动物营养学报 >

2025 , Vol. 37 >Issue 12: 8055 - 8066

DOI: https://doi.org/10.12418/CJAN2025.655

综述

中性粒细胞胞外陷阱在畜禽健康中的作用及其营养调控研究进展

  • 周佳 ,
  • 张丽 ,
  • 曾钰 ,
  • 董贤文 , *
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  • 重庆市畜牧科学院, 重庆 402460
* 董贤文,研究员,E-mail:

周 佳(1992—),男,湖北黄冈人,助理研究员,博士,主要从事反刍动物营养研究。E-mail:

收稿日期: 2025-05-30

  网络出版日期: 2025-12-13

基金资助

重庆市财政资金项目(25504C)

重庆市财政资金项目(24516C)

重庆市技术创新与应用发展专项面上项目(CSTB2024TIAD-LDX0006)

重庆市技术创新与应用发展专项面上项目(CSTB2025TIAD-GPX0002)

收起

Research Progress on Role of Neutrophil Extracellular Traps in Livestock and Poultry Health and Its Nutritional Regulation

  • ZHOU Jia ,
  • ZHANG Li ,
  • ZENG Yu ,
  • DONG Xianwen , *
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  • Chongqing Academy of Animal Sciences, Chongqing 402460, China
* professor, E-mail:

Received date: 2025-05-30

  Online published: 2025-12-13

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摘要

中性粒细胞胞外陷阱(NETs)作为中性粒细胞释放的DNA纤维网状结构,在病原防御中发挥双重角色:适度形成有助于捕获和清除病原体,而过度释放则通过释放蛋白酶、活性氧等效应分子加剧组织损伤和炎症反应。在畜禽生产中,多种因素如病原感染、代谢性疾病及生殖活动(如受孕)等均可诱发NETs的形成。近年来,利用营养策略调控NETs的形成逐渐受到关注。本文综述了NETs在畜禽健康中的生物学功能与作用机制,并探讨了营养干预策略调控NETs形成的应用前景,以期为防治NETs相关疾病和推动健康养殖实践提供理论依据。

关键词： 中性粒细胞胞外陷阱; 畜禽养殖; 免疫防御; 组织损伤; 营养调控

本文引用格式

周佳 , 张丽 , 曾钰 , 董贤文 . 中性粒细胞胞外陷阱在畜禽健康中的作用及其营养调控研究进展[J]. 动物营养学报, 2025 , 37(12) : 8055 -8066 . DOI: 10.12418/CJAN2025.655

Abstract

Neutrophil extracellular traps (NETs), which are DNA-based fibrous networks released by neutrophils, play a dual role in pathogen defense: moderate formation helps capture and eliminate pathogens, whereas excessive release exacerbates tissue damage and inflammatory responses by releasing effector molecules such as proteases and reactive oxygen species. In livestock and poultry production, various factors including pathogen infection, metabolic diseases, and reproductive activities (such as conception) can induce the formation of NETs. In recent years, there has been growing interest in utilizing nutritional strategies to regulate NETs formation. This review summarizes the biological functions and mechanisms of NETs in livestock and poultry health, and discusses the potential application of nutritional interventions in modulating NETs formation, aiming to provide a theoretical basis for preventing and treating NETs-related diseases and promoting healthy farming practices.

Key words： neutrophil extracellular traps; livestock and poultry farming; immune defense; tissue damage; nutrition regulation

在机体的免疫防御体系中,中性粒细胞作为先天性免疫的关键效应细胞,其经典抗菌功能主要包括吞噬作用和脱颗粒作用。2004年,Brinkmann等[1]在《Science》上首次报道了中性粒细胞的一种新型免疫防御机制:在病原体刺激下,中性粒细胞可通过一种独特的程序性死亡方式,释放由解聚染色质DNA、组蛋白以及多种抗菌蛋白[如髓过氧化物酶(myeloperoxidase,MPO)、中性粒细胞弹性蛋白酶(neutrophil elastase,NE)等]构成的纤维网状结构。该结构被命名为“中性粒细胞胞外诱捕网”(neutrophil extracellular traps,NETs),中文亦称“中性粒细胞胞外陷阱”。这一过程伴随特征性的细胞形态变化,既不同于细胞凋亡(表现为细胞皱缩和凋亡小体形成),也区别于坏死(以细胞膜破裂和内容物释放为特征)[2],后被学术界正式定义为一种炎性细胞死亡方式,即NETosis[3]。随着研究的不断深入,NETs的生物学功能已从最初的抗感染作用拓展至包括无菌性炎症、自身免疫性疾病、肿瘤转移及组织损伤等多种病理过程[4-5]。尽管目前关于NETs的形成机制及其病理效应的认识主要基于人类和模式动物的研究,但该现象在鸡[6]、猪[7]、羊[8]、牛[9]、骆驼[10]等畜禽动物中也已得到明确表型特征。畜禽动物在养殖过程中常面临病原感染、环境应激和营养代谢等多重免疫挑战。研究表明,NETs通过其既能捕获病原体又可介导炎症损伤的双重作用机制,在畜禽动物特异性免疫防御及相关病理性损伤中扮演关键角色[11-12]。近年研究发现,营养干预在体外可有效调控NETs的形成[13-14],为畜禽动物健康养殖提供了新的策略思路。本文就NETs在畜禽动物健康中的作用及营养调控的研究进展进行了综述,旨在为改善畜禽动物健康和生产性能提供参考。

1 NETs的形成机制及生物学特性

中性粒细胞在从病原体激活、组织迁移到免疫调节、微生物杀灭及NETs形成的整个免疫应答过程中表现出高度进化保守性[15]。中性粒细胞通过其表面模式识别受体(pattern recognition receptor,PRR)系统精确识别病原体相关分子模式(pathogen-associated molecular patterns,PAMPs)和损伤相关分子模式(damage-associated molecular patterns,DAMPs)。目前已明确的NETs形成机制可根据其对还原型烟酰胺腺嘌呤二核苷酸磷酸氧化酶(nicotinamide adenine dinucleotide phosphate oxidase, NOX)的依赖性分为NOX依赖性途径和非依赖性途径[16-17](图1)。在NOX依赖性途径中,病原体或炎性刺激可激活NOX,产生活性氧(reactive oxygen species,ROS),进而促使MPO和NE从颗粒中释放并进入细胞核,并通过肽酰精氨酸脱亚氨酶4(peptidylarginine deiminase 4,PAD4)介导的组蛋白瓜氨酸化导致染色质解聚和核膜破裂,最终释放NETs[18-19]。该过程中细胞内钙离子(Ca2+)浓度显著升高[20]。研究表明,NOX2介导产生的超氧阴离子及其衍生物过氧化氢在NETs形成的信号级联反应中发挥核心调控作用[21-22]。在ROS等效应分子的作用下,PAD4特异性催化组蛋白H3和H4的精氨酸残基发生瓜氨酸化修饰[23]。而非依赖性途径则通过Toll样受体(Toll-like receptors, TLR)感知信号,快速释放由氧化线粒体DNA构成的NETs[24-25]。NETs的释放方式根据刺激类型和强度可分为2种形式[26-27]:自杀型/溶解性NETosis以细胞膜完整性完全丧失、核膜破裂和细胞内容物彻底释放为特征,中性粒细胞死亡(图1-A);而活力型/非溶解性NETosis则能维持细胞核和细胞膜的完整性,通过囊泡运输途径选择性释放NETs成分(图1-B)。在NETs体外诱导模型中,佛波酯(phorbol 12-myristate 13-acetate,PMA)通过激活NOX2依赖性途径[28]、金黄色葡萄球菌通过PAMPs[8]以及脂多糖(lipopolysaccharide,LPS)通过TLR4信号通路[29],均可在适当条件下于2~4 h内高效诱导中性粒细胞形成NETs,为研究NETosis的分子机制和相关疾病治疗靶点提供了重要的体外模型。电子显微镜观察显示,成熟的NETs由直径15~17 nm的DNA纤维构成三维网状骨架,其上规律分布着直径约25 nm的球形蛋白聚集体,这种独特的网状结构可扩展至原细胞体积的15倍,在感染部位形成有效的物理屏障[30-31]。这些形成机制与结构特性是理解NETs在畜禽免疫防御与炎症损伤中的基础,也为通过营养策略靶向调控NETs以改善健康提供了关键理论依据。
图1 中性粒细胞胞外陷阱形成机制的示意图

A: 自杀型/溶解性NETosis suicidal/lytic NETosis; B:活力型/非溶解性NETosis vital/non-Lytic NETosis。Pathogens: 病原体;PMA: 佛波酯 phorbol 12-myristate 13-acetate; NADPH oxidase: 还原型烟酰胺腺嘌呤二核苷酸磷酸氧化酶 nicotinamide adenine dinucleotide phosphate oxidase; ROS: 活性氧 reactive oxygen species; NE: 中性粒细胞弹性蛋白酶 neutrophil elastase; MPO: 髓过氧化物酶 myeloperoxidase; PAD4: 肽酰精氨酸脱亚氨酶4 peptidylarginine deiminase 4; CitH3: 瓜氨酸组蛋白H3 citrullinated Histone H3; NETs: 中性粒细胞胞外陷阱 neutrophil extracellular traps; Neutrophil death: 中性粒细胞死亡; LPS: 脂多糖 lipopolysaccharide; E. coli: 大肠杆菌 Escherichia coli; S. aureus: 金黄色葡萄球菌 Staphylococcus aureus; C. albicans: 白色念球菌 Candida albicans; Activated PLTs: 活化的血小板 activated platelets; Neutrophil alive: 中性粒细胞存活。

Fig.1 Schematic illustration of formation mechanism of neutrophil extracellular traps[32]

2 畜禽生产中NETs形成的诱导因素

2.1 病原体感染

病原体感染是诱导畜禽动物体内NETs形成的主要因素之一,这一过程表现出明显的病原特异性。多种常见畜禽病原体(包括细菌、真菌、支原体、寄生虫和病毒)均可通过其特有的分子模式激活中性粒细胞,诱导NETs的生成[33]。细菌主要借助其表面结构成分激活中性粒细胞释放NETs。这些成分包括肽聚糖[1]、LPS[34]、脂磷壁酸[35]及鞭毛[36]等。真菌则通过细胞壁组分(如β-葡聚糖[37]、甘露聚糖[38])、信号分子(如法尼醇[39])及形态特征(如分生孢子和菌丝[40-41])诱导NETs形成。此外,寄生虫的细胞外囊泡[42]以及病毒与模式识别受体的相互作用[43-44]也被证实参与NETs的诱导过程。病原体的识别主要依赖于中性粒细胞表面的模式识别受体系统,包括TLR4[45-46]、核苷酸结合寡聚化结构域样受体(nucleotide-binding and oligomerization domain-like receptors,NLRs)[47]、C型凝集素受体(C-type lectin receptors,CLRs)[48]、补体受体(complement receptors,CRs)[49]以及蛋白酶激活受体2(protease-activated receptor 2,PAR2)[50]。不同病原体通过激活不同受体触发下游信号级联反应,其机制存在显著差异。研究表明,革兰氏阳性细菌(如金黄色葡萄球菌)通过协同激活CRs和TLR2受体,启动NOX2介导的ROS信号级联反应,促进导致染色质解聚和NETs释放[2,51];而革兰氏阴性菌(如大肠杆菌)则主要依赖TLR4途径诱导NETs形成[52]。通过查阅相关文献,本文总结了在畜禽动物特定感染模型中多种病原体诱导NETs形成的相关研究(表1)。
表1 畜禽动物模型中病原体诱导NETs形成的示例

Table 1 Examples of pathogen-induced NETs formation in livestock and poultry animal models

病原体类型
Pathogen type		 具体病原体
Specific pathogen		 宿主物种
Host species		 疾病或病理模型
Disease or
pathological mode		 文献
Reference
细菌
Bacteria		 乳房链球菌Streptococcus uberis 绵羊 乳腺炎 [11]
沃纳氏葡萄球菌Staphylococcus warneri 山羊 乳腺炎体外模型 [8]
无乳链球菌Streptococcus agalactiae 牛 乳腺炎 [53]
金黄色葡萄球菌Staphylococcus aureus 牛 乳腺炎 [54]
大肠杆菌Escherichia coli 牛 乳腺炎体外模型 [55]
耻垢分枝杆菌Mycobacterium smegmatis 牛 无 [56]
溶血性曼氏杆菌Mannheimia haemolytica 牛 体外感染模型 [57]
链球菌Streptococcus suis 猪 脑膜炎 [58]
胸膜肺炎放线杆菌Actinobacillus pleuropneumoniae 猪 肺炎 [59]
支原体
Mycoplasma		 无乳支原体Mycoplasma agalactiae 绵羊 乳腺炎 [60]
牛支原体Mycoplasma bovis 牛 体外感染模型 [61]
鸡败血支原体Mycoplasma gallisepticum 鸡 鸡毒支原体病 [62]
十二指肠贾第鞭毛虫Giardia duodenalis 山羊 体外感染模型 [63]
肝片吸虫Fasciola hepatica 绵羊 肝吸虫病 [64]
弓形虫Toxoplasma gondii 绵羊和牛 弓形虫感染体外模型 [65]
犬新孢子虫Neospora caninum 山羊、牛 体外感染模型 [66-67]
艾美耳球虫Eimeria arloingi 山羊、牛 体外感染模型 [68-69]
奥氏奥斯特线虫Ostertagia ostertagi 牛 感染体外模型 [9]
瘤胃吸虫Calicophoron daubneyi 牛 体外感染模型 [70]
伊氏锥虫Trypanosoma evansi 骆驼 体外感染模型 [10]
病毒
Virus		 呼吸道合胞病毒Respiratory syncytial virus 牛 合胞病毒感染 [71]
甲型流感病毒Influenza A viruses 猪 甲型流感病毒感染 [72]

2.2 代谢性疾病

目前,关于代谢性疾病诱导NETs形成的研究相对有限,且现有证据主要集中于反刍动物(主要是围产期奶牛),在单胃动物(如猪和家禽)中则鲜见相关报道。亚临床低血钙是高产奶牛在产犊后及泌乳初期常见的一种代谢性疾病,以血钙浓度持续低于正常生理水平(2 mmol/L)为主要特征。研究表明,亚临床低血钙奶牛可通过下调中性粒细胞内Ca2+浓度,导致线粒体发生损伤与ROS生成减少,最终抑制NETs的形成[73]。细胞内Ca2+浓度的变化是中性粒细胞激活的重要标志;在正常生理状态下,细胞受刺激后胞质Ca2+浓度会通过内质网等细胞内钙库的释放和/或细胞外Ca2+的内流而迅速升高[74]。然而,在亚临床低血钙奶牛中,由于钙释放激活钙调节蛋白1(calcium release-activated calcium modulator 1,ORAI1)基因的表达下调,中性粒细胞的钙摄取能力受损,导致其无法有效动员Ca2+,这可能是NETs形成受阻的重要原因[75]。酮病是围产期奶牛因能量代谢失衡所致的一种代谢性疾病,以血液中酮体(如β-羟丁酸、乙酰乙酸和丙酮)异常积累为主要表现,可显著增加奶牛罹患乳腺炎及其他传染病的风险[55]。体外研究表明,β-羟丁酸能够使大肠杆菌激活的奶牛中性粒细胞形成的NETs减少约10倍,并对其NETs相关杀菌活性产生类似程度的抑制[55]。另外,围产期奶牛由于脂质代谢活动加强而易发生能量负平衡,导致血液中游离脂肪酸(non-esterified fatty acids, NEFA)的浓度激增,而高浓度的NEFA会进一步参与机体氧化应激反应,并诱发失控的炎症反应[76]。NEFA通过激活核因子-κB(nuclear factor kappa-B,NF-κB)-NOD样受体热蛋白结构域相关蛋白3(NOD-like receptor thermal protein domain associated protein 3,NLRP3)-半胱天冬酶(cysteine-aspartic proteases,Caspase)1/4信号轴,促进Gasdermin D介导的孔道形成,同时伴随ROS含量升高,最终在体外诱导奶牛中性粒细胞释放NETs[77]。这些研究揭示了奶牛围产期代谢紊乱与NETs形成之间的复杂互作关系。

2.3 生殖受孕

在受孕过程中,精液在子宫内的沉积可激活中性粒细胞介导的免疫反应并诱导NETs的形成[78]。研究表明,牛精子与中性粒细胞在体外共培养15 min后即可观察到NETs的产生[79]。该过程依赖于NOX和PAD4的激活,并需要细胞内及细胞外Ca2+的协同流入[80]。值得注意的是,新鲜精液、冻融精液乃至单独的精浆均能诱导牛中性粒细胞形成NETs[78]。类似地,猪精子也可触发NETs形成,这不仅会损伤精子功能,还可导致NETs与胚胎发生纠缠[81],从而影响受孕结局。在生理状态下,精子诱导的NETs形成可能是一种进化上保守的防御机制,用于清除异常或缺陷精子[82]。然而,这一过程同时也会破坏精子质膜完整性、降低精子活力并削弱其受精能力,从而对繁殖效率产生负面影响[83]。另外,机体也形成相应的保护机制以维持生殖微环境的稳定。例如,牛输卵管可分泌多种活性因子[如前列腺素(E2)[84]、内皮素-1[85]和α1-酸性糖蛋白[86]],在保护精子免受NETs吞噬的同时,维持中性粒细胞对病原体的吞噬活性,从而有效遏制细菌传播[87],保障成功受孕。受孕过程中精液诱导NETs形成是影响动物繁殖效率的重要因素[88]。

2.4 其他诱导因素

霉菌毒素是畜禽养殖中常见的饲料污染物,其中伏马菌素B1在鸡体内抑制NETs形成[89],而在牛中性粒细胞中通过NOX依赖途径在体外诱导NETs产生[90];玉米赤霉烯酮则通过诱导线粒体ROS爆发,激活细胞外信号调节激酶(extracellular signal-regulated kinase,ERK)和p38信号通路,从而促进NETs释放[91]。重金属污染如铅暴露可通过抑制鸡内质网Ca2+回流减少NETs形成,铝污染则通过影响猪硒蛋白信号通路抑制溶酶体ROS爆发及NETs释放,而硒在一定程度上可拮抗铅和铝的抑制作用[92-93]。此外,微量元素如铜、锌和硒在体外均能显著诱导奶牛NETs形成[94]。某些抗生素如恩诺沙星可通过促进革兰氏阴性菌释放LPS,进而增强牛中性粒细胞NETs的形成[95]。这些研究表明,多种外源物质均可有效诱导NETs的形成,为理解其在畜禽免疫调控与健康中的作用提供了理论依据。

3 NETs在畜禽健康中的作用

3.1 病原体捕获和清除

NETs通过其特有的DNA-蛋白质复合网状结构,形成高效的物理捕获屏障,实现对病原体的快速固定和滞留。从结构基础来看,NETs由DNA纤维和球形蛋白组装形成的特征性球形功能区可物理性限制病原体运动,同时纤维上密集分布的抗菌蛋白(如MPO、NE等活性迅速提升)形成强效杀菌微环境[1,96]。在体外病原刺激下,经扫描电镜观察可见,牛中性粒细胞在受到十二指肠贾第鞭毛虫滋养体或犬新孢子虫速殖子体外刺激后,均能释放NETs,形成典型的网状纤维结构,并有效捕获与包裹病原体[67,97]。绵羊体内NETs也通过释放含抗菌蛋白的DNA复合物可高效捕获乳房链球菌[11]。有趣的是,牛、羊中性粒细胞在响应弓形虫速殖子时表现出显著的种属差异[65]:牛源中性粒细胞主要依赖高活性MPO实现对病原体的高效杀伤,而羊源中性粒细胞则主要通过形成NETs对虫体进行物理禁锢,限制其移动。类似地,阿氏艾美耳球虫可诱导山羊NETs形成并被其捕获,但NETs仅具物理束缚作用,无法显著影响虫体存活率[68]。值得注意的是,链球菌和支原体可分别通过分泌核酸酶A,降解猪源与牛源NETs的DNA骨架,进而逃逸NETs介导的免疫捕获作用[98-99]。金黄色葡萄球菌也可通过调控Ca2+通道,改变低血钙症奶牛外周血中性粒细胞内Ca2+浓度进而诱导自噬体形成并阻断自噬通量,最终实现免疫逃逸[99]。因此,NETs通过捕获病原体为宿主构筑先天防御屏障,但其效能常被病原体的逃逸机制所削弱,而未来针对逃逸因子的靶向中和策略或NETs稳定性的增强手段,有望成为防控畜禽疫病的新方向。

3.2 加剧组织损伤

NETs在免疫防御中发挥重要作用,但其过度或持续存在可能导致细胞和组织的炎症损伤。过量的NETs通过其细胞毒性组分(如组蛋白和NE)破坏细胞膜完整性及细胞间基质蛋白,导致内皮细胞和上皮细胞的连接结构损伤,从而加剧组织病理损伤[100-101]。在以猪为损伤模型的研究中发现,缺血再灌注通过诱导NETs浸润及PAD释放,催化纤维蛋白原等血浆蛋白瓜氨酸化,加重组织损伤[102];而清除NETs可改善吸入性肺损伤供肺的功能与形态,表明靶向NETs具有治疗潜力[103]。在畜禽健康方面,关于NETs介导的炎性损伤研究主要聚焦于奶牛乳腺炎模型,特别是NETs对乳腺上皮细胞的损伤机制及其分子通路[104-105]。NETs的主要组成成分组蛋白通过激活Caspase-1、NLRP3和Caspase-3等凋亡蛋白或炎症小体,诱导乳腺上皮细胞发生焦亡、凋亡和坏死,从而损伤乳腺上皮屏障[106]。NETs在奶牛子宫内膜炎发病过程中通过高迁移率族蛋白B1(high mobility group protein,HMGB1)/NLRP3/Caspase-1信号通路诱导子宫内膜上皮细胞焦亡,导致组织炎症和损伤;而抑制NETs形成可显著缓解上述病理变化[107]。此外,研究还发现,NETs的形成是母体营养不良诱发宫内生长发育受限绵羊胎儿肝脏损伤的一个新机制[12]。这些发现表明,NETs作为重要的免疫效应机制,在其调控失衡时会转化为关键致病介质,加剧组织损伤进程。

4 营养调控在畜禽NETs形成中的作用

营养调控对畜禽NETs的形成表现出双向调节作用。一方面,槲皮素和山奈酚可以降低ROS的形成,从而抑制NETs的形成并缓解其对山羊精子的损伤[108-109];油酸[110]和丁酸钠[14]则分别在体外通过抑制自噬通量和中性粒细胞自噬来减少奶牛中性粒细胞形成NETs。另一方面,在抗感染过程中,槲皮素可通过增强ROS介导的杀菌功能促进NETs释放[111];女贞子活性物质(齐墩果酸和红景天苷)[112]及烟酸[13]也可通过不同机制在体外诱导奶牛中性粒细胞形成NETs,增强病原体清除能力。这表明营养对NETs的调控具有多样性,既可通过抗氧化和抑制自噬等途径抑制过度炎症损伤,也可在免疫防御中增强NETs的抗菌功能。目前,关于营养调控在畜禽NETs中的作用研究仍处于初步阶段,相关报道多局限于体外试验,亟待更多体内研究进一步验证其调控机制与应用潜力。

5 小结

NETs作为先天免疫防御的重要组成部分,其适度形成对抵抗病原微生物感染至关重要,而过度激活则可能加剧组织炎症和损伤。在畜禽动物中,NETs最初被认为参与对抗各类病原体的免疫反应,而近年研究进一步揭示其与乳腺炎、子宫内膜炎等疾病相关的组织损伤机制密切相关。目前,一些植物提取物和脂肪酸等营养活性物质已在体外被证实可调控NETs的形成,但相关研究仍存在分子机制解析不足、NETs动态平衡调控研究缺乏以及实际生产应用效果待验证等局限。值得注意的是,过度抑制NETs可能削弱先天免疫防御功能,反而增加感染风险,因此在养殖生产中需权衡其调控策略。未来,深入探究营养干预对NETs形成与功能的调控机制,不仅可为自身免疫性疾病的防治提供新思路,也将为畜禽健康养殖提供理论依据。然而,NETs作为一种新兴的免疫机制,其调控网络及生理病理意义仍有待进一步阐明,相关研究尚处于起步阶段,需更多系统性探索以推动其在畜牧及兽医领域的应用。

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