖,为机体提供所需的能源物质[2-3]。充分研究谷物在单胃草食家畜胃肠道中的吸收转运和对肠道生长发育的影响,对指导实际生产中马驹饲料配方中谷物的选择与搭配、通过营养调控手段实现马产业的提质增效具有重要意义。胃肠道中的葡萄糖主要在钠依赖性葡萄糖转运载体(sodium dependent glucose transporters,SGLTs)或易化葡萄糖转运载体(facilitated glucose transporters,GLUTs)的介导下进入肠上皮细胞[4],参与机体的糖代谢。表皮生长因子(epidermal growth factor,EGF)、胰岛素样生长因子-1(insulin-like growth factor-1,IGF-1)和胰岛素样生长因子-1受体(insulin-like growth factor-1 receptor,IGF-1R)等是调控动物胃肠道结构功能、发育及健康的重要基因,具有促进肠道发育、完善肠道结构功能、提高肠道酶活性和肠道免疫功能等作用[5-6]。EGF可在转录水平调控钠依赖性葡萄糖转运载体1(sodium dependent glucose transporter 1,SGLT1)的表达,从而促进肠道中葡萄糖的吸收转运[7]。谷物淀粉酶解程度、血糖-胰岛素反应及供能效率受多种因素影响[8-9],包括谷物中淀粉直连和支链的比例以及机体葡萄糖转运载体的表达水平等。而葡萄糖转运载体的表达水平受自身发育阶段、饲粮营养水平、昼夜节律、激素水平及肠道内环境等因素影响[10-13]。Moran等[14]研究发现,给28日龄的断奶仔猪饲喂不同水平的碳水化合物饲粮,高水平碳水化合物饲粮的仔猪肠道中SGLT1 基因相对表达量显著增加,而对易化葡萄糖转运载体2(facilitated glucose transporters 2, GLUT2)基因相对表达量无影响。李永洙等[15]研究发现,给羔羊饲喂以玉米为主的代乳粉及开食料后,在羔羊25~75日龄,饲喂代乳粉及开食料的羔羊肠道中SGLT1和GLUT2 基因相对表达量高于对照组。孙大明等[16]研究发现,相比玉米和豌豆淀粉,给断奶前羔羊补饲木薯淀粉开食料可显著提高羔羊瘤胃上皮IGF-1和IGF-1R 基因相对表达量。
葡萄糖转运载体基因与胃肠道发育相关基因表达水平的高低受饲粮营养水平的限制,营养水平不足会降低其基因相对表达量[12]。因此,研究葡萄糖转运载体基因与胃肠道发育相关基因,一方面能够反映胃肠道发育机能和马驹对谷物中淀粉的利用情况;另一方面可以评价饲粮营养水平是否满足马驹生长发育的需要。不同谷物来源淀粉分子结构、直链和支链比例及含量等不同,其在同种动物体内消化代谢的程度不同,产生的效果不同。研究不同谷物来源的淀粉在马驹胃肠道消化代谢情况,明确其对马驹胃肠道相关基因表达的影响,对规范和科学断奶马驹饲粮组成和均衡营养水平具有重要的意义。因此,本研究在饲粮淀粉水平相同条件下,研究不同谷物(玉米、燕麦和大麦)来源淀粉对断奶后哈萨克马驹胃肠道葡萄糖转运载体及生长发育相关基因表达的影响,探讨不同谷物来源淀粉对断奶马驹胃肠道葡萄糖转运吸收及生长发育的作用,为断奶马驹的健康调控和科学饲养管理提供理论参考。

1 材料与方法

1.1 试验设计

试验选择出生日期相近、5月龄断奶的健康哈萨克马公马驹18匹,随机分为3组,根据精料补充料中谷物不同分别记为玉米组、燕麦组和大麦组,每组6匹。在相同的粗饲料条件下,以谷物淀粉摄入量相同为原则,以2 g/kg BW谷物淀粉为基础,计算马驹精料补充料补喂量,进行为期60 d的饲养试验。具体试验动物、试验设计、饲粮营养水平及饲养管理与本课题组黄新新等[17]研究相同。精料补充料补喂量及粗饲料采食量见表1。粗饲料按照苜蓿∶干草=2∶1进行饲喂。
表1 精料补充料补喂量及粗饲料采食量

Table 1 Concentrate supplement supplementation and roughage intakekg/d

项目
Items
玉米组
Corn group
燕麦组
Oats group
大麦组
Barley group
精料补充料 Concentrate supplement 0.71±0.00 0.72±0.00 0.72±0.00
粗饲料 Forages 5.62±0.62 5.52±0.55 5.67±0.37
总采食量 Total intake 6.33±0.62 6.24±0.55 6.39±0.37
干物质采食量 Dry matter intake 5.62±0.55 5.54±0.49 5.67±0.33

1.2 样品采集

采样当天马驹禁食12 h,禁水2 h。马驹初始体重和宰前活重见表2。马驹安乐死后,颈部放血后进行解剖,分离其胃、十二指肠、空肠、回肠、盲肠、结肠和直肠。从胃底部、十二指肠、空肠、回肠、盲肠和结肠固定部位分别取一段组织样本,手术剪纵行剪开后使用生理盐水冲洗样本中内容物,冲洗干净后置于一次性滤纸上。胃黏膜从胃底部区域刮取;小肠黏膜分别从十二指肠、空肠和回肠中段取15 cm刮取后混合均匀;盲肠黏膜从盲囊底部区域刮取;结肠黏膜从结肠中段区域刮取;将以上各部位黏膜立即分装于无RNA酶冻存管中,液氮速冻后转入-80 ℃冰箱储存,用于相关基因表达的测定。
表2 马驹初始体重和宰前活重

Table 2 Initial weight and live weight before slaughter of foalskg

项目
Items
玉米组
Corn group
燕麦组
Oats group
大麦组
Barley group
初始体重 Initial weight 112.50±3.37 112.36±12.10 112.30±6.34
宰前活重 Live weight before slaughter 143.83±7.94 139.13±6.02 147.08±4.86

1.3 样品中基因表达的测定

本研究使用试剂盒测定胃肠道中的5个目的基因,分别为SGLT1、易化葡萄糖转运载体1(facilitated glucose transporter 1,GLUT1)、EGFIGF-1和IGF-1R。使用TaKaRa反转录试剂盒将样品总RNA反转录成cDNA,使用实时荧光定量PCR(RT-qPCR)检测方法定量,引物序列见表3。以甘油醛-3-磷酸脱氢酶(glyceraldehyde-3-phosphate dehydrogenase,GAPDH)作为内参基因,采用2-△△Ct法计算目的基因相对表达量。
表3 引物序列

Table 3 Primer sequences

基因名称
Genes names
基因ID
Gene ID
引物序列
Primer sequences (5'—3')
产物大小
Produce size/bp
钠依赖性葡萄糖转运载体1
SGLT1
100033952 F:CATTGTGGTCGTCTCCCTC
R:TGCATCCAGGTCAATACGTTC
106
易化葡萄糖转运载体1
GLUT1
100034080 F:GTCTCTGGCTTCTCCAAC
R:TGAAGAACAGAACCAGGAGCA
121
表皮生长因子
EGF
106782874 F:TGTTCCAGCGATTATGACGGA
R:ATTGTCTGTTTTCTCCACGTT
122
胰岛素样生长因子-1
IGF-1
100034198 F:CTTGCTCACCTTCACCAG
R:CTCCTCAGATCACAGCTCC
198
胰岛素样生长因子-1受体
IGF-1R
100055698 F:TCCATCTTTGTGCCCAGACCT
R:CTCTCCTTGTTGTCCACTCTGC
176
甘油醛-3-磷酸脱氢酶
GAPDH
100033897 F:ACATCATCCCTGCTTCTACTGG
R:ATGCCTGCTTCACCACCT
183

1.4 数据处理

在Excel 2010中先将数据进行预处理,再使用SPSS 18.0软件进行单因素方差分析,采用Duncan氏法进行各组之间多重比较,试验结果均以平均值±标准差(mean±SD)表示。

2 结果与分析

2.1 不同谷物来源淀粉对断奶马驹胃和小肠葡萄糖转运载体基因表达的影响

2.1.1 不同谷物来源淀粉对断奶马驹胃黏膜葡萄糖转运载体基因表达的影响

不同谷物来源淀粉对断奶马驹胃黏膜SGLT1和GLUT1基因相对表达量的影响见图1。各组之间胃黏膜SGLT1基因相对表达量无显著差异(P>0.05)(图1-A)。燕麦组胃黏膜GLUT1基因相对表达量显著高于玉米组和大麦组(P<0.05),而玉米组和大麦组之间无显著差异(P>0.05)(图1-B)。这表明与饲喂玉米和大麦相比,饲喂燕麦可提高断奶马驹胃黏膜GLUT1基因相对表达量。
图1 不同谷物来源淀粉对断奶马驹胃黏膜葡萄糖转运载体基因表达的影响

数据柱无字母或标注相同字母表示差异不显著(P>0.05),不同小写字母表示差异显著(P<0.05),不同大写字母表示差异极显著(P<0.01)。下图同。

Fig.1 Effects of different grain sources of starch on gastric mucosa glucose transporter gene expression of weaned foals

Value columns with no letter or the same letter mean no significant difference (P>0.05), while with different small letters mean significant difference (P<0.05), and with different capital letters mean significant difference (P<0.01). The same as below.

2.1.2 不同谷物来源淀粉对断奶马驹小肠黏膜葡萄糖转运载体基因表达的影响

不同谷物来源淀粉对断奶马驹小肠黏膜SGLT1和GLUT1基因相对表达量的影响见图2。玉米组小肠黏膜SGLT1基因相对表达量显著高于燕麦组(P<0.05),且极显著高于大麦组(P<0.01);而燕麦组和大麦组之间无显著差异(P>0.05)(图2-A)。这表明与饲喂燕麦和大麦相比,饲喂玉米可提高断奶马驹小肠黏膜SGLT1基因相对表达量。大麦组小肠黏膜GLUT1基因相对表达量显著高于玉米组和燕麦组(P<0.05),而玉米组和燕麦组之间无显著差异(P>0.05)(图2-B)。这表明与饲喂玉米和燕麦相比,饲喂大麦可提高断奶马驹小肠黏膜GLUT1基因相对表达量。
图2 不同谷物来源淀粉对断奶马驹小肠黏膜葡萄糖转运载体基因表达的影响

Fig.2 Effects of different grain sources of starch on small intestinal mucosa glucose transporter gene expression of weaned foals

2.2 不同谷物来源淀粉对断奶马驹肠道生长发育相关基因表达的影响

2.2.1 不同谷物来源淀粉对断奶马驹小肠生长发育相关基因表达的影响

不同谷物来源淀粉对断奶马驹小肠黏膜EGFIGF-1和IGF-1R基因相对表达量的影响见图3。各组之间小肠黏膜EGF基因相对表达量无显著差异(P>0.05)(图3-A)。各组之间小肠黏膜IGF-1基因相对表达量无显著差异(P>0.05)(图3-B)。各组之间小肠黏膜IGF-1R基因相对表达量无显著差异(P>0.05)(图3-C)。
图3 不同谷物来源淀粉对断奶马驹小肠生长发育相关基因表达的影响

Fig.3 Effects of different grain sources of starch on small intestine growth and development related gene expression of weaned foals

2.2.2 不同谷物来源淀粉对断奶马驹盲肠生长发育相关基因表达的影响

不同谷物来源淀粉对断奶马驹盲肠黏膜EGFIGF-1和IGF-1R基因相对表达量的影响见图4。大麦组盲肠黏膜EGF基因相对表达量极显著高于玉米组和燕麦组(P<0.01),燕麦组显著高于玉米组(P<0.05)(图4-A)。这表明与饲喂玉米和燕麦相比,饲喂大麦可提高断奶马驹盲肠黏膜EGF基因相对表达量。各组之间盲肠黏膜IGF-1基因相对表达量无显著差异(P>0.05)(图4-B)。大麦组盲肠黏膜IGF-1R基因相对表达量极显著高于玉米组和燕麦组(P<0.01),而玉米组和燕麦组之间无显著差异(P>0.05)(图4-C)。这表明与饲喂玉米和燕麦相比,饲喂大麦可提高断奶马驹盲肠黏膜IGF-1R基因相对表达量。
图4 不同谷物来源淀粉对断奶马驹盲肠生长发育相关基因表达的影响

Fig.4 Effects of different grain sources of starch on cecum growth and development related gene expression of weaned foals

2.2.3 不同谷物来源淀粉对断奶马驹结肠生长发育相关基因表达的影响

不同谷物来源淀粉对断奶马驹结肠黏膜EGFIGF-1和IGF-1R基因相对表达量的影响见图5。大麦组结肠黏膜EGF基因相对表达量极显著高于燕麦组(P<0.01),且显著高于玉米组(P<0.05);而玉米组显著高于燕麦组(P<0.05)(图5-A)。这表明与饲喂玉米和燕麦相比,饲喂大麦可提高断奶马驹结肠黏膜EGF基因相对表达量。各组之间结肠黏膜IGF-1基因相对表达量无显著差异(P>0.05)(图5-B)。玉米组结肠黏膜IGF-1R基因相对表达量极显著高于燕麦组和大麦组(P<0.01),而燕麦组和大麦组之间无显著差异(P>0.05)(图5-C)。这表明与饲喂燕麦和大麦相比,饲喂玉米可提高断奶马驹结肠黏膜IGF-1R基因相对表达量。
图5 不同谷物来源淀粉对断奶马驹结肠生长发育相关基因表达的影响

Fig.5 Effects of different grain sources of starch on colon growth and development related gene expression of weaned foals

3 讨论

3.1 不同谷物来源淀粉对断奶马驹胃和小肠葡萄糖转运载体基因表达的影响

消化道中的葡萄糖需要通过葡萄糖转运载体介导转运和吸收,葡萄糖的吸收利用除了受到自身浓度的影响外,还受到葡萄糖转运载体活性及数量的影响[10,12,18]SGLT1和GLUT1是胃肠道中介导转运和细胞摄取葡萄糖的主要载体之一,其基因的表达主要受遗传与饲粮组成的影响[12,19-21]。饲粮中淀粉的含量、淀粉分子结构及淀粉在胃肠道中酶解生成葡萄糖的水平是影响葡萄糖转运载体表达的主要因素之一[19,22]。谢晨[23]研究表明,相比饲喂非蜡质玉米淀粉和豌豆淀粉饲粮,饲喂纯蜡质玉米淀粉饲粮可显著提高育肥猪空肠黏膜SGLT1和GLUT2基因相对表达量,这是由于纯蜡质玉米淀粉饲粮显著提高了空肠内葡萄糖含量及黏膜上甜味感受体表达引起的。此外,饲粮中谷物淀粉直链和支链的比例影响葡萄糖转运载体相关基因的表达。任文[24]研究发现,高直链和支链的比例的豌豆淀粉饲粮能够显著提高羔羊小肠中SGLT1和GLUT2基因相对表达量。本研究中,饲喂燕麦的马驹胃黏膜GLUT1基因相对表达量显著高于饲喂玉米和大麦的马驹;而饲喂大麦的马驹小肠GLUT1基因相对表达量显著高于饲喂玉米和燕麦的马驹;但饲喂玉米能够显著提高马驹小肠SGLT1基因相对表达量。研究表明,短链脂肪酸和乳酸杆菌属(Lactobacillus)可以促进胃肠道中胰高血糖素样肽-2(GLP-2)的表达[25],进而提高淀粉酶和麦芽糖酶等活性,促进肠道黏膜发育及增强葡萄糖转运载体表达和活性[26]。而直链淀粉可被马胃中Lactobacillus和链球菌属(Streptococcus)等淀粉降解菌利用降解,产生乳酸和短链脂肪酸[27]。相对于玉米和大麦,燕麦中直链淀粉含量较高,可达25.2%~29.4%[28-30]。同时,Li等[31]研究发现,饲喂燕麦谷物的马驹胃内容物中厚壁菌门(Firmicutes)、乳杆菌科(Lactobacillaceae)、LactobacillusLactobacillus hayakitensis相对丰度高于饲喂玉米和大麦的马驹。因此,燕麦中相对较高的直链淀粉含量改变了胃中细菌的种类和丰度,增加了短链脂肪酸浓度,是本研究中饲喂燕麦的马驹胃黏膜中GLUT1基因相对表达量显著升高的原因。本研究小肠黏膜SGLT1和GLUT1基因表达量差异可能与玉米和大麦直链淀粉含量低[29-30],而支链淀粉高有关。大量支链淀粉被小肠淀粉酶解为葡萄糖,葡萄糖浓度升高是刺激小肠SGLT1和GLUT1基因相对表达量升高的原因[14]

3.2 不同谷物来源淀粉对断奶马驹肠道生长发育相关基因表达的影响

EGFIGF-1与动物肠道组织结构与功能密切相关,能够改善动物对营养物质消化吸收,修复动物受损肠道组织,促进胃肠道生长发育和功能成熟[32-33]。同时,肠道IGF-1R基因相对表达量增加可增强IGF-1促进动物胃肠道发育和完善其生理功能的生理作用[5]
动物肠道发育相关基因(IGF-1、IGF-1R以及EGF)的表达受多种因素调控,如生长发育阶段、激素、介导信号通路及饲粮营养水平等[33-36]。Shen等[37]报道,营养水平是影响哺乳动物IGF-1基因表达的主要因素。饲粮营养物质对基因的表达调控主要是在转录和翻译水平,对翻译后的影响较小[36]。于洋洋[5]研究发现,给羔羊饲喂不同淀粉来源饲粮时,高直链和支链淀粉比例的豌豆淀粉饲粮可显著提高56日龄羔羊十二指肠和回肠IGF-1和IGF-1R基因相对表达量及空肠和回肠EGF基因相对表达量。相振田[25]研究表明,高直链和支链淀粉比例的豌豆淀粉饲粮可显著提高断奶仔猪十二指肠、空肠和回肠中IGF-1、IGF-1RGLP-2基因相对表达量。这表明高直链和支链淀粉比例的饲粮能够促进肠道中生长发育相关基因的表达,反之则抑制生长发育相关基因的表达[38]。本研究中,饲喂大麦的马驹盲肠黏膜EGFIGF-1R和结肠黏膜EGF基因相对表达量显著或极显著高于饲喂玉米和燕麦的马驹;饲喂燕麦的马驹盲肠黏膜EGF基因相对表达量显著高于饲喂玉米的马驹;饲喂玉米的马驹结肠黏膜EGF基因相对表达量显著高于饲喂燕麦的马驹,且IGF-1R基因相对表达量极显著高于饲喂燕麦和大麦的马驹。研究显示,谷物中非淀粉多糖会影响胃肠道的发育[39]。本研究马驹盲肠和结肠黏膜EGFIGF-1R基因表达差异可能与不同谷物中非淀粉多糖种类和含量有关。玉米中富含木聚糖和阿拉伯糖等非淀粉多糖[40],而大麦和燕麦中富含β葡聚糖[41]。这些非淀粉多糖在后肠中可被微生物利用降解产生短链脂肪酸,短链脂肪酸能够为肠道黏膜细胞提供能量而促进细胞增殖,进而促进与生长发育相关基因的表达[42]。Hedemann等[43]研究显示,β葡聚糖可促进大鼠结肠黏膜的生长。此外,β葡聚糖可被微生物利用降解产生短链脂肪酸,从而改变盲肠黏膜部分基因表达[44]。这可能是本研究中饲喂玉米的马驹盲肠和结肠黏膜中EGFIGF-1R基因相对表达量显著或极显著升高的原因。黄新新等[17]研究发现,相比于饲喂玉米和燕麦,给马驹饲喂大麦能够显著或极显著提高盲肠和结肠净重。肠道净重的增加表明其细胞增殖、DNA和蛋白质含量增加,进而可能上调生长发育相关基因的表达[26,33],这可能是本研究中饲喂大麦的马驹盲肠黏膜EGFIGF-1R和结肠黏膜EGF基因相对表达量显著或极显著高于饲喂玉米和燕麦的马驹的原因。

4 结论

① 胃黏膜中葡萄糖转运载体基因表达方面,饲喂燕麦能够提高GLUT1基因相对表达量。
② 小肠黏膜中葡萄糖转运载体基因表达方面,饲喂玉米能够提高SGLT1基因相对表达量,饲喂大麦能够提高GLUT1基因相对表达量,但对生长发育相关基因表达无显著影响。
③ 盲肠黏膜中生长发育相关基因表达方面,饲喂大麦能够提高EGFIGF-1R基因相对表达量。
④ 结肠黏膜中生长发育相关基因表达方面,饲喂玉米和大麦能够提高EGF基因相对表达量,饲喂玉米能够提高IGF-1R基因相对表达量。
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