Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 9: 6645 - 6655

DOI: https://doi.org/10.12418/CJAN2026.532

RESEARCH PAPER

Effects of Carboxymethyl Pachymaran on Growth Performance, Serum Indices and Intestinal Barrier Function of Broilers Challenged with Lipopolysaccharide

  • HUANG Minghai , 1 ,
  • CHEN Bo 1 ,
  • GAO Luyao 1 ,
  • XIONG Fuxuan 1 ,
  • WU Sichao 1 ,
  • XIONG Xiaojun 2 ,
  • PANG Yilin 2 ,
  • DENG Wei 3 ,
  • LYU Zonghao 1, 3 ,
  • HE Jianhua , 1, *
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  • 1 College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China
  • 2 Hunan Xiangnong Animal Pharmaceutical Co., Ltd., Changsha 410299, China
  • 3 Huaihua Academy of Agricultural Sciences, Huaihua 418000, China
* professor, E-mail:

Received date: 2026-01-17

  Online published: 2026-09-12

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Abstract

This experiment was conducted to investigate the effects of carboxymethyl pachymaran (CMP) on growth performance, serum indices and intestinal barrier function of broilers challenged with lipopolysaccharide (LPS). A 2×2 two-factor experimental design was adopted, with the two factors being LPS challenge (with or without) and CMP supplementation (with or without). A total of 240 one-day-old male Arbor Acres (AA) broilers were randomly assigned to four groups: CON group, CMP group, LPS group and CMPL group, with 6 replicates per group and 10 broilers per replicate. Broilers in CON and LPS groups were fed a basal diet, while those in CMP and CMPL groups received the basal diet supplemented with 200 mg/kg CMP. The trial lasted for 20 days, divided into the pre-challenge period (1 to 15 days of age) and challenge period (16 to 20 days of age). On 16, 18 and 20 days of age, broilers in LPS and CMPL groups were intraperitoneally injected with LPS at a dosage of 500 μg/kg BW, whereas broilers in CON and CMP groups received an equal volume of 0.86% sterile normal saline via intraperitoneal injection. The results showed as follows: 1) LPS challenge significantly decreased the average daily gain (ADG), average daily feed intake and final body weight of broilers during 16 to 20 days of age (P<0.05); dietary CMP supplementation significantly increased the ADG during 16 to 20 days of age (P<0.05). 2) LPS challenge significantly increased the spleen index of broilers (P<0.05). 3) LPS challenge significantly increased the serum alanine transaminase (ALT) activity and creatinine (CREA) content of broilers (P<0.05); dietary CMP supplementation significantly decreased the serum ALT activity (P<0.05). 4) LPS challenge significantly decreased the serum superoxide dismutase (SOD) and glutathione peroxidase (GPx) activities (P<0.05), and significantly increased the serum malondialdehyde (MDA) content of broilers (P<0.05); dietary CMP supplementation significantly increased the serum SOD and GPx activities (P<0.05), and significantly decreased the serum MDA content (P<0.05). 5) LPS challenge significantly increased the serum interleukin-1β (IL-1β) and tumor necrosis factor-α (TNF-α) contents, and significantly decreased the serum interleukin-10 (IL-10) content of broilers (P<0.05); dietary CMP supplementation significantly decreased the serum TNF-α content (P<0.05). There was a significant interactive effect between LPS challenge and dietary CMP supplementation on the serum TNF-α content of broilers (P<0.05). 6) LPS challenge significantly increased the serum D-lactic acid (D-LA) and endotoxin (ET) contents of broilers (P<0.05); dietary CMP supplementation significantly decreased the serum D-LA and ET contents (P<0.05). 7) LPS challenge significantly decreased the villus height and villus height/crypt depth ratio (V/C) in duodenum and jejunum (P<0.05), and significantly increased the crypt depth in duodenum of broilers (P<0.05); dietary CMP supplementation significantly increased the V/C in jejunum (P<0.05). In conclusion, dietary supplementation with 200 mg/kg CMP can enhance antioxidant and anti-inflammatory capacities, reduce intestinal permeability, optimize morphological structure of small intestinal mucosa and maintain the integrity of intestinal physical barrier, thereby alleviating growth retardation and adverse impacts induced by LPS challenge in broilers to a certain extent.

Key words： carboxymethyl pachymaran; lipopolysaccharide; broilers; antioxidant; immunity; intestinal barrier function

Cite this article

HUANG Minghai , CHEN Bo , GAO Luyao , XIONG Fuxuan , WU Sichao , XIONG Xiaojun , PANG Yilin , DENG Wei , LYU Zonghao , HE Jianhua . Effects of Carboxymethyl Pachymaran on Growth Performance, Serum Indices and Intestinal Barrier Function of Broilers Challenged with Lipopolysaccharide[J]. Chinese Journal of Animal Nutrition, 2026 , 38(9) : 6645 -6655 . DOI: 10.12418/CJAN2026.532

过去几十年中,鸡肉凭借营养优良、性价比高的优势广受消费市场欢迎,推动了肉鸡养殖产业的快速发展。然而,在现代集约化养殖模式中,肉鸡易受到高饲养密度、高温及外源致病菌等多重应激刺激,诱发机体发生氧化应激和炎症反应,导致生长性能下降[1-2]。因此,如何缓解不利条件下肉鸡生长性能的下降,成为养鸡业重点关注的问题。众多研究表明,从传统中药材茯苓的菌核中提取出来的茯苓多糖具有抗氧化[3]、抗炎[4]、抗肿瘤[5]等生物学功能。肉鸡饲养试验表明,饲粮中添加0.5~1.0 g/kg[6]或0.1%[7]茯苓多糖能够促进免疫器官发育,提升机体免疫功能,进而促进生长。多糖的生物活性直接受其化学结构的影响,如结构单元、糖苷键、相对分子质量和构象等。而通过羧甲基化[8]、乙酰化[9]、硫酸化[10]和磷酸化[11]等手段对多糖的结构进行适度修饰,可以增强多糖衍生物的抗氧化及抗肿瘤活性。研究发现,对茯苓多糖进行羧甲基修饰制成羧甲基茯苓多糖(carboxymethyl pachymaran,CMP),可使其溶解度与生物活性增加[12-13]。陈博等[14]研究发现,饲粮中添加200 mg/kg CMP可以提高肉仔鸡的生长性能。然而,目前关于CMP对免疫应激状态下肉鸡影响的研究较少。因此,本研究通过腹腔注射脂多糖(LPS)构建肉鸡免疫应激模型,探讨饲粮中添加CMP对LPS应激肉仔鸡生长性能、血清指标及肠道屏障功能的影响,以期为评估CMP在肉鸡健康养殖中的潜在应用价值提供参考。

1 材料与方法

1.1 试验材料

试验所用CMP及LPS(血清型O55∶B5)均为市购产品,CMP产品中CMP的含量不低于60%,取代度不低于0.55[15]。

1.2 试验设计

本试验通过湖南农业大学生物医学研究伦理委员会审查批准,批准号:伦审科2022第(051)号。采用2×2双因素试验设计,2个因素分别为LPS应激与否和CMP添加与否。选取1日龄爱拔益加(AA)肉仔鸡公雏240只,随机分为4组(CON组、CMP组、LPS组和CMPL组),每组6个重复,每个重复10只鸡。其中,CON组和LPS组饲喂基础饲粮,CMP组和CMPL组饲喂基础饲粮+200 mg/kg CMP。CMP添加剂量参照本课题组前期梯度试验结果[14]确定。试验期20 d,分为应激前期(1~15日龄)与应激期(16~20日龄)。于16、18、20日龄时,对LPS组和CMPL组肉仔鸡腹腔注射500 μg/kg BW LPS,对CON组和CMP组肉仔鸡腹腔注射等体积0.86%无菌生理盐水[16-17]。基础饲粮参照NRC(1994)[18]和《鸡饲养标准》(NY/T 33—2004)[19]中白羽肉鸡营养需要配制,其组成及营养水平参见本课题组之前研究[14]。

1.3 饲养管理

进雏前对鸡舍进行全面清扫与消毒处理,并空置通风48 h。试验采用3层笼养模式,育雏第1周舍温控制在33~34 ℃,此后每周下调2~3 ℃,直至26 ℃。舍内采用自然通风,每日根据环境温度调节通风量。采用人工连续照明系统,全天恒定光照。试验期间,肉仔鸡自由采食和饮水,并按常规免疫程序进行疫苗接种。

1.4 样品采集

于21日龄时,每组随机选取6只鸡(每重复1只,禁食12 h),翅静脉采集血液,静置30 min后,于4 ℃、4 000×g条件下离心15 min,分离血清并置于-80 ℃保存。采血结束后,将试验鸡颈部放血处死,解剖并完整分离胸腺、肝脏、脾脏及法氏囊,剔除附着脂肪与多余组织后进行称重,计算免疫器官指数[免疫器官指数(%)=100×免疫器官鲜重(g)/宰前空腹活重(g)]。同时,分别取十二指肠、空肠、回肠中段约2 cm肠段,置于4%多聚甲醛中固定保存。

1.5 测定指标与方法

1.5.1 生长性能

试验期间每日记录喂料量与剩料量,统计各重复采食量;分别于肉仔鸡1、15、20日龄时逐只空腹称重;并据此计算应激前期与应激期的平均日增重(ADG)、平均日采食量(ADFI)及料重比(F/G)。

1.5.2 血清指标

生化指标:采用卓越450型全自动生化分析仪(上海科华实验系统有限公司)测定血清总蛋白(TP)、白蛋白(ALB)、球蛋白(GLB)、葡萄糖(GLU)、总胆红素(T-Bil)、甘油三酯(TG)、肌酐(CREA)含量及谷丙转氨酶(ALT)、谷草转氨酶(AST)活性,并计算白球比(ALB/GLB)。
抗氧化指标:采用北京盒子生工科技有限公司生产的试剂盒测定血清总抗氧化能力(T-AOC)及超氧化物歧化酶(SOD)、谷胱甘肽过氧化物酶(GPx)、过氧化氢酶(CAT)活性和丙二醛(MDA)含量。
免疫指标:采用湖南艾方生物科技有限公司生产的酶联免疫吸附测定(ELISA)试剂盒测定血清白细胞介素-1β(IL-1β)、白细胞介素-2(IL-2)、白细胞介素-10(IL-10)和肿瘤坏死因子-α(TNF-α)含量。
肠道通透性指标:采用湖南艾方生物科技有限公司生产的ELISA试剂盒测定血清D-乳酸(D-LA)和内毒素(ET)含量。

1.5.3 肠道黏膜形态

取4%多聚甲醛固定后的肠道组织,经石蜡包埋、切片、苏木精-伊红(HE)染色处理,于光学显微镜(AF-610-5-6,湖南艾方生物科技有限公司)下拍照观测,测量绒毛高度(VH)和隐窝深度(CD),并计算绒隐比(V/C)。

1.6 数据统计与分析

所有数据均使用SPSS 26.0软件处理,通过一般线性模型进行双因素方差分析,分析LPS应激和添加CMP的主效应及其交互作用,并对各组测定指标进行单因素方差分析(one-way ANOVA),差异显著者采用Duncan氏法进行事后多重比较。结果以平均值和均值标准误(SEM)表示,P<0.05表示差异显著,0.05≤P<0.10表示差异有显著趋势。

2 结果与分析

2.1 CMP对LPS应激肉仔鸡生长性能的影响

由表1可知,在应激前期(1~15日龄),饲粮中添加CMP对肉仔鸡生长性能未产生显著影响(P>0.05)。在应激期(16~20日龄),LPS应激显著降低了肉仔鸡末重、ADG及ADFI(P<0.05);饲粮中添加CMP显著提高了ADG(P<0.05),且有增加末重的趋势(P=0.075)。LPS应激与饲粮中添加CMP对肉仔鸡各项生长性能指标无显著交互作用(P>0.05)。多重比较结果显示,在应激期,与LPS组相比,CMPL组肉仔鸡ADG显著提高(P<0.05)。
表1 CMP对LPS应激肉仔鸡生长性能的影响

Table 1 Effects of CMP on growth performance of broilers challenged with LPS

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
1~15日龄1 to 15 days of age
初重IBW/g 43.14 43.81 43.61 43.45 0.202 0.896 0.553 0.333
末重FBW/g 483.16 481.92 474.85 475.01 2.691 0.183 0.923 0.900
平均日增重ADG/(g/d) 29.33 29.20 28.74 28.77 0.182 0.185 0.889 0.843
平均日采食量ADFI/(g/d) 34.43 35.37 34.64 34.35 0.187 0.273 0.374 0.102
料重比F/G 1.17 1.21 1.20 1.19 0.007 0.633 0.359 0.105
16~20日龄16 to 20 days of age
末重FBW/g 828.80a 836.61a 784.61b 804.55b 5.488 0.001 0.075 0.421
平均日增重ADG/(g/d) 69.12ab 70.93a 61.95c 65.90b 0.889 0.001 0.020 0.359
平均日采食量ADFI/(g/d) 100.68 100.09 93.03 96.58 1.221 0.021 0.515 0.366
料重比F/G 1.45 1.41 1.50 1.46 0.020 0.219 0.325 0.902

LPS:LPS应激效应;CMP:添加CMP效应;LPS×CMP:LPS应激与添加CMP的交互作用。同行数据肩标无字母或相同小写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)。下表同。

LPS: effect of LPS challenge; CMP: effect of supplementation with CMP; LPS×CMP: interaction between LPS challenge and supplementation with CMP. In the same row, values with no letter or the same lowercase letter superscripts mean no significant difference (P>0.05), while with different lowercase letter superscripts mean significant difference (P<0.05). The same as below.

2.2 CMP对LPS应激肉仔鸡免疫器官指数的影响

由表2可知,LPS应激显著提高了肉仔鸡脾脏指数(P<0.05);饲粮中添加CMP及LPS应激与饲粮中添加CMP的交互作用对肉仔鸡各免疫器官指数均无显著影响(P>0.05)。
表2 CMP对LPS应激肉仔鸡免疫器官指数的影响

Table 2 Effects of CMP on immune organ indexes of broilers challenged with LPS

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
肝脏指数Liver index 2.32 2.30 2.53 2.35 0.049 0.185 0.321 0.443
脾脏指数Spleen index 0.08b 0.06b 0.14a 0.12a 0.009 0.001 0.148 0.939
胸腺指数Thymus index 0.20 0.26 0.20 0.27 0.018 0.913 0.102 0.977
法氏囊指数Bursa of Fabricius index 0.23 0.23 0.26 0.23 0.009 0.269 0.464 0.422

2.3 CMP对LPS应激肉仔鸡血清生化指标的影响

由表3可知,LPS应激显著提高了肉仔鸡血清ALT活性和CREA含量(P<0.05),有提高血清T-Bil含量的趋势(P=0.059);饲粮中添加CMP显著降低了血清ALT活性(P<0.05)。LPS应激与饲粮中添加CMP对肉仔鸡各项血清生化指标无显著交互作用(P>0.05)。
表3 CMP对LPS应激肉仔鸡血清生化指标的影响

Table 3 Effects of CMP on serum biochemical indices of broilers challenged with LPS

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
谷丙转氨酶ALT/(U/L) 7.73b 6.88b 9.45a 8.10ab 0.289 0.005 0.027 0.594
谷草转氨酶AST/(U/L) 251.38 245.56 244.38 239.16 3.758 0.402 0.489 0.970
总蛋白TP/(g/L) 21.72 19.90 22.22 21.60 0.385 0.147 0.111 0.421
白蛋白ALB/(g/L) 8.11 8.16 8.08 7.90 0.108 0.519 0.774 0.615
球蛋白GLB/(g/L) 13.77 13.22 13.85 13.70 0.153 0.372 0.273 0.529
白球比ALB/GLB 0.58 0.61 0.58 0.57 0.008 0.160 0.523 0.270
总胆红素T-Bil/(μmol/L) 9.29 8.86 10.15 10.31 0.297 0.059 0.821 0.612
肌酐CREA/(μmol/L) 23.45ab 19.28b 30.30a 31.06a 1.692 0.004 0.564 0.404
葡萄糖GLU/(mmol/L) 11.58 11.61 11.14 11.26 0.184 0.317 0.847 0.915
甘油三酯TG/(mmol/L) 0.28 0.26 0.30 0.27 0.011 0.544 0.338 0.642

2.4 CMP对LPS应激肉仔鸡血清抗氧化指标的影响

由表4可知,LPS应激显著降低了肉仔鸡血清SOD和GPx活性(P<0.05),显著提高了血清MDA含量(P<0.05);饲粮中添加CMP显著提高了血清SOD和GPx活性(P<0.05),显著降低了血清MDA含量(P<0.05)。LPS应激与饲粮中添加CMP对肉仔鸡各项血清抗氧化指标无显著交互作用(P>0.05)。多重比较结果显示,与LPS组相比,CMPL组肉仔鸡血清SOD活性显著提高(P<0.05),血清MDA含量显著降低(P<0.05)。
表4 CMP对LPS应激肉仔鸡血清抗氧化指标的影响

Table 4 Effects of CMP on serum antioxidant indices of broilers challenged with LPS

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
过氧化氢酶CAT/(U/mL) 46.45 58.29 43.34 49.23 3.673 0.425 0.249 0.695
总抗氧化能力T-AOC/(μmol/mL) 2.06 2.29 1.95 2.05 0.057 0.133 0.140 0.554
谷胱甘肽过氧化物酶GPx/(U/mL) 2 490.36ab 2 890.86a 2 132.59b 2 410.47ab 92.006 0.014 0.041 0.698
超氧化物歧化酶SOD/(U/mL) 54.83ab 60.29a 44.42c 51.41b 1.527 0.001 0.006 0.710
丙二醛MDA/(nmol/mL) 1.44b 1.17b 2.31a 1.56b 0.118 0.001 0.006 0.173

2.5 CMP对LPS应激肉仔鸡血清免疫指标的影响

由表5可知,LPS应激显著提高了肉仔鸡血清IL-1β和TNF-α含量(P<0.05),显著降低了血清IL-10含量(P<0.05),有提高血清IL-2含量的趋势(P=0.053);饲粮中添加CMP显著降低了血清TNF-α含量(P<0.05)。LPS应激与饲粮中添加CMP对肉仔鸡血清TNF-α含量具有显著交互作用(P<0.05)。多重比较结果显示,与LPS组相比,CMPL组肉仔鸡血清TNF-α含量显著降低(P<0.05)。
表5 CMP对LPS应激肉仔鸡血清免疫指标的影响

Table 5 Effects of CMP on serum immune indices of broilers challenged with LPS pg/mL

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
白细胞介素-1β IL-1β 51.10b 51.24b 61.54a 58.08a 1.189 0.001 0.302 0.265
白细胞介素-2 IL-2 12.05 12.59 13.37 13.02 0.223 0.053 0.822 0.305
肿瘤坏死因子-α TNF-α 79.33bc 77.20c 97.58a 85.47b 1.957 0.001 0.005 0.038
白细胞介素-10 IL-10 83.59a 85.67a 75.59b 77.98ab 1.474 0.006 0.398 0.953

2.6 CMP对LPS应激肉仔鸡血清肠道通透性指标的影响

由表6可知,LPS应激显著提高了肉仔鸡血清D-LA和ET含量(P<0.05);饲粮中添加CMP显著降低了血清D-LA和ET含量(P<0.05)。LPS应激与饲粮中添加CMP对肉仔鸡血清D-LA和ET含量无显著交互作用(P>0.05)。
表6 CMP对LPS应激肉仔鸡血清肠道通透性指标的影响

Table 6 Effects of CMP on serum intestinal permeability indices of broilers challenged with LPS

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
D-乳酸D-LA/(μg/mL) 67.97b 63.95b 74.49a 69.73ab 1.235 0.007 0.043 0.859
内毒素ET/(EU/mL) 12.44bc 11.72c 14.08a 13.29ab 0.240 0.001 0.034 0.913

2.7 CMP对LPS应激肉仔鸡肠道黏膜形态的影响

由表7可知,LPS应激显著降低了肉仔鸡十二指肠和空肠VH、V/C(P<0.05),显著提高了十二指肠CD(P<0.05),有降低回肠V/C的趋势(P=0.069);饲粮中添加CMP显著提高了空肠V/C(P<0.05),有提高十二指肠V/C(P=0.099)及空肠VH(P=0.086)的趋势。LPS应激与饲粮中添加CMP对肉仔鸡肠道黏膜形态指标无显著交互作用(P>0.05)。多重比较结果显示,与LPS组相比,CMPL组肉仔鸡空肠V/C显著提高(P<0.05)。
表7 CMP对LPS应激肉仔鸡肠道黏膜形态的影响

Table 7 Effects of CMP on intestinal mucosa morphology of broilers challenged with LPS

项目
Items		 组别Groups SEM P值P-value
CON CMP LPS CMPL LPS CMP LPS×CMP
十二指肠Duodenum
绒毛高度VH/μm 1 247.73a 1 277.51a 1 114.56b 1 141.40b 21.353 0.001 0.414 0.966
隐窝深度CD/μm 153.65ab 149.56b 161.90a 156.54ab 1.583 0.011 0.100 0.820
绒隐比V/C 8.12a 8.54a 6.89b 7.30b 0.175 0.001 0.099 0.986
空肠Jejunum
绒毛高度VH/μm 1 048.35ab 1 081.82a 880.44c 969.56bc 22.656 0.001 0.086 0.422
隐窝深度CD/μm 136.55 131.66 138.54 133.72 2.450 0.696 0.354 0.994
绒隐比V/C 7.74ab 8.26a 6.37c 7.26b 0.202 0.001 0.030 0.542
回肠Ileum
绒毛高度VH/μm 725.79 734.32 701.61 706.90 15.962 0.453 0.840 0.962
隐窝深度CD/μm 110.69 109.44 122.79 112.83 2.450 0.115 0.247 0.364
绒隐比V/C 6.60 6.80 5.70 6.30 0.191 0.069 0.290 0.589

3 讨论

3.1 CMP对LPS应激肉仔鸡生长性能的影响

大量研究表明,LPS应激会降低动物采食量,抑制机体对营养物质的消化与吸收,特别是通过改变营养物质在机体内的再分配,参与炎症因子释放和多种急性蛋白合成等免疫过程的发生,进而抑制动物生长[20-22]。研究发现,在肉仔鸡生长前期腹腔注射500 μg/kg BW LPS,能够显著降低其ADG、ADFI和末重[23]。本研究观察到,LPS应激显著降低了肉仔鸡应激期的ADFI、ADG和末重,与前人研究结果相符。研究表明,在5%葡聚糖硫酸钠(DSS)诱导的小鼠免疫应激模型中,灌胃300 mg/kg取代度为0.63或0.48的CMP,均能显著缓解小鼠的体重降低,减轻应激造成的生长抑制[24]。陈博等[14]研究表明,饲粮中添加200 mg/kg CMP可显著增加肉仔鸡ADG和末重。本研究结果表明,在应激前期,饲粮中添加200 mg/kg CMP并未使肉仔鸡生长性能指标发生显著改变;而在应激期,饲粮中添加200 mg/kg CMP显著提高了肉仔鸡ADG,这说明CMP可缓解LPS应激诱发的生长抑制,改善肉仔鸡生长表现。有研究证实,在肉仔鸡的饲粮中添加植物源多糖可通过提升机体抗氧化酶活性、降低血清炎症因子含量及保护肠道屏障完整,进而缓解应激对肉仔鸡生长性能的抑制[25-27]。因此本研究进一步探讨CMP对LPS应激肉仔鸡抗炎、抗氧化能力以及肠道屏障的影响。

3.2 CMP对LPS应激肉仔鸡血清生化指标的影响

血清生化指标可在不同程度上反映动物机体的生理状态、器官功能以及对营养物质的代谢状况。当肝脏发生炎症或肝细胞受损时,正常储存在肝细胞中的肝脏代谢酶ALT和AST会因细胞膜通透性增大而释放到血液中[28]。肝脏损伤还会干扰其对T-Bil的吸收与代谢过程,造成血清中T-Bil蓄积[29]。血清中CREA含量可作为评估机体肾脏功能的指标。有研究表明,LPS应激使小鼠血清ALT、AST活性与CREA含量显著提高,诱发肝肾损伤[30]。本研究血清生化指标结果表明,LPS应激显著提高了肉仔鸡血清ALT活性,并且有提高血清T-Bil含量的趋势,与前人研究结果相符。王灿红等[31]在氟尿嘧啶(5-Fu)诱导的小鼠肝脏损伤模型中发现,灌胃200 mg/kg CMP能够显著逆转5-Fu引发的血清ALT、AST活性升高,减轻5-Fu对小鼠肝脏的毒副作用。本研究发现,饲粮中添加CMP显著降低了血清ALT活性,表明CMP能够在一定程度上减轻LPS应激造成的肉鸡肝脏损伤。

3.3 CMP对LPS应激肉仔鸡血清抗氧化指标的影响

氧化应激被认为是导致病理性疾病的重要因素之一,其主要由过量活性氧自由基引发内在氧化还原系统紊乱所致[32]。LPS应激可诱导过量活性氧自由基生成并诱导小胶质细胞活化,降低机体抗氧化酶活性,扰乱脂质代谢平衡,同时产生大量MDA,进而引发细胞和组织氧化损伤[33]。MDA是评价脂质过氧化程度的生物标志物[34]。本研究结果表明,LPS应激会使肉仔鸡血清MDA含量显著升高,提示LPS应激导致了机体氧化与抗氧化系统的失衡,这与前人研究结论一致。研究表明,在草酸盐诱导的人近曲肾小管上皮细胞(HK-2细胞)氧化应激损伤模型中,CMP能够显著提升HK-2细胞内SOD活性,同时有效降低活性氧自由基水平、MDA含量以及8-羟基脱氧鸟苷(8-OHdG)的生成,从而改善细胞活力,减轻氧化损伤[35]。Tan等[24]研究表明,在DSS诱导的小鼠结肠炎模型中,通过灌胃300 mg/kg CMP能够增强小鼠血清SOD活性,降低氧化应激水平,从而有效缓解小鼠结肠炎症状。在白羽肉鸡上的研究也显示,饲粮中添加200 mg/kg CMP可显著提升肉仔鸡血清中抗氧化酶活性,同时降低MDA含量,有效增强机体抗氧化功能[14]。本研究结果与之相符,饲粮中添加200 mg/kg CMP既可显著提高肉仔鸡血清SOD和GPx活性,也能显著降低血清MDA含量,表明CMP具有缓解氧化应激的作用,原因可能是:1)CMP具有较强的氢原子或单电子供应能力[36];2)具有较强的过渡金属离子螯合能力[35];3)能够调控机体酶促抗氧化系统。

3.4 CMP对LPS应激肉仔鸡免疫功能的影响

家禽免疫系统在维持机体内环境稳定及抵御外界应激源刺激方面具有重要作用。免疫器官作为免疫系统的重要组成部分,其发育状况与机体免疫功能密切相关,通常以免疫器官指数加以评估。脾脏既是次级淋巴器官,也是抗原刺激后产生抗体的起始部位[37-38]。Yang等[39]研究表明,在肉仔鸡14~20日龄及28~34日龄,通过腹腔注射500 μg/kg BW LPS,能够显著增加21和35日龄肉鸡脾脏指数。本研究中发现,LPS攻毒后肉仔鸡的脾脏指数显著增加,与前人研究结果相似。LPS应激导致肉仔鸡脾脏肥大,可能是由于其诱导的免疫反应促进了促炎细胞因子产生,并引发炎性细胞向脾脏聚集,进而导致脾脏代偿性增生[40]。
促炎细胞因子的诱导和分泌对于激活宿主先天防御系统、调节适应性免疫反应至关重要。但是LPS应激会导致机体过度分泌促炎细胞因子,进而导致机体稳态失衡,造成广泛的组织损伤与器官功能障碍,严重威胁机体健康[41]。体外研究表明,通过LPS诱导构建巨噬细胞RAW264.7炎症模型,不同分子质量(5.93×104、11.58×104、26.02×104)和羧甲基取代度(0.13、0.32、0.95)的CMP都能够显著降低巨噬细胞中促炎因子TNF-α的释放[42]。体内研究发现,摄入300 mg/kg CMP能够显著降低结肠炎模型小鼠血清促炎因子IL-1β、TNF-α和白细胞介素-6(IL-6)含量,显著提高血清抗炎因子IL-4含量,缓解小鼠肠道损伤[24]。本研究结果与之类似,LPS应激显著提高了肉仔鸡血清IL-1β和TNF-α含量,显著降低了血清IL-10含量,而饲粮中CMP的添加能够降低肉仔鸡血清TNF-α含量,缓解了由LPS应激引发的肉仔鸡血清炎症因子分泌失衡。王峰等[43]研究发现,CMP可以通过抑制Toll样受体4(TLR4)、髓样分化因子88(MyD88)和核转录因子-κB p65(NF-κB p65)蛋白的表达,介导大鼠核转录因子-κB(NF-κB)信号通路的级联反应,从而有效缓解溃疡性结肠炎引起的炎症反应。综上所述,CMP可能通过抑制TLR4/NF-κB炎症信号通路的激活,介导NF-κB的表达,从而调控促炎细胞因子的诱导与分泌,进而缓解LPS应激造成的肉仔鸡炎症损伤。

3.5 CMP对LPS应激肉仔鸡肠道屏障功能的影响

D-LA在动物体内主要由肠道微生物发酵产生,ET是革兰氏阴性菌细胞壁裂解时释放的产物,具有较强毒性[44-45]。当肠道黏膜因缺血、缺氧或炎症受损时,肠道通透性增加,肠道黏膜完整性被破坏,D-LA和ET便会透过肠黏膜进入血液循环,直接损伤细胞,加剧全身炎症反应[46]。因此,血清中D-LA与ET含量不仅是衡量肠道通透性的核心指标,也是评估肠道物理屏障完整性的关键参数[47]。本研究结果表明,LPS攻毒后肉仔鸡血清D-LA和ET含量显著提高,说明LPS应激会造成肉仔鸡肠道通透性异常增加,为外源性有害物质透过肠黏膜进入循环系统提供了潜在通路;而饲粮中添加CMP能够显著降低血清D-LA与ET含量,表明CMP具有降低肠道通透性、保护肠道屏障功能的作用。
研究表明,LPS应激会导致肉仔鸡肠黏膜脱落、肠道上皮细胞坏死、肠道VH、V/C降低以及CD升高,从而破坏肠道形态结构,降低生长性能[48-49]。本研究发现,LPS应激显著降低了肉仔鸡十二指肠和空肠VH、V/C,显著提高了十二指肠CD,这与前人研究结果[50]一致。研究发现,饲粮中添加50和100 mg/kg茯苓提取物(主要有效成分为茯苓多糖β-1,3-D-葡聚糖,含量>60.0%),可显著提高肉仔鸡十二指肠、空肠及回肠VH和V/C,显著降低CD,改善肠道形态结构[51]。另有研究发现,CMP可通过减少肠道上皮细胞坏死,保护肠黏膜屏障,从而缓解DSS诱导的小鼠结肠损伤[24]。本研究发现,饲粮中添加CMP可显著提高肉仔鸡空肠V/C,且有提高十二指肠V/C及空肠VH的趋势。而CD的降低以及VH、V/C的提高,表明了肠道上皮细胞沿隐窝-绒毛轴向顶端的迁移速度加快,上皮细胞谱系的分化过程完整且有序,有利于增强机体消化和吸收能力[52]。由此推测,饲粮中添加200 mg/kg CMP可能通过促进肉仔鸡肠道上皮细胞的分化和增殖,减少了应激引起的肠道上皮细胞坏死,从而改善肠道形态结构。

4 结论

饲粮中添加200 mg/kg CMP可增强肉仔鸡抗氧化和抗炎能力,同时降低肠道通透性、优化小肠黏膜形态结构,维持肠道物理屏障完整,进而在一定程度上缓解LPS应激对肉仔鸡的生长抑制及不利影响。

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