Chinese Journal of Animal Nutrition >

2025 , Vol. 37 >Issue 11: 7388 - 7405

DOI: https://doi.org/10.12418/CJAN2025.602

REVIEW

Regulatory Mechanisms of Nutritional and Functional Components in Forage on Flavor of Ruminant Products

  • LIU Jindi , 1, 2 ,
  • SUN Mingyuan 3 ,
  • XIN Hangshu 4 ,
  • ZHONG Rongzhen , 1, *
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  • 1 Jilin Province Feed Processing and Ruminant Precision Breeding Cross regional Cooperation Technology Innovation Center, State Key Laboratory of Black Soils Conservation and Utilization, Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences, Changchun 130102, China
  • 2 College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 101408, China
  • 3 Jilin GerunjiaBiotechnology Co., Ltd., Changchun 130102, China
  • 4 College of Animal Science and Technology, Northeast Agricultural University, Harbin 150030, China
*professor, E-mail:

Received date: 2025-04-13

  Online published: 2025-11-14

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Abstract

Flavor is a core indicator of ruminant product quality and the primary sensory characteristic for consumers to assess the quality of animal-derived products, and its formation is closely related to volatile flavor substances and taste presenting substances. Improving the flavor of ruminant meat can create differentiated flavor profiles to meet diversified consumer demands. This paper reviewed the key volatile compounds and taste presenting substances in ruminant meat, and explored the molecular interactions among nutrients, active substances and volatile components in forage on the synthesis of flavor precursors, fat oxidation regulation and microbial metabolism, with a view to providing theoretical basis for the efficient use of forage and production of high-quality animal-derived products, and to promote the efficient conversion and utilization of forage resources and the synergistic development of the flavor industry under the combined mode of planting and raising.

Key words： ruminants; flavor substances; forage composition; nutritional regulation

Cite this article

LIU Jindi , SUN Mingyuan , XIN Hangshu , ZHONG Rongzhen . Regulatory Mechanisms of Nutritional and Functional Components in Forage on Flavor of Ruminant Products[J]. Chinese Journal of Animal Nutrition, 2025 , 37(11) : 7388 -7405 . DOI: 10.12418/CJAN2025.602

过去10年,全球肉类生产量持续增长[1],预测到2033年,全球动物存栏数将增加10%,基于供给蛋白质计算的产出将提升14%[2]。在人口、饮食习惯以及生活方式差异极大的背景下,未来几十年人们对不同风味肉类产品的需求也将呈现激增态势[3],并且更注重肉类品质对人类健康的影响[4]。饮食质量差与人类心血管疾病发病率和死亡率的升高密切相关[5]。富含优质蛋白质与微量营养元素的反刍动物产品是目前全球动物食品市场的重要增长点。但反刍动物肉类所具有的特征风味物质(如4-甲基辛酸)会产生膻味或腥味,这在一定程度上降低了部分消费者的购买欲。风味作为关键感官特性[6],也会直接影响到肉类的食用体验[7]。当前,面临反刍动物产品风味品质下降的问题,其主要受多重因素的影响,如饲养管理粗放,优质草场退化加剧草畜矛盾,兽药农药残留超标激发不良风味,核心畜群遗传潜力不足,进口冲击迫使养殖户依赖快速育肥,以及分级标准未进行市场化难以引导品质提升等[8-11]。不同饲养方式所得反刍动物产品品质存在差异,且各有利弊。研究表明,放牧羊肉氧化稳定性更高,苦味、涩味比舍饲羊肉低[12];但草场退化和市场需求促使快速育肥盛行,甚至出现“假放牧”产品,溯源体系不完善进一步削弱消费者的信任[13-14]。反刍动物产品风味的形成与动物品种、饲草成分、气候条件、饲喂方法、屠宰年龄等密切相关,其中饲草成分起着关键性作用[15]。饲草是反刍动物的主要食物来源[16],影响反刍动物肌肉的生理结构和肉中呈味物质的合成与代谢,进而影响消费者的食品选择行为以及公共政策[17]。反刍动物瘤胃和肠道是一个复杂的微生物生态系统[18],饲粮可以通过调节胃肠道微生物的组成对反刍动物的健康和产品质量产生重大影响[19]。研究饲草对反刍动物产品风味的改善可助力满足市场的需求[20]。本综述深入探讨了反刍动物肉类风味种类以及饲草中营养成分、活性成分、挥发性成分对反刍动物肉品风味的调控作用,以期为优质风味产品生产和饲草行业可持续发展提供参考。

1 反刍动物肉中风味物质种类

风味是气味和滋味的综合评价,反刍动物肉中的挥发性有机化合物(volatile organic compounds,VOCs)和非挥发性滋味呈味物质决定熟肉的香味和滋味。反刍动物肉品的风味有香味、滋味、腥味、膻味、金属味等多种,其中生肉不具有特征香味,只有轻微的血腥味、咸味和金属味[21],但是生肉中的风味前体物质决定熟肉特征风味的形成。风味前体物质为非挥发性成分,主要包括水溶性化合物和脂质[22]。肉类的特征香味主要来自熟肉,是生肉通过热诱导反应,包括美拉德反应和脂质、硫胺素降解以及二者互作产生大量滋味呈味物质和VOCs来产生[23]。

1.1 反刍动物肉中的滋味呈味物质

主要滋味呈味物质见表1。反刍动物肉中的滋味呈味物质来源于水溶性风味前体物质如氨基酸、肽、核苷酸、无机盐、有机酸和糖类等,影响熟肉的基本味道(酸味、甜味、苦味、咸味和鲜味)[24-25]。肉中的酸味主要来源于一些氨基酸及琥珀酸、乳酸等有机酸和磷酸[26];甜味主要源自糖类以及侧基小的氨基酸,如甘氨酸;苦味主要来源于苦味肽及氨基酸类(侧基较大并带碱基时,如亮氨酸)[27];咸味来源于无机盐、谷氨酸盐、天门冬氨酸盐[28];鲜味主要来源于氨基酸类、核苷酸类、多肽以及有机酸等,当2种或2种以上的鲜味物质同时存在会产生协同效应,显著增强风味[29]。
表1 主要滋味呈味物质

Table 1 Main taste flavor substances

滋味
Tastes		 呈味物质
Flavor materials		 参考文献
References
酸Sour 谷氨酸、天门冬氨酸、组氨酸、精氨酸、天门冬酰氨、琥珀酸、
乳酸、吡咯烷酮羧酸、磷酸		 [26]
甜Sweet 葡萄糖、核糖、果糖、丙氨酸、羟脯氨酸、脯氨酸、甘氨酸、
丝氨酸、苏氨酸、赖氨酸、半胱氨酸、蛋氨酸、天冬酰胺、谷酰胺		 [30]
苦Bitter 肌肽、鹅肌肽、次黄嘌呤、其他苦味肽类、肌苷酸、肌酸、精氨酸、
亮氨酸、赖氨酸、组氨酸、色氨酸、蛋氨酸、缬氨酸、异亮氨酸、苯丙氨酸、酪氨酸		 [27]
咸Salty 无机盐、谷氨酸盐、天门冬氨酸盐 [28]
鲜Fresh 谷氨酸钠、5'-肌苷酸、5'-鸟苷酸、多肽、乳酸、琥珀酸 [29]

1.2 决定反刍动物肉类香味的VOCs

决定反刍动物肉类香味的VOCs主要有醛类、酮类、醇类、酯类、烷烃、酸类,其次是呋喃、含硫化合物、含氮杂环化合物、含氧杂环化合物等[31]。基于气相色谱-质谱联用(gas chromatography- mass spectrometry,GC-MS)分析的挥发性物质,通常用气味活度值(OAV,浓度除以气味阈值的比率)≥1的气味物质来表征反刍动物肉类的特征气味物质[31-33]。研究发现,可以通过定量分析芳香活性化合物并比较其OAV,进一步阐明它们对整体香气的相对贡献[34-35],如百里香炖羊肉的特征香味物质(OAV≥1)是己醛、壬醛、辛醛、(E)-2-辛烯醛、(E,E)-2,4-癸二烯醛和二甲基三硫[36]。Watkins等[37]确定了羊肉中对风味影响最大的15种化合物的主要香味特征是羊肉味、蘑菇味、脂肪味、油炸味、焦糖味、蜡质味、坚果味、硫磺味、柠檬味、花香味、青草味等。Bueno等[38]检测出了新鲜羊肉和冷冻羊肉在烧烤后产生香味的最重要的8种香味物质——(Z)-2-庚烯醛、2,5-二甲基吡嗪、(Z)-2-癸烯醛、2-丁基-2-辛烯醛、2-乙酰基-2-噻唑啉、(E,E)-2,4-癸二烯醛和2种未知物质。12-甲基十三醛是水煮牛肉和烤牛肉的特征香味物质[39-40]。不同种类动物的肉类香气差异不完全源于香味物质质量的不同,而可能与特定化合物的数量差异有关,从而影响香气的感知。对于反刍动物,醛类对肉类香味起决定性的作用。牛、羊肉中的主要VOCs及其香气特征见表2。
表2 牛、羊肉中的主要挥发性有机化合物及其香气特征

Table 2 Main VOCs and their aroma characteristics of beef and mutton

类别
Categories		 挥发性有机化合物以及香气特征
VOCs and their aroma characteristics		 参考文献
References
醛类Aldehydes 戊醛(苦杏仁味、麦芽味、辛辣味)、己醛(青草味、花香味、脂肪香味)、庚醛
(脂肪香味、咸味、烤肉味)、辛醛(脂肪香味、果香味)、壬醛(脂肪香味、
花香味、柑橘味)、(E)-2-辛烯醛(脂肪香味)、苯甲醛(苦杏仁味)、(Z)-2-
癸烯醛(烤肉味)、癸醛(柠檬味、花香味、烤肉味)、(E)-2-癸烯醛(油炸味)、
(E)-2-庚烯醛(蔬菜味)、十二醛(肥皂味)、对甲基苯甲醛(苦杏仁味)		 [41-43]
醇类Alcohols 戊醇(果香味、香醋味)、己醇(酒精味、脂肪香味、果香味)、庚醇(
脂肪香味、柑橘味、酒香味、青草味)、2-乙基己醇(甜味、花香味)、
辛醇(脂肪香味、果香味、青草香味)、壬醇(花香味、柑橘味、脂肪香味、
青草香味)、1-辛烯-3-醇(青草香味、蘑菇味、金属味)		 [41-43]
酮类Ketones 苯乙酮(苦杏仁味)、2,3-辛二酮(甜奶油味、甘薯味)、2-庚酮
(果香味、辛辣味、肉桂味)、6-甲基-2-庚酮(丁香味、
薄荷脑味)、2-甲基环戊酮(烤牛肉味)、3-甲基环戊酮(烤牛肉味)		 [41-43]
酯类Esters 乙酸乙酯(水果味、甜香味)、甲酸庚酯(清新味)、甲酸辛酯
(黄瓜味)、丁酸乙酯(果香味、菠萝味、苹果味)		 [41-43]
呋喃类Furans 2-丁基呋喃(水果味)、3-戊基呋喃(青豆味) [41-43]
酸类Acids 己酸(羊肉味)、4-甲基辛酸(腥膻味) [41-43]
含硫化合物
Sulfur-containing compounds		 二甲基三硫(洋葱味、刺激性气味、硫磺味)、1-(甲硫基)
乙硫醇(肉香味、洋葱味)		 [30]

2 饲草成分的功能化利用

2.1 饲草中的营养成分

饲草富含蛋白质、脂肪酸、碳水化合物及微量元素等营养成分。常见的饲草有紫花苜蓿、红三叶草和紫云英等豆科饲草,甜高粱、黑麦草和墨西哥玉米草等禾本科饲草以及菊苣、鲁梅克斯、饲用甜菜和灯心草等其他科饲草。不同种类饲草的营养成分含量具有显著差异。豆科牧草粗蛋白质含量高于禾本科牧草,粗纤维、粗脂肪含量低于禾本科牧草[44]。豆科饲草适合提供蛋白质,禾本科适合提供能量。此外,小黑麦和狗牙根虽然为新型碳源饲草,但其粗蛋白质含量显著高于一般禾本科饲草[45]。籽粒苋是富含蛋白质的氮源饲草,其粗蛋白质含量可与苜蓿媲美[46-47]。
最佳觅食理论是研究动物以最小成本获取最大收益的觅食策略,其研究的深入有助于探究饲草摄入后的代谢机制[48-49]。动物对食物选择受到多种因素的影响,其中包括食物的感官特征(如气味、味道和质地)以及进食后的反馈体验(如积极或消极的感受)[50]。草食动物在寻找食物时,会通过视觉、触觉或化学线索来获取植物的营养或防御状态信息[51-52]。反刍动物偏好于能够快速提高饱腹效应的饲草,在等营养水平条件下,饲草本身的感官特性也可以刺激或抑制采食行为[53]。那些优先选择优质叶片的动物更能利用饲草中存在的营养异质性[54]。Geay等[55]研究发现,营养会影响反刍动物骨骼肌结构及其生化成分,进而影响肌肉结构组成和肉的感官品质(嫩度、色泽、风味和多汁性)。大量的事实证明,营养需求会影响动物对食物的偏好,而感官特性会调节动物对各种食物的辨别[56-58]。

2.1.1 蛋白质

蛋白质是维持反刍动物正常生命活动的必需物质[59]。周圻等[60]研究显示,随着粗蛋白质和微量元素含量的增加,海南坡鹿对饲草的喜食性显著提高。饲粮的蛋白质水平会影响肌内脂肪含量,进而影响肉品硬度、大理石花纹等[61]。在波杂羔羊上的研究发现,提高饲粮蛋白质水平,羊肉的肉色、眼肌面积、熟肉率呈增加趋势,且羊肉具有更好的鲜嫩多汁性,氨基酸含量增加,风味更佳[62],但饲粮的蛋白质水平过高则会使波杂羔羊肌肉过于紧实,影响口感[63]。另外,某些氨基酸可能在加热过程中与还原糖发生美拉德反应,产生香味物质,比如杂环化合物[30]。也有研究发现不同蛋白质水平饲粮对公牦牛肉品质的影响有限[64]。
目前,饲草蛋白质的高效利用与风味品质的协同调控主要涉及2个方面:一方面,饲草蛋白质的降解与氮排放密切相关[65];另一方面,瘤胃微生物对蛋白质的不完全代谢会产生硫化物、生物胺等异味前体物质,影响产品风味品质[66]。通过精准调控蛋白质供应,不仅能提高氮利用效率,减少环境污染,还能促进风味前体物质的合成,从而提升肉与乳产品的风味品质,这为实现环境友好型优质畜产品生产提供了思路。

2.1.2 脂肪酸

动物体内的脂肪酸主要有饱和脂肪酸(saturated fatty acid,SFA)、单不饱和脂肪酸(monounsaturated fatty acid,MUFA)和多不饱和脂肪酸(polyunsaturated fatty acid,PUFA),其中n-6和n-3系列的PUFA对动物产品风味形成起关键作用,而MUFA因抗氧化性是平衡风味的关键脂肪酸。反刍动物瘤胃微生物会部分氢化PUFA生成共轭亚油酸(conjugated linoleic acid,CLA)和SFA(如硬脂酸),并有可能增加衍生动物产品中异油酸的含量。饲草中脂肪酸的成分和含量能够调控羊肉肌内脂肪含量[67]。饲草中的脂肪酸以PUFA(如亚麻酸、亚油酸)为主[68]。其中,禾本科植物往往比豆科植物能提供更多的总脂肪酸和α-亚麻酸,而提供的亚油酸却比豆科植物少[69]。但这并不是绝对的,PUFA含量在不同饲草种类之间存在差异。例如,豆科牧草中白三叶草比红三叶草和苜蓿含有更高含量的亚麻酸,红三叶草的亚油酸含量较高,苜蓿的亚油酸含量最低[70]。除此之外,亚麻籽油、大豆油和亚麻籽油等植物油也都是α-亚麻酸的来源,动物食用后可以利用这些植物油合成二十碳五烯酸(eicosapentaenoic acid,EPA)、二十二碳五烯酸(docosapentanoic acid,DPA)和二十二碳六烯酸(docosahexaenoic acid,DHA),它们在炎症、免疫和风味的关键调控方面与花生四烯酸具有同等重要的作用[71],适量的EPA、DPA、DHA可增强畜产品鲜味,但需控制氧化。饲草中的脂肪酸作为风味前体物质直接参与风味代谢网络,成为调控反刍动物肉品风味品质的关键因素,建议通过优化饲粮脂肪酸比例和补充外源抗氧化物质来平衡PUFA代谢对肉品风味的影响。

2.1.3 碳水化合物

碳水化合物是反刍动物营养系统的基石,对反刍动物的健康、生长、消化功能以及生产性能至关重要。反刍动物的主要碳水化合物来源是含有纤维素和半纤维素的纤维饲料以及富含淀粉的谷物等[72]。纤维素是由葡萄糖组成的大分子多糖;半纤维素是由木糖和阿拉伯糖等聚合而成的异质多糖;可溶性糖主要包括单糖、寡糖等,在瘤胃发酵过程中可以平衡瘤胃pH,降低亚急性瘤胃酸中毒(subacute ruminal acidosis,SARA)发生的风险,还可以弥补乳脂合成的前体物质(如乙酸盐和丁酸盐)的不足[73]。虽然牧草中的木质素不是碳水化合物,但它对纤维素和半纤维素的消化率有显著影响;木质素会与结构碳水化合物和细胞壁蛋白质结合,降低养分的可用性[74]。饲草中的碳水化合物不仅影响动物的消化吸收过程,还通过代谢产物的生成、微生物发酵等途径影响反刍动物肉品特征香气的累积和口感体验,合理配制反刍动物饲草碳水化合物的比例,有助于提高饲草资源利用率。

2.2 饲草中的活性物质

风味不仅反映了畜产品的质量,也反映了人类对动物福利的关注[75]。植物制剂是最早用于防治人类和牲畜疾病的治疗方法,可以通过植物疗法寻找具有创新性的抗菌剂[76]。饲草含有酚类化合物、萜类、生物碱等生物活性化合物(plant bioactive compounds,PBC),具有显著的抗氧化、抗炎作用,这些成分可被提取并转化为功能性产品,通过增加PUFA含量和氧化稳定性对肉类风味带来有益作用[77]。国外学者研究发现,从水果和蔬菜副产品中提取的饲料添加剂具有抗菌、抗氧化和益生的功效,因其具有多重有益特性,这类添加剂有望成为传统抗生素的替代品[78]。抗生素的累积会让肉类产生不良风味,如硫磺味、苦味。饲草中的活性成分对反刍动物风味的影响多聚焦于其功能,如在反刍动物饲粮中使用天然化合物(丁香酚、百里香酚、香兰素)可改善肉色、嫩度,增加香草味、松香味、奶香味等风味[79]。除此之外,有研究发现沙葱活性成分提高了肉羊背最长肌中己醛(青草味)和1-辛烯-3-醇(蘑菇香、蔬菜香)的含量[80]。

2.2.1 酚类

酚类是指芳香烃通过莽草酸途径产生的化合物,其中苯环上的氢原子被羟基取代,是一类重要的次级代谢产物,主要包括类黄酮、酚酸等,具有抗氧化、抗菌及消炎等作用,可以减少抗生素累积带来的异味[81]。比如,红三叶含有鹰嘴豆芽素A和B、染料木素和大豆素等异黄酮类物质,具有作为抗生素替代品的潜力[82];芦丁和丁香酸可增加瘤胃体外发酵挥发性脂肪酸含量,而鞣花酸则降低挥发性脂肪酸含量[83]。饲草中酚类化合物的种类见表3。
表3 饲草中酚类化合物的种类

Table 3 Types of phenolic compounds in forages

类别Categories 化合物Compounds 典型饲草Typical forages
类黄酮
Flavonoids		 黄酮、黄酮醇、查尔酮、花青素、异黄酮、
黄烷、黄烷酮、黄烷醇、新类黄酮、噢哢、芦丁等		 高粱[86]、核桃叶[87]、紫花苜蓿[88]、
高羊茅[89]
酚酸
Phenolic acids		 咖啡酸、香豆酸、阿魏酸、芥子酸、鞣花酸、
绿原酸、原儿茶酸、没食子酸、香草酸、丁香酸、
龙胆酸、对羟基苯甲酸等		 核桃叶[87]、紫花苜蓿[88]、
高羊茅[89]、榛子叶[90]、黑麦草[91]
单宁
Tannins		 水解单宁、缩合单宁 沙柳[92]、银合欢[93]、
百脉根[94]、沙棘[95]
在畜牧业生产中使用抗生素有助于保持动物健康和生产力,但这一做法也加剧了耐药性病原体在牲畜和人类中的传播,对公共健康构成了严重威胁[84-85]。挖掘饲草的酚类化合物的药用潜力,不仅可以减少抗生素带来的异味积累,还能够减少氧化反应,保持肉质的新鲜和口感,酚类还可以与其他成分协同增强肉类的风味层次,提升消费者的食用体验。

2.2.2 萜类

萜类是最大类的植物天然产物,根据分子骨架中异戊二烯单元的数量可以分为单萜、倍半萜、双萜、三萜等。饲草可能通过将特定植物的萜类化合物直接转移到肉中来影响反刍动物肉品的最终风味,从而赋予肉品特征香气。例如,来自饲草的植醇、植烯、萜烯和倍半萜可能会在肌肉组织中积累,这些化合物一旦进入肉中,如果浓度足够高,会直接影响最终风味,或者在热加工过程中发生降解,形成新的风味化合物[37]。精准配制饲草中萜类化合物的种类和浓度,能够为不同类型的反刍动物产品生产提供个性化的风味调控方案,可以满足消费者日益增长的多样化需求。

2.2.3 生物碱

含生物碱成分的可饲用天然植物已被广泛应用。例如,研究发现从番茄植物不同部位提取的番茄碱能不同程度地抑制滴虫的生长,减轻病原体污染[96];刺檗(Berberis vulgaris)富含生物碱、黄酮类、花青素和单宁等植物次生化合物,其生物活性成分中含有大量异喹啉生物碱(小檗碱、巴马汀和药根碱)和双苄基异喹啉生物碱,对乳畜代谢障碍、脂质代谢紊乱、氧化和炎症应激有积极影响[97];对奶山羊补充小檗碱可改善血液、初乳和牛奶中的抗氧化标志物,还能增加干物质和代谢蛋白质的摄入量,改善能量平衡,提高初乳和牛奶产量[98-99]。

2.3 饲草中的挥发性物质

植物会释放出VOCs,其中一些会被动物感知,某些饲草中所含的芳香物质能够作为食品添加剂,通过刺激各种消化酶的活性,诱导动物采食[100],还能通过减少发酵产物和病原微生物的数量、加强前期营养物质的消化、改善肠道对重要营养物质的可及性以及改善抗氧化状态和免疫反应来提高风味[101]。Brenes等[102]强调了VOCs通过口鼻感应对脂质代谢的影响,尤其受到关注。
饲草中的VOCs主要通过植物次生代谢途径产生,包括萜类、醇类、酮类、酯类、醛类和酚类等,它们的组合和浓度决定了饲草的特定香气和风味。饲草中的VOCs在不同种属间存在差异[103-105],对动物肉品质的调控需要考虑品种带来的差异。适口性是增加草食动物对采食量的主要驱动力之一,这也进一步证实了饲草中VOCs与动物健康和饮食福利之间的密切关系。国外学者证实了喷洒黑麦草或白三叶芳香提取物会增加山羊对秸秆颗粒的偏好[106]。可在饲粮中添加VOCs,同时也有必要检测饲草中的活性成分,以确定饲草挥发性成分协同活性成分对反刍动物产品风味调控的作用[102]。

2.3.1 挥发性萜类

分子质量较小的萜类化合物如单萜和倍半萜多为有特殊气味的挥发性化合物,这有助于鉴别“牧场”动物产品,舍饲动物产品通常以酮和挥发性酸为特征[107-109]。Ueda[110]研究发现,从牧草植物叶绿素中提取的1-植物烯是放牧牛奶中特有的VOCs,其含量与牧草摄入量之间存在正相关性。不同种类饲草中挥发性萜类含量不同。例如,来自伞形科植物牧场的牛奶比来自禾本科植物牧场的牛奶含有更大量和更广泛种类的萜烯[111]。希腊牛至(Origanum vulgare L.)中挥发性萜烯(香芹酚、γ-萜品烯、对伞花烯、月桂烯、α-蒎烯、β-石竹烯、百里酚)含量最高[112],香芹酮和柠檬烯在葛缕子(Carum carvi L.)中含量最高[113]。事实上,动物源产品风味都具有饲草或青贮饲料味道的某些特征[114]。

2.3.2 醛类

醛类是一类含醛基(-CHO)的有机化合物,通常为挥发性或半挥发性化合物,分子质量较小,通过气孔或表皮释放到环境中,易与呼吸道接触,也会影响反刍动物的适口性。饲草中的醛类会影响牛奶、奶酪和肉类的风味[115-116]。醛类对反刍动物肉品风味的影响主要是通过醛类物质的沉积,加热时经美拉德反应生成吡嗪、呋喃等化合物,增强“野味”特征。己醛、正己醇、(Z)-3-己烯醛和(E)-2-己烯醛是饲草的“绿色气味”[117],绿色气味化合物占新鲜高羊茅牧草中VOCs的11.2%,占干草中VOCs的8.5%[118]。研究发现,己醛和壬醛可使熟化乳制品略带果香味、柠檬味、草药味和油脂氧化味,苯甲醛可以带来樱桃香气[119-120],正丁醛可以产生麦芽味或化学味道[114]。青贮饲料中的色胺、甲醛会抑制反刍动物的食欲[121]。醛类的化学性质活泼,易发生氧化、还原等反应,适量醛类可增强肉类“天然草饲”风味,但过量则引发氧化劣变[122]。

2.3.3 其他类

Mayland等[118]研究了高羊茅中VOCs与牛偏好的关系,发现牛可能会对6-甲基-1-5-庚烯-2-酮含量较高的饲草种类产生偏好,但对(Z)-3-己烯基丙酸酯或乙酸含量较高的饲草种类产生厌恶。除了对适口性的影响,VOCs本身也会赋予动物产品味道。酯类和醇类化合物赋予牛奶愉悦的味道,而丙酮和2-丁酮则带来不愉快的气味。正常牛奶中含有这些化合物,但绝对量或相对量的变化会产生异味。比如,青贮饲料中的(Z)-3-己烯醇会给牛奶带来异味[114]。最重要的是,除了高剂量危害以外,并不是所有的挥发性化合物对动物都是有利的,香豆素是黄花茅干草或鲜草精油中检测到的最丰富的化合物,必须考虑其毒性,尤其是对放牧动物的毒性[123]。

3 饲草成分对反刍动物产品风味的调控机制

3.1 饲草底物驱动的直接调控

3.1.1 脂肪酸前体供给

在所有物种中,肉类脂肪酸组成都可以通过饮食改变。新西兰草饲喂绵羊增加了羊肉中n-6或n-3 PUFA含量,二者会产生不同范围的风味前体物质,对肉品风味发展有积极的影响[124]。放牧体系下,反刍动物摄入大量新鲜饲草可提高乳制品中脂肪酸(如PUFA、MUFA、CLA)和抗氧化剂(α-生育酚的合成2R立体异构体除外)的含量[125]。n-3 PUFA对风味的影响主要体现在:1)分解产物挥发性强,气味阈值低;2)分解产物活性高,易参与美拉德反应;3)分解产物相互作用形成特有香味;4)易氧化,增加n-6、n-9 PUFA分解产物等[126]。适度增加n-3 PUFA含量有助于反刍动物肉品风味的改良。PUFA氧化生成的醛类(如己醛)和酮类(如2-壬酮)是肉香的重要表征化合物,但过度氧化会导致不良氧化风味。Yang等[127]发现,牧草组的滩羊胸最长肌中蛋氨酸、γ-丁内酯、苯甲醛和乙酸乙酯含量明显高于浓缩饲料组,而不良的氧化风味脂质衍生的饱和醛、醇和烯醛含量明显低于浓缩饲料组,说明牧草组羊肉的苦味、鲜味、浓味和甜味均优于浓缩饲料组,放牧改善羊肉风味。

3.1.2 挥发性萜类物质的转移

挥发性萜类物质容易通过动物呼吸道和胃肠道转移到动物源产品中,并且动物呼吸道接触比口服瘤胃接触影响更快[128-129],挥发性萜类物质主要通过气味转移性影响反刍动物风味。Vialloninsta等[130]研究发现,西洋蓍草(Achillea millefolium)中的单萜和倍半萜从饲草中转移到乳脂中的速度非常快,没有记忆效应,且改变饲草的植物种类组成会对乳脂中单萜烯和倍半萜烯的比例和数量产生很大影响。绵羊摄入与吸入柠檬烯会改变羊奶和奶酪的香气,增加柠檬香、柑橘香[131]。此外,柠檬烯的抗氧化活性可以延缓羊肉脂质氧化过程,减少变色过程、氧化代谢物的产生和酸败,相对高剂量的短期处理可诱导肌肉脂肪中的萜烯富集,改变反刍动物肉品风味[132]。

3.1.3 抗氧化物的作用

酚类对反刍动物风味的影响主要是通过清除自由基、减少肌肉中PUFA(如亚油酸、α-亚麻酸)的氧化、保护肌内脂肪的氧化稳定性、保留肉中天然风味前体物质、延缓脂质过氧化产生异味来实现。除此之外,维生素E,即α-生育酚,是动物组织中主要的脂溶性抗氧化剂,可以延缓氧化并延长肉的保质期。通过在饲粮中添加维生素E来增加其在肌肉和脂肪中的沉积,可以减少由氧化引起的氧合肌红蛋白转化为高铁肌红蛋白,以及组织膜中PUFA降解产生的酸败气味,减少陈腐味[133-134];此外,提高维生素E含量还可以减少牛奶中的氧化风味[135]。其他的,如类胡萝卜素具有维生素A功能和抗氧化特性,尽管β-胡萝卜素和叶黄素在牛奶中的含量较低,但它们通过着色和抗氧化作用,影响乳制品的风味[136]。
优质饲草是畜牧业不可或缺的生产资料,可以为动物提供丰富的营养物质,提高饲料利用率,也可以减少抗生素和化学添加剂的使用,减少外来异味对动物产品风味的影响,有助于提升畜产品的品质和产量,降低有害物质进入食物链的风险。

3.2 瘤胃微生物介导的代谢转化

3.2.1 挥发性脂肪酸的生成

纤维素和半纤维素可以被瘤胃微生物(如纤维降解菌、产乙酸菌)分解为挥发性脂肪酸。乙酸、丙酸和丁酸是挥发性脂肪酸的主要形式,主要由纤维素、半纤维素、淀粉和糖等植物材料发酵产生。乙酸和葡萄糖作为乙酰辅酶A的前体,参与氧化或脂肪生成过程。丁酸盐也会被迅速氧化用于脂肪生成或乳脂合成,二者均可提升肉品的香味及乳制品的浓郁风味。丙酸盐由肝脏清除,但大部分转化为葡萄糖[137]。葡萄糖、挥发性脂肪酸和氨基酸是反刍动物胰岛素分泌的潜在刺激物[138]。含有高浓度可降解淀粉的饲粮会导致瘤胃发酵产物偏向丙酸,丙酸可能诱导胰高血糖素释放,这反过来也会直接通过其对肝脏糖原分解和糖异生的影响刺激胰岛素分泌[139-140]。这会间接影响反刍动物肌肉中糖原含量,从而调控宰后肌肉的pH,影响肉的嫩度和风味。
碳水化合物还可通过微生物与代谢调控的相互作用,优化瘤胃微生物发酵,促进有益代谢、减少低效或有害代谢[141]。肠道微生物可将碳水化合物分解为短链脂肪酸作为宿主的能量来源,短链脂肪酸能与肠道上皮细胞G蛋白偶联受体结合,通过肠-脑轴调控中枢神经系统食欲相关的激素分泌,调节宿主的新陈代谢[142],影响反刍动物肉中风味物质的积累。Chen等[143]研究发现,相比饲喂全株水稻和玉米青贮饲料,饲喂水稻秸秆饲料的公牛瘤胃液样本微生物多样性更高,转运体在反刍动物饲喂系统中发挥着重要作用,其丰度越高,越有利于瘤胃微生物区系获取饲草能量,产生瘤胃微绒毛蛋白(为动物提供蛋白质),促进营养吸收,减少粪臭素积累。纤维含量较高的黑麦草-白三叶牧草处理的体外瘤胃液发酵培养物的产琥珀酸丝状杆菌(F. succinogenes)和厌氧真菌数量较高,而甜菜处理的原生动物数量、挥发性脂肪酸和氨态氮浓度较高,pH较低[144]。饲草成分对肠道菌群的重塑作用要强于遗传因素,植物纤维可通过提高水解细菌和刺激短链脂肪酸的产生来提高微生物的多样性[145]。在育肥羊上的研究显示,苜蓿纤维中的植物多糖成分与瘤胃和盲肠微生物多样性呈正相关,促进代谢物表达上调,改善碳水化合物消化、脂肪代谢、免疫和炎症反应,提升肌肉抗氧化能力,改善羊肉嫩度与风味[146]。特定的饲粮脂肪酸可通过改变细菌群落中纤维降解菌的比例来支持瘤胃中的纤维消化[147]。

3.2.2 微生物蛋白合成

饲草中的蛋白质可以在瘤胃中经过降解与微生物蛋白合成新的营养成分[148]。例如,蛋白质在瘤胃中被分解为游离氨基酸(如谷氨酸、天冬氨酸、含硫氨基酸等),是重要的风味前体物质来源,谷氨酸代谢产物(如谷氨酰胺)可提升肉类的鲜味,天冬氨酸参与合成鲜味肽,与谷氨酸协同作用强化风味层次感,含硫氨基酸在加热时分解产生硫醇、二甲基硫醚等化合物,可能带来“野味”或“腥膻”特征。饲喂豆科牧草白三叶草的绵羊比饲喂黑麦草的绵羊瘤胃中挥发性脂肪酸浓度和丙酸比例要高,其羊肉风味更浓郁[149]。研究证明,适宜的饲粮蛋白质水平能够提高妊娠期秦川母牛[150]的繁殖性能和育成期云上黑山羊[151]的生长性能,增加血清高密度脂蛋白、总蛋白含量,并提高瘤胃纤维素降解菌的相对丰度,促进碳水化合物代谢及糖的生物合成与代谢等过程。饲粮中添加中草药能促进动物机体蛋白质及氨基酸的合成,提高氮的代谢与利用,有助于提高肌肉中鲜味氨基酸、甜味氨基酸、半胱氨酸和肌苷酸的含量[152-153]。

3.2.3 甲烷代谢调控

高纤维饲草会促进反刍动物瘤胃中纤维降解菌的增殖,促进纤维物质的降解[154]。添加菊苣可以刺激盲肠中拟杆菌属(Bacteroides)的增殖,抑制包括理研菌属(Rikenella)在内的病原微生物增殖,增加肌肉中的总氨基酸和美味氨基酸含量[155]。原花青素又称缩合单宁(condensed tannin,CT),能与蛋白质结合,中等浓度的CT可以减少瘤胃微生物将蛋白质降解为氨的量,提高氨基酸的吸收,减少甲烷排放,降低羊肉膻味[156];此外,适量的CT还能减少寄生虫危害和牛腹胀风险,减少驱虫药和化学洗涤剂的使用,减少粪臭素沉积[157-160]。给反刍动物饲喂含CT的饲草,可通过减少蛋白质降解和抑制蛋白质分解及吲哚和鞣酸形成瘤胃微生物的活动,减少瘤胃中吲哚和鞣酸的形成,以减轻反刍动物肉品的田园风味[161]。植物皂苷是一类天然存在的三萜类或甾体苷,具有广泛的生物和药理活性,在免疫刺激、抗肝脏毒性、抗积食、杀灭软体动物、溶血、抗真菌和抗病毒等方面有积极作用[162]。研究发现,皂苷或含皂苷的植物作为饲料添加剂对瘤胃微生物、瘤胃发酵以及皂苷在反刍动物体内的代谢具有积极影响[163]。值得注意的是,含皂苷的植物(以豆科植物为主)常被用作动物饲料,而其他一些植物因含有毒性,通常需要从其特定部位提取皂苷后才能作为饲料添加剂使用。

3.3 反刍动物代谢通路响应

饲草中的n-3 PUFA可以通过激活过氧化物酶体增殖物激活受体,促进反刍动物脂肪细胞分化。放牧羊肉较舍饲羊肉具有更优的挥发性风味特征与脂肪酸组成,其机制在于牧草中亚麻酸、亚油酸和花生四烯酸作为脂氧合酶(lipoxygenase,LOX)参与脂肪氧化反应的主要底物,激活了LOX基因的表达,催化生成更多的风味化合物,赋予放牧羊肉较为丰富的风味[164]。除此之外,饲草中的小檗碱通过激活核因子E2相关因子2(nuclear factor E2-related factor 2,Nrf2)(协调细胞对氧化应激反应的重要转录因子)通路增强抗氧化酶活性,减少脂质过氧化产物,延缓肉中异味形成[165]。饲草功能成分在反刍动物肉品风味调控中发挥着重要作用,既能对代谢过程起到激活或者抑制作用,又能平衡氧化还原反应。

4 小结与展望

反刍动物肉品风味的形成是一个复杂的过程,受遗传及饲养等多方面的影响。本文总结了反刍动物肉品中风味物质种类,饲草营养成分、活性成分、挥发性成分对反刍动物肉品风味的调控机制。目前,关于饲草成分对动物风味的调控机制亟待深化,随着组学技术、测序技术以及人工智能(artificial intelligence,AI)的发展和进步,借助多平台技术的联用,有助于深入探究饲草成分、瘤胃代谢及风味特征的关系。
大数据和人工智能为风味研究带来了革命性的变化[166],在这一领域中,机器学习(machine learning,ML)和深度学习(deep learning,DL)等子领域不断取得新进展。例如,利用肉中的脂肪酸和挥发性化合物作为羔羊胴体分类的生物标记,可以提高ML在开发食物链可追溯系统方面的适用性和有效性[167]。联用技术与AI相结合的关键是要整合组学、仪器检测与模型建立,从经验驱动转向数据驱动,从小尺度扩展到区域尺度,从而更深入地分析利用反刍动物肉类的芳香规律性。综上所述,消费者对于反刍动物肉品风味的偏好存在差异,期待未来的研究能够深化揭示饲草成分对反刍动物肉品风味形成的调控机制,为消费者提供风味多样化、高品质的动物源产品,为饲草的养殖效果和产品转化提供参考。

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