动物营养学报 >

2025 , Vol. 37 >Issue 2: 709 - 722

DOI: https://doi.org/10.12418/CJAN2025.061

综述

营养调控改善猪肉品质研究进展

  • 熊云霞 ,
  • 杨雪芬 ,
  • 蒋宗勇 ,
  • 王丽 , *
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  • 广东省农业科学院动物科学研究所,猪禽种业全国重点实验室,农业农村部华南动物营养与饲料重点实验室,岭南现代农业科学与技术广东省实验室河源分中心,广东省畜禽育种与营养研究重点实验,广州 510640
*王 丽,研究员,E-mail:

熊云霞(1988—),女,江西南昌人,副研究员,硕士,研究方向为猪营养与饲料。E-mail:

Copy editor: 武海龙

收稿日期: 2024-08-14

  网络出版日期: 2025-02-16

基金资助

国家生猪产业技术体系(CARS-35)

河源市高层次人才引进项目

收起

Advances in Nutritional Regulation for Improving Pork Quality

  • XIONG Yunxia ,
  • YANG Xuefen ,
  • JIANG Zongyong ,
  • WANG Li , *
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  • Guangdong Provincial Key Laboratory of Animal Breeding and Nutrition, Heyuan Branch Center, Guangdong Provincial Laboratory of Lingnan Modern Agricultural Science and Technology, Key Laboratory of Animal Nutrition and Feed Science in South China of Ministry of Agriculture and Rural Affairs, State Key Laboratory of Swine and Poultry Breeding Industry, Institute of Animal Science, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China
*professor, E-mail:

Received date: 2024-08-14

  Online published: 2025-02-16

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摘要

猪肉品质是影响消费者购买猪肉意愿的主要因素之一,直接影响猪肉的经济价值。相较于育种手段,通过营养调控手段改善猪肉品质具有时间短、见效快、成本低、易于操作等优势。本文就近年有关营养调控猪肉品质的国内外研究进展进行综述,为养猪产业高质量发展提供一定的参考。

关键词： 猪; 肉品质; 营养调控

本文引用格式

熊云霞 , 杨雪芬 , 蒋宗勇 , 王丽 . 营养调控改善猪肉品质研究进展[J]. 动物营养学报, 2025 , 37(2) : 709 -722 . DOI: 10.12418/CJAN2025.061

Abstract

Pork quality is one of the primary factors influencing consumer purchasing decisions and directly impacts its economic value. Compare with breeding methods, nutritional regulation to improving pork quality has the advantages of shorter timeframes, quicker results, lower costs and easier implementation etc. This paper reviews recent domestic and international research progress on nutritional strategies to enhance pork quality, providing valuable insights for the high-quality development of the swine industry.

Key words： pig; pork quality; nutritional regulation

我国是世界第一大猪肉生产国和消费国。生长性能、繁殖性能和肉品质是猪的3个主要经济性状。在物质匮乏年代,对猪肉的追求还停留在产量上,因此早前的研究比较关注提高猪的生长性能和繁殖性能。但是,随着生活水平的提高,人们越来越关注猪肉的品质问题,肉品质直接影响猪肉的经济价值[1]。就猪肉加工者和猪肉终端消费者而言,比较关注的肉品质性状指的是猪的胴体性状、猪肉的感官品质及营养价值等,主要包括瘦肉率、背膘厚、肉色、pH、滴水损失、肌内脂肪(intramuscular fat,IMF)含量、肌纤维类型、肌肉风味前体物质(氨基酸和脂肪酸的组成)等。肌肉色泽主要取决于肌红蛋白和细胞色素含量,肌肉中肌红蛋白含量和氧化还原稳定性会影响猪肉最终的感官颜色[2]。肌肉嫩度指肉在咀嚼过程中的软硬程度和易咀嚼性,直接影响消费者的口感体验和满意度。IMF含量、pH及肌纤维类型均会一定程度影响肌肉的嫩度。一般认为,猪肉IMF含量达到2%左右可以达到比较好的食用品质,大多数IMF相关基因富集于脂质代谢途径,包括脂肪酸转运和摄取、脂肪酸β氧化、脂质合成、脂质储存和脂肪分解[3];宰后48 h最终pH与肉的嫩度存在强相关性[4];提高肌肉Ⅰ型肌纤维占比可以显著改善肌肉嫩度[5]。
猪肉品质受遗传特性、营养、饲养管理、屠宰方式等众多因素的影响。遗传因素决定了猪肉品质的基本潜力,而营养调控则在充分发挥这一潜力方面起着至关重要的作用。研究表明,可通过营养调控手段促进肌肉纤维类型的转化从而改善肉品质,提高Ⅰ型(慢速氧化型)肌纤维的比例可显著改善肉的品尝特性[6]。另外,环境应激、氧化应激等引发的蛋白质修饰、细胞凋亡等最终也会影响肉品质[7],缓解机体应激的营养物质可以一定程度缓解应激导致的肉品质劣化。改善猪肉品质的营养物质代谢路径主要涉及机体脂质调控代谢、蛋白质合成、肌纤维类型转变、应激缓解等。通过合理调整营养物质组成、添加功能性添加剂及选择不同饲料原料种类和适当的饲料原料加工,可以有效改善猪肉的感官品质、营养价值和加工特性。相较于遗传育种手段,通过营养调控手段改善猪肉品质具有时间短、见效快、成本低、易于操作等优势。本文将从调整营养物质组成、添加功能性添加剂、选择饲料原料种类和加工方式等改善猪肉品质营养调控手段的研究进展展开综述。

1 调整营养物质组成

1.1 能量

在调整饲粮能量时应密切关注能量与蛋白质、赖氨酸(Lys)的比值。有研究发现,采用等蛋白质氨基酸平衡饲粮[粗蛋白质(crude protein,CP)水平12.01%、标准回肠可消化赖氨酸(standard ileal digestibility-lysine,SID-Lys)水平0.60%、标准回肠可消化蛋氨酸(standard ileal digestibility-methionine,SID-Met)水平0.20%、标准回肠可消化苏氨酸(standard ileal digestibility-threonine,SID-Thr)水平0.44%、标准回肠可消化色氨酸(standard ileal digestibility-tryptophan,SID-Trp)水平0.13%)]饲喂育肥猪,当饲粮消化能(digestible energy,DE)从13.02 MJ/kg提高至15.22 MJ/kg,可显著提高肌肉IMF含量,降低蒸煮损失及嫩度,但同时也显著提高24 h黄度值;而当饲粮DE从13.02 MJ/kg降低至10.84 MJ/kg时,对肉品质无显著影响[8]。因此,饲粮有效能与Lys的比值对育肥猪的生长、胴体性状和肉品质特性以及蛋白质合成和脂肪沉积速率有很大影响,提高饲粮DE与蛋白质比值,可导致猪肌肉IMF含量升高。但是,建议育肥阶段饲粮能量水平不应过高,过高能量水平会影响最终的胴体性状[8],可能导致腹部大量脂肪堆积。

1.2 蛋白质和氨基酸

低蛋白质或低氨基酸饲粮增加了脂肪在皮下、内脏和IMF组织中的沉积,导致瘦肉率降低,这主要是由于饲粮中Lys限制和必需氨基酸不平衡所致[9]。在不同氮营养素饲粮模式下,降低生长育肥猪饲粮CP水平并补充前4种限制性氨基酸,可在不影响饲料转化效率和主要肌肉组织重量的前提下,改善肌肉肉色。为研究饲粮CP水平对肉品质的影响,采用等能低蛋白质氨基酸平衡饲粮试验设计,设置生长期(30~60 kg)饲粮:正常蛋白质(CP水平18%)、低蛋白质(CP水平16%)和极低蛋白质(CP水平13%)饲粮,即降低饲粮CP水平2~5个百分点;设置育肥前期(60~100 kg)饲粮:正常蛋白质(CP水平16%)、低蛋白质(CP水平13%)和极低蛋白质(CP水平10%)饲粮,即降低饲粮CP水平3~6个百分点。其中,生长期饲粮含DE 14.2 MJ/kg、SID-Lys 0.97%、SID-Met 0.57%、SID-Thr 0.60%、SID-Trp 0.17%,育肥前期饲粮含DE 14.2 MJ/kg、SID-Lys 0.72%、SID-Met 0.40%、SID-Thr 0.50%、SID-Trp 0.13%;结果表明,饲喂低蛋白质或极低蛋白质饲粮可显著提高猪肌肉IMF及单不饱和脂肪酸(monounsaturated fatty acids,MUFA)含量,显著降低肌肉多不饱和脂肪酸(polyunsaturated fatty acid,PUFA)含量及剪切力;并且饲喂低蛋白质饲粮可显著提高肌肉红度值及肌肉Ⅰ型和Ⅱa型肌纤维比例,饲喂极低蛋白质饲粮可显著提高肌肉中牛磺酸和鲜味氨基酸含量[10]。
也有研究发现,在70~100 kg体重阶段,当设置饲粮净能均为2 475 kcal/kg(1 kcal=4.184 kJ)时,分别用正常蛋白质(CP水平13.5%)、低蛋白质(CP水平10.76%)、极低蛋白质(CP水平8.02%)饲粮饲喂育肥猪,低蛋白质和极低蛋白质饲粮均可提高肌肉IMF含量,降低肌肉持水能力及剪切力;且饲喂极低蛋白质饲粮提高了肌肉PUFA含量,并降低饱和脂肪酸(saturated fatty acid,SFA)含量,显著降低肌肉中异味物质如组胺、亚精胺、精胺和酪胺含量[11]。但是,本实验室研究发现,在育肥后期(100~128 kg),将氨基酸平衡饲粮中的CP水平从16%降低至10%,会显著提高第1肋和最后肋的背膘厚,但是会提高肌肉的黄度值[12]。根据NRC(2012)[13]推荐,30~75 kg体重阶段猪,饲粮CP水平为17%;75~100 kg体重阶段猪,饲粮CP水平为12.13%;100~135 kg体重阶段猪,饲粮CP水平为10.44%。现有低蛋白质饲粮试验中设计的正常蛋白质饲粮组的CP水平往往高于NRC(2012)推荐量,因此在设计低蛋白质饲粮组的时候是有失偏颇的。并且,低蛋白质氨基酸平衡饲粮即使满足氨基酸需求,过度降低饲粮CP水平也可能因非必需氨基酸合成受限及蛋白质原料供应不足导致猪生长性能受损。因此,在确保育肥猪生长性能、屠宰性能和肉品质的前提下,应根据各体重阶段营养物质需要量精准配制饲粮,建议试验设计正常蛋白质饲粮组的CP水平应该使用NRC(2012)的推荐量。
作为哺乳动物的必需氨基酸,支链氨基酸[(branched chain amino acids,BCAA;包括亮氨酸(leucine,Leu)、异亮氨酸(isoleucine,Ile)、缬氨酸(valine,Val)]在能量稳态、脂质代谢、蛋白质合成中起着至关重要的作用。骨骼肌是体内BCAA转氨基代谢的主要部位,饲粮中BCAA添加水平及BCAA之间的比例均会影响机体脂质代谢。Leu可在体内转化为乙酰辅酶A(acetyl-CoA),并调节猪的脂肪酸代谢。Leu及其代谢产物α-酮异己酸酯(α-ketoisocaproate,KIC)和β-羟基-β-甲基丁酸酯(β-hydroxy-β-methyl butyrate,HMB)被认为是调节猪肉脂肪酸沉积的重要物质。KIC导致猪背最长肌中脂肪酸转运蛋白(fatty acid transport protein,FATP)、乙酰辅酶A羧化酶(acetyl-CoA carboxylase,ACC)、脂肪组织甘油三酯脂肪酶(adipose tissue triglyceride lipase,ATGL)、CCAAT增强子结合蛋白α(CCAAT-enhancer-binding protein α,C/EBPα)、过氧化物酶体增殖物激活受体γ(peroxisome proliferator-activated receptor γ,PPARγ)和固醇调节元件结合蛋白1c(sterol regulatory element-binding protein 1c,SREBP1c)基因表达降低;HMB主要通过腺苷依赖蛋白激酶(adenosine 5'-monophosphate-activated protein kinase,AMPK)-哺乳动物雷帕霉素靶蛋白(mammalian target of rapamycin,mTOR)信号通路调节骨骼肌中的脂质合成代谢和分解代谢[14]。Leu可通过脂肪因子-mTOR-沉寂信息调节因子1(silent information regulator 1,SIRT1)信号通路促进育肥猪脂肪组织脂肪分解[15]。高Ile水平(0.53%)可增加肌肉24 h pH、肌节长度,并有减少肌肉滴水损失的趋势[16]。Val∶Ile比例降低或提高(约为正常的0.5或者2.0倍)均导致背膘增厚、瘦肉率降低,肌肉滴水损失和蒸煮损失提高,IMF含量升高[17]。在低蛋白质饲粮(CP水平12%)中,Leu∶Ile∶Val比例设置为2∶1∶1或2∶1∶2可加速脂肪因子的分泌和脂肪组织中的脂肪酸氧化[18]。
除BCAA外,其他功能性氨基酸也会影响肉品质。饲粮中添加牛磺酸可改善育肥猪肉品质,主要是通过钙调磷酸酶(calcineurin,CaN)/活化T细胞核因子c1(nuclear factor of activated T cells c1,NFATc1)途径促进肌纤维类型从糖酵解纤维向氧化纤维的转化[19]。饲粮中添加1%谷氨酸(glutamate,Glu)和1.44%丙氨酸(alanine,Ala),可显著降低平均背膘厚(降低33.88%),并且降低肌肉和背脂中SFA含量[20]。本实验室研究发现,饲粮中添加1%精氨酸(arginine,Arg)可以降低肌肉48 h滴水损失,提高IMF含量,提高机体抗氧化能力[21]。Arg和Glu补充剂在养猪业中对提高猪肉品质具有相当大的互补优势,饲粮中添加1% Arg+1% Glu可显著降低平均背膘厚,降低肌肉和背脂中SFA含量,提高IMF含量[20]。通过额外添加0.57%甘氨酸(glycine,Gly)并减少豆粕用量,使低蛋白质饲粮(CP水平12%)的丝氨酸(serine,Ser)和Gly的比例调整为1∶2,总量为1.44%时,可提高59~100 kg体重阶段猪的肌肉IMF含量,并提高肌肉中氧化型肌纤维占比,从而改善肉品质[22]。

1.3 脂肪酸

饲粮脂肪酸参与猪骨骼肌和脂肪组织的脂滴生成和能量代谢。研究发现,随着饲粮中n-6/n-3 PUFA比率的降低,皮下脂肪组织和背最长肌的n-6/n-3 PUFA比例降低;相较于饲粮中n-6/n-3 PUFA比例2.5∶1、5∶1和10∶1,饲粮中n-6/n-3 PUFA的适宜比例为1∶1,可以增加猪的肌肉重量,降低猪的脂肪组织重量,说明饲粮中n-6/n-3 PUFA比例可显著影响体脂沉积[23]。Wang等[24]就饲粮中补充PUFA对猪肉品质的影响开展Meta分析发现,高和低水平的PUFA补充均可增加肌肉IMF含量。此外,补充PUFA对肉色亮度值和24 h pH的影响与饲粮PUFA添加水平和猪的生长阶段有关,饲粮中添加PUFA改善猪的肉品质,主要表现在肌肉IMF含量的提高。在补充PUFA的时候,值得注意的是,饲粮PUFA水平要控制在饲粮粗脂肪水平的50%以下,否则容易氧化,发生黄膘肉,饲粮PUFA水平上限推荐量为1.5%。
最常用的PUFA补充剂是共轭亚油酸(conjugated linoleic acid,CLA)。CLA是亚油酸的二级衍生物,在调节糖、脂代谢中起着重要的生理作用。饲喂添加0.75% CLA的饲粮时,猪的眼肌面积随体重增加而线性增加,但是第10肋、第1肋和最后肋的背膘厚线性降低,且眼肌组织和皮下脂肪组织中SFA和CLA异构体含量均线性增加[25]。同时,本实验室研究发现,用添加1.25%~2.50% CLA(纯度为80%,其中顺式-9、反式-11 CLA为36.91%,反式-10、顺式-12 CLA为37.46%,其他异构体为5.37%)饲粮饲喂60~100 kg生长育肥猪,可提高瘦肉率(提高3.5%~4.7%)和IMF含量,并降低第1肋和第10肋的背膘厚,且缓解机体脂质氧化情况[26]。进一步研究饲粮中添加CLA对猪背最长肌蛋白质组的影响发现,CLA显著影响了肌肉能量代谢、脂肪酸氧化与合成、氨基酸代谢、防御和转运等过程相关蛋白质的丰度,IMF含量的增加与碳酸酐酶3和天冬氨酸转氨酶丰度的增加呈正相关[27]。CLA主要通过激活过氧化物酶体增殖物激活受体α(peroxisome proliferator-activated receptor α,PPARα)信号通路增加脂肪组织脂肪酸结合蛋白(adipocyte fatty acid binding protein,A-FABP)的表达丰度,并调控与IMF沉积相关的基因和酶的表达,从而增加IMF含量[28]。
α-亚麻酸是一种n-3 PUFA,亚麻籽中富含α-亚麻酸。研究发现,饲粮中添加5%~10%亚麻籽能一定程度提高生长育肥猪生长性能,增加背最长肌中n-3 PUFA含量,降低n-6/n-3 PUFA比例,且对胴体性状及肉品质无负面影响[29]。也有研究发现,饲粮中添加3.0%亚麻籽油可提高肌肉n-3脂肪酸含量,并降低了n-6/n-3 PUFA比例,但是同时也显著提高了育肥猪背膘厚[30]。以上2个试验结果中背膘厚的不同变化可能是添加的亚麻籽、亚麻籽油中α-亚麻酸含量不同所致。短链脂肪酸(short chain fatty acid,SCFA)是后肠细菌发酵不消化碳水化合物的主要产物,可作为肠道上皮细胞的能量物质。生长猪回肠灌注SCFA可增加眼肌面积和屠体重,并降低肌肉滴水损失[31]。

1.4 中性洗涤纤维(neutral detergent fiber,NDF)

适度增加NDF摄入量可以改善猪的脂质代谢并改变肉品质[32]。饲粮中NDF水平由14%提高到16%,可以降低背膘厚,提高肌肉风味氨基酸含量[33]。

1.5 其他营养物质

体内研究指出,提高饲粮钙水平(0.9%升高至1.5%)对猪体重无影响,但改善了猪背最长肌的肉色,减少了背膘厚度,增加了肌肉IMF含量;体外研究发现,将培养基中钙含量从2 mmol/L升高至3 mmol/L对C2C12细胞增殖没有显著影响,但促进细胞分化为慢肌纤维,并增加细胞中甘油三酯的积累;钙通过降低线粒体β氧化关键酶活性调控细胞内脂肪代谢[34]。
维生素E(vitamin E,VE)作为一种常见的抗氧化剂,可以缓解运输应激,并有助于保持猪的肉品质。饲喂325 mg/kg维生素E可减少肌肉的蒸煮损失,增加总PUFA、C18∶2n-6和C18∶3n-3的比例[35]。
微量元素铁可以改善肌肉肌红蛋白含量,从而改善肌肉色泽。饲粮中添加45~60 mg/kg氨基酸络合铁可显著或极显著提高生长育肥猪背最长肌肉色红度和饱和度值[36]。
硒是机体必需微量元素之一。硒可提高动物机体抗氧化能力、调节免疫应答、促进生长发育并参与营养物质代谢和提高繁殖性能等。NRC(2012)[13]推荐,对于70~100 kg体重阶段猪,硒推荐量是0.152 mg/kg;对于100~130 kg体重阶段猪,硒推荐量是0.146 mg/kg。硒是畜禽饲粮中重要的抗氧化剂之一,饲粮中通常添加的是无机硒亚硒酸钠。有机硒比无机硒生物利用率更高,有机硒主要包括蛋氨酸硒、酵母硒。本实验室研究发现,在肥育猪饲粮中添加0.15~0.30 mg/kg蛋氨酸硒可提高硒在组织中的沉积,提高机体抗氧化能力[37]。相较于亚硒酸钠,饲粮中添加0.30 mg/kg酵母硒可使猪背膘厚下降7.76%,眼肌面积提高4.13%,肌肉pH显著提高[38]。同时,在研究不同酵母硒水平(0.3~5.0 mg/kg)对育肥猪肉品质的影响发现,低酵母硒水平(0.3~1.0 mg/kg)可提高肌肉红度值,提高肌肉酪氨酸含量,改善机体抗氧化能力,且高水平(5.0 mg/kg)酵母硒可引起18种脂质分子代谢上调[39]。

2 添加功能性添加剂

2.1 植物提取物

在养殖业减抗替抗的大背景下,包括中草药在内的植物提取物受到行业广泛关注。该类植物提取物包括多酚、黄酮等活性成分,主要通过调控机体蛋白质合成、脂质代谢及改善机体抗氧化活性等改善肉质。研究发现,饲粮中添加800 mg/kg苹果多酚可降低猪背膘厚和腹脂指数,降低肌肉的亮度值和黄度值,显著提高肌肉CP含量、必需氨基酸、风味氨基酸和总氨基酸比例以及氨基酸转运蛋白mRNA表达水平[40]。本实验室发现,饲粮中添加100 mg/kg甜叶菊绿原酸可改善生长育肥猪机体抗氧化能力[41]。同时,其他研究也发现,饲粮中添加0.05%绿原酸可显著提高屠宰率、瘦肉率和背最长肌pH,显著增加股二头肌粗脂肪含量,显著降低背最长肌红度值,改善机体抗氧化能力[42]。此外,有报道称,饲粮中添加绿原酸改善了猪血清游离氨基酸谱,并促进氨基酸转运到骨骼肌,提高了背最长肌和股二头肌中钠离子依赖的中性氨基酸转运蛋白2(sodium-coupled neutral amino acid transporter 2,SNAT2)mRNA表达水平,促进了磷酸化蛋白激酶B(p-protein kinase B,p-Akt)的表达,并激活了背最长肌肉中mTOR-p70核糖体S6激酶蛋白(p70 ribosomal s6 kinase protein,S6K1)-真核生物翻译起始因子4e结合蛋白1(eukaryotic translation initiation factor 4e-binding protein 1,4EBP1)轴,降低了肌肉萎缩F-box蛋白(muscle atrophy F-box,MAFbx)mRNA表达丰度,增强了肌肉蛋白质的生物合成[43]。
饲粮中添加75 mg/kg鞣花酸增加了猪肌肉24 h pH、红度值、IMF含量和风味氨基酸含量,降低了背最长肌的滴水损失和剪切力;鞣花酸通过激活NFATc1和增殖激活受体-γ辅激活因子-1α(proliferator-activated receptor-gamma coactivator-1alpha,PGC-1α)信号通路,参与肌红蛋白表达[44]。饲粮中添加50~200 mg/kg葡萄籽原花青素提取物可增加肌肉24 h pH、红度值、CP含量,并降低剪切力、48 h滴水损失、乳酸糖酵解电位,提高PUFA含量以及PUFA与SFA的比例[45]。饲粮中添加400~600 mg/kg白藜芦醇可提高肌肉红度值和IMF含量,并降低剪切力;白藜芦醇可激活Kelch样ECH相关蛋白1(Kelch-like ECH-associated protein 1,Keap1)-核因子红系2相关因子2(nuclear factor erythroid 2-related factor 2,Nrf2)抗氧化通路,直接与Keap1结合,降低Keap1-Nrf2蛋白互作的紧密性,从而发挥其优异的抗氧化活性[46]。有报道称,肥育猪后期饲粮中添加500 mg/kg大豆异黄酮(isoflavone,ISO)可提高其日增重,增加血液中脂联素含量,上调背最长肌中PPARγ的mRNA表达丰度,增加IMF的沉积,从而改善猪肉品质[47]。而本实验室发现,饲粮中添加超高剂量(640 mg/kg)大豆苷元虽然可改善肌肉中的抗氧化酶活性,但是在肝脏、血浆和脂肪组织中具有促氧化作用[48]。饲粮中添加200~400 mg/kg大蒜素可显著降低猪背膘厚度,提高背最长肌pH、肌苷酸含量,降低滴水损失、肌肉剪切力,提高最长肌中总氨基酸、总必需氨基酸及鲜味氨基酸含量[49]。饲粮中添加300 mg/kg姜黄素降低了金华猪的背膘厚和背部脂肪脂肪细胞大小,还提高了肌内粗脂肪含量和眼肌面积,下调了金华猪背部脂肪中脂肪生成相关基因PPARγ、C/EBPα的mRNA表达丰度,且提高了背最长肌中PPARγ的mRNA表达丰度[50]。饲粮中添加0.025%~0.050%黄芪、党参和大蒜素混合物,可显著提高育肥猪肌肉的红度值,并降低亮度值[51]。综上所述,植物提取物改善肉品质方面有较多研究,大部分集中在多酚和黄酮类物质,在肉品质改善上具有明显的剂量效应,过量添加往往不利于饲粮的适口性,且会大大提高成本。

2.2 益生菌及益生元

益生菌一方面可以通过改善机体肠道菌群从而进一步改善猪肉品质,另一方面可以通过调节仔猪相关基因和蛋白质的表达等方式来改善猪肉品质,包括调节肌纤维类型比例[52]、促进IMF沉积以及调节骨骼肌代谢[53]。本实验室研究发现,从仔猪阶段就开始在饲粮中添加5×1010 CFU/kg的罗伊氏乳杆菌1(Lactobacillus reuteri 1)可显著降低育肥猪背最长肌滴水损失和剪切力,提高肌肉中肌苷酸和谷氨酸含量,并改变肌纤维类型,从而改善肥育猪肉品质[54]。Chang等[55]研究发现,在饲粮中添加益生菌混合制剂可降低背最长肌的剪切力,并提高PUFA和ω脂肪酸(ω3和ω6)含量,而降低MUFA含量。β-葡聚糖是一种功能性多糖,广泛分布于真菌、细菌和谷类种子(燕麦、黑麦和大麦等)的细胞壁中,具有促进肠道益生菌生长、增加SCFA产生及改善肠道屏障等功能。研究发现,饲粮中添加100 mg/kg β-葡聚糖可提高猪肌肉pH、红度值、IMF含量,降低滴水损失[56]。

2.3 酶制剂

酶制剂常用于辅助改善畜禽营养物质消化吸收功能,猪生产中常用的酶制剂包括蛋白酶、碳水化合物酶、植酸酶等。添加外源蛋白酶是提高猪CP消化率并降解饲粮中的蛋白质抗营养物质的一种营养策略。研究发现,饲粮中补充蛋白酶制剂不仅可以改善营养物质消化率和生长性能,还可提高育肥猪的瘦肉率和眼肌面积[57];饲粮中添加木聚糖酶、甘露聚糖酶和蜘蛛酶(Arazyme®)混合物,可显著降低背最长肌的蒸煮损失和剪切力[58];在含有挤压大豆种子和菜籽粕的饲粮中添加植酸酶可显著改善肉色[59]。

2.4 其他功能性物质

作为新一代无毒、无害、安全的饲料添加剂,核苷酸在促进肠道成熟、肠黏膜发育、生长性能和脂质代谢方面发挥着重要作用。研究发现,饲粮中添加0.2%腺苷(adenosine,AMP)可降低肌肉滴水损失,提高红度值,但是也同时会降低肌肉中游离氨基酸含量[60]。二丁酰-环腺苷(db-cAMP)是一种具有口服活性的细胞渗透衍生物,具有与内源性环腺苷相同的功能,后者在许多生物活性中起着信号转导的关键作用,如调节细胞生长、酶活性、脂肪分解和基因表达。db-cAMP通过增加生长激素(growth hormone,GH)-胰岛素样生长因子-1(insulin-like growth factor-1,IGF-1)轴和前阿片黑素细胞皮质激素(pro-opiomelanocortin,POMC)系统的活性来发挥抑制脂肪合成和积累的功能,饲粮中添加10~15 mg/kg的db-cAMP可降低猪腹脂率、第1肋和第10肋的背膘厚,提高瘦肉率和眼肌面积,且育肥早期效果优于育肥后期[61-62]。
甜菜碱是Gly的三甲基衍生物,存在于大多数生物体中。在动物中,甜菜碱可用作有机渗透保护剂以保护细胞免受损伤,或通过转甲基化作为甲基供体,甜菜碱可通过上调脂肪酸转位酶(fatty acid translocase,FAT/CD36)、FATP和脂肪酸结合蛋白3(fatty acid binding protein 3,FABP3)等基因参与脂质代谢[63]。饲粮中添加2 500 mg/kg甜菜碱可降低育肥猪背膘厚和肌肉滴水损失,提高肌肉IMF含量,提升血清脂质代谢和抗氧化能力[64]。
γ-氨基丁酸(γ-aminobutyric acid,GABA)是一种在动物体内起抑制神经作用的非蛋白质氨基酸,是中枢神经系统中最主要的抑制性神经递质,同时对动物正常生理功能也具有重要调节作用。饲粮中添加200 g/t GABA可提高育肥猪的屠宰率、眼肌面积及肌肉CP含量和IMF含量,并降低平均背膘厚和肌肉滴水损失[65]。肌肽为β-丙氨酰基-L-组氨酸,本实验室研究发现,饲粮中添加100 mg/kg肌肽可提高宰后45 min、24 h和48 h肌肉pH,降低宰后48 h肌肉滴水损失,提高宰后45 min肌肉红度值,并提高机体抗氧化活性[66]。肌酸是一种含氮的有机酸,可由Arg、Gly和甲硫氨酸(methionine)3种氨基酸合成,主要在肌肉中储存和使用,为细胞提供能量。饲粮中添加一水肌酸使背最长肌宰后24 h pH显著增加,滴水损失和压榨损失显著降低[67];一水肌酸与α-硫辛酸联用(0.5 g/kg的一水肌酸和0.1 g/kg的α-硫辛酸)显著增加生长育肥猪眼肌面积,降低生长育肥猪背最长肌滴水损失和压榨损失[68]。
胍基乙酸是一种新型营养补充剂,是合成肌酸及磷酸肌酸在体内的直接前体。研究发现,补充胍基乙酸可以改善育肥猪的生长性能和肉品质,改变肌酸代谢,增加猪组织中的ATP负荷和肌肉AMP的循环[69-70]。饲粮中添加500 mg/kg胍基乙酸可显著提高屠宰率和眼肌面积,促进肌纤维分化[71]。氧化三甲胺(TMAO)是一种微生物群衍生的代谢产物。饲粮中补充1 g/kg TMAO有降低体重的趋势,但是显著增加背膘厚,增加背最长肌IMF含量,这是因为TMAO可显著上调肌细胞中脂肪酸合成酶(fatty acid synthase,FASN)和SREBP1的mRNA表达,增加脂肪细胞面积,下调皮下脂肪中肉碱棕榈酰基转移酶1B(carnitine palmitoyltransferase 1B,CPT1B)的表达[72]。

3 选择饲料原料种类和加工方式

3.1 饲料原料种类

前期有关改善肉品质的饲粮营养物质含量调整和添加剂添加大都基于玉米-豆粕型基础饲粮。近年,为响应农业农村部《饲用豆粕减量替代三年行动方案》,业内做了大量豆粕减量替代的研究工作。用黑水虻虫粉替代饲粮中的玉米、豆粕、豆油等,可提高眼肌面积、IMF含量、肌苷酸含量、Ⅱa型肌纤维mRNA表达[73]。用菜籽粕和蚕豆替代14.26%豆粕,可降低肌肉亮度和黄度值,改善机体抗氧化能力[74]。用苜蓿叶粉替代25%豆粕,可增加背最长肌IMF含量、熟肉率、红度值和抗氧化活性,改善肌肉氨基酸组成,增加MUFA含量,提高背最长肌脂质合成和抗氧化相关基因mRNA表达[75]。用桑叶粉部分替代饲粮中的玉米、豆粕和麦麸,可增强机体抗氧化能能力,提高肌肉PUFA含量[76]。饲粮淀粉来源也会影响肉品质,豌豆淀粉饲粮相较于木薯淀粉饲粮可显著提高眼肌面积和无脂瘦肉率,提高肌肉pH及IMF、肌苷酸含量,降低背膘厚及肌肉滴水损失、剪切力,提高肌肉中风味氨基酸和脂肪酸含量,降低n-6/n-3 PUFA比值[77]。这些基础研究为非常规饲料原料在生产中的应用提供了坚实的理论基础。

3.2 饲料原料加工方式

饲料原料经过处理可以改变原料的营养价值、消化率和利用效率,从而影响猪的生长性能和最终的肉品质。发酵饲料是利用微生物发酵工艺发酵饲料原料而得到的饲料,微生物发酵可降解饲料中的抗营养因子水平,促进营养物质消化吸收,特别是对于非常规饲料原料的应用具有重要意义[78]。同时,发酵用微生物及其代谢产物可改善机体肠道健康,从而进一步改善肉品质。本实验室研究发现,在低蛋白质饲粮(CP水平11.81%)的基础上,使用发酵菜籽粕部分替代豆粕(50%),对60~110 kg育肥猪的生长性能和胴体性状无负面影响,但可显著提高猪肉的肌苷酸含量和感官评分,并具有改善猪肉脂肪酸组成的作用,其中油酸、MUFA含量均显著提高,亚油酸、二十烷二烯酸及PUFA含量均显著下降[79]。复合益生菌固态发酵饲粮可通过提高胰岛素/蛋白激酶B(Akt)/mTORC1蛋白合成通路,激活叉头框蛋白O1(fork head box O1,Foxo1)/MAFbx通路,同时调节核糖体蛋白和参与肌肉收缩和肌肉肥大的蛋白,提高眼肌面积和瘦肉率,降低背膘厚,降低肌肉剪切力和滴水损失[80]。研究表明,饲喂发酵饲料能够提高猪肉风味前体物(如风味氨基酸、脂肪酸)含量,从而提高猪肉的风味和口感[81]。但是,饲料发酵涉及发酵过程,耗时耗力,大大提高了饲料成本;且受发酵工艺和发酵菌群的影响,饲喂效果可能不稳定。而且发酵饲料一般都是液体状态,目前产业上还没有大面积推广配套的液体饲喂系统。

4 复配方案

由于单一营养调控手段往往无法全面发挥其改善肉品质的功效,因此有很多应用研究将多种营养调控手段复配,如表1所示。营养调控复配方案的基本思路是将调节蛋白质合成及调节脂质代谢的物质组合,从抗应激及改善机体抗氧化活性着手,辅以添加抗氧化剂、抗应激剂等。但是,考虑到饲料成本问题,不建议复配3种以上的营养调控策略。
表1 改善猪肉品质营养复配方案

Table 1 Nutritional complex supplementation strategies for enhancing pork quality

项目
Items		 饲喂体重阶段
Feeding weight
stage/kg		 试验周期
Experimental
period/d		 效果
Effect		 文献来源
Literature source
1%谷氨酸+1.44%L-丙氨酸
1% Glu+1.44% L-Ala		 77~120 60 降低平均背膘厚,降低肌肉和
背脂中饱和脂肪酸含量		 [20]
1%精氨酸+1%谷氨酸
1% Arg+1% Glu		 77~120 60 降低平均背膘厚,降低肌肉和背脂中
饱和脂肪酸含量,提高肌内脂肪含量		 [20]
0.25%亮氨酸+0.025%绿原酸0.25%
Leu+0.025% chlorogenic acid		 68~88 25 降低背膘厚,提高瘦肉率、背最长肌
pH、股二头肌粗脂肪含量,
改善机体抗氧化能力		 [42]
100 mg/kg维生素E+0.3 mg/kg酵母硒
100 mg/kg VE+0.3 mg/kg yeast selenium		 60~110 60 提高屠宰率、肌肉红度,降低肌肉黄度,
缓解氧化鱼油导致的氧化应激反应		 [82]
200 mg/kg维生素E+0.3 mg/kg酵母硒+20 mg/kg大豆异黄酮
200 mg/kg VE+0.3 mg/kg yeast
selenium+20 mg/kg ISO		 100~130 28 提高眼肌面积,降低最后肋背膘厚、
滴水损失,改善机体抗氧化能力		 [83]
0.03%姜黄素+2.5%二十二碳六烯酸
0.03% curcumin+2.5% docosahexaenoic acid		 90~135 40 增加眼肌面积,降低背膘厚、肌肉滴水损
失,增加IMF含量,增加背最长肌甘氨酸、
脯氨酸、棕榈油酸和硬脂酸含量,提高肌球
蛋白重链Ⅱa、肌球蛋白重链Ⅱx和固醇
调节元件结合蛋白1的mRNA
相对表达水平,改善机体抗氧化活性		 [84]
0.5 g/kg一水肌酸+0.1 g/kg α-硫辛酸
0.5 g/kg creatine monohydrate+0.1 g/kg α-lipoic acid		 60~70 50 增加生长育肥猪眼肌面积,降低生长育肥
猪背最长肌滴水损失和压榨损失		 [68]
1 g/kg胍基乙酸+0.5 g/kg甜菜碱
1 g/kg guanidinoacetic acid+0.5 g/kg betaine		 15 降低育肥猪的平均背膘厚,降低肌肉
硬度、滴水损失、蒸煮损失、剪切力,
提高宰后45 min、24 h pH		 [85]
500 mg/kg胍基乙酸+200 mg/kg甜菜碱
500 mg/kg guanidinoacetic acid+
200 mg/kg betaine		 86~110 38 提高屠宰率和眼肌面积,促进肌纤维
分化,联合添加胍基乙酸与甜菜碱的
促生长效果优于单独添加胍基乙酸		 [71]
200 mg/kg维生素C+100 mg/kg
维生素E+1 500 mg/kg甜菜碱+
2 500 mg/kg牲血素
200 mg/kg VC+100 mg/kg VE+
1 500 mg/kg betaine+
2 500 mg/kg sanguin		 75~120 提高肌肉红度值,降低剪切力和滴水损失 [86]

5 小结与展望

综上所述,营养调控手段改善肉品质主要是指通过营养物质调控机体脂肪代谢、蛋白质合成、肌纤维类型转化、改善机体应激等方面从而改善猪的胴体性状及肉品质。目前,营养调控与遗传背景、环境因素的交互作用机制研究有限。机体能量代谢、蛋白质合成、脂肪酸合成、转运及脂质累积的协同调节机制尚不完全清楚,不同营养调控措施之间的交互作用研究不足。且有的营养物质在改善一些肉品质指标的同时可能会劣化另一些肉品质指标,如有的营养物质在促进IMF沉积的同时会提高背膘厚并降低瘦肉率,因此,这些营养物质在改善猪肉品质的程度上难以量化评估。开发新型、高效、安全的功能性饲料添加剂是未来产业的一个研究热点。同时,有些功能添加剂在改善肉品质效果上不稳定,可能与饲喂时间长短、添加物质的稳定性、生产工艺、添加剂量、分子结构、物质纯度等有关,部分研究结果在实际生产中的应用效果有待验证。有些功能添加剂虽然改善肉品质效果好,但是成本高,不利于产业推广,应加强新型功能添加剂在实际生产中的应用研究,评估其经济效益和可行性。目前,已有研究对于营养调控对猪肉品质的长期影响及其安全性评估不足。安全、高效、低成本的复配产品将是产业的发展方向之一。

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