动物营养学报 >

2024 , Vol. 36 >Issue 7: 4243 - 4257

DOI: https://doi.org/10.12418/CJAN2024.366

研究论文

标准全消化道可消化钙、磷含量对断奶仔猪肠-肝-肾轴钙、磷吸收的影响及机制

  • 孙焱杰 , 1, 2 ,
  • 韦洋洋 1, 2 ,
  • 苗振彦 2 ,
  • 熊云霞 2 ,
  • 吴绮雯 2 ,
  • 杨雪芬 2 ,
  • 王丽 2 ,
  • 蒋宗勇 2 ,
  • 易宏波 , 2, * ,
  • 黄艳娜 , 1, *
展开
  • 1 广西大学动物科学技术学院,南宁 530004
  • 2 广东省农业科学院动物科学研究所,猪禽种业全国重点实验室,农业农村部华南动物营养与饲料重点实验室,广东省畜禽育种与营养研究重点实验室,广州 510640
*易宏波,副研究员,硕士生导师,E-mail: ;
黄艳娜,教授,硕士生导师,E-mail:

孙焱杰(1999—),男,四川内江人,硕士研究生,研究方向为动物营养与饲料科学。E-mail:

Copy editor: 陈鑫

收稿日期: 2024-01-13

  网络出版日期: 2024-07-09

基金资助

国家“十四五”重点研发专项(2021YFD1300402)

国家现代农业产业技术体系(CAR-35)

广东省乡村振兴战略专项资金项目(2023TS-3)

科技创新战略专项资金-高水平农科院建设(202106TD)

科技创新战略专项资金-高水平农科院建设(R2023PY-JX014)

收起

Effects and Mechanism of Standard Total Tract Digestibility Calcium and Phosphorus Contents on Calcium and Phosphorus Absorption in Gut-Liver-Kidney Axis of Weaned Piglets

  • SUN Yanjie , 1, 2 ,
  • WEI Yangyang 1, 2 ,
  • MIAO Zhenyan 2 ,
  • XIONG Yunxia 2 ,
  • WU Qiwen 2 ,
  • YANG Xuefen 2 ,
  • WANG Li 2 ,
  • JIANG Zongyong 2 ,
  • YI Hongbo , 2, * ,
  • HUANG Yanna , 1, *
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  • 1 School of Animal Science and Technology, Guangxi University, Nanning 530004, China
  • 2 Guangdong Key Laboratory of Animal Breeding and Nutrition, Key Laboratory of Animal Nutrition and Feed Science in South China,Ministry of Agriculture and Rural Affairs, State Key Laboratory of Swine and Poultry Breeding Industry, Institute of Animal Science, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China
*YI Hongbo, associate professor, E-mail: ;
HUANG Yanna, professor, E-mail:

Received date: 2024-01-13

  Online published: 2024-07-09

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摘要

本试验旨在研究标准全消化道可消化钙(STTD Ca)和标准全消化道可消化磷(STTD P)含量对7~25 kg断奶仔猪钙、磷吸收的影响。选取21日龄、体重接近的杜长大杂交断奶仔猪144头,随机分为6组,每组6个重复,每个重复4头猪(阉公猪和母猪各2头)。第1~6组饲粮中STTD Ca和STTD P的比例均为1.2,STTD P含量分别为0.18%、0.26%、0.34%、0.42%、0.50%、0.58%,STTD Ca含量分别为0.216%、0.312%、0.408%、0.504%、0.600%、0.696%。试验期42 d。结果表明:1)随着饲粮中STTD Ca和STTD P的含量逐渐升高,血清碱性磷酸酶活性逐渐降低,但组间没有显著差异(P>0.05),血清降钙素(CT)、甲状腺素(T4)、1,25二羟维生素D3[1,25(OH)D3]、甲状旁腺素(PTH)含量逐渐升高,且6组显著高于1组(P<0.05)。1组血清中钙含量显著低于其他组(P<0.05),其中2、4和6组的血清钙含量较于其他组更高;1组血清磷含量显著高于5组(P<0.05)。2)饲粮中STTD Ca和STTD P的含量逐渐升高对胃内容物及盲肠内容物pH无显著影响(P>0.05)。3)所有组肠道结构完整,1和2组十二指肠绒毛高度/隐窝深度显著高于5和6组(P<0.05), 2组空肠绒毛高度/隐窝深度显著低于5组(P<0.05)。4)在十二指肠中,2和3组维生素D受体(VDR)的基因相对表达量显著高于1和5组(P<0.05),1和2组钙磷结合蛋白1(CALB1)的基因相对表达量显著高于4、5和6组(P<0.05),1组细胞色素P450家族成员24A1(CYP24A1)的基因相对表达量显著高于其他组(P<0.05);在空肠中,4和6组的钙离子通道6(TRPV6)的基因相对表达量显著高于2和5组(P<0.05),6组S100钙结合蛋白G(S100G)的基因相对表达量显著高于2组(P<0.05),4组ATP酶质膜钙离子转运蛋白1(ATP2B1)的基因相对表达量显著高于2组(P<0.05),5和6组溶质载体家族蛋白34成员A3(SLC34A3)的基因相对表达量显著高于1和2组(P<0.05),1组CYP24A1的基因相对表达量显著高于其他组(P<0.05);在回肠中,CYP24A1和SLC34A3的基因相对表达量逐渐降低,1和2组CYP24A1的基因相对表达量显著高于其他组(P<0.05),2组SLC34A3的基因相对表达量显著高于其他组(P<0.05);在结肠中,1组TRPV6、ATP2B1和CALB1的基因相对表达量显著高于其他组(P<0.05)。在肾脏中,甲状旁腺素1受体(PTH1R)和钙离子通道5(TRPV5)的基因相对表达量先升高后降低,当升高到NRC(2012)推荐STTD P添加量(STTD P含量为0.34%~0.42%)时到达最大;3和4组PTH1R的基因相对表达量显著高于其他组(P<0.05),4组TRPV5的基因相对表达量显著高于其他组(P<0.05)。在肝脏中,2组SLC34A3的基因相对表达量显著高于4组(P<0.05),1组CALB1的基因相对表达量显著高于其他组(P<0.05)。综上所述,饲粮中STTD Ca和STTD P的比例为1.2时,随着饲粮中STTD Ca和STTD P的含量逐渐升高,十二指肠和结肠对钙、磷的吸收能力逐渐减弱,空肠对钙、磷的吸收能力逐渐增强,肝脏对钙、磷吸收能力逐渐降低。当饲粮中STTD Ca和STTD P的含量增加到NRC(2012)推荐的STTD P添加量(饲粮STTD P含量为0.34%~0.42%)时,肾脏对钙、磷重吸收的能力最强。结合血清钙、磷含量数据,当饲粮中STTD Ca和STTD P的含量分别在0.216%~0.312%和0.18%~0.26%时,断奶仔猪能有效利用饲粮钙、磷。

关键词： 断奶仔猪; STTD Ca; STTD P; 肠道; 肾脏; 肝脏; 钙、磷吸收

本文引用格式

孙焱杰 , 韦洋洋 , 苗振彦 , 熊云霞 , 吴绮雯 , 杨雪芬 , 王丽 , 蒋宗勇 , 易宏波 , 黄艳娜 . 标准全消化道可消化钙、磷含量对断奶仔猪肠-肝-肾轴钙、磷吸收的影响及机制[J]. 动物营养学报, 2024 , 36(7) : 4243 -4257 . DOI: 10.12418/CJAN2024.366

Abstract

The objective of this study was to investigate the effects of standardized total tract digestibility Ca (STTD Ca) and standardized total tract digestibility P (STTD P) contents on calcium and phosphorus absorption in 7 to 25 kg weaned piglets. A total of 144 weaned piglets with a similar body weight at 21 days of age were randomly divided into 6 groups, with 6 replicates in each group and 4 pigs (two castrated boars and two sows) in each replicate. The proportions of STTD Ca and STTD P in the diet were 1.2, and the contents of STTD P were 0.18%, 0.26%, 0.34%, 0.42%, 0.50% and 0.58%, respectively, and the contents of STTD Ca were 0.216%, 0.312%, 0.408%, 0.504%, 0.600% and 0.696%, respectively. The trial period was 42 days. The results showed as follows: 1) with the gradual increase of the contents of STTD Ca and STTD P in the diet, the serum alkaline phosphatase activity gradually decreased, but there was no significant difference between the groups (P>0.05), and the serum contents of calcitonin (CT), thyroxine (T4), 1,25 dihydroxyvitamin D3 [1,25(OH)D3] and parathyroid hormone (PTH) increased gradually, and group 6 were significantly higher than group 1 (P<0.05). The serum calcium content in group 1 was significantly lower than that in other groups (P<0.05), and the serum calcium content in groups 2, 4 and 6 was higher than that in other groups. The serum phosphorus content in group 1 was significantly higher than that in group 5 (P<0.05). 2) The contents of STTD Ca and STTD P in the diet increased gradually, and there was no significant effect on the pH of gastric contents and cecal contents (P>0.05). 3) The intestinal structure of all groups was intact, the duodenal villus height/crypt depth in groups 1 and 2 was significantly higher than that in groups 5 and 6 (P<0.05), and the jejunal villus height/crypt depth in group 2 was significantly lower than that in group 5 (P<0.05). 4) In the duodenum, the relative expression levels of vitamin D receptor (VDR) gene in groups 2 and 3 were significantly higher than those in groups 1 and 5 (P<0.05), the relative expression levels of calcium and phosphorus binding protein 1 (CALB1) gene in groups 1 and 2 were significantly higher than those in groups 4, 5 and 6 (P<0.05), and the relative expression levels of cytochrome P450 family member 24A1 (CYP24A1) gene in group 1 were significantly higher than those in other groups (P<0.05). In the jejunum, the expression levels of calcium channel 6 (TRPV6) gene in groups 4 and 6 were significantly higher than those in groups 2 and 5 (P<0.05), the relative expression levels of S100G gene in group 6 was significantly higher than those in group 2 (P<0.05), the relative expression level of ATPase plasma membrane calcium ion transporter 1 (ATP2B1) gene in group 4 was significantly higher than that in group 2 (P<0.05), the relative expression levels of A3 (SLC34A3) member of solute carrier family protein 34 in groups 5 and 6 that significantly higher than that in groups 1 and 2 (P<0.05), and the relative expression level of CYP24A1 genes in group 1 was significantly higher than those in other groups (P<0.05). In the ileum, the relative expression levels of CYP24A1 and SLC34A3 genes decreased gradually, and the relative expression levels of CYP24A1 gene in groups 1 and 2 were significantly higher than those in other groups (P<0.05), and that of SLC34A3 gene in group 2 was significantly higher than that in other groups (P<0.05). In the colon, the relative expression levels of TRPV6, ATP2B1 and CALB1 genes in group 1 were significantly higher than those in other groups (P<0.05). In the kidney, the gene expression levels of parathyroid hormone 1 receptor (PTH1R) and calcium ion channel 5 (TRPV5) increased first and then decreased, and reached the maximum when it increased to the recommended STTD P addition in NRC (2012) diet (STTD P content was 0.34% to 0.42%). The relative expression level of PTH1R gene in groups 3 was significantly higher than that in the other groups (P<0.05), and the relative expression of TRPV5 gene in the group 4 was significantly higher than that in the other groups (P<0.05). In the liver, the relative expression level of SLC34A3 gene in group 2 was significantly higher than that in group 4 (P<0.05), and the relative expression level of CALB1 gene in group 1 was significantly higher than that in other groups (P<0.05). In summary, when the ratio of STTD Ca and STTD P in the diet is 1.2, with the gradual increase of the contents of STTD Ca and STTD P in the diet, the absorption capacity of calcium and phosphorus in duodenum and colon gradually weaken, the absorption capacity of calcium and phosphorus in jejunum gradually increase, and the absorption capacity of calcium and phosphorus in the liver gradually decrease. When the contents of STTD Ca and STTD P in the diet increase to the STTD P addition recommended by NRC (2012) (the content of STTD P in the diet is 0.34% to 0.42%), the ability of the kidney to reabsorb calcium and phosphorus is the strongest. Combined with serum calcium and phosphorus data, when the contents of STTD Ca and STTD P in the diet are 0.216% to 0.312% and 0.18% to 0.26%, respectively, weaned piglets can effectively use the calcium and phosphorus in the diet.

Key words： weaned piglets; STTD Ca; STTD P; intestine; kidney; liver; calcium and phosphorus absorption

钙、磷的吸收70%~90%发生在小肠,总吸收的10%发生在胃和大肠[1]。肾脏是钙、磷重吸收的主要场所。未被肠道吸收的钙、磷主要通过粪便排出体外,而细胞外液中未被吸收利用的钙、磷离子经过肾小球时,在近端小管和远端小管被重新吸收,多余的钙、磷则通过尿液排出体外[2]。钙、磷的吸收受到多种因素的影响,其中当饲粮中钙含量过高时,会与磷结合,形成不会被肠道吸收的聚合物,从而降低磷的吸收率,所以饲粮中钙、磷的比例至关重要[3⇓-5]。有数据表明,基于标准全消化道可消化钙(STTD Ca)配制的饲粮在节约资源和促进断奶仔猪生长发育方面是优于基于总钙配制的饲粮,同时与其他STTD Ca和标准全消化道可消化磷(STTD P)比例的饲粮相比,当STTD Ca和STTD P比例在1~1.35时,断奶仔猪的生长性能会有所增加[6-7]。进一步研究表明,当STTD Ca和STTD P比例约为1.2时,断奶仔猪的平均日增重和料重比较高[5,8]。在本团队前期研究中,对平均体重分别为10.38 kg(n=50)和24.36 kg(n=50)的断奶仔猪体成分中的钙、磷进行了测定,结果表明,断奶仔猪的体成分Ca和P比值约为1.2[9]。因此,本试验研究了饲粮中STTD Ca和STTD P比例为1.2对7~25 kg断奶仔猪血清生化指标、肠道形态、肠-肝-肾轴调控钙、磷吸收相关基因的影响,旨在为实际生产提供基础数据,以期能够实现降本增效的目的。

1 材料与方法

1.1 试验饲粮

在研究开始之前,用于试验的饲料原料的钙和磷含量分别按照GB/T 6432—2018、GB/T 6437—2018的方法测定。根据原料钙的标准全消化道消化率和磷的标准全消化道消化率的数据计算出原料的STTD Ca和STTD P的含量,以便进一步配制饲粮。饲料原料钙和磷含量见表1。同时,为了消除植酸酶对试验的影响,试验中没有添加植酸酶。试验分为2个阶段,7~11 kg阶段和12~25 kg阶段。2个阶段的饲粮STTD Ca和STTD P的比例是相同的,但不同阶段仔猪对STTD Ca和STTD P的需求不同。因此,使用NRC(2012)推荐的STTD P添加量作为中间值,2个阶段的推荐STTD P添加量差值作为浓度梯度[10]。最终确定饲粮中STTD Ca和STTD P的比例均为1.2,分别配制6种饲粮,STTD P含量分别为0.18%、0.26%、0.34%、0.42%、0.50%、0.58%,STTD Ca含量分别为0.216%、0.312%、0.408%、0.504%、0.600%、0.696%,其他参照NRC(2012)营养需要配制,制作为颗粒饲料。试验饲粮组成及营养水平见表2。
表1 饲料原料钙和磷含量及相关指标(风干基础)

Table 1 Calcium and phosphorus contents of feed raw materials and relate indexes (air-dry basis)%

项目
Items		 钙含量
Ca
content		 磷含量
P
content		 钙的标准全
消化道消化率
STTD of Ca		 磷的标准全
消化道消化率
STTD of P		 标准全消化道
可消化钙含量
STTD Ca
content		 标准全消化道
可消化磷含量
STTD P
content		 来源
Source
玉米 Corn 0.04 0.21 43.00 34.00 0.02 0.07
[11]
豆粕 Soybean meal 0.28 0.50 66.00 48.00 0.18 0.24
低蛋白乳清粉
Low protein whey powder		 0.37 0.49 82.00 92.00 0.30 0.45


[12-13]
浓缩蛋白乳清粉
Concentrate protein
whey powder		 0.25 0.26 88.00 92.00 0.30 0.45
石粉 Limestone 36.41 0.45 71.00 88.30 25.85 0.40 [14-16]
磷酸二氢钙 Ca(H2PO4)2 15.47 22.87 86.00 88.30 13.30 20.19
[17]
磷酸二氢钠 NaH2PO4 0.09 17.56 77.00 93.80 0.07 16.47

浓缩蛋白乳清粉作为非常规原料,含有11.70 MJ/kg 的净能,76.32%的粗蛋白质,1.33%的粗纤维,0.20%的粗脂肪。

Concentrate protein whey powder is an uncommon raw material, containing 11.70 MJ/kg net energy, 76.32% crude protein, 1.33% crude fiber, and 0.20% crude fat.

表2 试验饲粮组成及营养水平(风干基础)

Table 2 Composition and nutrient levels of experimental diets (air-dry basis)%

项目
Items		 阶段1 Phase 1 阶段 2 Phase 2
1组
Group
1		 2组
Group
2		 3组
Group
3		 4组
Group
4		 5组
Group
5		 6组
Group
6		 1组
Group
1		 2组
Group
2		 3组
Group
3		 4组
Group
4		 5组
Group
5		 6组
Group
6
原料 Ingredients
玉米 Corn 29.51 29.51 29.51 29.51 29.51 29.51 52.82 52.82 52.82 52.82 52.82 52.82
膨化玉米 Puffed corn 25.00 25.00 25.00 25.00 25.00 25.00 10.00 10.00 10.00 10.00 10.00 10.00
豆粕 Soybean meal 16.97 16.97 16.97 16.97 16.97 16.97 22.64 22.64 22.64 22.64 22.64 22.64
低蛋白乳清粉
Low protein whey powder		 10.00 10.00 10.00 10.00 10.00 10.00 3.00 3.00 3.00 3.00 3.00 3.00
浓缩蛋白乳清粉
Whey protein concentrate		 10.00 10.00 10.00 10.00 10.00 10.00 3.00 3.00 3.00 3.00 3.00 3.00
大豆油 Soybean oil 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00
50%氯化胆碱
50% choline chloride		 0.20 0.20 0.20 0.20 0.20 0.20 0.20 0.20 0.20 0.20 0.20 0.20
氯化钠 NaCl 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40
L-赖氨酸 L-lysine 0.37 0.37 0.37 0.37 0.37 0.37 0.46 0.46 0.46 0.46 0.46 0.46
DL-蛋氨酸 DL-methionine 0.14 0.14 0.14 0.14 0.14 0.14 0.14 0.14 0.14 0.14 0.14 0.14
L-苏氨酸 L-threonine 0.07 0.07 0.07 0.07 0.07 0.07 0.13 0.13 0.13 0.13 0.13 0.13
L-缬氨酸 L-valine 0.04 0.04 0.04 0.04 0.04 0.04
二氧化钛 TiO2 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40
氧化锌 ZnO 0.20 0.20 0.20 0.20 0.20 0.20
预混料 Premix1) 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00
石粉 Limestone 0.42 0.62 0.82 1.02 1.21 1.41 0.47 0.67 0.86 1.05 1.25 1.43
磷酸二氢钙 Ca(H2PO4)2 0.10 0.44 0.77 1.12 1.46 1.80 0.20 0.53 0.89 1.24 1.59 1.94
磷酸二氢钠 NaH2PO4 0.05 0.12 0.19 0.25 0.31 0.37 0.11 0.18 0.23 0.28 0.33 0.38
碳酸氢钠 NaHCO3 0.36 0.31 0.27 0.23 0.19 0.16 0.19 0.15 0.12 0.08 0.05 0.02
二氧化硅 SiO2 2.81 2.25 1.69 1.12 0.57 2.80 2.24 1.67 1.12 0.55
合计 Total 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00
营养水平 Nutrient levels2)
净能 NE/(MJ/kg) 10.69 10.69 10.69 10.69 10.69 10.69 10.47 10.47 10.47 10.47 10.47 10.47
粗蛋白质 CP 20.90 21.08 21.03 21.06 21.10 21.05 19.02 19.03 18.08 19.01 19.02 19.01
标准回肠可消化赖氨酸 SID Lys 1.48 1.48 1.48 1.48 1.48 1.48 1.27 1.27 1.27 1.27 1.27 1.27
标准回肠可消化苏氨酸 SID Thr 0.87 0.87 0.86 0.87 0.87 0.87 0.75 0.74 0.75 0.75 0.76 0.75
标准回肠可消化蛋氨酸 SID Met 0.47 0.48 0.48 0.47 0.48 0.48 0.41 0.41 0.42 0.41 0.41 0.41
标准回肠可消化色氨酸 SID Tyr 0.28 0.27 0.28 0.28 0.28 0.28 0.21 0.21 0.22 0.21 0.21 0.21
标准回肠可消化缬氨酸 SID Val 0.92 0.93 0.93 0.93 0.93 0.93 0.93 0.93 0.92 0.93 0.93 0.93
钙 Ca 0.330 0.391 0.528 0.626 0.738 0.882 0.410 0.423 0.512 0.638 0.799 0.894
磷 P 0.291 0.412 0.523 0.594 0.689 0.762 0.33 0.397 0.502 0.594 0.686 0.732

1) 预混料为每千克饲粮提供 Premix provided the following per kg of diets:VA 9 920 IU,VD3 2 240 IU,VE 24 mg,VK3 4 mg,VB1 2.4 mg,VB2 8 mg,VB6 6.4 mg,VB12 32 mg,烟酸 nicotinic acid 32 mg,泛酸 pantothenic acid 12 mg,叶酸 folic acid 0.8 mg,生物素 biotin 0.64 mg,Fe(as ferrous sulfate) 90 mg,Cu (as copper sulfate) 12 mg,Mn (as manganese sulfate) 52.5 mg,I (as potassium iodide) 0.525 mg,Se (as sodium selenite) 0.36 mg,Zn (as zinc sulfate) 60 mg。

2)粗蛋白质、钙、磷为实测值,其余为计算值。CP,Ca and P were measured values,while the rest were calculated values.

1.2 试验设计及饲养管理

本试验通过广东省农业科学院动物科学研究所实验动物福利伦理委员会的审查(批准编号GDIAS20221103)。试验选取21日龄杜长大杂交断奶仔猪144头,随机分为6组,分别饲喂以上6种饲粮,每组6个重复,每个重复4头猪(阉公猪和母猪各2头),每栏之间的体重没有显著差异(P>0.05)。试验期42 d。全期试验仔猪自由采食,充足饮水,所有猪只均按正常免疫程序进行免疫接种。

1.3 检测指标与方法

1.3.1 饲粮营养成分

依据NRC(2012)原料成分表中各原料净能计算饲粮净能;依据NRC(2012)原料成分表中各原料标准回肠可消化氨基酸与总氨基酸间的比例计算饲粮标准回肠可消化氨基酸含量;按照GB/T 6432—2018的方法,用Kjeltec 8400分析仪测定的总氮含量乘以系数6.25来估测粗蛋白质含量;按照GB/T 6432—2018的方法,用乙二胺四乙酸二钠络合滴定法测定钙含量;按照GB/T 6437—2018的方法,利用分光光度法测定磷含量。

1.3.2 血清生化指标

在试验第42天20:00仔猪空腹12 h,第43天08:00逐头称重后,待屠宰的36头仔猪前腔静脉采血10 mL置于无抗凝管中静置1 h,并置于离心机内1 360×g及4 ℃条件下离心15 min,分离得到血清,并使用无菌1.5 mL EP管分装,每管500 μL,-20 ℃保存备用,后续对血清生化指标进行测定。

1.3.3 胃、盲肠内容物pH

试验第43天,从每栏随机挑选1头仔猪进行屠宰,完整取下胃和盲肠,使用pH仪分别测定3次胃和盲肠内容物pH,在每次测定时用吸水纸擦净pH仪的探测针,然后记录数据。

1.3.4 肠道形态

试验第43天,从每栏随机挑选1头仔猪进行屠宰,环切约1 cm的十二指肠(近端)、空肠(中段)、回肠(远端),用注射器吸取磷酸盐缓冲液(PBS)轻轻冲洗后分别置于4%多聚甲醛样品瓶中固定保存,后续进行肠道形态学检测。经过冲水、脱水、透明、石蜡包埋等处理后,苏木精-伊红(HE)染色切片观察,每张切片测量9个完整的肠道绒毛及其伴生的隐窝,最后计算绒毛高度/隐窝深度(V/C)。

1.3.5 肠道、肾脏和肝脏的钙、磷吸收功能相关基因表达

用实时荧光定量PCR测定肠道、肾脏和肝脏中钙、磷吸收功能相关基因相对表达量,方法如下:利用Trizol Re-agent(Invitrogen, 美国)提取肠道、肾脏和肝脏中的总RNA,接着反转录合成cDNA。后续PCR扩增反应体系为10 μL: Premix Taq 5 μL,上、下游引物各0.5 μL, DNA模板2 μL,2 μL无酶水(DEPC水)。实时荧光定量PCR反应扩增程序为95 ℃ 30 s(预变性),95 ℃ 15 s(变性),60 ℃ 30 s(退火),72 ℃ 30 s(延伸),共39个循环。试验所需引物均由Primer Premier 5.0软件设计,由上海生工生物工程有限公司合成,引物序列信息见表3。内参基因为β-肌动蛋白(β-actin),目的基因相对表达量用2-△△Ct法计算。
表3 引物序列信息

Table 3 Primer sequence information

基因
Genes		 引物序列
Primer sequence (5'—3')		 产物长度
Product size/bp
维生素D受体
VDR		 F: AGGCTTCTTCAGACGGAGCATGAA
R: ACTCCTTCATGCCGATGTCCA		 140
钙离子通道5
TRPV5		 F: AGGGTCGGTTTCTCTCGCTA
R: GGCATAGGTGATGGTGATGACA		 75
钙离子通道6
TRPV6		 F: TCCAGACAGAGGACCCTAACAAG
R: GTGAGAAACAGCTCAAAGGTGCTA		 82
S100钙结合蛋白G
S100G		 F: CGCAACAGTCCCATTTAAGGA
R: TCAGCAGAGACATGGGTGGTT		 72
ATP酶质膜钙离子转运蛋白1
ATP2B1		 F: GGGCGGGCAGGTCATT
R: CCGCCGGGAGAAGATCA		 86
钙结合蛋白1
CALB1		 F: ACGCTGACGGAAGTGGTTAC
R: ATCCAGCCTTCTTTCGTGCC		 84
甲状旁腺素1受体
PTH1R		 F: GGCGTCCATTACATCGTCTT
R: AAAGTCCAGTGCCAATGTCC		 198
溶质载体家族蛋白34成员A1
SLC34A1		 F: TGGGCTTGTGTGACTGAGAG
R: CCCAGTCAGAGTTGTGCGTA		 198
溶质载体家族蛋白34成员A2
SLC34A2		 F: GAATCAGCCCGAAACAAGAG
R: AAACCATCCGTCCAACAGAG		 128
溶质载体家族蛋白34成员A3
SLC34A3		 F: TCGTCCTGGTCACAGTCC
R: CGGGGTTCTCATAGCAGTG		 192
溶质载体家族蛋白17成员A4
SLC17A4		 F: TTTTCAATTTCCACCCAACAAAT
R: GGGTGGGCAGAGCTGTGT		 76
细胞色素P450家族成员24A1
CYP24A1		 F: GCTGGACAACAAAATCAATGAG
R: CTCATAGAGCACAAGGCAGATG		 142
甲状腺受体α
THRA		 F: TGGAAAGCGAAAAAGAAAGAAC
R: TGAGTAGGTGGGATGGAGATTC		 202
β-肌动蛋白
β-action		 F: CACGCCATCCTGCGTCTGGA
R: AGCACCGTGTTGGCGTAGAG		 380

1.4 数据统计及分析

数据采用SPSS 25.0统计软件进行单因素方差分析(one-way ANOVA),结果用平均值和均值标准误(SEM)表示,GraphPad Prism软件作图,P<0.05表示差异显著。

2 结果与分析

2.1 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪血清生化指标的影响

由表4可知,4、5组的血清谷草转氨酶活性显著低于1、2和3组(P<0.05);随着饲粮中STTD Ca和STTD P的含量逐渐升高,血清碱性磷酸酶活性逐渐降低,但组间没有显著差异(P>0.05);1组的血清尿素含量显著高于2、3、4和5组(P<0.05)。
表4 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪血清生化指标的影响

Table 4 Effects of STTD Ca to STTD P contents in diets on serum biochemical indexes of 7 to 25 kg weaned piglets

项目
Items		 组别 Groups SEM P值
P-value
1 2 3 4 5 6
总蛋白 TP/(mmol/L) 55.55 53.37 57.63 57.00 51.29 47.50 2.77 0.057
谷丙转氨酶 ALT/(U/L) 65.82 65.43 67.07 46.68 65.39 56.63 6.19 0.491
肌酐 CRE/(mmol/L) 111.00 123.57 138.85 135.89 106.61 110.63 4.82 0.227
谷草转氨酶 AST/(U/L) 99.39a 82.96a 84.44a 53.84b 51.42b 70.55ab 4.56 0.006
碱性磷酸酶 ALP/(U/L) 249.42 241.36 232.29 228.84 217.61 210.45 6.79 0.608
白蛋白 ALB/(mmol/L) 24.35a 24.35a 25.35a 25.14a 24.55a 20.34b 0.14 0.005
尿素 UREA/(mmol/L) 3.94a 2.88b 2.73b 2.88b 2.38b 3.10ab 0.17 0.036

同行数据肩标不同字母表示差异显著(P<0.05)。下表同。

In the same row, values with different letter superscripts mean significant difference(P<0.05). The same as below.

由表5可知,1组血清中钙含量显著低于其他组(P<0.05),其第2、4、6组的血清钙含量较于其他组更高;1组血清磷含量显著高于5组(P<0.05);随着饲粮中STTD Ca和STTD P的含量逐渐升高,血清降钙素(CT)、甲状腺素(T4)、1,25二羟维生素 D3[1,25(OH)D3]、甲状旁腺素(PTH)含量逐渐升高,且6组显著高于1组(P<0.05)。
表5 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪血清钙、磷及调控激素含量的影响

Table 5 Effects of STTD Ca to STTD P contents in diets on serum calcium and phosphorus and regulatory hormone contents of 7 to 25 kg weaned piglets

项目
Items		 组别 Groups SEM P值
P-value
1 2 3 4 5 6
钙 Ca/(mmol/L) 0.65b 0.91a 0.83a 0.93a 0.91a 0.90a 0.02 0.001
磷 P/(mmol/L) 0.75a 0.63ab 0.68ab 0.66ab 0.49b 0.52ab 0.03 0.039
甲状腺素 T4/(mmol/L) 576.56d 974.70cd 1 379.15bc 1 729.88ab 1 723.56ab 2 396.60a 130.72 <0.001
甲状旁腺素 PTH/(mmol/L) 70.31c 90.85b 94.12b 109.02b 131.73a 150.99a 5.27 <0.001
降钙素 CT/(mmol/L) 81.87d 102.87cd 98.36d 124.04bc 144.35ab 158.92a 5.45 <0.001
1,25二羟维生素D3
1,25(OH)D3/(mmol/L)		 57.70c 65.18bc 81.13ab 75.45abc 85.15ab 93.70a 3.35 0.012
成纤维细胞生长因子-23
FGF-23/(mmol/L)		 420.22 485.37 469.08 418.80 451.37 491.74 12.16 0.356

2.2 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪胃、盲肠内容物pH的影响

由表6可知,随着饲粮中STTD Ca和STTD P的含量逐渐升高,各组间胃内容物pH和盲肠内容物pH均无显著差异(P>0.05),但是随着饲粮中STTD Ca和STTD P的含量逐渐升高,盲肠内容物pH呈现升高的趋势(P=0.085)。
表6 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪胃、盲肠内容物pH的影响

Table 6 Effects of STTD Ca to STTD P contents in diets on pH in stomach and cecal contents of 7 to 25 kg weaned piglets

项目
Items		 组别 Groups SEM P值
P-value
1 2 3 4 5 6
胃内容物 Stomach contents 2.76 3.93 2.48 3.18 3.23 3.36 0.23 0.598
盲肠内容物 Cecal contents 5.94 6.27 6.32 6.17 6.43 6.26 0.05 0.085

2.3 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道形态的影响

由表7可知,十二指肠隐窝深度1和2组显著低于5和6组(P<0.05),十二指肠绒毛高度/隐窝深度1和2组显著高于5和6组(P<0.05),空肠隐窝深度1和2组显著高于5和6组(P<0.05),空肠绒毛高度/隐窝深度2组显著低于5组(P<0.05)。十二指肠、空肠及回肠的HE染色切片如图2所示,所有试验组断奶仔猪小肠黏膜结构与形态完整。
表7 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道形态的影响

Table 7 Effects of STTD Ca to STTD P contents in diets on intestinal morphology of 7 to 25 kg weaned piglets

项目
Items		 组别 Groups SEM P值
P-value
1 2 3 4 5 6
十二指肠 Duodenum
绒毛高度 Villus height/μm 512.23a 472.03c 476.90bc 451.22cd 502.03ab 435.74d 3.82 <0.001
隐窝深度 Crypt depth/μm 250.27d 266.99cd 290.02bc 294.13b 350.52a 354.31a 3.76 <0.001
绒毛高度/隐窝深度
Villus height/crypt depth		 2.15a 1.88b 1.76b 1.76b 1.49c 1.30c 0.03 <0.001
空肠 Jejunum
绒毛高度 Villus height/μm 385.63a 363.73ab 373.93ab 366.24ab 346.06b 319.97c 3.83 <0.001
隐窝深度 Crypt depth/μm 324.44a 328.51a 295.15b 297.89b 263.21c 288.36bc 3.77 <0.001
绒毛高度/隐窝深度
Villus height/crypt depth		 1.37ab 1.15b 1.35ab 1.35ab 1.41a 1.16b 0.29 0.047
回肠 Ileum
绒毛高度 Villus height/μm 377.86bc 361.54cd 384.84bc 425.44a 389.74b 351.37d 3.54 <0.001
隐窝深度 Crypt depth/μm 282.96bc 288.92ab 291.24ab 310.67a 301.37ab 263.69c 2.89 <0.001
绒毛高度/隐窝深度
Villus height/crypt depth		 1.39 1.29 1.36 1.45 1.35 1.39 0.19 0.165
图1 肠道形态观察

Fig.1 Observation of intestinal morphology (5×)

图2 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道调控钙、磷旁吸收途径相关基因相对表达量的影响

TRPV6:钙离子通道6 transient receptor potential vanilloid 6;S100G:S100钙结合蛋白G S100 calcium binding protein G Gene;ATP2B1:ATP酶质膜钙离子转运蛋白1 ATPase plasma membrane Ca2+ transporting 1。
数据柱标注不同字母表示差异显著(P<0.05)。下图同。Different letters on the data column indicate significant differences (P<0.05). The same as below.

Fig.2 Effects of STTD Ca to STTD P contents in diets on relative expression levels of genes related to intestinal regulation of calcium-phosphorus absorption pathway of 7 to 25 kg weaned piglets

2.4 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道、肾脏和肝脏钙、磷吸收的影响

2.4.1 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道钙、磷吸收的影响

如图2、图3所示,随着饲粮中STTD Ca和STTD P的含量逐渐升高,十二指肠中,维生素D受体(VDR)、钙磷结合蛋白1(CALB1)、细胞色素P450家族成员24A1(CYP24A1)的基因相对表达量有逐渐降低的趋势,2和3组VDR的基因相对表达量显著高于1和5组(P<0.05),1和2组CALB1的基因相对表达量显著高于4、5和6组(P<0.05),1组CYP24A1的基因相对表达量显著高于其他组(P<0.05);在空肠中,钙离子通道6(TRPV6)、S100钙结合蛋白G(S100G)、ATP酶质膜钙离子转运蛋白1(ATP2B1)和溶质载体家族蛋白34成员A3(SLC34A3)的基因相对表达量呈现逐渐升高的趋势,4和6组的TRPV6基因相对表达量显著高于2和5组(P<0.05),6组S100G的基因相对表达量显著高于2组(P<0.05),4组ATP2B1的基因相对表达量显著高于2组(P<0.05),5和6组SLC34A3的基因相对表达量显著高于1和2组(P<0.05),CYP24A1的基因相对表达量有逐渐降低的趋势,1组CYP24A1的基因相对表达量显著高于其他组(P<0.05);在回肠中,CYP24A1和SLC34A3的基因相对表达量逐渐降低,1和2组CYP24A1的基因相对表达量显著高于其他组(P<0.05),2组SLC34A3的基因相对表达量显著高于其他组(P<0.05);在结肠中,TRPV6、S100G、ATP2B1、VDR、CALB1和CYP24A1的基因相对表达量逐渐降低,1组TRPV6、ATP2B1和CALB1的基因相对表达量显著高于其他组(P<0.05)。
图3 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道调控钙、磷激素及其他相关基因相对表达量的影响

VDR:维生素D受体 vitamin D receptor;CALB1:钙结合蛋白1 calbindin 1;CYP24A1:细胞色素P450家族成员24A1 cytochrome P450 24A1;SLC34A3:溶质载体家族蛋白34成员A3 solute carrier family 34 member 2。

Fig.3 Effect of STTD Ca and STTD P contents in diets on relative expression levels of genes related to calcium, phosphorus hormone and others in intestinal tract of 7 to 25 kg weaned piglets

2.4.2 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肾脏钙、磷重吸收的影响

如图4所示,随着饲粮中STTD Ca和STTD P的含量逐渐升高,肾脏CALB1、PTH1R、溶质载体家族蛋白17成员A4(SLC17A4)和TRPV5的基因相对表达量先升高后降低,3和4组PTH1R的基因相对表达量显著高于其他组(P<0.05),4组TRPV5的基因相对表达量显著高于其他组(P<0.05)。
图4 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肾脏钙、磷重吸收的影响

PTH1R:甲状旁腺素1受体 parathyroid hormone 1 receptor;CYP24A1:细胞色素P450家族成员24A1 cytochrome P450 24A1;SLC17A4:溶质载体家族蛋白17成员A4 Solute carrier family 17 member 4;CALB1:钙结合蛋白1 calbindin 1;SLC34A1:溶质载体家族蛋白34成员A1 solute carrier family 34 member 1;SLC34A2:溶质载体家族蛋白34成员A2 solute carrier family 34 member 2;SLC347A3:溶质载体家族蛋白347成员A3 solute carrier family 347 member 2;THRA:甲状腺受体α thyroid hormone receptor alpha;TRPV5:钙离子通道5 transient receptor potential vanilloid 5。

Fig.4 Effects of STTD Ca to STTD P contents in diets on calcium and phosphorus reabsorption in kidney of 7 to 25 kg weaned piglets

2.4.3 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肝脏钙、磷吸收的影响

如图5所示,随着饲粮中STTD Ca和STTD P的含量逐渐升高,肝脏中SLC34A3、CALB1和VDR的基因相对表达量呈现逐渐降低的趋势,2组SLC34A3的基因相对表达量显著高于4组(P<0.05),1组CALB1的基因相对表达量显著高于其他组(P<0.05)。
图5 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肝脏钙、磷吸收的影响

SLC34A1:溶质载体家族蛋白34成员A1 solute carrier family 34 member 1;SLC34A2:溶质载体家族蛋白34成员A2 solute carrier family 34 member 2; CALB1:钙结合蛋白1 calbindin 1;VDR:维生素D受体 vitamin D receptor。

Fig.5 Effects of STTD Ca to STTD P contents in diets on absorption of calcium and phosphorus in liver of 7 to 25 kg weaned piglets

3 讨论

3.1 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪血清生化指标的影响

在NRC(2012)推荐的饲粮STTD Ca与STTD P的比值为2.15[10],但这可能导致钙过量添加,而钙的过量添加会降低磷的吸收,还会造成资源的浪费。因此,为了达到提高生产利用率的目的,关于饲粮钙、磷的添加比例已经有了很多的推荐,所以本试验旨在研究饲粮中STTD Ca和STTD P比例为1.2时,对7~25 kg断奶仔猪体内钙、磷吸收的影响。本试验研究结果表明,随着饲粮中STTD Ca和STTD P的含量逐渐升高,血清碱性磷酸酶活性逐渐降低。碱性磷酸酶是一种多功能酶,它能水解多种磷酸酶单脂酶的底物,还在动物的骨骼矿化方面起着重要的作用。研究结果表明,在饲粮中含有较低的STTD Ca和STTD P时可以有效提高血清中碱性磷酸酶活性,提高血液钙、磷的吸收和骨骼矿化的能力[18]。有研究报道称,低水平钙、磷饲粮显著提高了血清碱性磷酸酶活性[19]。本试验研究结果与其研究报道结果基本一致。本试验中,断奶仔猪血清中CT、T4、1,25(OH)D3、PTH等激素的含量,随着饲粮中STTD Ca和STTD P的含量升高而逐渐升高,这一结果和以前的研究报道结果[20]基本一致。本试验中,2、4和6组血清钙含量较其他组更高,所以当饲粮钙、磷含量为这3个组的饲粮钙、磷含量时,能促进断奶仔猪对摄入钙的吸收。1组断奶仔猪血清中磷含量最高,说明当饲粮钙、磷含量低时,可能增强断奶仔猪对摄入磷的吸收。

3.2 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪胃、盲肠内容物pH的影响

有研究报道称,随着饲粮中无机钙源的增加,胃酸强度会降低,进而降低蛋白质、能量等营养物质的消化率[21]。本试验对胃和盲肠内容物的pH的研究结果表明,随着饲粮中STTD Ca和STTD P的含量逐渐升高,胃和盲肠内容物pH均无显著变化,但是随着饲粮中STTD Ca和STTD P的含量逐渐升高,胃内容物pH呈现升高的趋势。本研究结果与上述研究报道的饲粮中无机钙源的增加,胃酸强度会降低的结果一致。这说明在饲粮中含有较低的STTD Ca和STTD P时对胃和盲肠的pH影响较低。

3.3 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪肠道形态的影响

小肠表面具有手指状微尺度的绒毛结构,它能扩大肠道表面积以利于宿主有效吸收,同时,其与隐窝结构对于肠上皮的结构完整性和维持肠道稳态也至关重要[22-23]。同时,绒毛高度/隐窝深度能够反馈肠道的健康状态与吸收能力[24-25]。研究结果表明,饲粮中STTD Ca和STTD P的含量对回肠的绒毛高度/隐窝深度没有显著影响,而十二指肠绒毛高度/隐窝深度随着饲粮中STTD Ca和STTD P的含量升高而逐渐降低。研究结果表明,随着饲粮中STTD Ca和STTD P的含量逐渐升高,十二指肠的吸收能力逐渐降低。

3.4 饲粮STTD Ca和STTD P含量对7~25 kg断奶仔猪脏器和肠道钙、磷吸收的影响

在肠道中,RPV6[26-27]、S100G[28]、ATP2B1[29]等基因调控钙磷旁吸收、跨细胞吸收,CYP24A1虽然不直接参与肠道的主要功能,但与维生素D的代谢有关[30],维生素D在体内含量变化可能间接影响肠道吸收[31],CALB1可能参与调节钙的吸收和细胞内信号[32],VDR可直接调控钙、磷的吸收[33],SLC34A3编码的转运蛋白参与了磷的吸收[34]。在肝脏和肾脏中,CYP24A1基因编码的酶同样参与维生素D的降解,将活性形式的维生素D转化为不活跃形式,是维持体内维生素D平衡的关键,CALB1可能参与钙的运输和信号传导,VDR参与调节钙的代谢和维生素D的活化,SLC34A3则负责将磷酸盐从尿液中重吸收回血液。有研究报道称,TRPV5、TRPV6和CALB1在十二指肠表达量更高,覆盖了上皮钙转运中3步过程的所有组成部分[35-36]。本试验研究结果表明,从十二指肠、空肠、回肠、结肠4个肠段中TRPV6、S100G、ATP2B1的相对表达量是呈现递减趋势的,这与前人研究报道结果相符。随着饲粮中STTD Ca和STTD P的含量逐渐升高,在十二指肠中,与旁吸收有关的基因CALB1和激素调控的相关基因VDR、CYP24A1的相对表达量呈现逐渐降低的趋势,研究结果表明饲粮中过高的钙、磷可能会抑制钙、磷在十二指肠中的吸收;在空肠中,与旁吸收有关的基因TRPV6、S100G、CALB1,激素调控的相关基因VDR和与磷吸收相关的基因SLC34A3的相对表达量呈现逐渐升高的趋势,表明随着饲粮钙、磷含量的升高能一定程度提高空肠对钙、磷的吸收;在结肠中,与旁吸收有关的基因TRPV6、S100G、ATP2B1、CALB1和激素调控的基因VDR、CYP24A1的基因相对表达量均呈现逐渐减少的趋势,说明饲粮中过高的钙、磷可能会抑制结肠对钙、磷的吸收。综上所述,当饲粮中STTD Ca和STTD P的比值为1.2时,随着饲粮中STTD Ca和STTD P的含量逐渐升高,十二指肠和结肠对钙、磷的吸收能力逐渐减弱,空肠对钙、磷的吸收能力逐渐增强。
在肾脏中,随着饲粮中STTD Ca和STTD P的含量逐渐升高,CALB1、PTH1R、SLC17A4和TRPV5的基因相对表达量均呈现先升高后降低的趋势,而峰值主要在NRC(2012)推荐的STTD P添加量。研究结果表明,当饲粮中STTD Ca和STTD P的比值为1.2时,随着饲粮中STTD Ca和STTD P的含量逐渐升高,当增加到NRC(2012)推荐的STTD P添加量(饲粮STTD P含量为0.34%~0.42%)时,肾脏对钙、磷重吸收的能力最强。在肝脏中,SLC34A3、CALB1和VDR的基因相对表达量呈现逐渐降低的趋势。研究结果表明,当饲粮中STTD Ca和STTD P的比值为1.2时,随着饲粮中STTD Ca和STTD P的含量逐渐升高,肝脏对钙、磷吸收能力逐渐降低。

4 结论

当饲粮中STTD Ca和STTD P的比值为1.2时,随着饲粮中STTD Ca和STTD P的含量逐渐升高,十二指肠和结肠对钙、磷的吸收能力逐渐减弱,空肠对钙、磷的吸收能力逐渐增强,肝脏对钙、磷吸收能力逐渐降低。当饲粮中STTD Ca和STTD P的含量增加到NRC(2012)推荐的STTD P添加量(饲粮STTD P含量为0.34%~0.42%)时,肾脏对钙、磷重吸收的能力最强。结合血清钙、磷含量数据,当饲粮中STTD Ca和STTD P的含量分别在0.216%~0.312%和0.18%~0.26%时,断奶仔猪能有效利用饲粮钙、磷。

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