动物营养学报 >

2026 , Vol. 38 >Issue 5: 3324 - 3332

DOI: https://doi.org/10.12418/CJAN2026.266

综述

酒糟血管紧张素转化酶抑制肽调控动物抗氧化和抗炎作用及其高效制备技术研究进展

  • 张彩莹 , 1 ,
  • 张董燕 2 ,
  • 赵涵 1, 3 ,
  • 刘明 , 1, *
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  • 1 北京农学院, 蛋白质高效合成与生物智造北京市重点实验室, 北京 102206
  • 2 北京市农林科学院畜牧兽医研究所, 北京 100097
  • 3 河北弘科荣达生物技术有限公司, 河北 057650
*刘 明,教授,博士生导师,E-mail:

张彩莹(2000—),女,河北衡水人,硕士研究生,研究方向为非粮饲料资源开发与利用。E-mail:

Office editor: 武海龙

收稿日期: 2025-10-29

  网络出版日期: 2026-05-14

基金资助

北京农学院人才强教工程项目(5066516008-6)

河北省省级科技计划资助(2022HBQZYCXY016)

收起

Advances in Regulation of Antioxidant and Anti-Inflammatory Effects in Animals of Angiotensin-Converting Enzyme Inhibitory Peptides from Distillers’ Grains and Its Efficient Preparation Technologies

  • ZHANG Caiying , 1 ,
  • ZHANG Dongyan 2 ,
  • ZHAO Han 1, 3 ,
  • LIU Ming , 1, *
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  • 1 Beijing Key Laboratory of Efficient Protein Synthesis and Intelligent Biomanufacturing, Beijing University of Agriculture, Beijing 102206, China
  • 2 Institute of Animal Science and Veterinary Medicine, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, China
  • 3 Hebei Hongke Rongda Bio-Technology Company Limited, Hebei 057650, China
*professor, E-mail:

Received date: 2025-10-29

  Online published: 2026-05-14

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摘要

酒糟作为酿酒工业副产物,其富含蛋白质及多种生物活性成分,经酶解或微生物发酵等处理可释放血管紧张素转化酶抑制肽(ACEi)。ACEi通过抑制肾素-血管紧张素-醛固酮系统(RAAS)关键酶,减少血管紧张素Ⅱ(AngⅡ)生成,阻断“氧化应激-血管紧张素转化酶(ACE)-AngⅡ”恶性循环和AngⅡ介导的线粒体氧化损伤通路,以及阻断AngⅡ触发的核因子-κB(NF-κB)信号通路,并通过介导肿瘤坏死因子(TNF)、晚期糖基化终产物受体(RAGE)相关通路,减少炎症因子分泌,进而调控氧化应激和炎症反应,在缓解畜禽氧化应激方面展现出良好的应用潜力。为此,本文对不同酒糟ACEi及其制备方法、ACEi调控动物抗氧化应激和抗炎作用进行了综述,并进一步从酶解、微生物发酵及DNA重组角度分析了ACEi高效制备技术,以期为促进酒糟ACEi在动物生产中的应用及开发新型绿色饲料添加剂提供参考。

关键词： 酒糟; 氧化应激; 抗炎; 血管紧张素转化酶抑制肽

本文引用格式

张彩莹 , 张董燕 , 赵涵 , 刘明 . 酒糟血管紧张素转化酶抑制肽调控动物抗氧化和抗炎作用及其高效制备技术研究进展[J]. 动物营养学报, 2026 , 38(5) : 3324 -3332 . DOI: 10.12418/CJAN2026.266

Abstract

Distiller’s grains, a by-product of the brewing industry, are rich in protein and various bioactive components, through processing methods such as enzymatic hydrolysis or microbial fermentation, the angiotensin-converting enzyme inhibitory peptides (ACEi) can be released. The ACEi exert their effects by inhibiting a key enzyme in the renin-angiotensin-aldosterone system (RAAS), thereby reducing angiotensin Ⅱ (AngⅡ) generation, disrupts the vicious cycle of “oxidative stress-angiotensin-converting enzyme (ACE)-AngⅡ” and AngⅡ-mediated mitochondrial oxidative damage pathways, and disrupts the AngⅡ-triggered nuclear factor-κB (NF-κB) signaling pathway. Moreover, ACEi modulate tumor necrosis factor (TNF) and receptor of advanced glycation end products (RAGE) related pathways, reducing secretion of inflammatory factors and ultimately alleviating oxidative stress and inflammatory responses, which shows significant potential in mitigating oxidative stress in livestock and poultry. Therefore, this review summarizes ACEi from various distiller’s grain and their preparation methods, and elaborates on the role of ACEi in regulating antioxidant and anti-inflammatory responses in animals, furthermore, efficient preparation technologies for ACEi, including enzymatic hydrolysis, microbial fermentation and DNA recombinant technology, are analyzed. The aim of this review is to provide a theoretical foundation for promoting the application of ACEi from distiller’s grains in animal production and to support the development of novel green feed additives.

Key words： distiller’s grains; oxidative stress; anti-inflammatory; angiotensin-converting enzyme inhibitory peptides

血管紧张素转化酶抑制肽(angiotensin-converting enzyme inhibitory peptides,ACEi)是具有抑制血管紧张素转化酶(angiotensin-converting enzyme,ACE)活性的一种生物活性肽,能调节肾素-血管紧张素-醛固酮系统(renin-angiotensin-aldosterone system,RAAS),减少血管紧张素Ⅱ(angiotensin Ⅱ,AngⅡ)生成,从而降低血管收缩,以及缓解氧化应激、炎症和免疫反应等,维持机体稳态[1-3]。在现代畜禽规模化养殖中,氧化应激是是制约动物健康福利与生产性能的重要因素之一,应激状态会导致畜禽体内活性氧(reactive oxygen species,ROS)积累与抗氧化系统失衡,进而造成细胞结构损伤及功能下降,最终导致生产性能下降和养殖效益受损[4-5]。随着抗生素减量化和功能性饲料添加剂需求的增长,天然生物活性物质的开发成为动物营养学研究热点[6],葡萄酒糟中ACEi被证实能提升肝脏还原型谷胱甘肽含量及增加血浆一氧化氮代谢物含量,间接缓解RAAS过度激活引发的血管收缩与组织损伤以及氧化应激等[7],但目前主要在大鼠中有相关研究报道。在畜禽生产中,氧化应激条件下同样存在RAAS异常激活现象[8],而使用酒糟ACEi缓解动物应激的研究与应用极少。基于此,本文围绕酒糟ACEi及其制备方法、ACEi在畜禽体内抗氧化和抗炎作用机制进行了综述,并进一步分析了ACEi高效制备技术,以期为新型功能性饲料添加剂研发与畜禽健康养殖应用提供参考。

1 不同类型酒糟ACEi

酒糟作为酿酒主要副产物,在产业经济目标上,白酒行业产量80亿L,相应产生230万t酒糟[9]。酒糟中含丰富的蛋白质,在固态发酵微生物分泌的蛋白酶可逐步将蛋白质水解为小分子肽段和氨基酸,形成包括ACEi(葡萄酒糟来源)在内的多种高附加值产物,这使其具备成为生物活性肽理想前体原料的良好潜力[10-11]。有研究指出,酒糟ACEi来源于酿酒所用的谷物原料,以糯米为原料发酵的红曲黄酒糟为例,其干物质基础下蛋白质含量可达35%,采用混合酶酶解,可制备出高活性的天然ACEi组合物,对ACE活性的抑制率最高可提升80%以上[12]。从干酒糟及其可溶物(distillers dried grains with solubles,DDGS)的醇溶蛋白水解产物中可筛选得到ACEi,通过活性反相高效液相色谱法(RP-HPLC)馏分分析,共鉴定并定量出6种新型肽段,与其他肽段相比,丙氨酸-缬氨酸-谷氨酰胺(alanine-valine-glutamine,AVQ)和酪氨酸-脯氨酸-谷氨酰胺(tyrosine-proline-glutamine,YPQ)半数抑制浓度(IC50)值分别达到181和220 μmol/L,而缬氨酸-亮氨酸-脯氨酸-缬氨酸-亮氨酸-丝氨酸(valine-leucine-proline-valine-leucine-serine,VLPVLS)为该体系中含量最高的肽,以干物质基础为基准计算,达16.96 mg/g,表明干酒糟的醇溶蛋白水解物是ACEi的优质潜在来源[13]。Su等[14]采用碱性蛋白酶水解大米酒糟,经超滤获得低分子质量馏分后,通过液相色谱-四极杆飞行时间质谱(LC-Q-TOF-MS)与液相色谱质谱(LC-MS)联用技术,从中鉴定出4种体外ACE抑制活性极强的肽段。除白酒糟外,葡萄酒糟来源的ACEi也具有很大开发潜力,如Bravo等[15]利用风味酶对葡萄酒糟进行蛋白质水解,通过超滤、RP-HPLC对肽进行分离后,利用纳升级高效液相色谱-(轨道阱)串联质谱法[nano-HPLC-(orbitrap) MS/MS]对ACE抑制活性最高的馏分进行肽鉴定,成功识别出6种肽,证实葡萄酒糟中同样存在ACEi。目前,食源性ACEi容易被人体肠道吸收且没有副作用,能在机体代谢过程中可发挥调节功能而受到关注[16]。现有研究充分论证了从各类酒糟中制备ACEi的可行性及潜能,但该领域仍面临一些挑战,由于酒糟酿造及制备方式都有所不同,因高温多轮发酵导致蛋白质美拉德反应、形成结合态肽,需特殊预处理释放活性[17],以期未来能从实验室到实际应用的跨越。

2 ACEi调控动物抗氧化应激和抗炎作用

2.1 抗氧化应激作用

郭艳等[18]在浓香型白酒酒糟中鉴定出19个环肽,其中8个环二肽涉及274个与抗氧化作用相关的核心靶点,其环二肽的抗氧化应激作用机制集合多靶点、多成分、多通路的特征。体外研究证实,酚类化合物能提高抗氧化活性,还具有ACE抑制性[19],ROS被认为是血管细胞重要的信号分子,而烟酰胺腺嘌呤二核苷酸磷酸(nicotinamide adenine dinucleotide phosphate,NADPH)氧化酶1(NADPH oxidase 1,NOX1)已被证实可介导AngⅡ诱导的超氧阴离子生成,调控氧化还原依赖信号通路[20]。如图1所示,氧化应激与ROS活性密切相关,一方面,ROS升高会激活ACE活性,导致生成更多的AngⅡ;另一方面,AngⅡ通过激活NADPH氧化酶进一步诱导ROS产生,形成“氧化应激-ACE激活-AngⅡ”恶性循环[21],这一过程会加剧畜禽的健康损伤,因此抑制ACE活性以减少AngⅡ生成,是打破该循环、缓解氧化应激损伤的关键切入点[22]。酒糟中含有多酚、黄酮类化合物等天然的生物活性成分,具有抗氧化能力[23-24],与血管ACEi功能存在机制层面的协同性。
图1 ACEi抗氧化应激通路

ACE:血管紧张素转换酶 angiotensin converting enzyme;ACEi:血管紧张素转换酶抑制肽 angiotensin converting enzyme inhibitory peptide;NADPH:烟酰胺腺嘌呤二核苷酸磷酸 nicotinamide adenine dinucleotide phosphate。

Fig.1 ACEi anti-oxidative stress pathway

研究表明,酒糟中酚类物质含量的差异对ACE抑制活性产生重要影响,表明其在调控RAAS方面的潜力[25]。AngⅡ是通过激活AngⅡ 1型受体,在主动脉内皮细胞中整合了多条功能障碍信号,激活NADPH氧化酶,导致产生ROS水平显著升高[26];下调沉默信息调节因子3(silent information regulator transcript 3,SIRT3)转录及线粒体定位,导致线粒体中超氧化物歧化酶(superoxide dismutase,SOD)2和ROS过度乙酰化,超氧阴离子累积,造成线粒体氧化损伤,同时激活沉默信息调节因子6(silent information regulator transcript 6,SIRT6)-Rho相关激酶1(Rho-associated kinase 1,ROCK1)-动力相关蛋白1(dynamin-related protein 1,Drp1)信号轴,促进线粒体裂变,最终导致内皮功能障碍[27-28]。天然抗氧化肽在家畜体内更易吸收,且几乎不产生不良副作用,主要通过清除自由基、增强机体抗氧化酶活性以消耗ROS、抑制脂质过氧化等机制发挥保护作用[29],这些肽序列中富含疏水性氨基酸、碱性氨基酸(如精氨酸、赖氨酸)和芳香族氨基酸(如苯丙氨酸、酪氨酸、色氨酸)的残基,这种结构不仅有助于提升其抗氧化活性,也是高效ACEi的关键结构特征[30]。研究表明,酒糟采用响应面法优化复合酶解条件,最佳工艺为温度55 ℃、pH 7.5、时间2 h,此时ACEi水解度达64.25%,在酶解3 h条件下获得最高ACE抑制活性(79.05%),该酶解产物同时表现出对1,1-二苯基-2-三硝基苯肼(1,1-diphenyl-2-picylhydrazyl,DPPH)自由基(清除率2.078%)和2,2'-联氮双(3-乙基苯并噻唑啉-6-磺酸)二铵盐[2,2'-azinobis-(3-ethylbenzthiazoline-6-sulphonate),ABTS]自由基(清除率32.26%)的清除能力[31]。基于特定氨基酸结构特征筛选或设计的肽,有望成为调控畜禽氧化应激损伤的高效分子。热应激条件下,家畜为维持体温与体液平衡,激活RAAS导致醛固酮含量升高,并伴随氧化应激加剧[32]。因此,ACEi通过抑制AngⅡ生成,可能改善应激状态下的肾脏灌注不足与氧化损伤。

2.2 抗炎作用

酒糟ACEi抗炎作用并非单一靶点抑制,而是通过“RAAS-肿瘤坏死因子(tumor necrosis factor,TNF)-晚期糖基化终产物受体(receptor of advanced glycation end products,RAGE)”多靶点协同调控实现多层抑制。在细胞模型中,其黄铜提取物活性成分可剂量依赖性抑制肿瘤坏死因子-α(tumor necrosis factor-α,TNF-α)和干扰素-γ(interferon-gamma,IFN-γ),诱导白细胞介素-6(interleukin-6,IL-6)和血清白介素-1β(interleukin-1β,IL-1β)分泌,抑制率分别达到82.56%和62.24%[33]。机制层面上,此效应主要依赖于其对核因子-κB(nuclear factor kappa-B,NF-κB)信号通路的精准干预。一方面,ACEi通过抑制ACE活性,减少AngⅡ生成,阻断其与血管紧张素1型受体(angiotensin receptor type 1,AT1R)结合所触发的NADPH氧化酶激活→ROS积累→抑制性κB激酶β(inhibitor of kappa B kinase beta,IKKβ)磷酸化→NF-κB抑制蛋白(inhibitor of kappa B alpha,IκBα)降解→NF-κB p65核转位经典信号通路,从而下调NF-κB信号通路的基因表达;另一方面,TNF信号通路是促发炎症状的核心信号通路,以ACEi卡托普利为例,其能够通过抑制TNF信号通路,同时上调血管紧张素转换酶2(angiotensin converting enzyme 2,ACE2)的表达,减轻病毒或应激诱导的心肌细胞炎症反应和凋亡[34-35]。除TNF信号通路外,RAGE激活是NF-κB驱动炎症重要的上游机制,RAGE激活后可直接触发NF-κB活化,同时与RAAS过度激活、氧化应激协同致使肾脏炎症、损伤[36-37]。AngⅡ在此过程中与其他炎症次级介质(如细胞因子、一氧化氮、内皮素-1、前列腺素及Rho蛋白通路等)存在的“细胞内交叉调控”,这种介质可通过相互作用激活NF-κB、激活蛋白-1(activator protein-1,AP-1)等转录因子,实现信号转导与效应放大,加剧炎症反应[38-39]。研究表明,AngⅡ参与调节肾血管收缩及氧输送障碍,而采用AT1R阻断剂(如药物氯沙坦)可显著改善低血压模型中的肾脏血流恢复,提示拮抗AngⅡ信号通路在缓解组织损伤中的关键作用[40-41]。这一结果间接证明,ACE通过抑制AngⅡ生成实现的“RAAS-NF-κB”调控,是缓解组织炎症的核心路径之一。仔猪由于营养变化、环境适应压力等断奶应激会导致免疫力下降,破坏仔猪肠道正常生理结构,导致易受病原微生物侵袭进而引发肺炎[42-43]。断奶应激会激活仔猪下丘脑-垂体-肾上腺(hypothalamic-pituitary-adrenal,HPA)轴,导致皮质酮水平升高,进而诱发氧化应激与过度炎症反应,ACE2作为严重急性呼吸系统综合征冠状病毒2型(SARS-CoV-2)的主要宿主细胞受体,在病毒进入细胞中起着至关重要的作用,通过饲喂桂枝理中汤颗粒下调ACE2的表达,在缓解断奶仔猪呼吸系统中的呼吸道过敏和肺炎方面具有显著效果[44]。AngⅡ负向调节ACE2的表达与活性,而ACEi可通过干预该信号通路增强ACE2的表达[45],维持RAAS平衡和改善断奶仔猪呼吸道健康。

3 高效制备ACEi技术方法

酒糟具有高蛋白质、高纤维的显著特征[46],目前关于酒糟利用的研究多集中于饲料领域,如通过酸溶蛋白技术提高其蛋白质资源的利用效率,固态发酵工艺诱导产纤维素酶提升酒糟的消化率,以及借助微生物发酵手段改善酒糟的饲用品质与适口性[47-49]。在工艺优化方面,预处理技术的选择直接影响后续酶释放,例如水提法作为物理预处理,可用于初步溶出可溶性蛋白[50]。研究表明,以米酒糟为例,通常采用“水提→陶瓷膜→纳滤膜→石油醚萃取→多级超滤”的组合工艺,最后5 000 u膜透过液对ACE活性的抑制率达到46%,较常规方法提升6.43%[51]。但是,提高酒糟中功能性生物活性肽含量,特别是ACEi的研究仍然薄弱,本文探索了提高ACEi得率和活性的有效路径,主要包括酶解法和微生物发酵法2种方式。

3.1 酶解法

酶解法常选用多种蛋白酶进行水解,酶的选择至关重要,它直接关乎最终提取的ACEi的效率和活性。单一酶酶解过程中存在一定的局限性,但复合酶酶解时,不同酶有不同的酶切点可以互补,可以生成丰富的ACEi,提高ACE抑制活性[52]。侯蕊等[53]研究多种蛋白酶(碱性蛋白酶、胰蛋白酶、动物蛋白酶等8种)对酒糟蛋白的酶解效果及酶解液ACE抑制活性,动物蛋白酶与胰蛋白酶的酒糟蛋白酶解率最高;碱性蛋白酶与木瓜蛋白酶的酶解液ACE抑制活性最高,均为(0.27±0.01) mg/mL。林晓婕等[54]利用木瓜蛋白酶对红曲糟蛋白进行酶解生成ACEi,酶添加量为6 000 U/g、酶解90 min、45 ℃、pH 5.7,此条件下,酶解率为45.11%±0.12%,蛋白质浓度为2.7 mg/mL时ACE抑制率为60.49%±1.70%,活性成分为<3 000 u短肽,超滤后<1 000 u组分活性最优(抑制率为70.65%±1.51%),含48.22%高活性肽,模拟胃肠消化后活性保持率为58.94%。马思洋等[55]在枸杞籽粕蛋白研究中进一步验证,碱性蛋白酶酶解产物的ACE抑制率显著高于其他蛋白酶,与酒糟提取ACEi研究结论一致,表明碱性蛋白酶对植物源性蛋白质具有广泛适用性。

3.2 微生物发酵法

微生物发酵是提升食源性蛋白质生物活性、制备ACEi的重要手段,已在豆粕、乳清等蛋白质副产物的ACEi中形成成熟技术路径与理论体系。但是,该技术在酒糟这一重要谷物加工副产物中的应用仍处于空白;作为稻米、高粱、小麦等谷物酿酒后的固体废弃物,酒糟不仅保留原料中未完全降解的醇溶蛋白、谷蛋白等优质蛋白,还富集酿酒微生物代谢的酶系与小分子前体物质,具备转化为ACEi的天然基础。Wang等[56]采用乳酸菌对8种不同谷物进行发酵后,ACE抑制率较未发酵增加69.28%。李帅等[57]以藜麦为原料,采用菌株为副干酪乳杆菌,制备的藜麦ACE抑制发酵液效果最好,抑制率达到86.50%±0.25%。Keska等[58]通过超声预处理(40 kHz,2.5 W/cm2,15 min)结合乳酸菌(嗜酸乳杆菌鲍尔L0938和动物双歧杆菌乳酸亚种BB12)发酵猪肉,能够显著提升蛋白质水解物的ACE抑制率(74%)与自由基清除率(>90%)。在发酵过程中,除了菌种选择至关重要外,蛋白质水解能力决定肽链的裂解程度[59],单一酵母菌株发酵有时无法充分降解底物蛋白质,需辅以乳酸菌才能有效水解蛋白质,促进生物活性肽释放[60]。

4 小结与展望

随着养殖业减抗政策的推进以及食品安全要求的不断提高,开发天然、安全且高效的功能性添加剂已成为畜牧业可持续发展的迫切需求。酒糟ACEi作为一种潜力显著的天然活性物质,符合这一发展方向,但其实际应用仍面临诸多挑战。目前,酒糟ACEi的制备与纯化工艺复杂、成本较高,难以实现规模化生产,且产物活性易受环境影响,稳定性较差,严重限制了其产业化应用。因此,未来需重点开发高效、低成本的生物制造技术,如新型酶解工艺、高效分泌表达系统、易于扩产的分离纯化技术等,以提升得率、降低生产成本,并增强产品稳定性。在作用机制方面,现有研究多集中于表观效应,如生长性能、肠道形态的改善,而对ACEi与宿主及肠道微生物互作的分子机制、关键靶点及信号通路仍缺乏系统解析。未来应借助多组学、基因编辑等技术,从分子层面深入探究其作用机理,为实现精准应用提供理论依据。尽管酒糟ACEi在畜禽体内的直接研究尚属空白,但其通过RAAS调控血管收缩、抗氧化与抗炎等机制已在其他模型中得到验证,这为其在畜禽健康养殖中的应用奠定了理论基础。同时,酒糟ACEi的研发不仅有助于实现酿酒副产物的高值化利用,也是推动畜禽养殖减抗提质、农业循环经济发展的有效途径。

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