Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 9: 6591 - 6603

DOI: https://doi.org/10.12418/CJAN2026.528

RESEARCH PAPER

Effects and Mechanisms of Lemna minor Linnaeus Flavonoids Extract on Alleviating Heat Stress in Broilers

  • LIU Lindong , 1 ,
  • CHEN Xiuyun 2 ,
  • DONG Wei 1 ,
  • JIA Yongshuang 1 ,
  • DONG Cui , 1, *
Expand
  • 1 College of Pharmacy, Xinyang Agriculture and Forestry University, Xinyang 464000, China
  • 2 College of Agriculture, Xinyang Agriculture and Forestry University, Xinyang 464000, China
* associate professor, E-mail:

Received date: 2026-01-14

  Online published: 2026-09-12

Fold

Abstract

This study was conducted to investigate the alleviating effects of Lemna minor Linnaeus flavonoids extract (LMF) on heat-stressed broilers and its possible mechanism. A total of 210 healthy 21-day-old Arbor Acres (AA) broilers with similar initial body weight were randomly allocated into 5 groups with 6 replicates per group and 7 birds per replicate. The normal control (NC) group was reared in a constant-temperature zone at 22 ℃ and fed a basal diet. The other four heat-stress groups were housed together in a programmed variable-temperature zone under a cyclic temperature pattern of 22 ℃—32 ℃—22 ℃, and they were fed the basal diet (HS group), the basal diet + 200 mg/kg vitamin E (PC group), the basal diet + 0.735 g/kg LMF (equivalent to 50 mg/kg total flavonoids, L-LMF group), or the basal diet + 1.470 g/kg LMF (equivalent to 100 mg/kg total flavonoids, H-LMF group). The trial lasted for 21 days. The results showed as follows: compared with the NC group, the final body weight (FBW), average daily gain (ADG), and jejunal villus height (VH) and villus height to crypt depth ratio (V/C) of the HS group were significantly decreased (P<0.05), whereas the feed to gain ratio (F/G), jejunal crypt depth (CD), as well as drip loss and cooking loss of breast muscle and thiobarbituric acid reactive substances (TBARS) values in breast muscle refrigerated for 0, 3 and 7 d were significantly increased (P<0.05). Dietary LMF supplementation significantly reversed the changes in the above indices (P<0.05), and no significant differences were observed between the H-LMF group and the NC group for these indices (P>0.05). Compared with the NC group, the activities of superoxide dismutase (SOD) and glutathione peroxidase (GSH-Px) in serum and liver, the mRNA relative expression levels of Occludin, Claudin-1 and zonula occludens-1 (ZO-1) in jejunal mucosa, and the mRNA relative expression levels of hepatic nuclear factor erythroid 2-related factor 2 (Nrf2), heme oxygenase-1 (HO-1) and NAD(P)H:quinone oxidoreductase-1 (NQO-1) in the HS group were significantly decreased (P<0.05), while the malondialdehyde (MDA) content in liver, as well as the contents of MDA, interleukin-1β (IL-1β), tumor necrosis factor-α (TNF-α) and lipopolysaccharide (LPS) and the activity of diamine oxidase (DAO) in serum of the HS group were significantly increased (P<0.05). Dietary LMF supplementation significantly reversed the changes in the above indices (P<0.05), and no significant differences were observed between the H-LMF group and the PC group for these indices (P>0.05). In conclusion, LMF can alleviate oxidative damage and inflammation, improve intestinal health and meat quality, and enhance growth performance in heat-stressed broilers, and this effect may be associated with the up-regulation of gene expression related to the hepatic Nrf2 signaling pathway.

Key words： Lemna minor Linnaeus; flavonoids extract; heat stress; broilers; Nrf2 signaling pathway

Cite this article

LIU Lindong , CHEN Xiuyun , DONG Wei , JIA Yongshuang , DONG Cui . Effects and Mechanisms of Lemna minor Linnaeus Flavonoids Extract on Alleviating Heat Stress in Broilers[J]. Chinese Journal of Animal Nutrition, 2026 , 38(9) : 6591 -6603 . DOI: 10.12418/CJAN2026.528

热应激是制约集约化肉鸡养殖效益的关键环境因素,可导致采食抑制、生长迟缓、死亡率上升及肌肉品质下降等一系列生产问题,造成显著的经济损失[1-2]。其病理生理学基础与机体氧化还原稳态的破坏密切相关,持续的热应激触发系统性氧化应激,进而损害肠道黏膜结构的完整性[3-6],并启动全身性炎症反应,最终形成“氧化损伤-屏障功能障碍-生产性能下降”的恶性循环[7-8]。在饲料中全面禁止添加抗生素及倡导健康养殖的背景下,筛选与开发能够高效缓解氧化应激的天然饲料添加剂,已成为保障畜牧业绿色可持续发展的重要研究方向[9-11]。
黄酮类化合物作为广泛存在于植物中的次级代谢产物,因其抗氧化与抗炎的生物活性,在动物营养领域备受关注[12]。近年来,其作用机理研究已从早期的直接清除活性氧,深化为对细胞内源性防御网络的调控。其中,核因子E2相关因子2(nuclear factor erythroid 2-related factor 2,Nrf2)是介导植物化学物调控哺乳动物细胞抗氧化反应的核心分子枢纽[13-14]。多项研究表明,特定的植物黄酮(如黄芪黄酮、黄芩黄酮、染料木素等)可通过调节Nrf2等信号通路,有效缓解热应激及肝脏损伤[15-18]。浮萍(Lemna minor Linnaeus)作为一种繁殖迅速、生物量高的水生植物,其作为蛋白质源和功能性饲料组分在畜禽养殖中的应用已受到关注[19-22]。体外及动物研究表明,浮萍乙醇提取物可减轻神经细胞毒性[23],在体外表现出抗氧化与抗炎作用[24],可用于绿色合成具有抗氧化特性的纳米材料[25],为其在缓解动物应激方面的应用提供了初步依据。然而,目前的研究大多集中于浮萍在食品与医药相关领域的开发[26-27];在饲养领域,虽有研究报道了以浮萍部分替代蛋鸡饲粮中的小麦和豆粕可对产蛋性能、蛋品质及机体抗氧化状态产生积极影响[28],但仍主要停留在粗提物或全草粉的替代效应观察层面。关于标准化浮萍黄酮提取物(Lemna minor Linnaeus flavonoids extract,LMF)在集约化养殖中常见的热应激模型下的系统性功效评估,尤其是基于Nrf2信号通路调控的分子机制,仍缺乏直接且深入的试验证据。
基于上述研究现状与空白,本研究旨在系统评估LMF对热应激肉鸡的保护作用并阐明其分子机制。具体而言,研究将首先制备标准化LMF,继而通过建立肉鸡热应激模型,全面评价LMF对动物生长性能、全身抗氧化与炎症状态、肠道屏障功能以及肉品质的改善效应;并重点在肝脏组织水平上,探究Nrf2信号通路在其中的可能调控作用。本研究将为浮萍的高附加值饲料化利用提供直接的科学依据,并为开发机理相对明确的天然抗热应激添加剂提供新方案。

1 材料与方法

1.1 伦理声明

所有动物试验程序均已通过信阳农林学院实验动物管理与使用委员会审查和批准(批准号:XYAFU-2023-AL-028)。

1.2 试验材料

试验所用维生素E为DL-α-生育酚乙酸酯,纯度≥96%。
浮萍于2023年6月采自河南省信阳市天然水域,洗净后60 ℃烘干,粉碎过60目筛,备用。称取浮萍干粉,按照重量(g)∶体积(mL)=1∶30的比例加入30倍体积的50%(v/v)乙醇溶液,于360 W功率下超声提取36 min。提取液经50 ℃减压浓缩后,采用AB-8大孔吸附树脂富集,收集70%(v/v)乙醇洗脱组分,经浓缩后进行冷冻干燥,得到LMF粉末,-20 ℃避光保存。LMF中总黄酮含量采用硝酸铝-亚硝酸钠比色法(以芦丁为标准品)测定,结果为重复6次后的平均值。经测定,LMF中总黄酮含量为(6.8±0.2)%。

1.3 试验设计与饲养管理

选取240只1日龄体重相近的健康爱拔益加(Arbor Acres,AA)商品代肉公雏经常规饲养20 d。21日龄时,选取210只初始体重相近的健康AA肉鸡,随机分为5组,每组6个重复,每个重复7只鸡。常温对照(NC)组饲养于22 ℃恒温区,饲喂基础饲粮;其余4个热应激组集中饲养于程序变温区,执行22 ℃—32 ℃—22 ℃的周期性循环变温模式,分别饲喂基础饲粮(HS组)、基础饲粮+200 mg/kg维生素E(PC组)、基础饲粮+0.735 g/kg LMF(按总黄酮计,添加水平为50 mg/kg,L-LMF组)和基础饲粮+1.470 g/kg LMF(按总黄酮计,添加水平为100 mg/kg,H-LMF组)。试验期21 d。
基础饲粮参照《鸡饲养标准》(NY/T 33—2004)进行配制,其组成及营养水平见表1。饲粮中LMF添加量基于课题组前期剂量筛选试验结果并参考同类黄酮物质的相关文献[17]确定。所有添加剂均采用逐级扩大法预混入基础饲粮,充分混匀后制粒备用。
表1 基础饲粮组成及营养水平(风干基础)

Table 1 Composition and nutrient levels of the basal diet (air-dry basis)

原料Ingredients 含量Content 营养水平Nutrient levels2) 含量Content
玉米Corn 58.50 代谢能ME/(MJ/kg) 13.03
豆粕Soybean meal 31.00 粗蛋白质CP 20.50
大豆油Soybean oil 4.00 粗脂肪EE 5.80
石粉Limestone 1.30 钙Ca 1.00
磷酸氢钙CaHPO4 1.80 总磷TP 0.65
食盐NaCl 0.30 有效磷AP 0.45
L-赖氨酸L-Lys 0.20 赖氨酸Lys 1.15
DL-蛋氨酸DL-Met 0.20 蛋氨酸+胱氨酸Met+Cys 0.85
预混料Premix1) 2.70
合计Total 100.00

1)预混料为每千克饲粮提供 The premix provided the following per kg of the diet:VA 12 500 IU,VD3 3 750 IU,VE 30 IU,VK3 3 mg,VB1 2.5 mg,VB2 8 mg,VB6 4 mg,VB12 0.025 mg,烟酸 nicotinic acid 40 mg,泛酸钙 calcium pantothenate 15 mg,Cu (CuSO4·5H2O) 8 mg,Fe (FeSO4·7H2O) 80 mg,Zn (ZnSO4·7H2O) 60 mg,Mn (MnSO4·H2O) 80 mg,I (KI) 0.35 mg,Se (Na2SeO3) 0.15 mg,植酸酶 phytase 500 U。

2)代谢能和有效磷为计算值,参照《中国饲料成分及营养价值表(2023年第34版)》计算,其余为实测值。ME and AP were calculated values, calculated with reference to the Tables of Feed Composition and Nutritive Values in China (34th edition, 2023), while the others were measured values.

饲养试验于2023年7—8月在信阳市的标准化肉鸡舍内完成。根据AA肉鸡生长中后期(21~42日龄)的生理特点,其适宜环境温度为18~24 ℃。为模拟生产中常见的中重度热应激,本试验建立了周期性循环热应激模型。通过计算机环境控制系统,将饲养区划分为恒温区与程序变温区。恒温区全程维持(22±1) ℃;程序变温区采用周期性循环变温模式:每天10:00—18:00(共8 h)维持高温(32±1) ℃,其余时间恢复至(22±1) ℃。试验期间,所有肉鸡自由采食与饮水,每天通过料槽定时饲喂2次(08:00和16:00),并按常规程序进行免疫。

1.4 样品采集与指标检测

1.4.1 饲粮营养水平测定

饲粮中粗蛋白质(CP)含量按照GB/T 6432—2018的凯氏定氮法测定;粗脂肪(EE)含量按照GB/T 6433—2025测定;钙(Ca)和总磷(TP)含量分别按照GB/T 6436—2018和GB/T 6437—2018测定;氨基酸含量按照GB/T 18246—2019,采用氨基酸自动分析仪测定。

1.4.2 生长性能测定

试验期开始和结束当天晨饲前,以重复为单位对所有肉鸡进行空腹称重,并准确记录试验全期的耗料总量,计算平均日增重(average daily gain,ADG)、平均日采食量(average daily feed intake,ADFI)和料重比(feed to gain ratio,F/G)。

1.4.3 样品采集

试验结束当天,从每个重复中选取1只最接近该重复平均体重的肉鸡进行翅静脉采血。血液静置后,于4 ℃、2 000×g条件下离心15 min制备血清,分装于-80 ℃保存备用。采血后进行屠宰开腹,立即采集肝脏及左侧胸大肌,胸大肌样品立即按1.4.7所述方法进行肉品质指标测定。同时,截取2段长度约2 cm空肠中段组织,其中一段放入4%多聚甲醛固定,另外一段用预冷生理盐水轻柔冲洗内容物后,沿肠系膜对侧纵向剪开,使用无菌载玻片轻轻刮取黏膜层,装入冻存管,经液氮速冻后转存于-80 ℃备用。

1.4.4 血清抗氧化、炎症及肠道屏障功能相关指标测定

血清抗氧化指标包括总抗氧化能力(total antioxidant capacity,T-AOC)、丙二醛(malondialdehyde,MDA)含量及超氧化物歧化酶(superoxide dismutase,SOD)和谷胱甘肽过氧化物酶(glutathione peroxidase,GSH-Px)活性;血清炎症指标包括白细胞介素-1β(interleukin-1β, IL-1β)和肿瘤坏死因子-α(tumor necrosis factor-α,TNF-α)含量;血清肠道屏障功能相关指标包括脂多糖(LPS)含量、二胺氧化酶(diamine oxidase,DAO)活性。其中,T-AOC与MDA和LPS含量及SOD和GSH-Px活性采用购自南京建成生物工程研究所的试剂盒进行测定;IL-1β、TNF-α含量及DAO活性采用上海酶联生物科技有限公司的酶联免疫吸附测定(enzyme-linked immunosorbent assay,ELISA)试剂盒测定。

1.4.5 空肠形态学指标及紧密连接蛋白基因表达分析

空肠组织样品经4%多聚甲醛固定、常规石蜡包埋、切片(厚度4 μm)及苏木精-伊红(HE)染色。染色后的切片在BX53型正置光学显微镜(Olympus,日本)下观察,并利用其配套的DP73图像采集系统进行全片扫描。采用Image-Pro Plus 6.0图像分析软件,在每个样本的完整视野内随机选取10根结构清晰的绒毛,由研究人员进行盲法测量,精确获取绒毛高度(villus height,VH)和隐窝深度(crypt depth,CD),并计算绒隐比(villus height to crypt depth ratio,V/C)。所有测量均遵循统一的校准标准。
采用TRIzol试剂(Invitrogen,美国)通过一步法提取空肠黏膜总RNA,提取的RNA经NanoDrop One超微量分光光度计(Thermo Fisher Scientific,美国)检测纯度与浓度,检测合格后使用PrimeScriptTM RT Reagent Kit with gDNA Eraser(TaKaRa,日本)进行基因组DNA去除及反转录合成cDNA。以合成的cDNA为模板,采用TB Green® Premix Ex TaqTM Ⅱ(TaKaRa,日本)在QuantStudioTM 5型实时荧光定量PCR系统(Applied Biosystems,美国)上进行扩增反应,检测闭合蛋白(Occludin)、封闭蛋白-1(Claudin-1)及闭锁小带蛋白-1(ZO-1)mRNA相对表达量,引物序列由生工生物工程(上海)股份有限公司合成,引物序列见表2。以β-肌动蛋白(β-actin)为内参基因,采用2-ΔΔCt法计算目的基因mRNA相对表达量。
表2 实时荧光定量PCR引物序列

Table 2 Primer sequences for real-time quantitative PCR

基因Genes 引物序列Primer sequences (5'—3') 产物大小Product size/bp
β-肌动蛋白β-actin F:CCACCAGCACAATGAAGATCA
R:TCATCACCAGAGTCCATCACG		 150
闭合蛋白Occludin F:GCTGCTAACGGCTACATCCA
R:TGGCTTCTGATGGTGTCTTG		 118
封闭蛋白-1 Claudin-1 F:TGGAACAGCAGCATCAGAGG
R:TCCACAGTCGATGCCATTTC		 102
闭锁小带蛋白-1 ZO-1 F:AAGCAGCCAGAACAAGCAGA
R:GCTGGTCTGGTAGCACCTTC		 135
核因子E2相关因子2 Nrf2 F:TGTGTGTGATTCAACCCGACT
R:TTAATGGAAGCCGCACCACT		 151
血红素加氧酶-1 HO-1 F:TTGGCAAGAAGCATCCAGA
R:TCCATCTCAAGGGCATTCA		 148
醌氧化还原酶-1 NQO-1 F:GTTCAATGCCGTGCTCTCAC
R:CCGCTTCAATCTTCTTCTGC		 149

1.4.6 肝脏抗氧化指标及Nrf2信号通路相关基因表达分析

准确称取约0.1 g肝脏组织,加入9倍体积(0.9 mL)预冷生理盐水,于冰浴条件下机械匀浆,随后在4℃、1 600×g条件下离心10 min,取上清液待测。采用南京建成生物工程研究所试剂盒测定肝脏匀浆上清中SOD、GSH-Px活性及MDA含量。
肝脏组织总RNA的提取、反转录及实时荧光定量PCR反应操作过程同1.4.5,用于检测Nrf2、血红素加氧酶-1(heme oxygenase-1,HO-1)及醌氧化还原酶-1(quinone oxidoreductase-1,NQO-1)mRNA相对表达量,引物序列由生工生物工程(上海)股份有限公司合成,引物序列见表2。目的基因mRNA相对表达量计算方法同1.4.5。

1.4.7 胸肌肉品质指标测定

滴水损失:取约50 g胸肌样品称重(W1),悬挂于4 ℃冰箱中24 h后,用滤纸吸干表面水分再次称重(W2),按以下公式计算滴水损失。
滴水损失(%)= $\frac{({\mathit{W}}_{1}-{\mathit{W}}_{2})}{{\mathit{W}}_{1}}$×100。
蒸煮损失:取约30 g胸肌样品称重(W3),装入自封袋水浴加热至中心温度达75 ℃,冷却擦干后称重(W4),按以下公式计算蒸煮损失。
蒸煮损失(%)= $\frac{({\mathit{W}}_{3}-{\mathit{W}}_{4})}{{\mathit{W}}_{3}}$×100。
脂质氧化程度:取绞碎的胸肌肉样,于4 ℃冷藏0(初始)、3、7 d,参照硫代巴比妥酸反应物(TBARS)测定试剂盒(南京建成生物工程研究所)说明书进行操作,测定532 nm处的吸光度值,并计算TBARS值,用以评估贮藏期间的脂质氧化程度。

1.5 数据统计分析

试验数据采用SPSS 26.0统计软件进行单因素方差分析(one-way ANOVA),若差异显著,则进一步使用Tukey’s HSD检验进行事后多重比较。结果数据以“平均值±标准差”的形式表示,P<0.05表示差异显著。

2 结果与分析

2.1 LMF对热应激肉鸡生长性能的影响

由表3可知,与NC组相比,HS组肉鸡的终末体重(FBW)、ADG显著降低(P<0.05),F/G显著升高(P<0.05)。与HS组相比,PC组和H-LMF组的FBW和ADG显著提高(P<0.05),F/G显著降低(P<0.05)。各组间ADFI无显著差异(P>0.05)。
表3 LMF对热应激肉鸡生长性能的影响

Table 3 Effects of LMF on growth performance of heat-stressed broilers

项目
Items		 组别Groups P值
P-value
NC HS PC L-LMF H-LMF
初始体重IBW/g 745.2±15.3 742.8±16.1 743.5±14.9 744.6±15.7 743.9±14.8 0.997
终末体重FBW/g 2 190.8±32.2a 2 002.8±34.6c 2 125.3±29.0ab 2 071.1±32.3bc 2 129.9±29.1ab 0.001
平均日增重ADG/(g/d) 68.8±2.2a 60.0±2.6c 65.8±2.0ab 63.2±2.3bc 66.2±2.1ab 0.001
平均日采食量ADFI/(g/d) 132.8±3.9 129.2±4.4 131.5±3.7 130.6±4.0 132.0±3.6 0.175
料重比F/G 1.93±0.05d 2.11±0.06a 2.00±0.05bc 2.07±0.06ab 1.97±0.05cd <0.001

同行数据肩标无字母或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)。下表同。

In the same row, values with no letter or the same letter superscripts mean no significant difference (P>0.05), while with different small letter superscripts mean significant difference (P<0.05). The same as below.

2.2 LMF对热应激肉鸡血清抗氧化、炎症及肠道屏障功能相关指标的影响

由表4可知,与NC组相比,HS组血清抗氧化能力(T-AOC及SOD和GSH-Px活性)显著降低(P<0.05),而氧化损伤产物(MDA)、促炎细胞因子(IL-1β、TNF-α)含量及肠道屏障损伤标志物(LPS含量、DAO活性)均显著升高(P<0.05)。饲粮补充LMF可剂量依赖性地逆转上述指标的变化,H-LMF组血清T-AOC及SOD和GSH-Px活性均显著高于HS组(P<0.05),显著低于NC组(P<0.05),与PC组相比无显著差异(P>0.05);H-LMF组血清MDA、IL-1β、TNF-α、LPS含量及DAO活性均显著低于HS组(P<0.05),显著高于NC组(P<0.05),与PC组相比无显著差异(P>0.05)。
表4 LMF对热应激肉鸡血清抗氧化、炎症及肠道屏障功能相关指标的影响

Table 4 Effects of LMF on serum antioxidant, inflammatory and intestinal barrier function-related indices of heat-stressed broilers

项目
Items		 组别Groups P值
P-value
NC HS PC L-LMF H-LMF
总抗氧化能力
T-AOC/(U/mL)		 12.9±0.8a 8.2±0.6d 11.6±0.7ab 9.9±0.5c 11.1±0.6b <0.001
超氧化物歧化酶
SOD/(U/mL)		 190±11a 142±9d 176±10ab 159±8c 172±9b <0.001
谷胱甘肽过氧化物
GSH-Px/(U/mL)		 670±38a 478±32d 620±35ab 548±30c 605±33b <0.001
丙二醛MDA/(nmol/mL) 5.1±0.3d 7.5±0.4a 5.7±0.3cd 6.4±0.3b 5.9±0.3c <0.001
白细胞介素-1β
IL-1β/(pg/mL)		 20.0±1.5d 48.0±3.0a 28.0±2.0c 37.5±2.5b 30.0±2.0c <0.001
肿瘤坏死因子-α
TNF-α/(pg/mL)		 48.5±3.0d 91.0±5.0a 59.0±4.0c 74.0±4.5b 63.0±3.5c <0.001
脂多糖LPS/(EU/mL) 0.26±0.02d 0.51±0.03a 0.34±0.02c 0.44±0.03b 0.36±0.02c <0.001
二胺氧化酶DAO/(U/L) 12.5±1.0c 25.3±1.8a 16.8±1.2b 21.4±1.5ab 17.6±1.3b 0.002

2.3 LMF对热应激肉鸡空肠形态学指标的影响

由表5可知,与NC组相比,HS组空肠VH和V/C均显著降低(P<0.05),空肠CD显著增加(P<0.05)。饲粮补充LMF有效逆转了这一趋势,H-LMF组的VH和V/C均显著高于HS组(P<0.05),与NC组和PC组相比均无显著差异(P>0.05)。
表5 LMF对热应激肉鸡空肠形态学指标的影响

Table 5 Effects of LMF on jejunal morphological indices of heat-stressed broilers

项目
Items		 组别Groups P值
P-value
NC HS PC L-LMF H-LMF
绒毛高度VH/μm 492±38a 405±32c 468±35ab 432±34bc 462±36ab <0.001
隐窝深度CD/μm 106±9b 119±11a 109±10ab 116±10a 111±9ab 0.002
绒隐比V/C 4.64±0.36a 3.40±0.27c 4.29±0.33ab 3.72±0.29bc 4.16±0.32ab <0.001

2.4 LMF对热应激肉鸡空肠黏膜紧密连接蛋白基因表达的影响

由表6可知,与NC组相比,HS组空肠黏膜紧密连接蛋白基因Occludin、Claudin-1及ZO-1的mRNA相对表达量显著降低(P<0.05)。L-LMF组和H-LMF组上述3个基因的mRNA相对表达量均显著高于HS组(P<0.05),与PC组相比无显著差异(P>0.05)。H-LMF组上述3个基因的mRNA相对表达量与NC组相比均无显著差异(P>0.05)。
表6 LMF对热应激肉鸡空肠黏膜紧密连接蛋白基因表达的影响

Table 6 Effects of LMF on gene expression of tight junction protein in jejunal mucosa of heat-stressed broilers

项目
Items		 组别Groups P值
P-value
NC HS PC L-LMF H-LMF
闭合蛋白Occludin 1.00±0.08a 0.52±0.06c 0.85±0.07ab 0.68±0.06b 0.82±0.07ab <0.001
封闭蛋白-1 Claudin-1 1.00±0.09a 0.48±0.05c 0.88±0.08ab 0.71±0.07b 0.90±0.08ab <0.001
闭锁小带蛋白-1 ZO-1 1.00±0.07a 0.61±0.06c 0.92±0.08ab 0.79±0.07b 0.94±0.08a <0.001

2.5 LMF对热应激肉鸡肝脏抗氧化指标及Nrf2信号通路相关基因表达的影响

由表7可知,与NC组相比,HS组肝脏SOD和GSH-Px活性显著降低(P<0.05),MDA含量显著升高(P<0.05)。H-LMF组肝脏SOD和GSH-Px活性显著高于HS组(P<0.05),MDA含量显著低于HS组(P<0.05),而均与PC组相比无显著差异(P>0.05)。
表7 LMF对热应激肉鸡肝脏抗氧化指标及Nrf2信号通路相关基因表达的影响

Table 7 Effects of LMF on hepatic antioxidant indices and Nrf2 signaling pathway-related gene expression of heat-stressed broilers

项目
Items		 组别Groups P值
P-value
NC HS PC L-LMF H-LMF
超氧化物歧化酶
SOD/(U/mg prot)		 185±10a 138±8d 170±9ab 152±8c 165±9b <0.001
谷胱甘肽过氧化物
GSH-Px/(U/mg prot)		 65.2±3.5a 46.5±3.0d 60.1±3.2ab 52.8±2.9c 58.6±3.1b <0.001
丙二醛
MDA/(nmol/mg prot)		 1.05±0.08d 1.68±0.12a 1.20±0.09cd 1.45±0.10b 1.25±0.09c <0.001
核因子E2相关因子2 Nrf2 1.00±0.08a 0.55±0.05c 0.91±0.07ab 0.73±0.06b 0.93±0.08a <0.001
血红素加氧酶-1 HO-1 1.00±0.09a 0.58±0.06c 0.89±0.08ab 0.76±0.07b 0.95±0.08a <0.001
醌氧化还原酶-1 NQO-1 1.00±0.07a 0.62±0.05c 0.94±0.07a 0.81±0.06b 0.96±0.07a <0.001
由表7可知,在Nrf2信号通路相关基因表达层面,与NC组相比,HS组肝脏Nrf2、HO-1和NQO-1的mRNA相对表达量均显著降低(P<0.05)。L-LMF组和H-LMF组肝脏中这3个基因的mRNA相对表达量均显著高于HS组(P<0.05)。H-LMF组肝脏中这3个基因的mRNA相对表达量与NC组和PC组相比均无显著差异(P>0.05)。

2.6 LMF对热应激肉鸡胸肌肉品质的影响

由表8可知,与NC组相比,HS组胸肌的滴水损失和蒸煮损失及冷藏0、3、7 d的TBARS值显著增加(P<0.05)。H-LMF组滴水损失和蒸煮损失及冷藏0、3、7 d的TBARS值显著低于HS组(P<0.05),与PC组和NC组相比无显著差异(P>0.05)。
表8 LMF对热应激肉鸡胸肌肉品质的影响

Table 8 Effects of LMF on breast meat quality of heat-stressed broilers

项目
Items		 组别Groups P值
P-value
NC HS PC L-LMF H-LMF
滴水损失Drip loss/% 2.48±0.15c 3.32±0.21a 2.72±0.17bc 3.05±0.19b 2.78±0.16bc 0.003
蒸煮损失Cooking loss/% 21.5±1.0b 25.8±1.2a 22.7±1.1b 24.2±1.1a 22.9±1.0b 0.001
硫代巴比妥酸反应物值TBARS value/(mg/kg)
冷藏0 d Refrigerated for 0 d 0.28±0.02c 0.35±0.03a 0.30±0.02bc 0.33±0.03ab 0.31±0.02bc 0.002
冷藏3 d Refrigerated for 3 d 0.38±0.02c 0.56±0.04a 0.42±0.03bc 0.50±0.03b 0.44±0.03bc 0.001
冷藏7 d Refrigerated for 7 d 0.54±0.03c 0.84±0.04a 0.59±0.03bc 0.73±0.04b 0.61±0.03bc <0.001

3 讨论

热应激通过诱发氧化应激,对肉鸡健康与生产造成多重危害。本研究首次系统证实,LMF能够剂量依赖性地缓解热应激损伤,其可能核心机制与激活肝脏的Nrf2内源性抗氧化信号通路密切相关。

3.1 LMF改善热应激肉鸡生长性能的综合效应

本试验结果表明,饲粮中添加1.470 g/kg LMF(按总黄酮计,添加水平为100 mg/kg)能显著逆转热应激导致的ADG下降和F/G升高,其效果与200 mg/kg维生素E相当。这一促生长作用与多种天然添加剂在热应激肉鸡上的报道一致,证实了通过营养干预缓解热应激、恢复生长的可行性[29-31]。该效应并非通过刺激采食实现(各组ADFI无显著差异),而是源于对机体生理状态的系统性改善。热应激引起的氧化损伤和肠道屏障功能障碍是抑制生长的核心病理环节[7,32]。本研究发现,与HS组相比,饲粮添加高剂量LMF在显著提升血清与肝脏SOD、GSH-Px等抗氧化酶活性、降低血清与肝脏氧化损伤产物MDA含量的同时,也显著降低了反映肠道屏障通透性的血清LPS与DAO含量。更重要的是,本研究证实LMF可显著缓解热应激导致的空肠绒毛萎缩和绒隐比下降及空肠黏膜紧密连接蛋白基因Occludin、Claudin-1和ZO-1的mRNA相对表达量的降低,为养分吸收的关键结构提供了支持。这与壳寡糖、薄荷提取物等通过保护肠道功能来改善生长性能的路径相似[33-34]。因此,LMF可能是通过“缓解氧化应激→保护肠道结构与屏障功能→从而为养分吸收创造有利条件”的途径,最终实现了生长性能的恢复。

3.2 LMF对热应激肉鸡肝脏Nrf2信号通路基因表达的上调作用

本研究中,与HS组相比,饲粮添加LMF可上调肝脏Nrf2及其下游靶基因HO-1、NQO-1的mRNA相对表达量。Nrf2是细胞防御氧化应激的中枢调节因子。在静息状态下,Nrf2与其抑制蛋白Kelch样环氧氯丙烷关联蛋白1(Kelch-like epichlorohydrin-associated protein 1,Keap1)结合并被泛素化降解;而当受到氧化压力或特定激活剂刺激时,Nrf2被释放并转运入核,启动下游HO-1、NQO-1等一系列编码Ⅱ相解毒酶和抗氧化蛋白基因的转录[9]。本试验中,与NC组相比,热应激可显著抑制肝脏Nrf2、HO-1及NQO-1基因的表达,而饲粮补充LMF,尤其是高剂量组,能显著逆转这一抑制,上调这些关键基因的mRNA相对表达量。这与Wang等[10]报道的杜仲黄酮通过调控Nrf2通路缓解猪氧化应激的机制相符。这些发现提示,LMF的抗氧化作用可能与调节Nrf2信号通路相关基因的转录有关[35]。其潜在的分子机制在于,LMF中富含的黄酮类成分可通过多种途径影响Keap1-Nrf2信号轴:它们可作为亲电体与Keap1蛋白的关键半胱氨酸残基发生共价结合,或通过调控上游激酶信号通路,解除Keap1对Nrf2的抑制作用,促进Nrf2蛋白核转位,诱导下游抗氧化基因的转录表达[36-38]。这提示LMF可能不仅直接中和自由基,其作用还与Nrf2信号通路相关基因的转录上调有关。其他研究也表明,绿原酸可通过影响Nrf2/HO-1信号通路保护热应激肉鸡肠道屏障[39],提示黄酮类化合物的作用存在多靶点特性。

3.3 LMF对热应激肉鸡全身性炎症与肉品质的改善作用

氧化应激与炎症反应紧密相连。本研究发现,LMF在提升热应激肉鸡抗氧化能力的同时,能显著降低其血清中关键促炎细胞因子IL-1β和TNF-α含量。IL-1β和TNF-α是激活下游炎症级联反应的核心信号分子,其含量的显著降低强烈提示机体全身性炎症状态得到缓解。更重要的是,这种抗炎效应与肠道屏障功能的改善同步发生:LMF同时降低了血清LPS和肠道损伤标志物DAO的含量,并改善了空肠形态学指标。本研究结果提示,LMF的抗炎作用可能与其上调Nrf2信号通路相关基因表达、缓解肝脏及全身氧化应激有关;氧化应激的减轻有助于保护肠道黏膜结构的完整性,从而减少肠道内毒素易位(血清LPS含量降低);最终,内毒素负荷的减少削弱了对全身免疫系统的持续刺激,导致了血清关键促炎细胞因子含量的下降。这一“抗氧化-屏障保护-抗炎”的协同路径,与先前关于壳寡糖[40]和白藜芦醇[41]等其他植物提取物的研究报道相一致。
肉品质的劣化与宰后肌肉的氧化进程密切相关。本试验发现,LMF能显著降低热应激肉鸡胸肌的滴水损失和冷藏0、3、7 d的TBARS值。滴水损失与肌原纤维蛋白的氧化变性及细胞膜完整性有关,而TBARS值直接反映脂质氧化程度。动态数据显示,H-LMF组胸肌TBARS值在开始冷藏时即显著低于HS组,且该差异持续至冷藏7 d,表明LMF在动物活体内建立的强效抗氧化状态,能在宰后贮藏期内持续保护肌肉细胞组分,延缓氧化酸败。这与裂殖壶藻粉[42]、斑蝥黄[43]等添加剂改善热应激肉鸡肉品质的报道一致。这表明LMF的应用价值可从养殖环节延伸至产品品质控制,有助于提升商品附加值[44]。
本研究为浮萍资源的高值化利用开辟了新路径。相较于组成复杂的粗提物,(6.8±0.2)%黄酮含量的标准化LMF产品更具开发可控性。其在按总黄酮计100 mg/kg添加水平下表现出的与200 mg/kg维生素E相当的综合性功效,展示了其作为潜在替代或互补型天然抗氧化剂的应用前景。一项在蛋鸡上的研究显示,用浮萍部分替代饲粮未对蛋鸡健康产生负面影响[28],为本研究剂量下的安全性提供了侧面支持。本研究也观察到血清肝功能指标(ALT、AST)在各组间无异常,进一步初步表明该应用剂量下的安全性。
未来研究可进一步深入:1)鉴定LMF中关键黄酮单体及其协同效应,可借鉴从浮萍中鉴定新型生物活性肽的思路[27];2)在不同动物种类或应激模型(如运输应激、免疫应激)中验证其功效;3)开展长期饲喂试验,全面评估其长期安全性、耐受性及对繁殖性能的影响。此外,本研究聚焦于Nrf2信号通路,未来可探索LMF是否也参与调控热应激下的其他关键通路,如与代谢(如胍基乙酸[45])、食欲调节(如牛磺酸[46])、神经内分泌(如γ-氨基丁酸[47])相关的路径,以构建更全面的作用网络,为其最终产业化应用提供更加全面的数据支持。

4 结论

综上所述,LMF可缓解热应激导致的肉鸡氧化损伤、炎症反应及肠道屏障功能障碍,进而改善生长性能与肉品质,该作用可能与上调肝脏Nrf2信号通路相关基因表达有关。

References

Publishing order | Descend order by publishing year | Descend order by cited within
[1]
MANUELIAN C L, PITINO R, RIGHI F, et al. Plant feed additives as natural alternatives to the use of synthetic antioxidant vitamins on poultry performances,health,and oxidative status:a review of the literature in the last 20 years[J]. Antioxidants, 2021, 10(5):659.

DOI

[2]
张少帅, 柴浩靓, 武洪志, 等. 热应激对鸡体热调节、生产性能和机体健康影响的研究进展[J]. 中国畜牧兽医, 2026, 53(1):39-48.

ZHANG S S, CHAI H L, WU H Z, et al. Heat stress impacts on thermoregulation,production performance,and organismal health in chickens:research advance[J]. China Animal Husbandry & Veterinary Medicine, 2026, 53(1):39-48. (in Chinese)

[3]
ZHANG S H. From challenge to opportunity:addressing oxidative stress in animal husbandry[J]. Antioxidants, 2023, 12(8):1543.

DOI

[4]
胡阳阳. 锌通过GPR39/PLC β与A20介导的相关信号通路缓解热应激诱导的肉仔鸡小肠黏膜损伤的分子机制研究[D]. 博士学位论文. 扬州: 扬州大学, 2025.

HU Y Y. Molecular mechanisms of zinc alleviating heat stress-induced small intestinal mucosal injury in broilers via GPR39/PLC β- and A20-mediated signaling pathways[D]. Ph.D.Thesis. Yangzhou: Yangzhou University, 2025. (in Chinese)

[5]
周慧爽, 李艳君, 林树乾, 等. 热应激对鸡肠道屏障的影响及损伤机制研究进展[J]. 中国畜牧杂志, 2024, 60(2):71-77.

ZHOU H S, LI Y J, LIN S Q, et al. Research progress on heat stress on intestinal barrier injury in poultry[J]. Chinese Journal of Animal Science, 2024, 60(2):71-77. (in Chinese)

[6]
邓谭杰, 荀文娟, 侯冠彧, 等. 热应激对鸡肠黏膜屏障的影响及其营养调控研究进展[J]. 中国家禽, 2023, 45(5):92-99.

DENG T J, XUN W J, HOU G Y, et al. Research progress on the effect of heat stress on intestinal mucosal barrier and nutritional manipulation in chickens[J]. China Poultry, 2023, 45(5):92-99. (in Chinese)

[7]
黄雪媛. 黄精须根多糖锌的制备及其对热应激肉仔鸡生长性能与免疫的影响[D]. 硕士学位论文. 雅安: 四川农业大学, 2024.

HUANG X Y.Preparation of polysaccharide-zinc complex from Polygonatum sibiricum fibrous root and its influence on the growth performance and immune response of heat-stressed broilers[D].Master’s Thesis. Ya’an: Sichuan Agricultural University, 2024. (in Chinese)

[8]
陆灿强, 舒邓群, 臧一天. 热应激诱导畜禽氧化应激、热休克反应与免疫和炎症反应的机制及营养调控措施[J]. 动物营养学报, 2021, 33(6):3115-3124.

DOI

LU C Q, SHU D Q, ZANG Y T. Mechanism and nutritional regulation of oxidative stress,heat shock response,immunity and inflammation induced by heat stress in livestock and poultry[J]. Chinese Journal of Animal Nutrition, 2021, 33(6):3115-3124. (in Chinese)

[9]
TSIPLAKOU E, PITINO R, MANUELIAN C L, et al. Plant feed additives as natural alternatives to the use of synthetic antioxidant vitamins in livestock animal products yield,quality,and oxidative status:a review[J]. Antioxidants, 2021, 10(5):780.

DOI

[10]
WANG J, DENG L F, CHEN M X, et al. Phytogenic feed additives as natural antibiotic alternatives in animal health and production:a review of the literature of the last decade[J]. Animal Nutrition, 2024, 17:244-264.

DOI

[11]
韦莹, 段叶辉, 邓近平. 风味氨基酸在猪、鸡生产中的应用研究进展[J]. 中国畜牧兽医, 2025, 52(3):1089-1101.

DOI

WEI Y, DUAN Y H, DENG J P. Research progress on the application of flavor amino acids in pig and chicken production[J]. China Animal Husbandry & Veterinary Medicine, 2025, 52(3):1089-1101. (in Chinese)

[12]
唐若韵. 柚皮素通过降低RIPK3和PDC的结合缓解热应激诱导的宁都黄鸡肝脏损伤[D]. 硕士学位论文. 南昌: 江西农业大学, 2024.

TANG R Y.Naringenin alleviates heat stress-induced liver injury in Nindu yellow chickens by decreasing RIPK3 and PDC binding[D].Master’s Thesis. Nanchang: Jiangxi Agricultural University, 2024. (in Chinese)

[13]
程玉花, 曲玉功, 朱秀高, 等. 硒代蛋氨酸及其衍生物的生物学作用及在猪鸡上的应用[J]. 饲料工业, 2025, 46(14):7-13.

CHENG Y H, QU Y G, ZHU X G, et al. Biological function of selenomethionine and its derivatives and its application in pigs and chickens[J]. Feed Industry, 2025, 46(14):7-13. (in Chinese)

[14]
李涛. 桑叶肽功能评价及缓解鸡胚热应激损伤的作用机制研究[D]. 博士学位论文. 镇江: 江苏科技大学, 2025.

LI T. Functional evaluation of mulberry leaf peptide and mechanism of alleviating heat stress injury in chicken embryos[D]. Ph.D.Thesis. Zhenjiang: Jiangsu University of Science and Technology, 2025. (in Chinese)

[15]
刘芸, 周蔓, 木其尔, 等. 黄芪黄酮的生物学功能及其在动物生产中的应用研究进展[J]. 饲料研究, 2026, 49(6):144-149.

LIU Y, ZHOU M, MU Q E, et al. Research progress on biological functions of Astragalus flavonoids and their applications in animal production[J]. Feed Research, 2026, 49(6):144-149. (in Chinese)

[16]
王一冰, 牛开元, HASSAN M, 等. 饲粮中添加黄芩黄酮对黄羽肉鸡生长性能、肉品质和抗氧化能力的影响[J]. 动物营养学报, 2026, 38(2):1015-1024.

DOI

WANG Y B, NIU K Y, HASSAN M, et al. Effects of dietary supplementation with Scutellaria flavonoids on growth performance, meat quality and antioxidant capacity of yellow-feathered broilers[J]. Chinese Journal of Animal Nutrition, 2026, 38(2):1015-1024. (in Chinese)

[17]
戚秋蓉. 染料木素通过调控Mfn2与PLIN1互作缓解热应激引起的肉鸡肝脏脂质沉积[D]. 硕士学位论文. 南昌: 江西农业大学, 2024.

QI Q R. Genistein reduces liver lipid deposition in broilers under heat stress by regulating the interaction between Mfn2 and PLIN1[D]. Master’s Thesis. Nanchang: Jiangxi Agricultural University, 2024. (in Chinese)

[18]
肖华钦, 熊云霞, 曹舒婷, 等. 黄芩黄酮的生物学功能及其在畜禽生产中的应用研究进展[J]. 动物营养学报, 2026, 38(2):926-935.

DOI

XIAO H Q, XIONG Y X, CAO S T, et al. Research progress on biological functions of Scutellaria baicalensis flavonoids and their application in livestock and poultry production[J]. Chinese Journal of Animal Nutrition, 2026, 38(2):926-935. (in Chinese)

[19]
贺洁, 丁志烟, 谭曾蕾, 等. 浮萍作为饲料的生物学功能及其在养殖业中的应用进展[J]. 饲料工业, 2024, 45(16):42-46.

HE J, DING Z Y, TAN Z L, et al. Biological function of duckweed and its application in feed[J]. Feed Industry, 2024, 45(16):42-46. (in Chinese)

[20]
孙炜妍, 马玉彬, 李习龙, 等. 浮萍的生物学功能及其在畜禽生产中的应用[J]. 动物营养学报, 2024, 36(4):2199-2207.

DOI

SUN W Y, MA Y B, LI X L, et al. Biological functions of duckweed and its application in livestock and poultry production[J]. Chinese Journal of Animal Nutrition, 2024, 36(4):2199-2207. (in Chinese)

DOI

[21]
MULLER T, COURNOYER A, BAZINET L. Emerging potentials of duckweed (Lemnaceae):from composition to protein uses in food and nutraceuticals:a review[J]. Food Research International, 2025, 219:116777.

DOI

[22]
SEFERLI M, LEFKAKI M, KOTANIDOU C, et al. Bioactives of the freshwater aquatic plants,Nelumbo nucifera and Lemma minor,for functional foods,cosmetics and pharmaceutical applications,with antioxidant,anti-inflammatory and antithrombotic health promoting properties[J]. Applied Sciences, 2024, 14(15):6634.

DOI

[23]
JEONG Y H, JUNG Y, LI W, et al. Spirodelae herba ethanol extract attenuates neurotoxicity in hippocampal cells and improves scopolamine-induced cognitive impairment in mice[J]. Frontiers in Pharmacology, 2025, 16:1638068.

DOI

[24]
SEFERLI M, LEFKAKI M, MANOUSAKIS V, et al. Amphiphilic bioactives of freshwater aquatic plants Nelumbo nucifera (Indian lotus) and Lemma sp. with antioxidant,anti-inflammatory and antithrombotic activities:in vitro study[J]. Pharmaceuticals, 2025, 18(6):835.

DOI

[25]
SHUKLA Y K, PANDEY P, PANDEY J P, et al. Duckweed extract-mediated green synthesis of ZnO nanoparticles and its antibacterial,antioxidant,and photocatalytic properties[J]. Biomass Conversion and Biorefinery, 2025, 15(14):21477-21491.

DOI

[26]
XIAO M G, LEUNG A Y K, ZHU Z H, et al. Optimization of the extraction of duckweed protein and its application as an antioxidant composite membrane in blueberry preservation[J]. International Journal of Biological Macromolecules, 2025, 311(Pt 2):143847.

DOI

[27]
COURNOYER A, BERNIER M È, ABOUBACAR H, et al. Machine learning-driven discovery of bioactive peptides from duckweed (Lemnaceae) protein hydrolysates:identification and experimental validation of 20 novel antihypertensive,antidiabetic,and/or antioxidant peptides[J]. Food Chemistry, 2025, 482:144029.

DOI

[28]
BAGHBAN-KANANI P, OTERI M, HOSSEINTABAR-GHASEMABAD B, et al. The effects of replacing wheat and soyabean meal with duckweed (Lemna minor) and including enzymes in the diet of laying hens on the yield and quality of eggs,biochemical parameters,and their antioxidant status[J]. Animal Science Journal, 2023, 94(1):e13888.

DOI

[29]
林思婷. 柚皮素调节线粒体动力学缓解热应激致宁都黄鸡十二指肠肠道屏障损伤[D]. 硕士学位论文. 南昌: 江西农业大学, 2025.

LIN S T. Naringenin modulates mitochondrial dynamics to alleviate heat stress-induced duodenal intestinal barrier damage in Ningdu yellow chickens[D]. Master’s Thesis. Nanchang: Jiangxi Agricultural University, 2025. (in Chinese)

[30]
谭相振, 林树乾, 杨世发, 等. 锦灯笼提取物缓解热应激致鸡肠道菌群紊乱的作用机理[J/OL]. 南方农业学报, 2025:1-14(2025-12-08)[2026-03-02].

TAN X Z, LIN S Q, YANG S F, et al. The mechanism of Physalis Calyx seu Fructus extracts relieving chicken intestinal flora disorder caused by heat stress[J/OL]. Journal of Southern Agriculture, 2025:1-14(2025-12-08)[2026-03-02]. in Chinese)

[31]
ZMRHAL V, HAMPEL D, LICHOVNIKOVA M. Curcuma and Scutellaria baicalensis extracts reduced thermoregulatory behavior in chickens subjected to moderate heat stress[J]. Applied Animal Behaviour Science, 2025, 282:106471.

DOI

[32]
王玉蕾, 赵淑娟, 公苹, 等. 基于ADMET预测和Keap1分子对接探究表儿茶素和染料木素缓解肉鸡热应激的作用研究[J/OL]. 南京农业大学学报, 2026:1-13(2026-01-19)[2026-01-23].

WANG Y L, ZHAO S J, GONG P, et al. Combining ADMET prediction and Keap1 molecular docking to investigate the effects of (-)-epicatechin and genistein on alleviating heat stress in broilers[J/OL]. Journal of Nanjing Agricultural University, 2026:1-13(2026-01-19)[2026-01-23]. in Chinese)

[33]
张迦, 王浩轩, 高金稔, 等. 壳寡糖对热应激肉鸡生长性能及肠道消化吸收功能的影响[J]. 动物营养学报, 2025, 37(7):4408-4421.

DOI

ZHANG J, WANG H X, GAO J R, et al. Effects of chitosan oligosaccharides on growth performance and intestinal digestion and absorption function of broilers under heat stress[J]. Chinese Journal of Animal Nutrition, 2025, 37(7):4408-4421. (in Chinese)

DOI

[34]
廖婵娟, 李丹, 阳巧梅, 等. 薄荷提取物对热应激肉仔鸡生长性能、血清生化及肠道免疫指标的影响[J]. 中国饲料, 2025(14):141-144.

LIAO C J, LI D, YANG Q M, et al. Effects of peppermint extract on growth performance,serum biochemical and intestinal immune indexes of broilers under heat stress[J]. China Feed, 2025(14):141-144. (in Chinese)

[35]
吴怡佳, 杨茹洁, 邓谭杰, 等. 豆蔻明对热应激儋州鸡肝脏抗氧化和抗炎性能的影响[J]. 中国饲料, 2025(3):99-105.

WU Y J, YANG R J, DENG T J, et al. Effect of cardamonin on antioxidant and anti-inflammatory properties of heat stress Danzhou chicken liver[J]. China Feed, 2025(3):99-105. (in Chinese)

[36]
HUANG W K, ZHONG Y, GAO B T, et al. Nrf2-mediated therapeutic effects of dietary flavones in different diseases[J]. Frontiers in Pharmacology, 2023, 14:1240433.

DOI

[37]
KIM M, JEE S C, SUNG J S. Hepatoprotective effects of flavonoids against benzo[a]pyrene-induced oxidative liver damage along its metabolic pathways[J]. Antioxidants, 2024, 13(2):180.

DOI

[38]
ZULKEFLI N, CHE ZAHARI C N M, SAYUTI N H, et al. Flavonoids as potential wound-healing molecules:emphasis on pathways perspective[J]. International Journal of Molecular Sciences, 2023, 24(5):4607.

DOI

[39]
孟立峰, 赵桂省, 张明江. 绿原酸对慢性热应激肉仔鸡肠道屏障保护作用的研究[J]. 饲料工业, 2024, 45(14):50-55.

MENG L F, ZHAO G S, ZHANG M J. Study on protective effect of chlorogenic acid on intestinal barrier of broiler under chronic heat stress[J]. Feed Industry, 2024, 45(14):50-55. (in Chinese)

[40]
高金稔, 王浩轩, 张迦, 等. 壳寡糖调控循环热应激肉鸡宰后胸肌能量代谢、抗氧化能力及脂肪酸组成对肉品质的影响[J/OL]. 动物营养学报, 2026:1-14(2026-02-13)[2026-03-02].

GAO J R, WANG H X, ZHANG J, et al. Regulation of chitosan oligosaccharides on postmortem energy metabolism,antioxidant capacity and fatty acid composition in breast muscle of broilers under cyclic heat stress: implications for meat quality[J/OL]. Chinese Journal of Animal Nutrition, 2026:1-14(2026-02-13)[2026-03-02]. in Chinese)

[41]
ZHAO Y Y, LI Z, WANG X C, et al. Resveratrol attenuates heat stress-induced impairment of meat quality in broilers by regulating the Nrf2 signaling pathway[J]. Animals, 2022, 12(15):1889.

DOI

[42]
郭旭明, 陈静龙, 雷闯闯, 等. 饲粮中添加裂殖壶藻粉对热应激黄羽肉鸡生长性能、肉品质、器官发育、血清生化指标、氧化应激及炎症反应的影响[J]. 动物营养学报, 2025, 37(9):5897-5910.

DOI

GUO X M, CHEN J L, LEI C C, et al. Effects of dietary Schizochytrium powder on growth performance,meat quality,organ development,serum biochemical indexes,oxidative stress and inflammatory response of yellow-feathered broilers under heat stress[J]. Chinese Journal of Animal Nutrition, 2025, 37(9):5897-5910. (in Chinese)

[43]
刘元, 谢婷婷, 尹朗庭, 等. 斑蝥黄对热应激怀乡鸡抗氧化能力与肠道黏膜结构的影响[J]. 中国兽医学报, 2023, 43(10):2171-2180.

LIU Y, XIE T T, YIN L T, et al. Effect of canthaxanthin on antioxidant capacity and small intestinal mucosal structure of Huaixiang chickens under heat stress[J]. Chinese Journal of Veterinary Science, 2023, 43(10):2171-2180. (in Chinese)

[44]
王玥, 白秀玉, 李强, 等. 环境热应激对畜禽养殖及肉质影响的研究进展[J]. 肉类研究, 2023, 37(7):71-78.

WANG Y, BAI X Y, LI Q, et al. Progress in research on the effects of environmental heat stress on livestock and poultry farming and meat quality[J]. Meat Research, 2023, 37(7):71-78. (in Chinese)

[45]
彭旭艺, 邢通, 李蛟龙, 等. 慢性热应激对肉鸡盲肠菌群结构和代谢的影响及日粮添加胍基乙酸作用效果研究[J]. 南京农业大学学报, 2024, 47(4):731-740.

PENG X Y, XING T, LI J L, et al. Effects of chronic heat stress on the cecal microflora structure and metabolism of broiler and effects of dietary supplementation guanidinoacetic acid[J]. Journal of Nanjing Agricultural University, 2024, 47(4):731-740. (in Chinese)

[46]
何晓芳, 张林, 李蛟龙, 等. 饲粮中添加牛磺酸对慢性热应激肉鸡血清内分泌激素水平、下丘脑形态及食欲相关基因表达的影响[J]. 畜牧与兽医, 2022, 54(12):24-30.

HE X F, ZHANG L, LI J L, et al. Effects of dietary taurine supplementation on the serum endocrine hormone level,hypothalamus morphology,appetite-related genes expression in broilers under chronic heat stress[J]. Animal Husbandry & Veterinary Medicine, 2022, 54(12):24-30. (in Chinese)

[47]
郭长征, 陈静龙, 雷闯闯, 等. γ-氨基丁酸对热应激黄羽肉鸡生长性能、器官指数、血清生化指标及肠道形态的影响[J]. 动物营养学报, 2024, 36(5):2938-2947.

DOI

GUO C Z, CHEN J L, LEI C C, et al. Effects of γ-aminobutyric acid on growth performance,organ indexes,serum biochemical indicators and intestinal morphology of yellow-feathered broiler chickens under heat stress[J]. Chinese Journal of Animal Nutrition, 2024, 36(5):2938-2947. (in Chinese)

Options
Outlines

/

〈 〉

京ICP备17050989号-2
Copyright © Chinese Journal of Animal Nutrition, All Rights Reserved.
Tel: 010－62817823　E-mail: cjan@vip.163.com
Powered by Beijing Magtech Co., Ltd.