Regulation of Chitosan Oligosaccharides on Postmortem Energy Metabolism, Antioxidant Capacity and Fatty Acid Composition in Breast Muscle of Broilers under Cyclic Heat Stress: Implications for Meat Quality

GAO Jinren; WANG Haoxuan; ZHANG Jia; LAN Ruixia

doi:10.12418/CJAN2026.085

2026 , Vol. 38 >Issue 2: 1061 - 1074

DOI: https://doi.org/10.12418/CJAN2026.085

RESEARCH PAPER

Regulation of Chitosan Oligosaccharides on Postmortem Energy Metabolism, Antioxidant Capacity and Fatty Acid Composition in Breast Muscle of Broilers under Cyclic Heat Stress: Implications for Meat Quality

GAO Jinren ,
WANG Haoxuan ,
ZHANG Jia ,
LAN Ruixia ^,^*

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College of Coastal Agriculture Sciences, Guangdong Ocean University, Zhanjiang 524088, China

^* lecturer, E-mail: Lanrx@gdou.edu.cn

Received date: 2025-07-31

Online published: 2026-02-12

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Abstract

The aim of this study was to evaluate the effects of dietary chitosan oligosaccharide (COS) supplementation on meat quality, postmortem muscle energy metabolism, antioxidant capacity and fatty acid composition in breast muscle of broilers under cyclic heat stress. A total of 192 male Arbor Acres (AA) broilers aged at 28 days with comparable body weight were selected and randomly divided into 4 groups, with 6 replicates per group and 8 chickens per replicate. The control group (CON group) was offered a basal diet, and the heat stress treatment groups received the basal diet supplemented with 0 (HS group), 200 (HSC₂₀₀ group) and 400 mg/kg (HSC₄₀₀ group) COS, respectively. The housing temperature in CON group was maintained at (24±1) ℃; for the heat stress treatment groups, the housing temperature was set at (34±1) ℃ from 10:00 to 18:00 and at (24±1) ℃ during the remainder of the day. The relative humidity was maintained at 65% to 75% throughout the trial. The experiment lasted for 14 days. The results showed as follows: 1) compared with CON group, the dressing percentage and half-eviscerated yield percentage in HS group were significantly decreased (P<0.05). Compared with HS group, the half-eviscerated yield percentage and abdominal fat percentage in HSC₄₀₀ group were significantly decreased (P<0.05). 2) Compared with CON group, the pH at 45 min and 24 h after slaughter and crude protein content in breast muscle in HS group were significantly decreased (P<0.05), and the drip loss and moisture content in breast muscle were significantly increased (P<0.05). Compared with HS group, the pH at 45 min and 24 h after slaughter in breast muscle both in HSC₂₀₀ group and HSC₄₀₀ group was significantly increased (P<0.05), and crude fat content in breast muscle in HSC₂₀₀ group was significantly increased (P<0.05). 3) Compared with CON group, the glycogen and glucose (GLU) contents in breast muscle in HS group were significantly decreased (P<0.05), and the lactic acid content and hexokinase (HK) activity in breast muscle were significantly increased (P<0.05). Compared with HS group, the glycogen and GLU contents in breast muscle both in HSC₂₀₀ group and HSC₄₀₀ group were significantly elevated (P<0.05), and the lactic acid content and HK activity in breast muscle were significantly decreased (P<0.05). 4) Compared with CON group, the malondialdehyde (MDA) content in breast muscle in HS group was significantly increased (P<0.05), and the activities of catalase (CAT), total superoxide dismutase (T-SOD) and glutathione peroxidase (GSH-Px) in breast muscle were significantly decreased (P<0.05). Compared with HS group, the MDA content in breast muscle both in HSC₂₀₀ group and HSC₄₀₀ group was significantly decreased (P<0.05), and the activities of CAT, T-SOD and GSH-Px in breast muscle were significantly increased (P<0.05). 5) Compared with CON group, the contents of caproic acid, behenic acid, γ-linolenic acid, docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA)+DHA, the ratios of polyunsaturated fatty acids to saturated fatty acids (PUFA/SFA) and unsaturated fatty acids to saturated fatty acids (UFA/SFA), as well as the fatty acid unsaturation index (UI), peroxidation trend index (PI), nutrition value index (NVI), hypocholesterolemic to hypercholesterolemic ratio (HHR) and health-promoting index (HPI) in breast muscle in HS group were significantly decreased (P<0.05); furthermore, the contents of myristic acid and palmitoleic acid, along with the index of atherogenicity (IA) and index of thrombogenicity (IT) in breast muscle in HS group were significantly increased (P<0.05). Compared with HS group, the contents of myristic acid and palmitoleic acid in breast in both HSC₂₀₀ group and HSC₄₀₀ group were significantly decreased (P<0.05), and the contents of γ-linolenic acid, DHA and EPA+DHA, as well as the NVI, HHR and HPI were significantly increased (P<0.05); additionally, the UFA/SFA in breast muscle in HSC₂₀₀ group was significantly increased (P<0.05). In summary, the heat stress impairs the slaughter performance and breast muscle meat quality of broilers. COS ameliorates these detrimental effects by mitigating postmortem glycolysis, enhancing antioxidant capacity, and improving fatty acid composition and health indices in breast muscle. Under the experimental conditions of this trial, dietary supplementation of COS at 200 mg/kg demonstrates superior efficacy in enhancing breast muscle meat quality in heat-stressed broilers.

Key words： chitosan oligosaccharides; heat-stressed broilers; meat quality; energy metabolism; antioxidant capacity; fatty acid composition

Cite this article

GAO Jinren , WANG Haoxuan , ZHANG Jia , LAN Ruixia . Regulation of Chitosan Oligosaccharides on Postmortem Energy Metabolism, Antioxidant Capacity and Fatty Acid Composition in Breast Muscle of Broilers under Cyclic Heat Stress: Implications for Meat Quality[J]. Chinese Journal of Animal Nutrition, 2026 , 38(2) : 1061 -1074 . DOI: 10.12418/CJAN2026.085

据报道,2024年中国鸡肉产量达1 500万t,预计2025年产量将继续增长至1 530万t^[1]。与此同时,中国肉鸡产业贸易格局发生显著转变,相较于2023年,2024年肉鸡进口量锐减32.54%,出口量则猛增22.74%,实现净出口,并预计2025年进口量将进一步下降而出口量保持高位^[1]。随着居民生活水平的提升以及消费习惯和健康理念的转变,鸡肉已成为主要的优质动物蛋白质来源,消费者对鸡肉的消费需求也朝更高营养、更优品质和更佳风味的方向转变^[2-3]。在肉鸡生产过程中,高温环境、集约化养殖、饲粮营养水平以及肉鸡品种等因素都会影响鸡肉品质^[4-5]。其中,热应激易导致鸡出现生理性紊乱,包括全身性免疫失调、内分泌紊乱和呼吸性碱中毒等,从而影响健康和生产性能^[6]。此外,热应激会诱发氧化损伤及皮质酮介导的代谢紊乱,加速宰后肌肉糖酵解,导致鸡肉品质下降,且因现代肉鸡遗传选育的体温调节缺陷以及鸡本身无汗腺的生理特性,使热应激成为影响鸡肉品质的主要环境应激源^[7]。我国禽肉主产区主要分布在广东、广西、福建等亚热带和热带地区,常年高温多雨、夏季时间长,易造成肉鸡热应激。一方面,肉鸡因背覆羽毛、缺乏汗腺及高强度遗传选育,本身就对热应激敏感;另一方面,全球气候变暖及集约化养殖,使这些地区的家禽养殖业更易受高温影响,导致肉品质和经济效益下降^[7-9]。此外,热应激引起的氧化应激,易引起胸肌的抗氧化能力下降,导致脂质和肌红蛋白氧化^[7],影响肉色、嫩度、风味和营养价值,甚至产生有毒化合物^[10-12]。因此,笔者推测提高热应激肉鸡胸肌的抗氧化能力,可能对改善热应激肉鸡的肉品质有积极作用。

功能性饲料添加剂的使用能有效缓解热应激对肉鸡的危害,对提高生长性能和抗氧化能力,缓解肠道屏障损伤、炎症反应及免疫抑制,以及提升屠宰性能和肉品质有一定改善作用^[13-15]。因此,营养调控是缓解肉鸡热应激的有效途径^[9,16-17]。壳寡糖(chitosan oligosaccharide,COS)是由2~10个氨基葡萄糖通过β-1,4-糖苷键连接而成的低聚寡糖,具有促生长、抗应激、抗氧化、抗炎、调节肠道菌群及糖和脂代谢等多种生物学功能^[18]。Lan等^[19]研究发现,COS通过提高肉鸡胸肌和腿肌自由基清除能力和抗氧化能力,进而提高肉品质。Chang等^[20-21]研究表明,COS改善慢性或急性循环热应激肉鸡肉品质与其调控宰后肌肉能量代谢和提高抗氧化能力相关。此外,COS还可通过改善肌肉脂肪酸组成来提高禽肉品质^[22-23]。目前,关于COS在肉鸡生产上的应用已有较多研究,关于对生长性能的影响有不同的结果^[20,24-25]。本课题组前期研究结果表明,COS能改善热应激肉鸡生长性能,热应激肉鸡饲粮中添加200和400 mg/kg COS分别提高7.46%和1.78%的平均日增重以及1.61%和3.45%的平均日采食量,并分别降低12.66%和4.80%的料重比^[26],但针对COS对循环热应激肉鸡肉品质改善作用的系统研究相对较少。因此,本试验旨在通过研究COS对循环热应激肉鸡胸肌肉品质、宰后肌肉能量代谢、抗氧化能力及脂肪酸组成的影响,探究COS是否通过调节宰后肌肉能量代谢、抗氧化能力及脂肪酸组成改善肉品质。

1 材料与方法

1.1 试验设计

本试验在广东海洋大学滨海农业学院动物科学系家禽试验基地进行,肉鸡的使用和试验方案经广东海洋大学滨海农业学院动物福利委员会批准(批准编号:SYXK-2018-0147)。试验选取192只28日龄、体重相近的雄性爱拔益加(AA)肉鸡为研究对象,随机分为4组,每组6个重复,每个重复8只鸡。对照组(CON组)饲喂基础饲粮,参照NRC(1994)进行配制,其组成及营养水平见表1;热应激处理组分别饲喂在基础饲粮的基础上添加0(HS组)、200(HSC₂₀₀组)和400 mg/kg(HSC₄₀₀组)COS的饲粮。COS为市购产品,纯度为93.6%,平均分子质量为1 768 u,脱乙酰度≥90%。试验期14 d。

表1 基础饲粮组成及营养水平(风干基础)

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

项目Items	含量Content
原料Ingredients
玉米Corn	55.00
豆粕Soybean meal	34.82
小麦麸Wheat bran	2.00
植物油Vegetable oil	5.00
石粉Limestone	0.50
磷酸氢钙CaHPO₄	1.60
食盐NaCl	0.30
DL-蛋氨酸DL-Met	0.18
L-赖氨酸L-Lys	0.10
氯化胆碱Choline chloride	0.10
维生素预混料Vitamin premix¹⁾	0.20
矿物质预混料Mineral premix²⁾	0.20
合计Total	100.00
营养水平Nutrient levels³⁾
代谢能ME/(MJ/kg)	12.82
粗蛋白质CP	19.92
钙Ca	0.93
总磷TP	0.44
蛋氨酸Met	1.30
赖氨酸Lys	0.45

1)维生素预混料为每千克饲粮提供 Vitamin premix provided the following per kilogram of the diet:VA 9 000 IU,VD₃ 3 240 IU,VE 6 IU,VK₃ 0.75 mg,VB₁ 1.5 mg,VB₂ 4.5 mg,VB₆ 1.5 mg,VB₁₂ 10 mg,生物素 biotin 0.2 mg,烟酰胺 nicotinamide 10.5 mg,叶酸 folic acid 0.45 mg,泛酸 pantothenic acid 9 mg。

2)矿物质预混料为每千克饲粮提供 Mineral premix provided the following per kilogram of the diet:Cu (as copper sulfate) 5~10 mg,Fe (as ferrous sulfate) 45~120 mg,Mn (as manganese sulfate) 45~85 mg,Zn (as zinc sulfate) 50~80 mg,Se (as sodium selenite) 0.2 mg,I (as potassium iodide) 0.15 mg.

3)粗蛋白质、钙和总磷含量为实测值,其余为计算值。The contents of CP, Ca and TP were analyzed values, while the others were calculated values.

试验期间,CON组肉鸡饲养于恒温环境,通过空调调节鸡舍温度为(24±1) ℃;热应激处理组肉鸡则暴露于循环热应激环境,参照本课题组之前的方法^[20],通过加热器调节鸡舍每日温度循环为:每日08:00—10:00将温度由24 ℃逐步升至34 ℃,10:00—18:00维持在(34±1) ℃,18:00—20:00由34 ℃逐步降至24 ℃,20:00至次日08:00维持在(24±1) ℃。同时,CON组和热应激处理组通过加湿器将鸡舍相对湿度均控制在65%~75%。整个试验期的饲养管理遵循《商品肉鸡生产技术规程》(GB/T 19664—2005)的规定执行。

1.2 检测指标及方法

1.2.1 饲粮营养水平

饲粮代谢能和氨基酸含量参照《中国饲料成分及营养价值表(2020年第31版)》计算,饲粮粗蛋白质、钙和总磷含量分别参照《饲料中粗蛋白的测定凯氏定氮法》(GB/T 6432—2018)、《饲料中钙的测定》(GB/T 6436—2018)和《饲料中总磷的测定分光光度法》(GB/T 6437—2018)中方法测定。

1.2.2 屠宰性能和样品采集

试验结束当天24:00禁食不断水,次日08:00从每个重复中选择1只体重接近于该组平均体重的肉鸡,称重后屠宰,根据《家禽生产性能名词术语和度量计算方法》(NY/T 823—2020)要求进行屠宰性能指标的测定,包括屠宰率、半净膛率、全净膛率、胸肌率和腹脂率。采集2份1 g左右左侧胸肌样品,迅速用液氮冷冻后,置于-80 ℃的超低温冰箱冷冻保存,分别用于胸肌能量代谢、抗氧化能力和脂肪酸组成的测定;另采集10 g左右左侧胸肌样品,置于-20 ℃保存,用于营养成分含量的测定;采集右侧胸肌样品,置于4 ℃冰箱保存24 h后,用于肉品质分析。

1.2.3 胸肌肉品质

根据《畜禽肉质的测定》(NY/T 1333—2007)要求进行胸肌肉品质的测定。肉鸡屠宰后,测定右侧胸肌宰后45 min的pH(pH_{45 min});4 ℃冰箱保存24 h后,测定宰后24 h的pH(pH_{24 h})、蒸煮损失、滴水损失和剪切力。在肉鸡宰后24 h,使用CR-10 Plus色差计(柯尼卡美能达)测定胸肌肉色,包括亮度(lightness,L^*)、红度(redness,a^*)和黄度(yellowness,b^*)值。每个样品重复测定3次,并取平均值。

1.2.4 胸肌营养成分含量

对胸肌样本进行营养成分的定量分析,分别依据《食品安全国家标准食品中水分的测定》(GB 5009.3—2016)、《食品安全国家标准食品中蛋白质的测定》(GB 5009.5—2016)、《食品安全国家标准食品中脂肪的测定》(GB 5009.6—2016)和《食品安全国家标准食品中灰分的测定》(GB 5009.4—2016)中方法,测定其水分、粗蛋白质、粗脂肪和粗灰分含量。

1.2.5 胸肌能量代谢和抗氧化能力

取1 g冻存肌肉样品,按1∶9[重量(g)∶体积(mL)]比例加入预冷的生理盐水并匀浆,于4 ℃、3 200×g离心15 min后,取上清液用于检测。参照试剂盒说明书测定糖原(A043-1-1)、葡萄糖(GLU,A154-1-1)、乳酸(A019-2-1)和丙二醛(MDA,A003-1-2)含量以及己糖激酶(HK,A077-3)、丙酮酸激酶(PK,A076-1-1)、乳酸脱氢酶(LDH,A020-2)、过氧化氢酶(CAT,A007-1-1)、总超氧化物歧化酶(T-SOD,A001-1-2)和谷胱甘肽过氧化物酶(GSH-Px,A005-1-2)活性,以上试剂盒均购自南京建成生物工程研究所。采用TaKaRa BCA Protein Assay Kit(T9300A)测定蛋白质浓度,试剂盒购自宝日医生物技术(北京)有限公司。

1.2.6 胸肌脂肪酸组成和健康指数

取1 g左右胸肌样品,液氮速冻后送样,委托上海美吉生物医药科技有限公司采用气相色谱质谱联用仪(8890-7000D,Agilent Technologies, Inc.,美国)测定胸肌脂肪酸含量。根据测定结果,参考Guo等^[27]的方法通过计算脂肪酸不饱和指数(fatty acid unsaturation index,UI)、过氧化趋势指数(peroxidation trend index,PI)、营养价值指数(nutrition value index,NVI)、动脉粥样硬化指数(index of atherogenicity,IA)、血栓形成指数(index of thrombogenicity,IT)、低高密度脂蛋白胆固醇血症指数[低血胆固醇与高血胆固醇比值(hypocholesterolemic to hypercholesterolemic ratio,HHR)]和促健康水平指数(health-promoting index,HPI)来评价胸肌的健康指数。

1.3 数据统计分析

试验数据采用SAS 9.4软件进行统计学处理与分析,其中CON组与HS组间数据采用t检验进行显著差异性分析,热应激处理组(HS组、HSC₂₀₀组和HSC₄₀₀组)间数据采用单因素方差分析(one-way ANOVA)和Duncan氏多重比较检验进行差异显著性分析,结果数据以“平均值±标准误”以及均值标准误(SEM)表示,P<0.05表示差异显著。

2 结果

2.1 COS对循环热应激肉鸡屠宰性能的影响

由表2可知,与CON组相比,HS组肉鸡屠宰率和半净膛率显著降低(P<0.05)。与HS组相比,HSC₄₀₀组半净膛率和腹脂率显著降低(P<0.05),且HSC₄₀₀组腹脂率显著低于HSC₂₀₀组(P<0.05)。

表2 COS对循环热应激肉鸡屠宰性能的影响

Table 2 Effects of COS on slaughter performance of broilers under cyclic heat stress %

项目 Items	组别Groups				均值标准误 SEM	P值P-value
项目 Items	CON	HS	HSC₂₀₀	HSC₄₀₀	均值标准误 SEM	P₁	P₂
屠宰率Dressing percentage	92.03±1.11	89.11±1.55	91.80±3.95	91.29±2.09	2.555	0.004	0.215
半净膛率 Half-eviscerated yield percentage	84.52±1.12	80.18±2.16^a	79.07±2.52^ab	77.14±1.34^b	3.269	0.002	0.037
全净膛率 Eviscerated yield percentage	69.60±0.70	69.30±1.22	68.88±2.93	69.21±2.73	1.997	0.621	0.952
胸肌率 Breast muscle percentage	25.95±1.71	24.30±2.28	26.38±1.41	25.52±2.06	1.935	0.185	0.166
腹脂率 Abdominal fat percentage	1.48±0.28	1.75±0.30^a	1.64±0.39^a	1.25±0.25^b	0.347	0.129	0.018

P₁代表CON组与HS组间t检验的P值,P₂代表热应激处理组(HS组、HSC₂₀₀组和HSC₄₀₀组)间单因素方差分析的P值;热应激处理组间同行数据肩标无字母或相同字母表示差异不显著(P>0.05),不同字母表示差异显著(P<0.05)。下表同。

P₁ represented the P-value between CON group and HS group by t-test, and P₂ represented the P-value among the heat stress treatment groups (HS group, HSC₂₀₀ group and HSC₄₀₀ group) by one-way analysis of variance; among the heat stress treatment groups, values in the same row with no letter or the same letter superscripts indicated no significant differences (P>0.05), while with different letter superscripts indicated significant differences (P<0.05). The same as below.

2.2 COS对循环热应激肉鸡胸肌肉品质和营养成分含量的影响

由表3可知,与CON组相比,HS组肉鸡胸肌pH_{45 min}和pH_{24 h}以及粗蛋白质含量显著降低(P<0.05),胸肌滴水损失和水分含量显著提高(P<0.05)。与HS组相比,HSC₂₀₀组和HSC₄₀₀组胸肌pH_{45 min}和pH_{24 h}显著提高(P<0.05);此外,HSC₂₀₀组胸肌粗脂肪含量显著提高(P<0.05)。

表3 COS对循环热应激肉鸡胸肌肉品质和营养成分含量的影响

Table 3 Effects of COS on meat quality and nutrient contents in breast muscle of broilers under cyclic heat stress

项目 Items	组别Groups				均值标准误 SEM	P值P-value
项目 Items	CON	HS	HSC₂₀₀	HSC₄₀₀	均值标准误 SEM	P₁	P₂
肉品质Meat quality
宰后45 min的pH pH_{45 min}	6.37±0.11	6.11±0.11^b	6.45±0.06^a	6.40±0.23^a	0.190	0.002	0.005
宰后24 h的pH pH_{24 h}	5.91±0.07	5.31±0.04^b	5.87±0.09^a	5.94±0.09^a	0.273	<0.001	<0.001
亮度L^*	48.50±3.51	47.80±3.18	48.07±3.31	49.28±2.95	3.077	0.725	0.765
红度a^*	5.68±1.71	5.30±1.37	6.39±1.41	6.28±1.86	1.561	0.678	0.498
黄度b^*	6.55±2.69	5.87±1.85	6.38±1.76	6.23±1.20	1.836	0.621	0.885
滴水损失Drip loss/%	4.34±0.38	5.20±0.64	4.55±0.55	4.77±0.50	0.587	0.021	0.158
蒸煮损失Cooking loss/%	33.75±2.04	35.57±1.24	35.11±1.97	34.74±0.98	1.662	0.096	0.719
剪切力Share force/N	26.83±2.71	27.57±2.05	27.15±6.13	24.85±2.99	3.709	0.607	0.254
营养成分含量Nutrient contents/%
水分Moisture	67.17±1.27	69.68±1.46	67.99±1.51	68.43±1.26	1.587	0.011	0.164
粗蛋白质Crude protein	22.79±1.44	21.04±0.87	21.83±1.47	21.97±1.28	1.359	0.033	0.552
粗脂肪Crude fat	4.71±0.63	4.18±0.59^b	4.91±0.23^a	4.45±0.24^b	0.516	0.162	0.020
粗灰分Crude ash	5.32±0.36	5.09±0.84	5.27±0.49	5.15±0.58	0.560	0.538	0.928

2.3 COS对循环热应激肉鸡胸肌能量代谢的影响

由表4可知,与CON组相比,HS组肉鸡胸肌糖原和GLU含量显著降低(P<0.05),胸肌乳酸含量和HK活性显著提高(P<0.05)。与HS组相比,HSC₂₀₀组和HSC₄₀₀组胸肌糖原和GLU含量显著提高(P<0.05),胸肌乳酸含量和HK活性显著降低(P<0.05)。

表4 COS对循环热应激肉鸡胸肌能量代谢的影响

Table 4 Effects of COS on energy metabolism in breast muscle of broilers under cyclic heat stress

项目 Items	组别Groups				均值标准误 SEM	P值P-value
项目 Items	CON	HS	HSC₂₀₀	HSC₄₀₀	均值标准误 SEM	P₁	P₂
糖原Glycogen/(mg/g)	1.15±0.16	0.84±0.07^b	1.18±0.16^a	1.14±0.13^a	0.187	0.004	0.001
葡萄糖GLU/(mmol/g)	1.51±0.34	0.66±0.17^b	1.32±0.25^a	1.11±0.34^a	0.420	0.001	0.004
乳酸 Lactic acid/(mmol/g)	1.04±0.11	1.55±0.17^a	1.08±0.20^b	0.96±0.23^b	0.291	<0.001	<0.001
己糖激酶HK/(U/mg)	100.14±7.46	129.70±7.54^a	104.72±10.64^b	106.76±18.28^b	16.069	<0.001	0.018
丙酮酸激酶PK/(U/g)	139.25±19.09	205.16±69.14	161.22±60.90	146.06±31.54	53.176	0.067	0.112
乳酸脱氢酶LDH/(U/g)	951.05±136.65	968.16±83.69	925.73±18.79	921.22±82.07	86.631	0.800	0.500

2.4 COS对循环热应激肉鸡胸肌抗氧化能力的影响

由表5可知,与CON组相比,HS组肉鸡胸肌MDA含量显著提高(P<0.05),胸肌CAT、T-SOD和GSH-Px活性显著降低(P<0.05)。与HS组相比,HSC₂₀₀组和HSC₄₀₀组胸肌MDA含量显著降低(P<0.05),胸肌CAT、T-SOD和GSH-Px活性显著提高(P<0.05)。

表5 COS对循环热应激肉鸡胸肌抗氧化能力的影响

Table 5 Effects of COS on antioxidant capacity in breast muscle of broilers under cyclic heat stress

项目 Items	组别Groups				均值标准误 SEM	P值P-value
项目 Items	CON	HS	HSC₂₀₀	HSC₄₀₀	均值标准误 SEM	P₁	P₂
丙二醛 MDA/(nmol/mg prot)	0.12±0.03	0.17±0.04^a	0.06±0.04^b	0.10±0.05^b	0.055	0.034	0.015
过氧化氢酶 CAT/(U/mg prot)	0.28±0.08	0.11±0.02^c	0.21±0.04^b	0.27±0.10^a	0.092	0.002	<0.001
总超氧化物歧化酶 T-SOD/(U/mg prot)	170.70±6.74	129.56±16.66^b	154.78±18.91^a	167.59±18.43^a	22.267	0.001	0.002
谷胱甘肽过氧化物酶 GSH-Px/(U/mg prot)	8.39±2.22	5.21±2.42^b	12.43±6.28^a	12.40±3.66^a	4.833	0.039	0.031

2.5 COS对循环热应激肉鸡胸肌脂肪酸组成和健康指数的影响

由表6可知,与CON组相比,HS组肉鸡胸肌己酸、山嵛酸、γ-亚麻酸、二十二碳六烯酸(DHA)、二十碳五烯酸(EPA)+DHA含量和多不饱和脂肪酸(PUFA)/饱和脂肪酸(SFA)、不饱和脂肪酸(UFA)/SFA以及UI、PI、NVI、HHR、HPI显著降低(P<0.05),胸肌肉豆蔻酸、棕榈油酸含量以及IA、IT显著提高(P<0.05)。与HS组相比,HSC₂₀₀组和HSC₄₀₀组胸肌肉豆蔻酸和棕榈油酸含量显著降低(P<0.05),胸肌γ-亚麻酸、DHA、EPA+DHA含量以及NVI、HHR、HPI显著提高(P<0.05);此外,HSC₂₀₀组胸肌UFA/SFA显著提高(P<0.05)。

表6 COS对循环热应激肉鸡胸肌脂肪酸组成和健康指数的影响

Table 6 Effects of COS on fatty acid composition and health indices in breast muscle of broilers under cyclic heat stress

项目 Items	组别Groups								均值标准误 SEM		P值P-value
项目 Items	CON		HS		HSC₂₀₀		HSC₄₀₀		均值标准误 SEM		P₁	P₂
脂肪酸组成Fatty acid composition
己酸 C6∶0/(μg/g)		0.44 ±0.09		0.30 ±0.03		0.35 ±0.09		0.37 ±0.11		0.095	0.010	0.163
辛酸 C8∶0/(μg/g)		0.44 ±0.05		0.41 ±0.05		0.44 ±0.04		0.44 ±0.06		0.045	0.309	0.501
癸酸 C10∶0/(μg/g)		0.99 ±0.10		1.11 ±0.51		1.22 ±0.43		1.11 ±0.45		0.387	0.578	0.443
月桂酸 C12∶0/(μg/g)		1.68 ±0.44		1.48 ±0.29		1.69 ±0.17		1.71 ±0.31		0.311	0.396	0.319
十三烷酸 C13∶0/(μg/g)		0.81 ±0.03		0.83 ±0.03		0.82 ±0.03		0.83 ±0.02		0.028	0.311	0.490
肉豆蔻酸 C14∶0/(μg/g)		15.58 ±3.76		23.66 ±3.67^a		19.69 ±1.67^b		17.06 ±3.59^b		4.385	0.004	0.009
十五烷酸 C15∶0/(μg/g)		3.39 ±1.01		2.69 ±0.59		3.32 ±0.59		3.28 ±0.67		0.743	0.178	0.211
棕榈酸 C16∶0/(μg/g)		712.71 ±277.80		920.61 ±118.20		773.82 ±203.54		802.71 ±274.85		226.271	0.137	0.479
十七烷酸 C17∶0/(μg/g)		22.95 ±1.81		21.15 ±0.82		22.15 ±1.63		22.44 ±1.49		1.538	0.062	0.146
硬脂酸 C18∶0/(μg/g)		660.33 ±251.93		419.62 ±144.88		543.53 ±191.86		592.70 ±248.25		218.725	0.077	0.240
花生酸 C20∶0/(μg/g)		2.74 ±0.78		2.14 ±0.46		2.49 ±0.51		2.65 ±0.67		0.621	0.141	0.301
二十一烷酸 C21∶0/(μg/g)		1.27 ±0.12		1.36 ±0.20		1.40 ±0.13		1.30 ±0.24		0.177	0.380	0.386
山嵛酸 C22∶0/(μg/g)		2.09 ±0.41		1.64 ±0.16		1.71 ±0.17		1.68 ±0.14		0.295	0.040	0.750
饱和脂肪酸 SFA/(μg/g)		1 425.43 ±528.36		1 397.00 ±161.80		1 372.63 ±349.39		1 448.30 ±514.08		388.872	0.904	0.926
肉豆蔻烯酸 C14∶1/(μg/g)		4.40 ±2.39		3.42 ±1.48		4.21 ±1.10		4.18 ±1.36		1.592	0.417	0.477
棕榈油酸 C16∶1/(μg/g)		217.85 ±62.82		310.66 ±53.56^a		235.50 ±42.27^b		228.49 ±53.73^b		62.441	0.021	0.026
十七碳烯酸 C17∶1/(μg/g)		20.17 ±1.16		20.06 ±0.65		20.18 ±0.54		20.50 ±0.98		0.827	0.841	0.637
油酸 C18∶1n9c/(μg/g)		1 837.20 ±952.54		1 118.87 ±572.35		1 822.82 ±495.97		1 755.78 ±833.45		752.401	0.151	0.203
二十碳烯酸 C20∶1n9/(μg/g)		23.38 ±2.79		22.54 ±1.11		23.00 ±1.08		22.24 ±1.15		1.644	0.518	0.297
二十四碳烯酸 C24∶1n9/(μg/g)		3.34 ±0.81		4.06 ±1.23		3.70 ±1.35		3.78 ±0.90		1.055	0.260	0.815
单不饱和脂肪酸 MUFA/(μg/g)		2 106.33 ±1 014.31		1 479.62 ±578.46		2 109.42 ±517.35		2 034.96 ±881.75		771.942	0.225	0.301
亚油酸 C18∶2n6/(μg/g)		2 509.40 ±1 205.01		1 409.74 ±656.91		2 165.72 ±771.48		2 235.49 ±1 060.29		978.149	0.087	0.245
γ-亚麻酸 C18∶3n6/(μg/g)		25.04 ±3.33		18.47 ±2.87^b		24.98 ±1.39^a		25.72 ±3.32^a		4.012	0.005	<0.001
α-亚麻酸 C18∶3n3/(μg/g)		182.37 ±91.51		105.62 ±27.00		163.36 ±59.95		162.85 ±75.21		69.665	0.097	0.186
二十碳二烯酸 C20∶2/(μg/g)		28.95 ±3.67		26.44 ±1.69		28.25 ±2.68		26.69 ±1.53		2.602	0.171	0.263
二十碳三烯酸 C20∶3n6/(μg/g)		29.73 ±5.11		27.55 ±1.36		27.77 ±2.80		27.78 ±1.09		2.977	0.353	0.964
二十碳三烯酸 C20∶3n3/(μg/g)		24.56 ±1.48		23.23 ±1.89		23.11 ±1.01		23.81 ±2.36		1.740	0.207	0.432
花生四烯酸 C20∶4n6/(μg/g)		108.98 ±14.76		91.84 ±24.28		94.20 ±18.47		99.08 ±29.94		22.129	0.177	0.614
二十碳五烯酸 C20∶5n3/(μg/g)		4.24 ±1.02		3.41 ±0.65		3.54 ±0.64		3.49 ±0.66		0.786	0.127	0.938
二十二碳六烯酸 C22∶6n3/(μg/g)		9.27 ±0.84		6.60 ±1.35^b		8.95 ±0.33^a		8.63 ±0.68^a		1.346	0.003	0.004
多不饱和脂肪酸 PUFA/(μg/g)		2 922.55 ±1 319.40		1 712.89 ±670.46		2 539.88 ±848.92		2 613.54 ±1 136.50		1 059.567	0.083	0.225
不饱和脂肪酸 UFA/(μg/g)		7 951.44 ±3 624.87		3 192.51 ±1 221.45		4 649.29 ±1 242.61		4 648.51 ±2 013.34		1 793.948	0.126	0.241

3 讨论

3.1 COS对循环热应激肉鸡屠宰性能的影响

肉鸡的屠宰性能反映其产肉性能,是评价肉鸡生产经济效益的重要指标。研究表明,热应激降低肉鸡屠宰率,并提高腹脂率^[28]。本研究中,与CON组相比,HS组肉鸡屠宰率和半净膛率显著降低,但腹脂率仅在数值上提高,这与Teyssier等^[29]的研究结果相吻合,表明循环热应激导致肉鸡屠宰率和半净膛率显著降低,而对腹脂率并未产生显著影响。本课题组先前在黄羽肉鸡上的研究发现,COS可显著提高全净膛率和胸肌率,并显著降低腹脂率^[19]。腹脂过度沉积会降低胴体产量和饲料转化效率^[30]。腹部脂肪沉积的减少表明COS对体内脂肪沉积有抑制作用,多项研究表明COS能调控脂质代谢,对缓解腹脂过度沉积有积极作用^[31-32]。COS可降低腹脂率,主要与COS的降脂功能、调控肝脏脂质代谢以及调控脂肪细胞增殖分化相关^[33]。与之前的研究结果一致,本研究中,与HS组相比,HSC₄₀₀组肉鸡腹脂率显著降低。

3.2 COS对循环热应激肉鸡胸肌肉品质的影响

肌肉的pH、肉色、滴水损失、蒸煮损失和剪切力均是衡量肉品质优劣的重要指标。pH与系水力、颜色、多汁性和嫩度相关;滴水损失和蒸煮损失是衡量系水力的指标,反映多汁性;肉色反映肉品新鲜度^[5]。大量研究表明,热应激显著降低肉鸡肉品质,主要表现为加速宰后胸肌糖酵解,肌肉pH快速下降;肌肉系水力降低,滴水损失和蒸煮损失增加;肌浆蛋白质变性,肉色、系水力发生改变^[20-21,34]。本研究结果同样表明,与CON组相比,在热应激条件下的肉鸡胸肌pH_{45 min}和pH_{24 h}显著降低,并且滴水损失显著提高。宰后肌肉能量对肉品质的形成具有重要作用,Chang等^[20]研究表明,热应激加速宰后胸肌糖酵解,这一效应加速了胸肌中糖原的分解。在糖酵解过程中,GLU转化为乳酸,导致胸肌中乳酸的累积,最终导致pH快速下降。宰后肌肉糖原和GLU含量以及糖酵解过程中的关键酶(HK、PK、LDH)活性都会直接或间接影响肉品质^[35]。本研究中,与CON组相比,HS组胸肌糖原和GLU含量显著降低,胸肌HK活性显著提高,最终导致乳酸积累,pH快速下降,表明热应激导致的肉品质下降与加速宰后肌肉糖酵解密切相关。本研究中,热应激肉鸡饲粮中添加COS显著提高胸肌pH_{45 min}、pH_{24 h}以及糖原和GLU含量,显著降低胸肌乳酸含量和HK活性,且COS添加组胸肌滴水损失在数值上有所下降。这与先前的研究结果一致,在热应激条件下,COS能对肉鸡宰后胸肌品质产生积极影响,通过改善热应激肉鸡宰后胸肌能量代谢、pH、肉色和系水力来提升肉品质^[20-21]。pH是评价肉品质的重要指标,直接反映宰后肌肉的糖酵解^[36]。此外,pH的快速下降影响肌肉的系水力、肉色和嫩度^[5]。宰后肌肉pH的变化,主要与宰后肌肉糖酵解的乳酸积累相关,因此,笔者推测COS改善热应激肉鸡肉品质可能是通过缓解宰后胸肌的糖酵解。

肌肉氧化损伤是热应激降低肉品质的主要原因。热应激引起骨骼肌氧化损伤,与活性氧(ROS)的过量产生密切相关,ROS水平升高引起氧化损伤并引发脂质过氧化^[37]。本研究中,与CON组相比,HS组肉鸡胸肌MDA含量显著升高,胸肌CAT、T-SOD和GSH-Px活性显著降低,表明热应激引起了胸肌氧化损伤,这与Lu等^[38]的报道一致,慢性热应激使肉鸡胸肌ROS和MDA含量升高,胸肌超氧化物歧化酶(SOD)活性降低。Chen等^[39]报道,当肉鸡体内氧化剂水平超过抗氧化剂水平时,自由基的过量生成就会导致氧化应激。当肉鸡肌肉氧化稳定性降低时,ROS的过量产生、脂质过氧化加剧、自由基清除能力下降以及抗氧化酶活性降低等多重机制导致肉品质下降^[21]。具有抗氧化作用的功能性饲料添加剂被广泛应用于改善热应激导致的肉品质下降^[40-41]。COS在C2、C3和C6位置上有对自由基清除作用的氨基、羧基和伯、仲羟基,具有抗氧化和自由基清除能力,通过提高GSH-Px、SOD和CAT活性,发挥自由基清除活性,减少脂质过氧化^[42]。本研究中,热应激肉鸡饲粮中添加COS显著提高胸肌CAT、T-SOD和GSH-Px活性,这与Chang等^[20]在黄羽肉鸡上的研究结果一致,表明COS可缓解热应激引起的胸肌氧化损伤。此外,胸肌滴水损失与抗氧化能力呈显著负相关^[43]。本研究中,热应激肉鸡饲粮中添加COS降低胸肌滴水损失可能与胸肌抗氧化能力的提高相关,表明COS可通过提高热应激肉鸡胸肌的抗氧化能力,进而提高肉品质。

3.3 COS对循环热应激肉鸡胸肌脂肪酸组成和健康指数的影响

肌肉脂肪酸组成与肌内脂肪含量和肉品质密切相关^[44]。脂肪酸的种类和含量不仅能影响鸡肉的营养价值和风味,还影响鸡肉的加工贮存和消费者健康^[7,45]。胸肌中的脂肪酸组成包括SFA和UFA,其中UFA可细分为单不饱和脂肪酸(MUFA)和PUFA,而PUFA还能进一步分为n-6 PUFA和n-3 PUFA。肉类是消费者摄入脂肪酸的主要途径,过多摄入将增加患心血管疾病和Ⅱ型糖尿病的风险,PUFA的适当摄入能降低血液中胆固醇水平并降低心血管疾病的发病率;不过,需要注意的是,如果n-6 PUFA/n-3 PUFA过高,容易导致炎症反应,增加心血管疾病的发病率^[45-47]。热应激改变肉鸡胸肌脂肪酸组成,但目前关于热应激对肉鸡肌肉脂肪酸组成影响的研究结果不完全一致。Salah等^[48]研究表明,热应激显著提高肉鸡胸肌肉豆蔻酸和棕榈酸含量,显著降低肉豆蔻烯酸、棕榈油酸、油酸、亚油酸、EPA和DHA含量。Zhao等^[49]研究表明,热应激显著提高肉鸡胸肌硬脂酸和SFA含量,显著降低胸肌亚油酸、α-亚麻酸、二十碳三烯酸、二十碳五烯酸、PUFA和n-6 PUFA含量以及PUFA/SFA。Ebrahim等^[50]研究表明,热应激显著提高肉鸡胸肌棕榈油酸和亚油酸含量及n-6 PUFA/n-3 PUFA,显著降低胸肌十七烷酸含量。本研究发现,热应激显著降低肉鸡胸肌己酸、山嵛酸、γ-亚麻酸、DHA、EPA+DHA含量以及PUFA/SFA、UFA/SFA,且显著提高胸肌肉豆蔻酸和棕榈油酸含量,与Zhao等^[49]的研究结果相似。不同研究发现热应激对肉鸡胸肌脂肪酸组成的影响不同,推测可能与鸡的品种、性别、日龄、饲粮组成及热应激处理方式不同相关,这还需做进一步研究。同时,本试验发现,热应激提高肉鸡胸肌IA和IT,IA和IT的升高反映了胸肌中促炎性和促血栓形成的脂肪酸比例增加。

先前的研究表明,COS可影响家禽肌肉脂肪酸组成。Zhou等^[23]研究报道,COS显著降低肉鸡胸肌肉豆蔻酸、棕榈酸、硬脂酸、SFA和棕榈油酸含量,显著提高胸肌油酸、二十碳烯酸和MUFA含量以及PUFA/SFA。Lan等^[51]研究报道,COS能够显著降低肉鸡腿肌棕榈酸、硬脂酸、SFA、二十碳三烯酸和DHA含量。Miao等^[22]研究指出,壳聚糖可以对豁眼鹅胸肌肉豆蔻酸、棕榈酸、硬脂酸、SFA和PUFA含量以及n-6 PUFA/n-3 PUFA产生显著降低的作用,对胸肌山嵛酸、棕榈油酸、油酸、二十碳烯酸、二十碳二烯酸、EPA和MUFA含量以及PUFA/SFA产生显著提高的作用。本研究发现,COS显著降低热应激肉鸡胸肌肉豆蔻酸和棕榈油酸含量,提高胸肌γ-亚麻酸、DHA、EPA+DHA含量以及NVI、HHR、HPI。摄入富含MUFA和PUFA而低SFA的食物,可通过改善血脂谱来降低心血管疾病风险^[52]。γ-亚麻酸和DHA为PUFA,分子结构中分别含3和6个双键,易被氧化生成羰基化物和氢过氧化物^[45]。COS具有抗氧化活性的成分,能够通过增强胸肌的抗氧化能力,降低PUFA中不饱和键的氧化水平,从而使γ-亚麻酸和DHA含量得到提高,同时也减轻胸肌的氧化损伤,其他具有抗氧化活性的添加剂也有类似的效果^[53]。因此,COS改善热应激肉鸡胸肌脂肪酸组成与其具有抗氧化的特性相关。PUFA为必需脂肪酸,人体不能自身合成,只能从食物中摄入。PUFA/SFA是一个用于评估饮食对心血管健康(CVH)影响的指标,它假设饮食中的所有PUFA都能降低血清低密度脂蛋白胆固醇和胆固醇含量,而所有SFA都能导致血清胆固醇含量升高^[54]。因此,这个比值越高,效果就越积极,比值的升高会降低食物带给人类心脏病和中风的风险,与消费者的健康呈正相关^[55]。COS提高热应激肉鸡胸肌PUFA/SFA,可提高鸡肉的营养价值,有降低心血管疾病发生的潜在作用^[45]。同时,摄入高水平EPA+DHA食品可以增强心血管保护功能^[56],COS提高热应激肉鸡胸肌EPA+DHA含量与其HPI升高的效应相一致。此外,HSC₂₀₀组在降低胸肌IA、IT以及提高MUFA/SFA和UFA/SFA方面效果优于HSC₄₀₀组,表明饲粮添加200 mg/kg COS在改善热应激肉鸡胸肌脂质健康指数方面更具优势。

4 结论

热应激降低肉鸡屠宰性能和胸肌肉品质,COS通过缓解宰后胸肌的糖酵解、提高抗氧化能力以及改善脂肪酸组成和健康指数,从而提高热应激肉鸡胸肌肉品质。在本试验条件下,COS提高热应激肉鸡胸肌肉品质以200 mg/kg添加量效果较好。

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