鸽早期小肠发育及碳水化合物对其调控的研究
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摘要
本研究以泰平王鸽为研究对象,通过研究孵化后期及出壳后两周内小肠形态、消化酶活性和小肠消化吸收相关功能基因表达的变化,揭示鸽早期发育阶段小肠的消化生理特性。在此基础上,采用孵化后期向羊膜腔补充外源营养物质的方法(胚蛋注射营养,In ovo feeding),探讨其对鸽小肠发育的早期营养调控作用。主要研究内容和结果如下：
1.鸽早期小肠形态和消化酶活性的发育规律
肠绒毛在孵化期12d已初步形成,绒毛高度和绒毛表面积在孵化期16d时迅速增加,其中十二指肠和空肠一直持续到14d；绒毛宽度从孵化期14d到出壳后24h内增加显著(P<0.05)；小肠隐窝在出壳当天开始形成内陷结构,并在出壳后3d内生长迅速(P<0.05)；十二指肠和空肠绒毛形态变化显著大于回肠(P<0.05)；肠上皮细胞密度从孵化期12d到出壳后3d显著增加(P<0.05)；粘膜DNA含量从孵化期12d到出壳当天呈线性增加(P<0.05)；蛋白质和DNA的比值出壳3d时开始显著增加(P<0.05)；孵化期12d小肠可检测到较低水平的麦芽糖酶、氨基肽酶-N、碱性磷酸酶和Na+-K+-ATP酶活性,孵化期14d可检测到蔗糖酶活性；二糖酶比活力在出壳后8d时达到最大值,且空肠增长速度显著高于回肠和十二指肠(P<0.05)；氨基肽酶-N比活力在3d后不再随日龄变化而变化；碱性磷酸酶比活力出壳前显著增加(P<0.05);Na+-K+-ATP酶比活力从孵化期12d到出壳后14d内随日龄呈线性增加(P<0.05)；空肠和回肠ATP酶比活力增长速度显著高于十二指肠(P<0.05)。小肠粘膜酶总活力从出壳后3-5d开始显著增加,并与体重随日龄变化显著线性正相关(P<0.05)；孵化期14d可检测到胰蛋白酶和胰凝乳蛋白酶活性,孵化期16d可检测到淀粉酶活性,胰腺酶活性随日龄变化的发育规律与小肠相似。
2.小肠刷状缘酶及相关营养物质转运载体基因的克隆及分析
采用RT-PCR方法克隆得到鸽小肠钠葡萄糖共转运载体SGLT1、葡萄糖转运载体GLUT2、寡肽转运载体PepTl、氨基肽酶APN和蔗糖-异麦芽糖酶SI基因片段长度分别为946bp、762bp、946bp、985bp和700bp。通过NCBI基因数据库Blast搜索结果表明这些序列为目的基因片段,提交到Genbank后获得登陆号分别为JN887478、JN887480、JN887476、JN887477和JN887479;鸽SGLT1基因翻译的氨基酸序列与斑胸草雀、火鸡和鸡一致性均在88%以上,与斑胸草雀的一致性最高,与人、鼠、羊等哺乳类动物也达到80%以上,在系统发育树上与哺乳类动物处于同一分支；鸽GLUT2、PepT1、APN和SI氨基酸序列与鸡的一致性最高,与哺乳类动物一致性较差,在系统发育树上与哺乳类动物处于不同分支。
3.小肠刷状缘酶及相关营养物质转运载体基因表达的时空变化
利用实时荧光定量PCR方法,分析PepT1、SGLT1、GLUT2、APN和SI mRNA相对表达量在十二指肠、空肠和回肠(孵化期12、14和16d,出壳当天,出壳后1、3、5、8和14d)以及卵黄囊膜(孵化期12、14、16d和出壳当天)上的变化规律。结果表明,孵化后期12d,目的基因nRNA在小肠已经开始表达；小肠营养物质转运载体基因nRNA相对表达量随日龄呈一次线性增加(P<0.05),APN mRNA相对表达量随日龄呈二次曲性增加(P<0.05),于孵化期16d显著下降(P<0.05),出壳当天再次增加(P<0.05), SI mRNA相对表达量随日龄呈三次曲性增加(P<0.05),于出壳当天和出壳后8d时分别显著上调(P<0.05)；十二指肠PepT1mRNA相对表达量最高,空肠SGLT1和GLUT2mRNA相对表达量最高,回肠APN mRNA相对表达量最高,小肠各段SI mRNA相对表达量无显著差异(P>0.05)；目的基因mRNA在卵黄囊膜各时期均有表达,且在出壳当天显著下调(P<0.05)。
4.胚蛋注射碳水化合物对鸽生长和能量贮存状态的影响
通过分析鸽胚蛋不同孵化日龄羊水体积变化,确定胚蛋注射时间为孵化期14.5d。选择发育良好的含活胚蛋200枚,编号后随机分成5个处理组,每个处理4个重复(每个重复分列于孵化箱内的不同托盘上)。对照组不注射任何物质,四个处理组注射营养液如下：(Ⅰ)蔗糖15g/L,麦芽糖15g/L, NaCl7.5g/L;(Ⅱ)蔗糖25g/L,麦芽糖25g/L, NaCl7.5g/L;(Ⅲ)蔗糖35g/L,麦芽糖35g/L, NaCl7.5g/L;(Ⅳ)蔗糖45g/L,麦芽糖45g/L, NaCl7.5g/L.预实验表明假注射和生理盐水注射均不会显著抑制鸽胚的生长发育,故正式试验不再设假注射和生理盐水注射作对照。于孵化期16d和出壳当天进行取样分析,通过研究胚蛋注射不同水平碳水化合物溶液对鸽出壳率、体重以及能量贮存状态的影响,探讨胚蛋注射营养技术在鸽中应用的效果。结果表明,从孵化期16d到出壳当天,各组中G6pase活性和血糖浓度均表现出上升的特点,而胸肌和糖原储备均表现出下降的特点(P<0.05)；与对照组相比,试验Ⅱ组显著提高了雏鸽孵化率(P<0.05),但随着碳水化合物注射溶液浓度的继续增加,孵化率呈线性下降(P<0.05)；胚蛋注射48h后,试验Ⅰ和Ⅱ组鸽胚体重显著高于其它各组(P<0.05),试验Ⅱ组鸽胚胸肌重显著高于对照组(P<0.05),试验Ⅱ和Ⅲ组鸽胚小肠重显著高于其它各组(P<0.05)；出壳当天,试验Ⅱ组雏鸽体重和绝对体重显著高于对照组(P<0.05),试验Ⅱ和Ⅲ组卵黄重显著低于其它各组(P<0.05),试验Ⅱ组雏鸽肝脏、胸肌、胃、和小肠重显著高于对照组,且显著提高了小肠相对重(P<0.05)；胚蛋注射48h后,血糖浓度随碳水化合物浓度的增加呈线性增长,G6pase活性则显著下降；胚蛋注射48h后,试验Ⅰ和Ⅱ组鸽胚肝糖原含量均显著高于对照组(P<0.05),出壳当天,试验Ⅱ组肌糖原浓度显著高于对照组(P<0.05)。
5.胚蛋注射碳水化合物对鸽小肠早期发育的调控作用
选择发育良好的含活胚蛋160枚,编号后随机分成2个处理组,每个处理4个重复(每个重复分列于孵化箱内的不同托盘上)。于孵化期14.5d进行胚蛋注射。对照组不注射任何物质,试验组注射营养液成分：蔗糖25g/L,麦芽糖25g/L,NaCl7.5g/L。于孵化期16d和出壳当天进行取样分析,研究胚蛋注射碳水化合物溶液对鸽空肠绒毛形态,刷状缘酶活性和消化吸收相关基因表达的影响。结果表明孵化期16d和出壳当天,胚蛋注射组空肠绒毛表面积比对照组分别增加了38%和23%(P<0.05),空肠麦芽糖酶比活力比对照组分别增加了43%和52%(P<0.05),蔗糖酶比活力比对照组分别增加了39%和26%(P<0.05),空肠碱性磷酸酶比活力比对照组分别增加了44%和35%(P<0.05)；孵化期16d,胚蛋注射组空肠DNA含量比对照组提高了20%(P<0.05)；出壳当天,胚蛋注射组空肠刷状缘氨基肽酶-N比活力比对照组增加了27%(P<0.05)；胚蛋注射碳水化合物显著提高了孵化期16d空肠SGLT1、GLUT2和APN mRNA相对表达量(P<0.05)。
综上所述,孵化后期及出壳后两周内鸽小肠形态、消化酶活性和消化吸收关键基因表达变化显著,且随日龄变化在小肠各段的发育模式各不相同,其中肠粘膜水解作用是鸽小肠早期发育阶段消化吸收营养物质的限速步骤；鸽出壳时小肠发育程度比肉鸡等家禽低,表现为鸽小肠粘膜水解酶比活力的增加和绒毛的生长一直持续到8-14d,而肉鸡等家禽则在出壳后72h内完成；鸽出壳后的早期发育过程中,小肠粘膜及胰腺中碳水化合物类水解酶变化比蛋白质类水解酶显著,表明碳水化合物类水解酶是限制鸽早期小肠消化功能发育成熟的主要因素；孵化期14.5d,向鸽胚羊膜腔中注射碳水化合物(蔗糖25g/L,麦芽糖25g/L,NaCl7.5g/L)可以增加鸽胚机体能量储备、提高鸽出壳体重和孵化率,并可以通过刺激肠上皮细胞增殖,增加绒毛表面积,提高刷状缘水解酶活性和上调消化吸收相关基因表达促进鸽孵化后期小肠的发育,从而提高了鸽出壳时小肠功能成熟程度,表明胚蛋注射可作为鸽小肠发育早期营养调控的有效手段。
This study was conducted to evaluate the digestive capacity in domestic pigeons (Columba livia) by determining the intestinal morphology, mucosal and pancreatic enzyme activities, and mRNA expression of the digestion/absorption related genes during pre-and posthatch development. Furthermore, the nutritional regulation on intestinal development prior to hatching in pigeons by in ovo feeding, a method of supplementing exogenous nutrients into the amnion of the late-term avian embryo, was also evaluated. The main results are listed as follows:
1. Early development of intestinal morphology and digestive enzyme activities in pigeons
Villi were rudimentary at12d of incubation (E12) and rapid increase of villus height and area was observed from E16. In the duodenum and jejunum, villus size continued to increase through14d posthatch (D14). Villus width increased dramatically from E14to24h after hatch (P<0.05). At day of hatch (DOH), crypts invagination was not complete throughout the small intestine, and increased intensively during the first3d posthatch (P<0.05). Villus area and crypt depth increased in parallel in the duodenum and jejunum and more slowly in the ileum (P<0.05). Enterocyte density peaked at3d after hatching in all segments. The mucosa DNA content increased linearly from E12to DOH, whereas the protein/DNA increased from D3(P<0.05). Mucosa enzyme activities were detectable for maltase, aminopeptidase-N (APN), Na+K+ATPase, and alkaline phosphatase (ALP) at E12, and for sucrase at E14. The disaccharidase mass specific activity peaked at D8and increased most dramatically in the jejunum as compared with the duodenum and ileum (P<0.05). Na+K+ATPase activity increased linearly from E12to D14, while APN changed slightly after D3(P<0.05). The ALP activity increased most dramatically before hatching (P<0.05). Intestinal total enzyme activities exhibited a steady increase after3-5d posthach, which was highly correlated with BW (P<0.05). Trypsin and chymotrypsin were detectable at E14, while amylase activity was detectable at E16. Pancreatic enzymes indicated patterns somewhat similar to those for intestinal enzymes.
2. Cloning and analyzing of pigeon brush border enzymes and nutrient transporter genes
The cDNA fragments for sodium glucose transporter SGLT1, glucose transporter GLUT2, oligopeptide transporter PepTl, aminopeptidase-N APN and sucrase-isomaltase SI were isolated and cloned using reverse transcription PCR, containing946bp,762bp,946bp, and985bp and700bp nucleotides, respectively. These cDNA fragments were submitted to Genbank with accession number JN887478, JN887480, JN887476, JN887477, and JN887479, respectively. The predicted amino acid sequence for pigeon SGLT1was more than88%identical to Zebra Finch, turkey and chicken, and showed highest identical to Zebra Finch. The pigeon SGLT1amino acid sequence also showed high identical to mammalias, such as human, sheep and mice, and they were located in the same branch in phylogenetic trees. The predicted amino acid sequences for pigeon GLUT2, PepTl, APN and SI were highly identical to those of chicken, and were located in different branches in phylogenetic trees as compared to mammalias.
3. The ontogeny of nutrient transporter and brush border enzymes gene expression
To better understand the digestive capacity in pigeons, this part was conducted to evaluate nutrient transporters and digestive enzymes gene expression in small intestine and yolk sac membrane (YSM) during pre-and posthatch development. Intestine was collected at embryo d12,14and16, day of hatch, and d1,3,5,8and14posthatch. YSM was collected at embryo d12,14,16and day of hatch. The mRNA expression of each gene was assayed using real-time PCR. The examined genes were all detectable in pigeon small intestine at E12. Expression of intestinal nutrient transporters increased linearly (P<0.05) with age, whereas that of APN increased quadratically (P<0.05) and SI increased cubically (P<0.05). The APN mRNA expression showed a decline from E14to E16, and subsequent increase to D14 (P<0.05), while the greatest mRNA quantities for SI were observed at day of hatch and D8after hatch (P<0.05). Levels of PepTl mRNA were greatest in the duodenum, GLUT2and SGLT1mRNA were greatest in the jejunum, and APN were greatest in the ileum (P<0.05). The YSM expressed all the examined genes. The YSM-expressed genes decreased between embryo d16and day of hatch, whereas intestine-expressed genes increased (P<0.05).
4. Effects of in ovo feeding of carbohydrates on growth and energy status in pigeons
The optimum time for in ovo injection was identified as14.5d of incubation by determining changes of pigeon amniotic fluid volume.200fertile eggs with viable embryos were identified by number and weighed. The eggs were then randomly distributed into5groups of4replications. The injection solutions were:(Ⅰ)15g/L maltose (M)+15g/L sucrose (S),(Ⅱ)25g/L M+25g/L S,(Ⅲ)35g/L M+35g/L%S,(Ⅳ)45g/L M+45g/L S, all dissolved in7.5g/L saline. Preliminary experiments conducted in our laboratory demonstrated that sham injection (shell perforated but without solution injection) or injection of200μL0.75%saline did not affect embryo growth and development. Thus, the other group was not injected and served as the control group in this study. Results showed that, G6pase activity and serum glucose level increased from E6to DOH in all groups, whereas those of pectoral muscle and energy status decreased (P<0.05). Treatment Ⅱ increased (P<0.05) the hatchability as compared with the control. However, the hatchability decreased linearly as the carbohydrate level continued to increase (P<0.05).48h after in ovo injection, treatment Ⅰ and Ⅱ showed highest BW, while treatment Ⅱ and Ⅲ showed highest intestinal weight (P<0.05). Pectoral muscle weight in treatment Ⅱ increased significantly as compared with the control group (P<0.05) at E16. At day of hatch, treatment Ⅱ increased pigeon hatching weight and the yolk sac free BW, while the yolk sac weight decreased in treatment Ⅱ and Ⅲ (P<0.05). The weight of liver, pectoral muscle, stomach and small intestine, as well as the relative intestinal weight, increased significantly as compared with that of the control (P<0.05).48h after in ovo injection, G6pase activity decreased while serum glucose increased compared to the control (P<0.05). The liver glycogen concentration increased in treatment Ⅰ and Ⅱ at E16, and the muscle glycogen increased in treatment Ⅱ at day of hatch (P<0.05).
5. Effects of in ovo injection of carbohydrates on small intestine development in pigeons
At day14.5of incubation, fertile eggs were injected with200μL of carbohydrate solution containing25g/L maltose+25g/L sucrose, all dissolved in7.5g/L saline. Jejunal villus surface area, activity of the brush border enzymes sucrase, maltase, aminopeptidase-N and alkaline phosphatase, and mRNA expression of the digestion/absorption related genes were examined at day16of incubation and day of hatch. Results showed that in ovo injection of carbohydrates caused a villus surface area increase of38%on day16of incubation and23%on day of hatch relative to controls (P<0.05). The in ovo injected pigeons exhibited higher (P<0.05) activities of jejunal sucrase, maltase and alkaline phosphatase per gram of tissue from day16of incubation to day of hatch, compared with the controls. At16d of incubation, the mucosa DNA content increased by20%by in ovo injection as compared to the control group (P<0.05). At day of hatch, aminopeptidase-N activity per gram of tissue in embryos injected in ovo was approximately27%greater (P<0.05) than control embryos. Enhanced expressions of the jejunal SGLT1, GLUT2and APN mRNA were found at day16of incubation in embryos that received carbohydrate solution into the amniotic fluid in comparison with the control group (P<0.05).
In conclusion, changes of pigeon intestinal morphology, enzyme activities, and mRNA expression of the digestion/absorption related genes were rapidly during pre-and posthatch development, although rates of development were different in the three intestinal segments. The results indicated that intestinal hydrolysis may be a determining step in digestion and young pigeons may hatch with a less mature digestive tract as compared to precocial chicks. Changes in carbohydrate-digesting enzymes activity in both small intestine and pancreas are more pronounced than for protein-digesting enzymes activity, suggesting that carbohydrate-digesting enzymes may be a determining step in pigeon intestinal development. In ovo injection of carbohydrate (25g/L maltose+25g/L sucros dissolved in7.5g/L saline) on14.5d of incubation can enhance the hatchability, hatching BW and glycogen reserves. Besides, the in ovo injection of carbohydrate improved villi size, brush border enzymes activity, and digestion/absorption-related intestinal genes mRNA expression and hence hatching intestinal maturity in pigeons. Therefore, the in ovo feeding may serve as a tool to regulate the pigeon intestinal development prior to hatching.
1.鸽早期小肠形态和消化酶活性的发育规律
肠绒毛在孵化期12d已初步形成,绒毛高度和绒毛表面积在孵化期16d时迅速增加,其中十二指肠和空肠一直持续到14d；绒毛宽度从孵化期14d到出壳后24h内增加显著(P<0.05)；小肠隐窝在出壳当天开始形成内陷结构,并在出壳后3d内生长迅速(P<0.05)；十二指肠和空肠绒毛形态变化显著大于回肠(P<0.05)；肠上皮细胞密度从孵化期12d到出壳后3d显著增加(P<0.05)；粘膜DNA含量从孵化期12d到出壳当天呈线性增加(P<0.05)；蛋白质和DNA的比值出壳3d时开始显著增加(P<0.05)；孵化期12d小肠可检测到较低水平的麦芽糖酶、氨基肽酶-N、碱性磷酸酶和Na+-K+-ATP酶活性,孵化期14d可检测到蔗糖酶活性；二糖酶比活力在出壳后8d时达到最大值,且空肠增长速度显著高于回肠和十二指肠(P<0.05)；氨基肽酶-N比活力在3d后不再随日龄变化而变化；碱性磷酸酶比活力出壳前显著增加(P<0.05);Na+-K+-ATP酶比活力从孵化期12d到出壳后14d内随日龄呈线性增加(P<0.05)；空肠和回肠ATP酶比活力增长速度显著高于十二指肠(P<0.05)。小肠粘膜酶总活力从出壳后3-5d开始显著增加,并与体重随日龄变化显著线性正相关(P<0.05)；孵化期14d可检测到胰蛋白酶和胰凝乳蛋白酶活性,孵化期16d可检测到淀粉酶活性,胰腺酶活性随日龄变化的发育规律与小肠相似。
2.小肠刷状缘酶及相关营养物质转运载体基因的克隆及分析
采用RT-PCR方法克隆得到鸽小肠钠葡萄糖共转运载体SGLT1、葡萄糖转运载体GLUT2、寡肽转运载体PepTl、氨基肽酶APN和蔗糖-异麦芽糖酶SI基因片段长度分别为946bp、762bp、946bp、985bp和700bp。通过NCBI基因数据库Blast搜索结果表明这些序列为目的基因片段,提交到Genbank后获得登陆号分别为JN887478、JN887480、JN887476、JN887477和JN887479;鸽SGLT1基因翻译的氨基酸序列与斑胸草雀、火鸡和鸡一致性均在88%以上,与斑胸草雀的一致性最高,与人、鼠、羊等哺乳类动物也达到80%以上,在系统发育树上与哺乳类动物处于同一分支；鸽GLUT2、PepT1、APN和SI氨基酸序列与鸡的一致性最高,与哺乳类动物一致性较差,在系统发育树上与哺乳类动物处于不同分支。
3.小肠刷状缘酶及相关营养物质转运载体基因表达的时空变化
利用实时荧光定量PCR方法,分析PepT1、SGLT1、GLUT2、APN和SI mRNA相对表达量在十二指肠、空肠和回肠(孵化期12、14和16d,出壳当天,出壳后1、3、5、8和14d)以及卵黄囊膜(孵化期12、14、16d和出壳当天)上的变化规律。结果表明,孵化后期12d,目的基因nRNA在小肠已经开始表达；小肠营养物质转运载体基因nRNA相对表达量随日龄呈一次线性增加(P<0.05),APN mRNA相对表达量随日龄呈二次曲性增加(P<0.05),于孵化期16d显著下降(P<0.05),出壳当天再次增加(P<0.05), SI mRNA相对表达量随日龄呈三次曲性增加(P<0.05),于出壳当天和出壳后8d时分别显著上调(P<0.05)；十二指肠PepT1mRNA相对表达量最高,空肠SGLT1和GLUT2mRNA相对表达量最高,回肠APN mRNA相对表达量最高,小肠各段SI mRNA相对表达量无显著差异(P>0.05)；目的基因mRNA在卵黄囊膜各时期均有表达,且在出壳当天显著下调(P<0.05)。
4.胚蛋注射碳水化合物对鸽生长和能量贮存状态的影响
通过分析鸽胚蛋不同孵化日龄羊水体积变化,确定胚蛋注射时间为孵化期14.5d。选择发育良好的含活胚蛋200枚,编号后随机分成5个处理组,每个处理4个重复(每个重复分列于孵化箱内的不同托盘上)。对照组不注射任何物质,四个处理组注射营养液如下：(Ⅰ)蔗糖15g/L,麦芽糖15g/L, NaCl7.5g/L;(Ⅱ)蔗糖25g/L,麦芽糖25g/L, NaCl7.5g/L;(Ⅲ)蔗糖35g/L,麦芽糖35g/L, NaCl7.5g/L;(Ⅳ)蔗糖45g/L,麦芽糖45g/L, NaCl7.5g/L.预实验表明假注射和生理盐水注射均不会显著抑制鸽胚的生长发育,故正式试验不再设假注射和生理盐水注射作对照。于孵化期16d和出壳当天进行取样分析,通过研究胚蛋注射不同水平碳水化合物溶液对鸽出壳率、体重以及能量贮存状态的影响,探讨胚蛋注射营养技术在鸽中应用的效果。结果表明,从孵化期16d到出壳当天,各组中G6pase活性和血糖浓度均表现出上升的特点,而胸肌和糖原储备均表现出下降的特点(P<0.05)；与对照组相比,试验Ⅱ组显著提高了雏鸽孵化率(P<0.05),但随着碳水化合物注射溶液浓度的继续增加,孵化率呈线性下降(P<0.05)；胚蛋注射48h后,试验Ⅰ和Ⅱ组鸽胚体重显著高于其它各组(P<0.05),试验Ⅱ组鸽胚胸肌重显著高于对照组(P<0.05),试验Ⅱ和Ⅲ组鸽胚小肠重显著高于其它各组(P<0.05)；出壳当天,试验Ⅱ组雏鸽体重和绝对体重显著高于对照组(P<0.05),试验Ⅱ和Ⅲ组卵黄重显著低于其它各组(P<0.05),试验Ⅱ组雏鸽肝脏、胸肌、胃、和小肠重显著高于对照组,且显著提高了小肠相对重(P<0.05)；胚蛋注射48h后,血糖浓度随碳水化合物浓度的增加呈线性增长,G6pase活性则显著下降；胚蛋注射48h后,试验Ⅰ和Ⅱ组鸽胚肝糖原含量均显著高于对照组(P<0.05),出壳当天,试验Ⅱ组肌糖原浓度显著高于对照组(P<0.05)。
5.胚蛋注射碳水化合物对鸽小肠早期发育的调控作用
选择发育良好的含活胚蛋160枚,编号后随机分成2个处理组,每个处理4个重复(每个重复分列于孵化箱内的不同托盘上)。于孵化期14.5d进行胚蛋注射。对照组不注射任何物质,试验组注射营养液成分：蔗糖25g/L,麦芽糖25g/L,NaCl7.5g/L。于孵化期16d和出壳当天进行取样分析,研究胚蛋注射碳水化合物溶液对鸽空肠绒毛形态,刷状缘酶活性和消化吸收相关基因表达的影响。结果表明孵化期16d和出壳当天,胚蛋注射组空肠绒毛表面积比对照组分别增加了38%和23%(P<0.05),空肠麦芽糖酶比活力比对照组分别增加了43%和52%(P<0.05),蔗糖酶比活力比对照组分别增加了39%和26%(P<0.05),空肠碱性磷酸酶比活力比对照组分别增加了44%和35%(P<0.05)；孵化期16d,胚蛋注射组空肠DNA含量比对照组提高了20%(P<0.05)；出壳当天,胚蛋注射组空肠刷状缘氨基肽酶-N比活力比对照组增加了27%(P<0.05)；胚蛋注射碳水化合物显著提高了孵化期16d空肠SGLT1、GLUT2和APN mRNA相对表达量(P<0.05)。
综上所述,孵化后期及出壳后两周内鸽小肠形态、消化酶活性和消化吸收关键基因表达变化显著,且随日龄变化在小肠各段的发育模式各不相同,其中肠粘膜水解作用是鸽小肠早期发育阶段消化吸收营养物质的限速步骤；鸽出壳时小肠发育程度比肉鸡等家禽低,表现为鸽小肠粘膜水解酶比活力的增加和绒毛的生长一直持续到8-14d,而肉鸡等家禽则在出壳后72h内完成；鸽出壳后的早期发育过程中,小肠粘膜及胰腺中碳水化合物类水解酶变化比蛋白质类水解酶显著,表明碳水化合物类水解酶是限制鸽早期小肠消化功能发育成熟的主要因素；孵化期14.5d,向鸽胚羊膜腔中注射碳水化合物(蔗糖25g/L,麦芽糖25g/L,NaCl7.5g/L)可以增加鸽胚机体能量储备、提高鸽出壳体重和孵化率,并可以通过刺激肠上皮细胞增殖,增加绒毛表面积,提高刷状缘水解酶活性和上调消化吸收相关基因表达促进鸽孵化后期小肠的发育,从而提高了鸽出壳时小肠功能成熟程度,表明胚蛋注射可作为鸽小肠发育早期营养调控的有效手段。
This study was conducted to evaluate the digestive capacity in domestic pigeons (Columba livia) by determining the intestinal morphology, mucosal and pancreatic enzyme activities, and mRNA expression of the digestion/absorption related genes during pre-and posthatch development. Furthermore, the nutritional regulation on intestinal development prior to hatching in pigeons by in ovo feeding, a method of supplementing exogenous nutrients into the amnion of the late-term avian embryo, was also evaluated. The main results are listed as follows:
1. Early development of intestinal morphology and digestive enzyme activities in pigeons
Villi were rudimentary at12d of incubation (E12) and rapid increase of villus height and area was observed from E16. In the duodenum and jejunum, villus size continued to increase through14d posthatch (D14). Villus width increased dramatically from E14to24h after hatch (P<0.05). At day of hatch (DOH), crypts invagination was not complete throughout the small intestine, and increased intensively during the first3d posthatch (P<0.05). Villus area and crypt depth increased in parallel in the duodenum and jejunum and more slowly in the ileum (P<0.05). Enterocyte density peaked at3d after hatching in all segments. The mucosa DNA content increased linearly from E12to DOH, whereas the protein/DNA increased from D3(P<0.05). Mucosa enzyme activities were detectable for maltase, aminopeptidase-N (APN), Na+K+ATPase, and alkaline phosphatase (ALP) at E12, and for sucrase at E14. The disaccharidase mass specific activity peaked at D8and increased most dramatically in the jejunum as compared with the duodenum and ileum (P<0.05). Na+K+ATPase activity increased linearly from E12to D14, while APN changed slightly after D3(P<0.05). The ALP activity increased most dramatically before hatching (P<0.05). Intestinal total enzyme activities exhibited a steady increase after3-5d posthach, which was highly correlated with BW (P<0.05). Trypsin and chymotrypsin were detectable at E14, while amylase activity was detectable at E16. Pancreatic enzymes indicated patterns somewhat similar to those for intestinal enzymes.
2. Cloning and analyzing of pigeon brush border enzymes and nutrient transporter genes
The cDNA fragments for sodium glucose transporter SGLT1, glucose transporter GLUT2, oligopeptide transporter PepTl, aminopeptidase-N APN and sucrase-isomaltase SI were isolated and cloned using reverse transcription PCR, containing946bp,762bp,946bp, and985bp and700bp nucleotides, respectively. These cDNA fragments were submitted to Genbank with accession number JN887478, JN887480, JN887476, JN887477, and JN887479, respectively. The predicted amino acid sequence for pigeon SGLT1was more than88%identical to Zebra Finch, turkey and chicken, and showed highest identical to Zebra Finch. The pigeon SGLT1amino acid sequence also showed high identical to mammalias, such as human, sheep and mice, and they were located in the same branch in phylogenetic trees. The predicted amino acid sequences for pigeon GLUT2, PepTl, APN and SI were highly identical to those of chicken, and were located in different branches in phylogenetic trees as compared to mammalias.
3. The ontogeny of nutrient transporter and brush border enzymes gene expression
To better understand the digestive capacity in pigeons, this part was conducted to evaluate nutrient transporters and digestive enzymes gene expression in small intestine and yolk sac membrane (YSM) during pre-and posthatch development. Intestine was collected at embryo d12,14and16, day of hatch, and d1,3,5,8and14posthatch. YSM was collected at embryo d12,14,16and day of hatch. The mRNA expression of each gene was assayed using real-time PCR. The examined genes were all detectable in pigeon small intestine at E12. Expression of intestinal nutrient transporters increased linearly (P<0.05) with age, whereas that of APN increased quadratically (P<0.05) and SI increased cubically (P<0.05). The APN mRNA expression showed a decline from E14to E16, and subsequent increase to D14 (P<0.05), while the greatest mRNA quantities for SI were observed at day of hatch and D8after hatch (P<0.05). Levels of PepTl mRNA were greatest in the duodenum, GLUT2and SGLT1mRNA were greatest in the jejunum, and APN were greatest in the ileum (P<0.05). The YSM expressed all the examined genes. The YSM-expressed genes decreased between embryo d16and day of hatch, whereas intestine-expressed genes increased (P<0.05).
4. Effects of in ovo feeding of carbohydrates on growth and energy status in pigeons
The optimum time for in ovo injection was identified as14.5d of incubation by determining changes of pigeon amniotic fluid volume.200fertile eggs with viable embryos were identified by number and weighed. The eggs were then randomly distributed into5groups of4replications. The injection solutions were:(Ⅰ)15g/L maltose (M)+15g/L sucrose (S),(Ⅱ)25g/L M+25g/L S,(Ⅲ)35g/L M+35g/L%S,(Ⅳ)45g/L M+45g/L S, all dissolved in7.5g/L saline. Preliminary experiments conducted in our laboratory demonstrated that sham injection (shell perforated but without solution injection) or injection of200μL0.75%saline did not affect embryo growth and development. Thus, the other group was not injected and served as the control group in this study. Results showed that, G6pase activity and serum glucose level increased from E6to DOH in all groups, whereas those of pectoral muscle and energy status decreased (P<0.05). Treatment Ⅱ increased (P<0.05) the hatchability as compared with the control. However, the hatchability decreased linearly as the carbohydrate level continued to increase (P<0.05).48h after in ovo injection, treatment Ⅰ and Ⅱ showed highest BW, while treatment Ⅱ and Ⅲ showed highest intestinal weight (P<0.05). Pectoral muscle weight in treatment Ⅱ increased significantly as compared with the control group (P<0.05) at E16. At day of hatch, treatment Ⅱ increased pigeon hatching weight and the yolk sac free BW, while the yolk sac weight decreased in treatment Ⅱ and Ⅲ (P<0.05). The weight of liver, pectoral muscle, stomach and small intestine, as well as the relative intestinal weight, increased significantly as compared with that of the control (P<0.05).48h after in ovo injection, G6pase activity decreased while serum glucose increased compared to the control (P<0.05). The liver glycogen concentration increased in treatment Ⅰ and Ⅱ at E16, and the muscle glycogen increased in treatment Ⅱ at day of hatch (P<0.05).
5. Effects of in ovo injection of carbohydrates on small intestine development in pigeons
At day14.5of incubation, fertile eggs were injected with200μL of carbohydrate solution containing25g/L maltose+25g/L sucrose, all dissolved in7.5g/L saline. Jejunal villus surface area, activity of the brush border enzymes sucrase, maltase, aminopeptidase-N and alkaline phosphatase, and mRNA expression of the digestion/absorption related genes were examined at day16of incubation and day of hatch. Results showed that in ovo injection of carbohydrates caused a villus surface area increase of38%on day16of incubation and23%on day of hatch relative to controls (P<0.05). The in ovo injected pigeons exhibited higher (P<0.05) activities of jejunal sucrase, maltase and alkaline phosphatase per gram of tissue from day16of incubation to day of hatch, compared with the controls. At16d of incubation, the mucosa DNA content increased by20%by in ovo injection as compared to the control group (P<0.05). At day of hatch, aminopeptidase-N activity per gram of tissue in embryos injected in ovo was approximately27%greater (P<0.05) than control embryos. Enhanced expressions of the jejunal SGLT1, GLUT2and APN mRNA were found at day16of incubation in embryos that received carbohydrate solution into the amniotic fluid in comparison with the control group (P<0.05).
In conclusion, changes of pigeon intestinal morphology, enzyme activities, and mRNA expression of the digestion/absorption related genes were rapidly during pre-and posthatch development, although rates of development were different in the three intestinal segments. The results indicated that intestinal hydrolysis may be a determining step in digestion and young pigeons may hatch with a less mature digestive tract as compared to precocial chicks. Changes in carbohydrate-digesting enzymes activity in both small intestine and pancreas are more pronounced than for protein-digesting enzymes activity, suggesting that carbohydrate-digesting enzymes may be a determining step in pigeon intestinal development. In ovo injection of carbohydrate (25g/L maltose+25g/L sucros dissolved in7.5g/L saline) on14.5d of incubation can enhance the hatchability, hatching BW and glycogen reserves. Besides, the in ovo injection of carbohydrate improved villi size, brush border enzymes activity, and digestion/absorption-related intestinal genes mRNA expression and hence hatching intestinal maturity in pigeons. Therefore, the in ovo feeding may serve as a tool to regulate the pigeon intestinal development prior to hatching.
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