Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 1: 335 - 345

DOI: https://doi.org/10.12418/CJAN2026.026

RESEARCH PAPER

Effects of α-Lipoic Acid on Muscle Quality, Texture Characteristics, Antioxidant Capacity and Muscle Fiber Types in White-Feathered Broilers

  • JIN Shuning ,
  • YIN Yue ,
  • LI Xiang ,
  • ZHANG Xiwen ,
  • LUO Qi ,
  • FENG Xinhui ,
  • ZHAO Yurong , *
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  • College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China
* professor, E-mail:

Received date: 2025-07-04

  Online published: 2026-01-13

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Abstract

This experiment was conducted to investigate the effects of α-lipoic acid (ALA) on muscle quality, textural characteristics, antioxidant capacity and muscle fiber types in white-feathered broilers. A total of 180 one-day-old healthy Arbor Acres (AA) broiler chicks with similar body weight were selected and randomly divided into 3 groups, with 6 replicates per group and 10 birds per replicate. The control group was fed a basal diet, while test groups Ⅰ and Ⅱwere supplemented with 100 and 300 mg/kg ALA in the basal diet, respectively. The trial lasted for 42 days. The results showed as follows: 1) compared with the control group, dietary supplementation with 100 and 300 mg/kg ALA significantly reduced drip loss, shear force, and yellowness (b*) value of the breast muscle (P<0.05); dietary supplementation with 100 mg/kg ALA significantly increased the adhesiveness of breast and thigh muscles as well as the redness (a*) value of breast muscle (P<0.05); dietary supplementation with 300 mg/kg ALA significantly decreased the lightness (L*) value of breast muscle and the shear force of thigh muscle (P<0.05), significantly improved the gumminess and chewiness of breast muscle (P<0.05), and significantly reduced the hydroxyproline (Hyp) content in breast muscle (P<0.05). 2) Dietary supplementation with different levels of ALA had no significant effects on the common nutritional component contents in breast and thigh muscles (P>0.05). 3) Compared with the control group, dietary supplementation with 100 and 300 mg/kg ALA significantly increased the glutathione peroxidase (GSH-Px) activity in both breast and thigh muscles (P<0.05), as well as the superoxide dismutase (SOD) activity in thigh muscle (P<0.05), while significantly reduced the malondialdehyde (MDA) content in thigh muscle (P<0.05). 4) Compared with the control group, dietary supplementation with 100 and 300 mg/kg ALA significantly upregulated the mRNA relative expression levels of myosin heavy chain Ⅰ (MyHC Ⅰ) and myosin heavy chain Ⅱa (MyHC Ⅱa) in breast muscle (P<0.05). Additionally, dietary supplementation with 100 mg/kg ALA significantly downregulated the mRNA relative expression level of myosin heavy chain Ⅱb (MyHC Ⅱb) in breast muscle (P<0.05). 5) Compared with the control group, dietary supplementation with 100 mg/kg ALA significantly up-regulated the mRNA relative expression levels of AMP-activated protein kinase (AMPK)α1, AMPKα2, sirtuin 1 (SIRT1), peroxisome proliferator-activated receptor-gamma coactivator-1α (PGC-1α), nuclear respiratory factor 1 (NRF1), and myocyte enhancer factor 2 (MEF2) in breast muscle (P<0.05); dietary supplementation with 300 mg/kg ALA significantly up-regulated the mRNA relative expression levels of AMPKα1, AMPKα2, SIRT1, and PGC-1α in breast muscle (P<0.05), while significantly down-regulated the mRNA relative expression level of AMPKγ3 in breast muscle (P<0.05). It is concluded that ALA can improve the muscle quality of white-feathered broiler by modifying textural characteristics, reducing collagen deposition, enhancing antioxidant capacity, upregulating expression of key genes in the AMPK/SIRT1/PGC-1α signaling pathway, and promoting the transition from glycolytic to oxidative muscle fiber types. Considering both the efficacy and cost-effectiveness, the addition of 100 mg/kg ALA to the diet of white-feathered broilers is recommended.

Key words： α-lipoic acid; white-feathered broilers; meat quality; antioxidant; muscle fiber types

Cite this article

JIN Shuning , YIN Yue , LI Xiang , ZHANG Xiwen , LUO Qi , FENG Xinhui , ZHAO Yurong . Effects of α-Lipoic Acid on Muscle Quality, Texture Characteristics, Antioxidant Capacity and Muscle Fiber Types in White-Feathered Broilers[J]. Chinese Journal of Animal Nutrition, 2026 , 38(1) : 335 -345 . DOI: 10.12418/CJAN2026.026

近年来,白羽肉鸡的肉品质下降问题日益凸显。研究发现,养殖过程中的氧化应激是导致该问题的重要原因:它会引发肌肉萎缩、胶原蛋白过度交联、肌纤维间脂肪沉积异常及炎性细胞浸润等一系列肌肉组织病变,从而严重影响肉品质[1]。这些肌肉组织病变主要表现为肉的硬度增加、色泽变暗、pH骤降,其表型与PSE肉极为相似[2]。因此,在肉鸡生产中寻找安全高效的天然抗氧化剂以缓解氧化应激,已成为当前的研究热点。
α-硫辛酸(ALA)作为一种天然抗氧化剂,在改善肉品质方面展现出潜力。研究表明,ALA能够通过调控肌肉氧化稳定性[3-4]与促进肌纤维类型转化,有效改善育肥猪[6]、羔羊[5]及肉牛[7]的肉品质。杨潇等[8]研究发现,在肉鸡饲粮中添加ALA可提升肉品质,并有效防止PSE肉的发生,但其具体作用机制尚未阐明。本团队前期研究证实,饲粮中添加100与300 mg/kg的ALA能改善白羽肉鸡的肠道形态、优化菌群结构,从而提高饲料转化率与生长性能[9]。基于此,本研究旨在进一步探究ALA对白羽肉鸡肉品质的影响,并从抗氧化层面及基因水平深入揭示其调控机制,以期为白羽肉鸡的健康养殖提供参考。

1 材料与方法

1.1 试验材料

爱拔益加(AA)肉仔鸡购自湖南顺成实业有限公司;ALA(纯度为99%)购自江苏润丰合成科技有限公司。

1.2 试验方法

1.2.1 试验设计

动物试验已经过湖南农业大学生物医学研究伦理委员会审批,审批号:伦审科2025第(151)号。选用遗传背景相同、体况健康的1日龄AA肉仔鸡180只,随机分为3组,每组6个重复,每个重复10只(公母各占1/2)。对照组饲喂玉米-豆粕型基础饲粮,试验Ⅰ组和试验Ⅱ组则在基础饲粮基础上分别添加100和300 mg/kg ALA。试验期为42 d(1~42日龄),期间鸡只自由采食和饮水,按照常规程序免疫。基础饲粮参照我国《鸡饲养标准》(NY/T 33—2004)中白羽肉鸡的营养需要量分阶段配制,其组成及营养水平见表1。
表1 基础饲粮组成及营养水平(风干基础)

Table 1 Composition and nutrient levels of basal diets (air-dry basis) %

项目
Items		 含量Content
1~21日龄
1 to 21
days of age		 22~42日龄
22 to 42
days of age
原料Ingredients
玉米Corn 58.01 62.23
豆粕Soybean meal (43%) 32.02 27.04
鱼粉Fish meal (67%) 3.24 2.98
大豆油Soybean oil 2.92 3.99
DL-蛋氨酸DL-Met (98%) 0.23 0.27
磷酸氢钙CaHPO4 1.30 1.10
石粉Limestone 0.98 1.09
食盐NaCl 0.30 0.30
预混料Premix1) 1.00 1.00
合计Total 100.00 100.00
营养水平Nutrient levels2)
代谢能ME/(MJ/kg) 12.54 12.96
粗蛋白质CP 21.50 19.52
钙Ca 1.00 1.00
非植酸磷NPP 0.47 0.41
赖氨酸Lys 1.16 1.02
蛋氨酸Met 0.58 0.59
蛋氨酸+半胱氨酸Met+Cys 0.93 0.91

1)预混料为每千克饲粮提供 The premix provided the following per kg of diets: VA 8 000 IU,VB1 2.25 mg,VB2 8.2 mg,VB6 2.8 mg,VB12 0.015 mg,VD3 2 000 IU,VE 20 IU,VK3 15 mg,生物素 biotin 0.18 mg,烟酸 nicotinic acid 35 mg,叶酸 folic acid 0.95 mg,泛酸 pantothenic acid 10 mg,Fe (as ferrous sulfate) 80 mg,Cu (as copper sulfate) 9 mg,Zn (as zinc sulfate) 100 mg,Mn (as manganese sulfate) 80 mg,Se (as sodium selenite) 0.30 mg,I (as potassium iodide) 0.40 mg。

2)粗蛋白质(GB/T 6432—2018)和钙(GB/T 6436—2018)为实测值,其他均为参照《中国饲料成分及营养价值表(2023年第34版)》所得计算值。CP (GB/T 6432—2018) and Ca (GB/T 6436—2018) were measured values, while others were calculated values with reference to China Feed Ingredient and Nutritional Value Tables (34th edition, 2023).

1.2.2 样品采集

饲养试验结束当天,在肉鸡禁食8 h后从每个重复中随机选择1只屠宰,采集右侧胸肌和腿肌样品,一部分直接用于品质、质构特性和常规营养成分含量的检测,一部分-20 ℃保存待测生化指标,还有一部分-80 ℃保存用于肌纤维类型及调节因子相关基因表达的测定。

1.2.3 肌肉品质的测定

参照《畜禽肉质的测定》(NY/T 1333—2007)测定胸肌和腿肌的pH、肉色[亮度(L*)、红度(a*)和黄度(b*)值]、失水率、剪切力和滴水损失。

1.2.4 肌肉质构特性的测定

取胸肌和腿肌样品,分别切割成1 cm×1 cm×1 cm大小的肉块,采用质构分析仪(TMS-Touch型,美国FTC公司)测定硬度、弹性、胶黏性、黏附性、内聚性和咀嚼性。

1.2.5 肌肉常规营养成分含量的测定

胸肌和腿肌中水分含量的测定采用直接干燥法(GB 5009.3—2016),蛋白质含量的测定采用凯氏定氮法(GB 5009.5—2016),脂肪含量的测定采用索氏抽提法(GB 5009.6—2016)。

1.2.6 肌肉中生化指标的测定

使用南京建成生物工程研究所生产的相应试剂盒,按照试剂盒说明书测定胸肌和腿肌中谷胱甘肽过氧化物酶(GSH-Px)和超氧化物歧化酶(SOD)活性、总抗氧化能力(T-AOC)、丙二醛(MDA)含量,以及胸肌中羟脯氨酸(Hyp)、糖原含量。

1.2.7 肌肉中肌纤维类型及调节因子相关基因表达的测定

将胸肌制备成匀浆液后提取总RNA,并采用琼脂糖凝胶电泳检测总RNA的完整性和纯度。使用反转录预混型试剂盒将RNA反转录为cDNA,将得到的cDNA存于-80 ℃冰箱待用。以β-肌动蛋白(β-actin)作为内参基因,登录NCBI网站用GenBank寻找肌球蛋白重链Ⅰ(MyHCⅠ)、肌球蛋白重链Ⅱa(MyHCⅡa)、肌球蛋白重链Ⅱb(MyHCⅡb)、腺苷酸活化蛋白激酶γ3(AMPKγ3)、腺苷酸活化蛋白激酶α1(AMPKα1)、腺苷酸活化蛋白激酶α2(AMPKα2)、沉默信息调节因子1(SIRT1)、辅助激活因子-1α(PGC-1α)、核呼吸因子1(NRF1)、肌细胞增强因子2(MEF2)等目的基因的序列,利用Primer Premier 5.0软件设计引物(表2)。以cDNA为模板,使用SYBR Green试剂盒,按照试剂盒说明书进行实时荧光定量PCR扩增。目的基因的mRNA相对表达量采用2-△△Ct法计算。所用到的试剂盒均购自湖南艾科瑞生物工程有限公司。
表2 引物序列

Table 2 Primer sequences

基因Genes 引物序列Primer sequences (5'—3') 登录号Accession No.
肌球蛋白重链Ⅰ
MyHCⅠ		 F:GGGCGCTATCATGCACTTTG
R:CTGCTCCACGTTCTGACCTT		 NM_204587.4
肌球蛋白重链Ⅱa
MyHCⅡa		 F:GACTTGGCCAGCAACATGGA
R:ATGCCTGTTTGCCTCTTGATAAC		 NM_001113709.2
肌球蛋白重链Ⅱb
MyHCⅡb		 F:ATCGACAACCTGCAACGAGT
R:ACGAGCTCTTTGAGCATTAACA		 NM_001319015.2
腺苷酸活化蛋白激酶γ3
AMPKγ3		 F:GGATGCTCACCATCACTG
R:TCTCCACGTCTCAATCTTGT		 XM_046921428.1
腺苷酸活化蛋白激酶α1
AMPKα1		 F:GTGGCATTTGGGGATACGGA
R:ATTCTTCCGTCGAACACGCA		 NM_001039603.2
腺苷酸活化蛋白激酶α2
AMPKα2		 F:CTTCGGCAAAGTCAAGGTTGG
R:GAAGAAGTCTGTTGGCGTGC		 NM_001039605.2
沉默信息调节因子1
SIRT1		 F:AGTAGTAGCGAAAGCGGCTC
R:TCGTTCCCTGCAGCTTCATT		 NM_001004767.2
辅助激活因子-1α
PGC-1α		 F:TGCAACCAGGACTCTGTATGG
R:GATGACCTCAGACTGGTCGC		 NM_001006457.2
核呼吸因子1
NRF1		 F:CCCATCCATCCGTAAGAGGC
R:ATTCTCCAAGGGTGCAGCTC		 NM_001030646.2
肌细胞增强因子2
MEF2		 F:TGTGTGACTGCGAGATTGCT
R:CGGGGCTTTCACAACCTTTG		 XM_046905690.1
β-肌动蛋白
β-actin		 F:TGCGTGACATCAAGGAGAAG
R:TGCCAGGGTACATTGTGGTA		 L08165

1.3 数据分析

试验数据使用Excel 2021进行汇总和整理后,采用SPSS 27.0软件进行单因素方差分析(one-way ANOVA),方差分析显著时采用Duncan氏法进行组间多重比较,以P<0.05作为差异显著标准,结果以平均值和均值标准误(SEM)表示。

2 结果

2.1 ALA对白羽肉鸡肌肉品质的影响

由表3可知,与对照组相比,饲粮中添加100和300mg/kgALA对胸肌和腿肌的pH45min、pH24 h和失水率均无显著影响(P>0.05),但显著降低了胸肌的滴水损失、剪切力和b*值(P<0.05);此外,饲粮中添加100 mg/kg ALA显著增加了胸肌的a*值,添加300 mg/kg ALA显著降低了胸肌的L*值及腿肌的剪切力(P<0.05)。
表3 α-硫辛酸对白羽肉鸡肌肉品质的影响

Table 3 Effects of ALA on muscle quality of white-feathered broilers

项目
Items		 对照组
Control group		 试验Ⅰ组
Test group Ⅰ		 试验Ⅱ组
Test group Ⅱ		 SEM P值
P-value
胸肌Breast muscle
pH45 min 6.67 6.53 6.63 0.071 0.181
pH24 h 5.97 5.98 5.98 0.042 0.975
滴水损失Drip loss/% 2.12a 1.60b 1.56b 0.186 0.014
剪切力Shear force/N 19.89a 17.28b 15.78b 0.898 <0.001
失水率Water loss rate/% 35.85 35.68 34.42 1.523 0.603
亮度值L* value 54.40a 53.92a 52.11b 0.444 <0.001
红度值a* value 6.99b 8.81a 6.04b 0.492 <0.001
黄度值b* value 12.38a 10.59b 9.34c 0.585 <0.001
腿肌Thigh muscle
pH45 min 6.54 6.41 6.57 0.076 0.114
pH24 h 6.42 6.37 6.52 0.063 0.086
滴水损失Drip loss/% 1.46 1.28 1.22 0.190 0.419
剪切力Shear force/N 19.98a 21.77a 14.67b 1.025 <0.001
失水率Water loss ratio/% 37.47 35.69 37.66 0.881 0.257
亮度值L* value 57.25 55.53 54.78 1.063 0.091
红度值a* value 9.06 8.32 7.89 0.677 0.181
黄度值b* value 10.72 10.66 10.19 0.409 0.388

同行数据肩标无字母或相同字母表示差异不显著(P>0.05),不同小写字母表示差异显著(P<0.05)。下表同。

In the same row, values with no letter or the same letter superscripts mean no significant difference (P>0.05), while with different small letter superscripts mean significant difference (P<0.05). The same as below.

2.2 ALA对白羽肉鸡肌肉质构特性的影响

由表4可知,与对照组相比,饲粮中添加100 mg/kg ALA显著提高了胸肌和腿肌的黏附性(P<0.05),添加300 mg/kg ALA显著提高了胸肌的胶黏性和咀嚼性(P<0.05)。
表4 α-硫辛酸对白羽肉鸡肌肉质构特性的影响

Table 4 Effects of ALA on muscle texture characteristics of white-feathered broilers

项目
Items		 对照组
Control group		 试验Ⅰ组
Test group Ⅰ		 试验Ⅱ组
Test group Ⅱ		 SEM P值
P-value
胸肌Breast muscle
硬度Hardness/N 21.30 20.35 23.56 1.343 0.080
黏附性Adhesiveness/(N/m) 0.17b 0.20a 0.16b 0.009 0.002
内聚性Cohesiveness 0.21 0.28 0.28 0.030 0.056
弹性Springiness/mm 1.01 1.10 1.10 0.149 0.804
胶黏性Gumminess/N 5.80b 6.27b 8.79a 0.768 0.003
咀嚼性Chewiness/mJ 5.21b 6.34ab 8.45a 1.051 0.042
腿肌Thigh muscle
硬度Hardness/N 76.13 64.59 64.24 9.909 0.273
黏附性Adhesiveness/(N/m) 0.20b 0.32a 0.24b 0.039 0.014
内聚性Cohesiveness 0.32 0.35 0.33 0.034 0.853
弹性Springiness/mm 1.39 1.18 1.18 0.137 0.237
胶黏性Gumminess/N 24.44 24.72 24.80 0.377 0.995
咀嚼性Chewiness/mJ 30.37 30.98 29.68 0.935 0.793

2.3 ALA对白羽肉鸡肌肉常规营养成分含量的影响

由表5可知,饲粮中添加不同水平ALA对胸肌和腿肌中水分、蛋白质和脂肪含量均没有产生显著影响(P>0.05)。
表5 α-硫辛酸对白羽肉鸡肌肉常规营养成分含量的影响(鲜样基础)

Table 5 Effects of ALA on common nutritional component contents in muscle of white-feathered broilers (fresh sample basis) %

项目
Items		 对照组
Control group		 试验Ⅰ组
Test group Ⅰ		 试验Ⅱ组
Test group Ⅱ		 SEM P值
P-value
胸肌Breast muscle
水分Moisture 76.44 76.45 76.25 0.350 0.813
蛋白质Protein 17.69 17.47 18.70 0.649 0.164
脂肪Fat 1.50 1.62 1.74 0.276 0.691
腿肌Thigh muscle
水分Moisture 77.63 77.99 78.15 0.266 0.175
蛋白质Protein 16.09 15.78 15.47 0.252 0.078
脂肪Fat 1.50 1.65 1.66 0.188 0.638

2.4 ALA对白羽肉鸡肌肉中生化指标的影响

由表6可知,与对照组相比,饲粮中添加100和300 mg/kg ALA显著提高了胸肌和腿肌中GSH-Px活性,并显著提高了腿肌中SOD活性(P<0.05),同时显著降低了腿肌中MDA含量(P<0.05);饲粮中添加300 mg/kg ALA显著降低了胸肌中Hyp含量(P<0.05);此外,饲粮中添加不同水平ALA对胸肌中糖原含量没有产生显著影响(P>0.05)。
表6 α-硫辛酸对白羽肉鸡肌肉中生化指标的影响

Table 6 Effects of ALA on biochemical indexes in muscle of white-feathered broilers

项目
Items		 对照组
Control group		 试验Ⅰ组
Test group Ⅰ		 试验Ⅱ组
Test group Ⅱ		 SEM P值
P-value
胸肌Breast muscle
丙二醛MDA/(nmol/mg prot) 0.11 0.09 0.07 0.038 0.618
谷胱甘肽过氧化物酶GSH-Px/(U/mg prot) 15.88c 17.10a 16.56b 0.072 <0.001
超氧化物歧化酶SOD/(U/mg prot) 19.83 20.94 21.47 0.810 0.152
总抗氧化能力T-AOC/(μmol/g prot) 22.68 21.22 22.74 0.749 0.103
羟脯氨酸Hyp/(μg/mg) 0.13a 0.12a 0.05b 0.006 <0.001
糖原Glycogen/(mg/g) 0.92 0.91 0.95 0.046 0.694
腿肌Thigh muscle
丙二醛MDA/(nmol/mg prot) 0.95a 0.39b 0.46b 0.144 0.003
谷胱甘肽过氧化物酶GSH-Px/(U/mg prot) 11.88b 13.57a 13.09a 0.439 0.005
超氧化物歧化酶SOD/(U/mg prot) 17.07b 22.78a 22.05a 1.127 <0.001
总抗氧化能力T-AOC/(μmol/g prot) 74.18 78.39 79.10 2.996 0.239

2.5 ALA对白羽肉鸡胸肌中肌纤维类型及相关调节因子基因表达的影响

由上述结果分析可知,ALA对白羽肉鸡胸肌品质的改善效果优于腿肌。因此,本试验选择胸肌为研究对象挖掘ALA改善白羽肉鸡肉品质的机制。由表7可知,与对照组相比,饲粮中添加100和300 mg/kg ALA显著提高了胸肌中MyHCⅠ和MyHCⅡa的mRNA相对表达量(P<0.05),显著提高了胸肌中AMPKα1、AMPKα2、SIRT1、PGC-1α的mRNA相对表达量(P<0.05),而添加100 mg/kg ALA显著降低了胸肌中MyHCⅡb的mRNA相对表达量(P<0.05);试验Ⅱ组胸肌中AMPKγ3的mRNA相对表达量显著低于对照组和试验Ⅰ组(P<0.05);此外,试验Ⅰ组胸肌中NRF1、MEF2的mRNA相对表达量最高,显著高于对照组和试验Ⅱ组(P<0.05)。
表7 α-硫辛酸对白羽肉鸡胸肌中肌纤维类型及相关调控因子基因表达的影响

Table 7 Effects of ALA on gene expression of muscle fiber types and associated regulatory factors in breast muscle of white-feathered broilers

项目
Items		 对照组
Control group		 试验Ⅰ组
Test group Ⅰ		 试验Ⅱ组
Test group Ⅱ		 SEM P值
P-value
肌球蛋白重链Ⅰ MyHCⅠ 1.00b 2.26a 2.05a 0.175 <0.001
肌球蛋白重链Ⅱa MyHCⅡa 1.00b 6.60a 6.84a 0.694 <0.001
肌球蛋白重链Ⅱb MyHCⅡb 1.00a 0.30b 0.87a 0.067 <0.001
腺苷酸活化蛋白激酶α1 AMPKα1 1.00c 2.23a 1.73b 0.160 <0.001
腺苷酸活化蛋白激酶α2 AMPKα2 1.00c 2.78a 1.51b 0.140 <0.001
腺苷酸活化蛋白激酶γ3 AMPKγ3 1.00a 1.17a 0.55b 0.185 0.012
沉默信息调节因子1 SIRT1 1.00c 3.25a 1.40b 0.159 <0.001
辅助激活因子-1α PGC-1α 1.00b 2.06a 2.03a 0.166 <0.001
核呼吸因子1 NRF1 1.00b 2.15a 1.07b 0.307 0.003
肌细胞增强因子2 MEF2 1.00b 2.22a 1.31b 0.159 <0.001

3 讨论

3.1 ALA对白羽肉鸡肌肉品质的影响

肉的颜色、pH、持水性及营养成分含量常作为评定肉品质优良的标准,反映肉的经济价值[10]。动物屠宰后,肌肉中贮存的糖原经糖酵解生成乳酸导致pH降低,糖原含量及糖酵解潜力会影响肌肉pH下降速度及持水性[11]。常江[5]研究发现,饲粮中添加ALA能通过降低羔羊肌肉中糖原含量和乳酸脱氢酶(LDH)活性来抑制糖酵解,减少乳酸堆积。本研究并未发现ALA对白羽肉鸡肌肉中糖原和pH产生显著影响,但与对照组相比,添加ALA减缓了胸肌和腿肌pH的下降速度。有研究表明,ALA可抑制屠宰后肌肉中腺苷酸活化蛋白激酶(AMPK)及糖原磷酸化酶的活性,减少肌肉在贮藏过程中发生的糖酵解,提高pH24 h[12]。本试验结果显示,饲粮中添加100和300 mg/kg ALA显著降低了白羽肉鸡胸肌的滴水损失,提高了持水性,与卢敏等[13]所得结果一致。肌肉较好的持水性与肉色中的L*值存在相关性,由于光反射作用,持水性越好,水分渗出越少,L*值越低,肉色越正常,越不易形成PSE肉[14]。本试验发现,饲粮中添加ALA显著降低了白羽肉鸡胸肌的L*和b*值,并显著提高了a*值。剪切力反映了肉的嫩度,饲粮中添加ALA显著降低了白羽肉鸡胸肌和腿肌的剪切力。嫩度与肌纤维直径和胶原蛋白含量呈负相关,与肌纤维密度呈正相关[15]。肌肉中Hyp含量反映了胶原蛋白含量,其含量越低,肌肉的嫩度越高。本试验中,饲粮中添加ALA显著降低了白羽肉鸡胸肌中Hyp含量,与El-Senousey等[16]所得结果一致。张勇等[17]的研究也显示,ALA降低了肉鸡胸肌和腿肌的剪切力,还显著降低了肌纤维直径,并且提升了肌纤维密度。

3.2 ALA对白羽肉鸡肌肉质构特性的影响

肌肉质构特性与其品质紧密相关,反映肌肉的组织结构和物理机械特性。质构剖面分析(TPA)作为一种模拟口腔咀嚼的双重压缩测试方法,可同时获得硬度、弹性、内聚性、胶黏性及咀嚼性等参数,相较于Warner-Bratzler剪切测试,TPA能更好地模拟咀嚼效果而非切割效果,更适用于生肉的质构特性检测[18]。硬度是消费者食用肉制品时首先感受到的口感特性,是描述食物抵抗变形或穿透所需的力,会影响肉的嫩度和商业价值[19]。本研究发现,添加ALA有降低白羽肉鸡腿肌硬度的趋势,并伴随有剪切力的显著下降,Shackelford等[20]在牛肉上也得出了硬度与剪切力呈正相关结果。ALA可能通过保持肌肉正常pH环境,维持蛋白水解酶活性,促进肌原纤维有序降解,缓解因肌纤维交联引起的硬度增加[21]。咀嚼性是对硬度、弹性和内聚性的综合性评价,弹性指食物受力变形后恢复原状的能力,弹性与咀嚼性越高,肉质口感越好,越有嚼劲[22];内聚性表示食物在被咀嚼时产生的抵抗受损并紧密连接的力,反映食物内部结合键的强度[19]。本试验结果显示,添加ALA对白羽肉鸡胸肌的咀嚼性有所提高,ALA可能通过增强细胞间的结合力,增加肌纤维间的紧密程度,使肉质更加紧实。胶黏性指肌肉组织细胞间结合力的大小,黏附性是指食物在测试或感官咀嚼时,食物对测试探头或者口腔的吸附能力。有报道表明,肉品的胶黏性与脂肪含量呈正相关,黏附性与蛋白质含量呈负相关[23]。本研究中,添加ALA后白羽肉鸡肌肉中脂肪含量有上升趋势,蛋白质含量有下降趋势,这可能是导致黏附性和胶黏性上升的原因。

3.3 ALA对白羽肉鸡肌肉抗氧化能力的影响

畜禽屠宰后发生的脂质和蛋白质氧化是肉质劣变的重要因素。脂质氧化会使肉制品产生酸败气味并产生有害物质MDA;蛋白质氧化会导致巯基丢失并生成羰基,破坏蛋白质结构稳定性,还会促使高铁肌红蛋白的生成[24]。因此,添加抗氧化剂改善氧化反应带来的肉质感官品质变差、营养价值降低,延长货架期,提升食品健康,是肉制品工业调控的关键措施。ALA作为天然抗氧化剂,能激活内源性抗氧化系统,提高GSH-Px、SOD和过氧化氢酶(CAT)活性[25],通过清除羟自由基和活性氧,降低血清、肌肉和肝脏中MDA含量,并能再生内源性抗氧化剂维生素E和谷胱甘肽[4]。李艳[26]的试验表明,添加ALA使肉鸡胸肌中MDA含量显著下降,腿肌SOD活性上升,胸肌腿肌中谷胱甘肽显著增加;Chen等[25]试验中显著增加了肉鸡胸肌SOD、GSH-Px活性,降低腿肌中MDA含量,与本试验结果相似。ALA还能缓解高温引起的氧化应激,提高肉鸡血清中SOD活性和T-AOC[27]。同时,本研究发现,各组白羽肉鸡腿肌中MDA含量均高于胸肌,ALA降低腿肌中MDA含量的效果更为显著,这可能是由于腿肌中氧化型肌纤维占比更高,有更多的线粒体和肌红蛋白,更容易发生氧化损伤[28]。

3.4 ALA对白羽肉鸡肌纤维类型的影响

肉品质与肌纤维特性密切相关。研究表明,氧化型肌纤维较酵解型肌纤维的直径更细且剪切力更低,酵解型肌纤维比例高的肌肉的系水力较低,屠宰后pH下降的更快[29]。MyHCⅠ代表的是慢速氧化型肌纤维,MyHCⅡa代表的是快速氧化型肌纤维,MyHCⅡb代表的是快速酵解型肌纤维。本研究发现,饲粮中添加ALA显著增加了MyHCⅠ和MyHCⅡa的mRNA相对表达量,同时降低了MyHCⅡb的mRNA相对表达量,提示ALA可促进酵解型肌纤维向氧化型肌纤维转化。这一过程可能涉及对AMPK的调控,AMPK是一种细胞能量感受器,可调控ATP的合成代谢,由1个催化亚基(α)和2个调节亚基(β和γ)组成[30],其中AMPKγ3高度特异性表达于骨骼肌中,其在家禽骨骼肌中的表达量与肌纤维中MyHCⅠ的比例呈显著负相关[31],在大鼠和小鼠骨骼肌中与MyHCⅡb的比例呈显著正相关[30],是调控肌纤维类型转化的重要节点基因。本试验中,ALA显著抑制白羽肉鸡胸肌中AMPKγ3的表达,表明ALA介导AMPKγ3调控酵解型肌纤维向氧化型肌纤维的转化。常江[5]研究发现,ALA通过激活AMPK/SIRT1/PGC-1α信号通路促进线粒体生物合成,改善滩羊骨骼肌中肌纤维类型。AMPK通过增加细胞内烟酰胺腺嘌呤二核苷酸(NAD+)激活SIRT1,SIRT1则通过脱乙酰化作用调控PGC-1α的活性,进而影响线粒体功能和合成的转录途径,最终诱导肌纤维类型的转变[32]。此外,NRF1可与PGC-1α相互作用,上调线粒体转录因子A的表达,进而激活线粒体蛋白合成,以此提升线粒体功能,促进肌纤维的发育[33]。PGC-1α还通过结合并协同激活肌肉选择性转录因子MEF2,调控葡萄糖转运蛋白4的表达,促进葡萄糖的摄取和利用[34]。本试验中,饲粮中添加ALA显著上调了白羽肉鸡胸肌中AMPKα1、AMPKα2、AMPKγ3、SIRT1、PGC-1α、NRF1和MEF2的mRNA相对表达量,表明ALA对白羽肉鸡肌纤维类型的调节是通过调控AMPK/SIRT1/PGC-1α信号通路实现的。

4 结论

饲粮中添加ALA可改变肌肉的质构特性,减少胶原蛋白沉积,增强抗氧化能力,上调AMPK/SIRT1/PGC-1α信号通路关键基因的表达,促进酵解型肌纤维向氧化型肌纤维转化,进而改善白羽肉鸡的肌肉品质。综合考虑作用效果和添加成本,推荐在白羽肉鸡饲粮中添加100 mg/kg的ALA。

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