Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 6: 4188 - 4196

DOI: https://doi.org/10.12418/CJAN2026.336

RESEARCH PAPER

Effects of Adding Thymol-Carvacrol Cocrystal in Drinking Water on Lipid Metabolism of Laying Hens in Late Laying Period

  • KONG Linglian , 1, 2 ,
  • ZHANG Xinran 2 ,
  • ZHANG Guoxin 3 ,
  • ZHANG Qi 4 ,
  • SONG Zhigang , 2, *
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  • 1 Jining Polytechnic College, Jining 272037, China
  • 2 College of Animal Science and Technology, Shandong Agricultural University, Tai’an 271018, China
  • 3 Department of Medical Laboratory Zoology, Hebei Medical University, Shijiazhuang 050017, China
  • 4 Cocrystal Health Industry (Zhejiang) Co., Ltd., Jiaxing 314109, China
* professor, E-mail:

Received date: 2025-11-07

  Online published: 2026-06-13

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Abstract

This study aimed to investigate the effects of adding thymol-carvacrol cocrystal in drinking water on lipid metabolism of laying hens in late laying period. Firstly, a total of 96 sixty-week-old Hy-Line brown laying hens were randomly assigned into 2 groups with 6 replicates per group and 8 hens per replicate. Hens in control group were given normal drinking water, while those in experimental group received drinking water added with 30 mg/L thymol-carvacrol cocrystal. The experiment lasted for 4 weeks following a 1-week pre-feeding period. Serum and hepatic lipid metabolism indicators in were measured at the end of the trial. Secondly, network pharmacology was employed to screen key targets of thymol-carvacrol cocrystal in regulating lipid metabolism in laying hens, their binding characteristics were validated by molecular docking technology, and the core pathways were identified through pathway enrichment analysis. The results showed as follows: 1) at end of week 4, compared with the control group, the experimental group had significantly lower serum triglyceride and total cholesterol contents (P<0.05), as well as significantly lower liver steatosis score and hepatic triglyceride content (P<0.05). 2) Prostaglandin-endoperoxide synthase 2 (PTGS2) and albumin (ALB) were identified as the core targets through which the thymol-carvacrol cocrystal regulated lipid metabolism in laying hens. 3) Molecular docking revealed that the binding energies of the main active components of the thymol-carvacrol cocrystal with PTGS2 and ALB were all below -20.92 kJ/mol. 4) Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway enrichment analysis identified 8 related signaling pathways, among which the calcium signaling pathway was the most significantly enriched. In conclusion, thymol-carvacrol cocrystal may improve lipid metabolism of laying hens in late laying period by acting on core targets such as PTGS2 and ALB, and by regulating the calcium signaling pathway.

Key words： thymol; carvacrol; laying hens; lipid metabolism; network pharmacology

Cite this article

KONG Linglian , ZHANG Xinran , ZHANG Guoxin , ZHANG Qi , SONG Zhigang . Effects of Adding Thymol-Carvacrol Cocrystal in Drinking Water on Lipid Metabolism of Laying Hens in Late Laying Period[J]. Chinese Journal of Animal Nutrition, 2026 , 38(6) : 4188 -4196 . DOI: 10.12418/CJAN2026.336

蛋鸡脂肪肝出血综合征(fatty liver hemorrhagic syndrome,FLHS)严重危害禽类健康,导致产蛋性能下降[1-2]。目前,功能性添加剂已用于改善FLHS相关肝脏及血液指标,为预防蛋鸡FLHS发生提供了新策略[3]。植物精油是从植物中提取的挥发性芳香化合物,具有抗氧化、抗炎及促生长等多种生理功能。研究表明,产蛋后期蛋鸡饲粮中添加复合精油可调节肠道脂质吸收,改善脂质代谢[4]。然而,精油的高挥发性、低水溶性及化学性质不稳定等特性,限制了其在家禽生产中的广泛应用。共晶技术通过构建分子间非共价键结合的稳定结构,可显著改善活性成分的水溶性、稳定性及生物利用度[5]。近年来,该技术在家禽营养领域的研究日益深入,展现出解决实际生产问题的潜力。研究表明,百里香酚-香芹酚共晶以剂量依赖性方式显著降低42日龄肉鸡血清甘油三酯(triglycerides,TG)含量,并减轻肠道脂质过氧化[6];大豆苷元-甜菜碱共晶则可缓解高能低蛋白质饲粮诱导的蛋鸡肝脏脂肪过度沉积[7]。上述研究主要基于饲粮添加途径。考虑到家禽的饮水量通常为采食量的1.75~2.00倍[8],开发基于共晶技术、适用于饮水给药的功能性添加剂,对改善蛋鸡健康养殖具有重要意义。
植物精油成分复杂,具有多靶点、多通路协同作用的特征,传统研究方法难以系统揭示其整体作用机制。网络药理学作为系统生物学与多向药理学交叉的新兴学科,能够从分子层面系统阐释活性成分与疾病相关靶点间的复杂相互作用关系,这与中药及植物提取物多成分协同作用的特点高度吻合。分子对接技术可通过计算活性成分与核心靶点间的结合能,预测二者结合亲和力及作用模式,为网络药理学筛选结果提供初步验证。本研究通过动物试验结合网络药理学与分子对接技术,探究饮水添加百里香酚-香芹酚共晶对产蛋后期蛋鸡脂质代谢的影响,并解析其潜在关键靶点及作用通路,以期为该制剂在蛋鸡健康养殖中的应用提供理论依据。

1 材料与方法

1.1 试验材料

百里香酚-香芹酚共晶由等比例的百里香酚共晶和香芹酚共晶混合而成,由共晶科技(嘉兴)有限责任公司提供。其中,百里香酚共晶由百里香酚与L-脯氨酸按摩尔比1∶1制备;香芹酚共晶
由香芹酚与L-脯氨酸按摩尔比1∶1制备。百里香酚和香芹酚均提取自牛至精油,有效含量均为25%,其余组分为碳酸钙载体。

1.2 动物试验

1.2.1 试验设计与饲养管理

本次动物试验在山东农业大学畜牧科技试验站完成,并获山东农业大学动物实验伦理审查委员会批准(批准号:2001002)。
选取96只60周龄海兰褐蛋鸡,随机分为2组,每组设6个重复,每个重复8只鸡。对照组正常饮水,试验组在饮水中添加30 mg/L百里香酚-香芹酚共晶(按产品推荐浓度添加至饮水)。预试期1周,正试期4周。
试验鸡饲养于双层A式蛋鸡笼中,自由采食与饮水,各组饲喂相同的基础饲粮,饲粮组成及营养水平见表1。饮水采用饮水桶单笼供应,每日定时更换,饮水量参考《海兰褐在线管理手册(国际标准)》设定,即60~65周龄蛋鸡每日每只165~236 mL。
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 61.00 代谢能 ME/(MJ/kg) 11.29
豆粕 Soybean meal 24.00 粗蛋白质 CP 16.50
豆油 Soybean oil 1.00 钙 Ca 3.50
石粉 Limestone 9.00 总磷 TP 0.58
预混料 Premix1) 5.00 有效磷 AP 0.39
合计 Total 100.00 赖氨酸 Lys 0.89
蛋氨酸 Met 0.40

1)预混料为每千克饲粮提供 Premix provided the following per kg of the diet:VA 12 500 IU,VB1 1 mg,VB2 8.5 mg,VB6 8 mg,VB12 0.02 mg,VD3 4 000 IU,VE 28 mg,VK3 2 mg,生物素 biotin 0.2 mg,叶酸 folic acid 2.2 mg,D-泛酸 D-pantothenic acid 50 mg;烟酸 nicotinic acid 32.5 mg,Cu (as copper sulfate) 10 mg,Fe (as ferrous sulfate) 20 mg,Mn (as manganese sulfate) 80 mg,Zn (as zinc sulfate) 80 mg,I (as potassium iodide) 0.30 mg,氯化胆碱 choline chloride 500 mg,NaCl 3 000 mg,DL-蛋氨酸 DL-Met 1 400 mg,L-赖氨酸盐酸盐 L-Lys·HCl 2 000 mg,植酸酶 phytase 250 U,玉米芯粉 corncob powder 43 800 mg。
2)代谢能、有效磷和氨基酸为计算值,依据《中国饲料成分及营养价值表(2021年第32版)》计算;粗蛋白质、钙和总磷为实测值,分别参照GB/T 6432—2018、GB/T 6436—2018和GB/T 6437—2018进行测定。ME, AP and AA were measured values, calculated with reference to the Tables of Feed Composition and Nutritive Values in China (32nd edition, 2021); CP, Ca and TP were measured values, determined in accordance with GB/T 6432—2018, GB/T 6436—2018 and GB/T 6437—2018, respectively.

1.2.2 指标测定与方法

1.2.2.1 血清脂质代谢指标

在试验第2周和第4周末,每组随机取12只蛋鸡(n=12)进行翅下静脉采血,采集的血液在室温静置30 min,4 ℃、3 000×g条件下离心10 min分离血清,保存于-20 ℃。使用全自动生化分析仪(HITACHI-7170,日立,日本)检测血清TG、总胆固醇(total cholesterol,TC)、高密度脂蛋白胆固醇(high density lipoprotein cholesterol,HDL-C)和低密度脂蛋白胆固醇(low density lipoprotein cholesterol,LDL-C)含量。

1.2.2.2 肝脏病理学切片和脂肪变性评分

在试验第4周末,每个重复随机取2只蛋鸡处死,采集肝脏组织样品,用4%多聚甲醛溶液固定24 h,经脱水处理后石蜡包埋,切片厚度为4 μm,并进行苏木精-伊红(HE)染色。染色切片在显微镜(Nikon Eclipse 80i,尼康,日本)下观察,以200×倍率拍摄图像。参照美国肝病协会提出的非酒精性脂肪性肝病(nonalcoholic fatty liver disease activity score,NAS)评分系统进行脂肪变性评分[9]。具体方法:每张肝脏切片随机选取5个视野,计算肝细胞脂肪变性的平均百分比。评分标准为:脂肪变性比例<5%计0分,5%~33%计1分,>33%~66%计2分,>66%计3分。

1.2.2.3 肝脏TG指标

取适量肝脏样品,按质量体积比加入生理盐水制备成10%组织匀浆,于4 ℃、1 000×g条件下离心10 min,收集上清液,采用比色法测定TG含量。使用BCA法检测肝脏组织匀浆上清液中的蛋白质含量,并使用蛋白质浓度对TG含量的结果进行归一化。

1.3 网络药理学分析

以“thymol”“carvacrol”和“L-proline”为关键词在中药系统药理学数据库(traditional Chinese medicine systems pharmacology database and analysis platform,TCMSP)(https://www.tcmsp-e.com/tcmsp.php)和SwissTargetPrediction数据库(http://swisstargetprediction.ch)中检索百里香酚-香芹酚共晶的作用靶点,合并后去除重复值。使用UniProt数据库(https://www.uniprot.org)将上述靶点名称标准化为基因名称,设置物种为“Gallus gallus”,仅保留经UniprotKB/Swiss-Prot审查的条目。以“lipid metabolism”为关键词在GeneCards数据库(https://www.genecards.org)和人类孟德尔遗传数据库(online mendelian inheritance in man,OMIM)(https://www.omim.org)中检索脂质代谢相关靶标。将上述靶点进行交集分析,利用微生信在线生物信息学分析与可视化云平台(https://www.bioinformatics.com.cn)绘制韦恩图,确定交集靶点,导入STRING数据库(https://cn.string-db.org)进行蛋白质-蛋白质相互作用(protein-protein interaction,PPI)网络分析,物种设置为“Gallus gallus”,置信度分数设置为0.4,使用Cytoscape 3.10.2中的CentiScaPe-2.2插件分析接近中心性(closeness centrality)、度中心性(degree centrality)和介数中心性(betweenness centrality),选择3项中心性值均高于均值的靶点作为核心靶点。使用DAVID数据库(https://davidbioinformatics.nih.gov)对交集靶点进行基因本体(gene ontology,GO)功能富集分析以及京都基因与基因组百科全书数据库(Kyoto encyclopedia of genes and genomes,KEGG)通路富集分析,设置物种为“Gallus gallus”,以P<0.01为阈值确定显著富集的GO条目和KEGG通路。

1.4 分子对接验证

在AlphaFold蛋白结构数据库(https://alphafold.com)中检索获得核心靶点的3D结构文件,同时在PubChem数据库(https://pubchem.ncbi.nlm.nih.gov)下载百里香酚、香芹酚和L-脯氨酸的3D结构文件。使用AutoDock Tools 1.5.6软件以PDBQT格式制备受体和配体文件,设置对接盒子的尺寸能够充分覆盖整个蛋白结构域,使用AutoDock Vina 1.1.2软件进行分子对接和结合能计算。最后通过PyMOL 3.1.0软件对分子对接结果进行分析和可视化处理,根据结合能大小和氢键数量选择最佳结合构象。

1.5 数据统计分析

试验数据使用GraphPad Prism 8.0.2软件进行统计分析并绘图,采用双尾学生t检验比较差异显著性。结果数据以平均值±标准误表示,P<0.05为差异显著。

2 结果

2.1 饮水中添加百里香酚-香芹酚共晶对产蛋后期蛋鸡血清和肝脏脂质代谢指标的影响

由表2可知,第2周末,饮水中添加百里香酚-香芹酚共晶对产蛋后期蛋鸡血清TG、TC、HDL-C和LDL-C含量均无显著影响(P>0.05)。第4周末,与对照组相比,试验组蛋鸡血清TG和TC含量显著降低(P<0.05),血清HDL-C和LDL-C含量无显著变化(P>0.05);试验组蛋鸡肝脏TG含量和脂肪变性评分显著降低(P<0.05)。
表2 饮水中添加百里香酚-香芹酚共晶对产蛋后期蛋鸡血清和肝脏脂质代谢指标的影响

Table 2 Effects of adding thymol-carvacrol cocrystal in drinking water on serum and hepatic lipid metabolism indicators of laying hens in late laying period

项目
Items		 时间
Time		 对照组
Control group		 试验组
Experimental group		 P值
P-value
血清 Serum/(mmol/L)

甘油三酯 TG		 第2周末 End of week 2 14.37±5.56 13.06±4.59 0.605
第4周末 End of week 4 17.67±4.78 13.54±4.33 0.042

总胆固醇 TC		 第2周末 End of week 2 3.19±1.27 3.15±0.94 0.930
第4周末 End of week 4 3.52±0.56 2.72±0.59 0.003

高密度脂蛋白胆固醇 HDL-C		 第2周末 End of week 2 0.77±0.21 0.88±0.22 0.236
第4周末 End of week 4 0.80±0.13 0.82±0.32 0.826

低密度脂蛋白胆固醇 LDL-C		 第2周末 End of week 2 0.89±0.16 0.83±0.21 0.714
第4周末 End of week 4 0.82±0.18 0.69±0.16 0.092
肝脏 Liver
甘油三酯 TG/(mmol/mg prot) 第4周末 End of week 4 13.37±1.83 11.70±1.13 0.023
脂肪变性评分 Steatosis score/分 第4周末 End of week 4 2.33±0.49 1.83±0.57 0.032

2.2 PPI网络构建及核心靶点筛选

如图1所示,去除未连接的节点后,得到由20个节点和24条边组成的PPI网络图。以介数中心性值≥32.20,接近中心性值≥0.02,度中心性值≥2.40进行筛选,得到核心靶点2个,即前列腺素内过氧化物合酶2(prostaglandin-endoperoxide synthase 2,PTGS2)和白蛋白(albumin,ALB)。
图1 百里香酚-香芹酚共晶调控蛋鸡脂质代谢的核心靶点

A:百里香酚-香芹酚共晶-脂质代谢靶点PPI网络图 PPI network diagram of targets between thymol-carvacrol cocrystal and lipid metabolism;B:核心靶点 core targets。

ACE:血管紧张素转换酶 angiotensin converting enzyme;ACHE:乙酰胆碱酯酶 acetylcholinesterase; ALB:白蛋白 albumin;ANPEP:丙氨酰氨基肽酶 alanyl aminopeptidase;AR:雄激素受体 androgen receptor;AURKA:极光激酶A aurora kinase A;BCHE:丁酰胆碱酯酶 butyrylcholinesterase;CCNA2:细胞周期蛋白A2 cyclin A2;CDK2:细胞周期蛋白依赖性激酶2 cyclin dependent kinase 2;CHRM3:胆碱能受体毒蕈碱3 cholinergic receptor muscarinic 3;CTSD:组织蛋白酶D cathepsin D;CYP19A1:细胞色素P450家族19亚家族A成员1 cytochrome P450 family 19 subfamily A member 1;F2:凝血因子Ⅱ coagulation factor Ⅱ;JAK:两面神激酶 Janus kinase;LYZ:溶菌酶 lysozyme;PTGS:前列腺素内过氧化物合酶 prostaglandin-endoperoxide synthase;REN:肾素 renin;TYR:酪氨酸酶 tyrosinase。

Fig.1 Core targets of thymol-carvacrol cocrystal in the regulation of lipid metabolism in laying hens

2.3 GO功能富集分析和KEGG通路富集分析

如图2所示,通过GO功能富集分析得到,显著富集的生物学过程(biological processes,BP)主要涉及腺苷酸环化酶激活肾上腺素能受体信号通路、丝裂原活化蛋白激酶(MAPK)级联反应的正向调节、蛋白质水解、蛋白质翻译后修饰、血压调节以及G蛋白偶联受体信号通路过程;细胞成分(cellular components,CC)主要与细胞质、质膜、细胞外间隙、突触、胞膜小窝和投射神经元有关;分子功能(molecular functions,MF)主要与金属离子结合、血红素结合、锌离子结合、α1-肾上腺素能受体活性和金属肽酶活性有关。
图2 GO功能富集分析

Fig.2 GO functional enrichment analysis

如图3所示,通过KEGG富集分析,共获得8个相关信号通路,包括钙信号通路、神经活性配体-受体相互作用、激素信号通路、心肌细胞肾上腺素能信号通路、精氨酸和脯氨酸代谢、血管平滑肌收缩、花生四烯酸代谢和血管内皮生长因子(vascular endothelial growth factor,VEGF)信号通路。
图3 KEGG通路富集分析

Fig.3 KEGG pathway enrichment analysis

2.4 百里香酚-香芹酚共晶主要成分与核心靶点的分子对接

如图4-A所示,PTGS2与百里香酚、香芹酚和L-脯氨酸之间的结合能分别为-25.95、-26.31和-19.19 kJ/mol,ALB与百里香酚、香芹酚和L-脯氨酸之间的结合能分别为-23.12、-26.23和-18.87 kJ/mol。如图4-B和图4-C所示,百里香酚与PTGS2的THR-185残基以及与ALB的GLN-243残基各形成1个氢键,键长分别为1.9和2.5 Å。如图4-D和图4-E所示,香芹酚与PTGS2残基HIS-374以及与ALB残基MET-458各形成1个氢键,键长分别为2.5和2.0 Å。
图4 百里香酚-香芹酚共晶的主要成分与核心靶点的分子对接

A:百里香酚-香芹酚共晶的主要成分与核心靶点的结合能 binding energy of the primary components from thymol-carvacrol cocrystal and core targets;B:PTGS2-thymol PTGS2-百里香酚 prostaglandin-endoperoxide synthase 2-thymol;C:ALB-thymol ALB-百里香酚 albumin-thymol;D:PTGS2-carvacrol PTGS2-香芹酚 prostaglandin-endoperoxide synthase 2-carvacrol;E:ALB-carvacrol ALB-香芹酚 albumin-carvacrol。PTGS2:前列腺素内过氧化物合酶2 prostaglandin-endoperoxide synthase 2;ALB:白蛋白 albumin。

Fig.4 Molecular docking between the primary components from thymol-carvacrol cocrystal and core targets

3 讨论

肝脏是蛋鸡脂质代谢的主要场所,合成机体内超过90%的脂肪酸[10]。肝脏脂肪过多可能引发蛋鸡脂质代谢障碍和FLHS。因此,有效调节蛋鸡的脂质代谢对于延长其高产寿命和保持健康至关重要[11]。血清和肝脏TG含量异常升高是肝脏脂肪变性的关键病理标志之一[12]。Çimrin等[13]在48周龄蛋鸡饲粮中补充1 000 mg/kg含百里香酚和香芹酚的植物提取物,饲养8周后发现,试验组蛋鸡血清TG和TC含量显著降低。本研究结果与之吻合,通过饮水中添加百里香酚-香芹酚共晶4周,同样显著降低了产蛋后期蛋鸡血清TG和TC含量。此外,通过饮水中添加百里香酚-香芹酚共晶4周,显著降低产蛋后期蛋鸡肝脏中的TG含量,减轻肝脏脂肪变性评分,提示该共晶对调节蛋鸡肝脏脂质代谢具有积极作用。
网络药理学作为系统生物学与多向药理学交叉的新兴学科,聚焦于构建“药物-靶点-通路”多维网络,系统揭示复杂生物系统内各要素间的内在联系[14]。为最大限度减少物种间基因功能差异可能导致的推断偏差,本研究在进行网络药理学靶点预测时,通过数据库将预测物种明确限定为“Gallus gallus”,确保所筛选出的核心靶点及其富集通路,均基于鸡的基因组注释信息,从而增强从生物信息学分析到蛋鸡体内应用的逻辑相关性与预测的可靠性。本研究利用网络药理学分析发现,PTGS2和ALB是百里香酚-香芹酚共晶调节蛋鸡脂质代谢的关键靶点。PTGS2编码环氧合酶-2蛋白,通过前列腺素E2介导的旁分泌机制抑制脂肪组织扩张,并防止小鼠肥胖和代谢紊乱的发展[15-16]。ALB是一种肝脏产生的血浆蛋白,ALB基因缺失显著降低高脂饮食小鼠血浆游离脂肪酸浓度,口服ALB则能减少高脂饮食小鼠的肝脏中性脂质的积累和脂肪变性[17-18]。分子对接结果显示,百里香酚和香芹酚均对PTGS2和ALB表现出较强的结合亲和力,并能在各自的结合口袋内形成稳定的氢键。相比之下,百里香酚-香芹酚共晶中的L-脯氨酸对核心靶点的结合亲和力较低,提示百里香酚和香芹酚是该共晶发挥调节蛋鸡脂质代谢作用的主要活性成分。本研究重点关注了百里香酚与香芹酚的核心作用,而L-脯氨酸在共晶中的角色亦不可忽视。除作为共晶载体改善活性成分的理化性质,L-脯氨酸自身也可能参与代谢调节。研究表明,脯氨酸作为一种功能性氨基酸,本身具有缓解氧化应激、参与能量代谢等广泛的生理功能[19]。尽管本研究的分子对接结果显示L-脯氨酸与核心靶点PTGS2和ALB的直接结合亲和力较弱,但其作为代谢前体或通过调节整体蛋白质合成[20],为百里香酚和香芹酚发挥核心药理作用提供了更有利的代谢内环境,从而在改善脂质代谢中起到协同或辅助调节作用。
钙信号可通过增强脂肪细胞的新陈代谢和分化增加能量消耗,并通过调节神经元兴奋性减少食物摄入,从而有效减少肥胖的发生[21]。其中,MAPK信号通路是一种典型的钙信号通路;脂肪细胞中MAPK信号通路的激活可以促进细胞内钙转运,调节脂肪细胞代谢[22-24]。研究证实,包括百里香酚和香芹酚在内的多种精油成分可靶向MAPK信号通路发挥抗炎作用[25-26]。在本研究中,交集靶点在MAPK级联反应的正向调节和钙信号通路中显著富集,提示钙信号通路可能参与了百里香酚-香芹酚共晶对蛋鸡脂质代谢的调控。以上结果表明,百里香酚-香芹酚共晶可能主要通过作用于PTGS2和ALB这2个关键靶点,并通过调控钙信号通路影响脂质代谢,从而改善蛋鸡FLHS。

4 结论

综上所述,饮水中添加30 mg/L百里香酚-香芹酚共晶可能通过调控PTGS2、ALB等关键靶点及钙信号通路,显著降低血清TG和TC含量,并减轻肝脏脂肪变性程度及降低肝脏TG含量,进而改善产蛋后期蛋鸡的脂质代谢。

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