动物营养学报 >

2026 , Vol. 38 >Issue 5: 3838 - 3850

DOI: https://doi.org/10.12418/CJAN2026.306

研究论文

不同精粗比饲粮中添加番茄红素对绵羊体外瘤胃发酵的影响

  • 申玉萍 ,
  • 崔崇乾 ,
  • 牟春堂 ,
  • 张建新 ,
  • 郝小燕 , *
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  • 山西农业大学动物科学学院, 太谷 030801
*郝小燕,教授,硕士生导师,E-mail:

申玉萍(2000—),女,云南镇雄人,硕士研究生,从事反刍动物营养与饲料科学研究。E-mail:

Office editor: 菅景颖

收稿日期: 2025-11-05

  网络出版日期: 2026-05-14

基金资助

山西省基础研究计划项目(202303021221102)

山西省现代农业产业技术体系建设专项(2026CYJSTX14)

现代农业产业技术体系(CARS-38)

收起

Effects of Lycopene Supplementation in Diets with Different Concentrate-Forage Ratios on in Vitro Ruminal Fermentation of Sheep

  • SHEN Yuping ,
  • CUI Chongqian ,
  • MU Chuntang ,
  • ZHANG Jianxin ,
  • HAO Xiaoyan , *
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  • College of Animal Science, Shanxi Agricultural University, Taigu 030801, China
*professor, E-mail:

Received date: 2025-11-05

  Online published: 2026-05-14

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摘要

为评估番茄红素在不同精粗比条件下对瘤胃发酵的调控作用,本试验采用2×5双因素试验设计,设置2个精粗比(55∶45与80∶20)和5个番茄红素添加量(0、10.3、20.6、30.9和41.2 mg/kg),共配制10种试验饲粮作为体外发酵底物。通过体外模拟瘤胃发酵48 h后,测定产气动力学参数、甲烷(CH4)产量、瘤胃发酵参数及消化酶活性。结果显示:1)饲粮精粗比与番茄红素添加量及二者的交互作用对24、48 h累积产气量均有显著影响(P<0.05),在高、低精料饲粮中均以添加20.6 mg/kg番茄红素时24、48 h累积产气量最高。与55∶45组相比,80∶20组的慢速降解部分产气量和潜在产气量均显著提高(P<0.05);与未添加番茄红素相比,添加20.6和30.9 mg/kg番茄红素可显著提高潜在产气量(P<0.05)。2)饲粮精粗比与番茄红素添加量的交互作用对有机物消化率、消化能和代谢能有显著影响(P<0.05),在高、低精料饲粮中均以添加20.6 mg/kg番茄红素时有机物消化率、消化能和代谢能最高。3)与55∶45组相比,80∶20组的CH4产量、乙丙比显著下降(P<0.05),总挥发性脂肪酸(TVFA)、丙酸和丁酸浓度显著升高(P<0.05)。番茄红素添加量30.9 mg/kg时,丙酸浓度显著高于添加量为0、10.3、41.2 mg/kg时(P<0.05);番茄红素添加量为20.6、30.9 mg/kg时,TVFA浓度显著高于添加量为0、41.2 mg/kg时(P<0.05)。4)与55∶45组相比,80∶20组pH显著降低(P<0.05),氨态氮(NH3-N)浓度显著升高(P<0.05);随着番茄红素添加量的增加,pH先升高后降低,在添加量为20.6 mg/kg时达到最高;各添加番茄红素组NH3-N浓度均显著低于未添加组(P<0.05),且以添加量为20.6 mg/kg时最低。5)与55∶45组相比,80∶20组α-淀粉酶、蛋白酶活性显著升高(P<0.05);随着番茄红素添加量的增加,羧甲基纤维素酶、α-淀粉酶活性先升高后降低,20.6 mg/kg组显著高于未添加组和41.2 mg/kg组(P<0.05);β-葡萄糖苷酶活性在20.6和30.9 mg/kg组均显著高于未添加组和41.2 mg/kg组(P<0.05)。综上所述,提高饲粮精粗比可显著提高体外瘤胃发酵产气量和TVFA浓度,降低CH4产量,有利于提高能量利用率;番茄红素通过提高消化酶活性促进瘤胃发酵,并降低NH3-N浓度,有利于提高氮利用率。本试验条件下,在高、低精粗比(80∶20和55∶45)饲粮中均以添加20.6 mg/kg番茄红素对绵羊体外瘤胃发酵的促进效果最佳。

关键词： 精粗比; 番茄红素; 绵羊; 体外瘤胃发酵

本文引用格式

申玉萍 , 崔崇乾 , 牟春堂 , 张建新 , 郝小燕 . 不同精粗比饲粮中添加番茄红素对绵羊体外瘤胃发酵的影响[J]. 动物营养学报, 2026 , 38(5) : 3838 -3850 . DOI: 10.12418/CJAN2026.306

Abstract

To evaluate the regulatory effects of lycopene on rumen fermentation under different concentrate-forage ratios, this experiment adopted a 2×5 two-factor experimental design. Two concentrate-forage ratios (55∶45 and 80∶20) and five lycopene supplemental levels (0, 10.3, 20.6, 30.9 and 41.2 mg/kg) were set, resulting in a total of 10 experimental diets prepared as substrates for in vitro fermentation. After 48 hours of simulated rumen fermentation in vitro, gas production kinetic parameters, methane (CH4) production, rumen fermentation parameters and digestive enzyme activities were determined. The results showed as follows: 1) the concentrate-forage ratio, lycopene supplemental level and their interaction all had significant effects on the 24 and 48 h cumulative gas production (P<0.05). In both high- and low-concentrate diets, the 24 and 48 h cumulative gas productions were highest when lycopene supplemental level was 20.6 mg/kg. Compared with the 55∶45 group, slowly degraded fraction gas production and potential gas production in the 80∶20 group were significantly increased (P<0.05); compared with the non-added group, the potential gas production was significantly increased by the supplemental levels of 20.6 and 30.9 mg/kg lycopene (P<0.05). 2) The interaction between the concentrate-forage ratio and lycopene supplemental level had significant effects on organic matter digestibility, digestible energy and metabolizable energy (P<0.05). In both high- and low-concentrate diets, organic matter digestibility, digestible energy and metabolizable energy were highest when lycopene was added at 20.6 mg/kg. 3) Compared with the 55∶45 group, the 80∶20 group showed significantly decreased CH4 production and acetate/propionate ratio (P<0.05), while the concentrations of total volatile fatty acids (TVFA), propionate and butyrate were significantly increased (P<0.05). When lycopene was added at 30.9 mg/kg, the propionate concentration was significantly higher than that at supplemental levels of 0, 10.3 and 41.2 mg/kg (P<0.05). When lycopene was added at 20.6 and 30.9 mg/kg, the TVFA concentration was significantly higher than that at supplemental levels of 0 and 41.2 mg/kg (P<0.05). 4) Compared with the 55∶45 group, the 80∶20 group exhibited a significantly lower pH (P<0.05) and a significantly higher ammonia nitrogen (NH3-N) concentration (P<0.05). With lycopene supplemental level increasing, pH initially increased and then decreased, reaching the highest value at the supplemental level of 20.6 mg/kg. The NH3-N concentration in all lycopene-added groups was significantly lower than that in the non-added group (P<0.05), with the lowest value observed at the supplemental level of 20.6 mg/kg. 5) Compared with the 55∶45 group, the activities of α-amylase and protease in the 80∶20 group were significantly increased (P<0.05). With lycopene supplemental level increasing, the activities of carboxymethyl cellulase and α-amylase first increased and then decreased, and were significantly higher in the 20.6 mg/kg group than in the non-added group and the 41.2 mg/kg group (P<0.05). The β-glucosidase activity in the 20.6 and 30.9 mg/kg groups was significantly higher than that in the non-added group and the 41.2 mg/kg group (P<0.05). In conclusion, increasing the concentrate-forage ratio in the diet can significantly enhance the gas production and TVFA concentration in rumen fermentation in vitro, reduce CH4 production, and is conducive to improving energy utilization efficiency. Lycopene promotes rumen fermentation by increasing digestive enzyme activities and reduced NH3-N concentration, which is beneficial for improving nitrogen utilization efficiency. Under this experiment conditions, the supplementation of 20.6 mg/kg lycopene in diets with both high and low concentrate-forage ratios (80∶20 and 55∶45) exerts the best promoting effect on sheep in vitro rumen fermentation.

Key words： concentrate-forage ratio; lycopene; sheep; in vitro rumen fermentation

随着全球对可持续农业和粮食安全的重视,通过绿色饲料添加剂调控瘤胃发酵过程,已成为提升反刍动物饲料效率和生产性能的重要途径。反刍动物瘤胃发酵是影响其营养物质消化和能量代谢的关键环节。研究表明,饲粮精粗比对瘤胃内环境和养分降解率有显著影响,提高饲粮精粗比可显著提升干物质、有机物及粗蛋白质的瘤胃降解率,且在精粗比为50∶50的条件下,各养分的降解率均达到峰值[1]。但在实际生产中,为追求更快的增重速度,肉羊育肥阶段普遍采用高比例精料饲喂方式。然而,长期饲喂过高比例精料的饲粮,易破坏瘤胃内环境稳态,降低碳水化合物、氮源等养分的转化效率;同时,还会通过干扰瘤胃微生物的正常代谢,间接诱发动物的氧化应激与炎症反应,进而制约生产潜力的发挥,影响畜产品品质[2-4]。因此,开发能够协同调控瘤胃发酵与养分利用效率并兼具改善动物健康与产品品质功能的绿色饲料添加剂,具有重要的现实意义。
番茄红素(lycopene,LYC)是一种天然、安全的类胡萝卜素,其分子式为C40H56,因含11个共轭双键的独特结构,具备强大的抗氧化活性。研究表明,番茄红素清除单线态氧的效率是β-胡萝卜素的2倍、维生素C的100倍[5],可通过捕获单线态氧、清除活性氧(ROS)及激活核因子E2相关因子2(Nrf2)信号通路诱导抗氧化酶表达等机制,增强动物机体抗氧化防御能力[6-7]。已有研究证明,番茄红素可改善酮病奶牛[8]、围产期奶牛[9]和育肥期肉羊[10]的抗氧化状态,提高生产性能,还能改善羊肉品质[11];另有研究表明,在羊饲粮中添加干番茄果渣可以提高氮的利用效率,减少粪便和尿液中氮的排泄量,并改善瘤胃发酵环境[12-13]。番茄红素具有强效抗氧化活性,可能通过清除瘤胃发酵过程中微生物代谢产生的大量活性自由基,优化瘤胃微生物区系,进而影响反刍动物的瘤胃发酵和营养物质消化率。然而,目前关于番茄红素对反刍动物瘤胃发酵及营养物质消化代谢影响的研究较少,其在不同精粗比饲粮条件下对瘤胃发酵参数的影响尚不明确。
本试验运用体外模拟瘤胃发酵技术,系统探究在不同精粗比饲粮条件下添加不同剂量番茄红素对绵羊瘤胃发酵的影响,旨在为番茄红素作为功能性绿色添加剂在羊生产中的应用提供理论支撑。

1 材料与方法

1.1 试验材料

试验用番茄红素产品购于陕西森弗天然制品有限公司,为棕红色粉末制剂,纯度为10.3%。

1.2 发酵底物制备

采用2×5双因素试验设计,设2个精粗比,分别为55∶45(A1)与80∶20(A2);并在每个精粗比饲粮中分别添加5个梯度的番茄红素产品,添加量分别为0、100、200、300和400 mg/kg,对应的番茄红素实际添加量依次为0(B1)、10.3(B2)、20.6(B3)、30.9(B4)、41.2 mg/kg(B5)。按照试验设计,共配制出10种试验饲粮作为体外发酵底物。2种精粗比饲粮均参照《肉羊营养需要量》(NY/T 816—2021)[14]中体重20 kg、日增重300 g的绵羊公羔营养需要量,以玉米、豆粕、玉米胚芽粕、花生壳及大豆皮为主要原料进行配制,其组成及营养水平见表1。
表1 不同精粗比饲粮组成及营养水平(干物质基础)

Table 1 Composition and nutrient levels of diets with different concentrate-forage ratios (DM basis)%

项目
Items		 精粗比
Concentrate-forage ratios
55∶45 (A1) 80∶20 (A2)
原料 Ingredients
玉米 Corn 27.0 47.5
豆粕 Soybean meal 10.8 10.8
玉米胚芽粕 Corn germ meal 8.0 14.5
花生壳粉 Peanut shell powder 20.0 10.0
大豆皮 Soybean peel 25.0 10.0
次粉 Secondary powder 5.0 3.0
预混料 Premix1) 2.0 2.0
石粉 Limestone 1.0 1.0
磷酸氢钙 CaHPO4 0.5 0.5
食盐 NaCl 0.7 0.7
合计 Total 100.0 100.0
营养水平 Nutrient levels2)
消化能 DE/(MJ/kg) 12.62 13.99
粗蛋白质 CP 14.26 14.26
粗脂肪 EE 3.68 3.11
粗灰分 Ash 7.70 6.75
中性洗涤纤维 NDF 34.00 23.84
酸性洗涤纤维 ADF 22.82 13.28
钙 Ca 0.87 0.87
磷 P 0.41 0.41

1)预混料为每千克饲粮提供The premix provided the following nutrients per kg of diets:VA 20 000 IU,VD 4 000 IU,VE 40 IU,Cu 15 mg,Fe 55 mg,Mn 40 mg,Se 0.3 mg,I 0.5 mg,Co 0.2 mg。

2)消化能参照《肉羊营养需要量》(NY/T 816—2021)[14]计算,其余均为实测值。DE was calculated in accordance with Nutrient Requirements of Meat-Type Sheep and Goat (NY/T 816—2021)[14], while the rest were all measured values.

1.3 供体动物与瘤胃液采集

本试验经山西农业大学动物伦理委员会批准(伦理审批编号:SXAU-EAW-2025S.IY.005021463),所有动物饲养、管理及采样操作均符合动物福利与伦理要求。试验用瘤胃液取自3只安装瘤胃瘘管的杜泊羊与小尾寒羊杂交的F1代公羊,体重为(50.0±4.5) kg。供体羊单栏饲养,每天07:00及19:00自由采食精粗比为55∶45的颗粒型全混合日粮,自由饮水。预试期15 d,第16天早上饲喂前采集瘤胃液,用4层纱布过滤,装入预热至39 ℃、充满二氧化碳(CO2)的暖壶中混匀,迅速带回实验室备用。

1.4 体外发酵试验

培养液的制备:将过滤后的瘤胃液与人工唾液[15]按1∶2体积比混合均匀,移入已预热的Fortuna Polifix分液装置(Poulten & Graf Ltd.,德国),全程维持39 ℃恒温并持续通入CO2保持厌氧环境。
体外发酵试验操作参照Menke等[15]的方法进行。精确称取0.200 0 g发酵底物(提前粉碎至1.0 mm粒径),与对应剂量的番茄红素充分混合后,装入长4.0 cm、宽1.5 cm、孔径38~40 μm的尼龙袋并封口,分别放入发酵管(100 mL注射器,Haberle公司,德国)中,同时设置6个仅放置空尼龙袋的空白对照管。将所有发酵管经39 ℃预热后,向每管加入30 mL上述培养液,倒置轻轻振荡发酵管以彻底排尽管内气体,立即记录初始刻度值(mL),将发酵管置于39 ℃恒温振荡培养箱中,恒温培养48 h,本试验全程设置3次重复。

1.5 样品采集

在体外发酵0、3、6、9、12、18、24、36、48 h时,依次读取并记录注射器活塞刻度值;发酵结束后,将发酵管放在冰水中冷却终止发酵反应,然后将发酵液转移到50 mL离心管中,使用酸度计(PHS-3C,上海雷磁仪器有限公司)立即测定pH,pH测定完后将发酵液分装,放-20 ℃冰箱保存待测。

1.6 测定指标与方法

1.6.1 常规营养成分测定

饲粮中干物质、粗蛋白质、粗脂肪及粗灰分含量分别参照GB/T 6435—2014、GB/T 6432—2018、GB/T 6433—2006和GB/T 6438—2007测定;中性洗涤纤维(NDF)、酸性洗涤纤维(ADF)分别参照GB/T 20806—2022和NY/T 1459—2022测定;钙含量参照GB/T 6436—2018采用高锰酸钾滴定法测定;磷含量参照GB/T 6437—2018采用钒钼黄分光光度法测定。

1.6.2 甲烷(CH4)产量、瘤胃发酵参数和消化酶活性的测定

利用气相色谱仪测定发酵48 h内收集气体的CH4产量[16]。发酵液中挥发性脂肪酸(VFA)浓度参照Wang等[17]报道的气相色谱法测定;氨态氮(NH3-N)浓度采用苯酚-次氯酸钠比色法[18]测定;酶活性参照Agarwal等[19]介绍的3,5-二硝基水杨酸(DNS)比色法测定,所有酶活性单位均定义为在39 ℃、30 min条件下,每毫升瘤胃液产生1 μmol还原糖所需的酶量[20]。

1.6.3 累积产气量和产气参数计算

累积产气量计算公式如下:

某时间点累积产气量(发酵底物0.200 0 g)(mL)=

该时间内产气量(mL)-对应时间内

3 支空白管平均产气量(mL)。

采用以下动态发酵参数模型[21]计算产气参数:
GP=a+b(1-exp-ct)。
式中:GP为t时刻的累积产气量(mL);a为快速降解部分产气量(mL);b为慢速降解部分产气量(mL);c为产气速率(%/h);a+b为潜在产气量(mL)。模型中的a、b、c基于非线性最小二乘法计算。

1.6.4 有机物消化率、消化能和代谢能计算

根据钱元诚等[22]给出的回归方程计算有机物消化率、消化能和代谢能:

DOM=0.760 2 Gb+0.636 5CP+22.5;

DE=0.138 4Gb+0.142CP+0.111EE+2.86;

ME=0.145 6Gb+0.076 75CP+

0.146 2EE+1.198。

式中:DOM为有机物消化率(%);DE为消化能(MJ/kg DM);ME为代谢能(MJ/kg DM);Gb为24 h累积产气量(mL);CP为发酵底物中粗蛋白质含量(% DM);EE为发酵底物中粗脂肪含量(% DM)。

1.7 数据处理与统计分析

试验数据经Excel 2019整理后,用SPSS 25.0开展双因素方差分析。差异显著时,采用Tukey法(方差齐)或Tambane法(方差不齐)进行多重比较。以P<0.05为差异显著标准。

2 结果与分析

2.1 不同精粗比饲粮中添加番茄红素对体外发酵产气量和产气参数的影响

由表2可知,饲粮精粗比由55∶45提高到80∶20,24、48 h累积产气量及慢速降解部分产气量和潜在产气量均显著增加(P<0.05)。随着番茄红素添加量的增加,24、48 h累积产气量及潜在产气量均先升高后降低,B3组显著高于其他各组(P<0.05)。饲粮精粗比与番茄红素添加量的交互作用对24和48 h累积产气量有显著影响(P<0.05),在2种精粗比饲粮中均以添加20.6 mg/kg番茄红素时24和48 h累积产气量最高。
表2 不同精粗比饲粮中添加番茄红素对体外发酵累积产气量和产气参数的影响

Table 2 Effects of lycopene supplementation in diets with different concentrate-forage ratios on cumulative gas production and gas production parameters of in vitro fermentation

项目
Items		 24 h累积
产气量
Cumulative
GP24 h/mL		 48 h累积
产气量
Cumulative
GP48 h/mL		 快速降解部
分产气量
GP of rapidly
degraded
fraction/mL		 慢速降解
部分产气量
GP of slowly
degraded
fraction/mL		 潜在
产气量
Potential
GP/mL		 产气速率
GP rate/
(%/h)
精粗比
Concentrate-
forage ratios		 番茄红素添加量
Lycopene supplemental
levels(mg/kg)
0 (B1) 28.75f 37.00d 6.66 32.31 38.87 0.05
10.3 (B2) 34.25de 42.75c 6.22 36.83 43.04 0.07

55∶45 (A1)		 20.6 (B3) 38.75b 48.25b 8.67 46.67 55.34 0.04
30.9 (B4) 37.00bc 46.50b 10.84 39.06 49.90 0.05
41.2 (B5) 32.25e 42.25c 7.46 38.49 45.94 0.05
0 (B1) 37.50bc 47.75b 4.62 44.41 49.03 0.06
10.3 (B2) 37.50bc 48.00b 5.48 45.36 50.83 0.05

80∶20 (A2)		 20.6 (B3) 41.00a 53.25a 7.80 49.68 57.48 0.05
30.9 (B4) 36.00cd 47.00b 3.20 47.86 51.05 0.05
41.2 (B5) 35.50cd 45.25bc 7.32 41.02 48.34 0.05
均值标准误 SEM 0.671 1.118 1.810 3.099 2.186 0.006

精粗比
Concentrate-forage ratios		 (55∶45) A1 34.20b 43.35b 7.97 38.65b 46.62b 0.05
(80∶20) A2 37.50a 48.25a 5.68 45.67a 51.35a 0.05
番茄红素添加量
Lycopene supplemental
levels/(mg/kg)		 0 (B1) 33.13c 42.38d 5.64 38.31 43.95c 0.06
10.3 (B2) 35.88b 45.38bc 5.84 41.09 46.94bc 0.06
20.6 (B3) 39.88a 50.75a 8.23 48.18 56.41a 0.05
30.9 (B4) 36.50b 46.75b 7.02 43.46 50.48b 0.05
41.2 (B5) 33.88c 43.75cd 7.39 39.75 47.14bc 0.05
P值
P-value		 A <0.001 <0.001 0.074 0.005 0.007 0.637
B <0.001 <0.001 0.598 0.067 0.002 0.104
A×B <0.001 0.012 0.291 0.504 0.327 0.233

A:精粗比;B:番茄红素添加量(以有效成分计,本试验所用番茄红素产品纯度为10.3%,有效成分添加量=产品添加量×10.3%);A×B:精粗比与番茄红素添加量的交互作用。同列数据中不同精粗比或不同番茄红素添加量处理下肩标字母相同或无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。

A: concentrate-forage ratio; B: lycopene supplemental level (on an active ingredient basis, the purity of the lycopene product used in this experiment is 10.3%, and the addition amount of active ingredient=product addition amount×10.3%); A×B: the interaction between concentrate-forage ratio and lycopene supplemental level. Data in the same column with the same or no letters in the shoulder label under different concentrate-to-forage ratios or lycopene supplemental levels indicate no significant difference (P>0.05), while with different lowercase letters indicate a significant difference (P<0.05). The same as below.

2.2 不同精粗比饲粮中添加番茄红素对体外发酵有机物消化率、消化能和代谢能的影响

由表3可知,饲粮精粗比由55∶45提高到80∶20,有机物消化率、消化能和代谢能均显著提高(P<0.05)。随着番茄红素添加量的增加,有机物消化率、消化能和代谢能均先升高后降低,B3组显著高于其他组(P<0.05)。饲粮精粗比与番茄红素添加量的交互作用对有机物消化率、消化能和代谢能有显著影响(P<0.05),在2种精粗比饲粮中均以添加20.6 mg/kg番茄红素时有机物消化率、消化能和代谢能最高,且在低精料饲粮条件下对有机物消化率、消化能和代谢能的提高效果更好。
表3 不同精粗比饲粮中添加番茄红素对体外发酵有机物消化率、消化能和代谢能的影响

Table 3 Effects of lycopene supplementation in diets with different concentrate-forage ratios on OMD, DE and ME of in vitro fermentation

项目
Items		 有机物消化率
OMD/%		 消化能
DE/(MJ/kg DM)		 代谢能
ME/(MJ/kg DM)
精粗比
Concentrate-forage ratios		 番茄红素添加量
Lycopene supplemental levels/(mg/kg)
0 (B1) 53.43f 9.27f 7.02f
10.3 (B2) 57.61de 10.03de 7.82de
55∶45 (A1) 20.6 (B3) 61.03b 10.66b 8.47b
30.9 (B4) 59.70bc 10.41bc 8.22bc
41.2 (B5) 56.09e 9.76e 7.53e
0 (B1) 60.08bc 10.42bc 8.21bc
10.3 (B2) 60.08bc 10.42bc 8.21bc
80∶20 (A2) 20.6 (B3) 62.75a 10.91a 8.72a
30.9 (B4) 58.94cd 10.21cd 7.99cd
41.2 (B5) 58.56cd 10.14cd 7.92cd
均值标准误 SEM 0.510 0.093 0.098
精粗比
Concentrate-forage ratios		 55∶45 (A1) 57.58b 10.03b 7.81b
80∶20 (A2) 60.08a 10.42a 8.21a
番茄红素添加量
Lycopene supplemental
levels/(mg/kg)		 0 (B1) 56.76c 9.85c 7.61c
10.3 (B2) 58.85b 10.23b 8.01b
20.6 (B3) 61.89a 10.78a 8.60a
30.9 (B4) 59.32b 10.31b 8.10b
41.2 (B5) 57.33c 9.95c 7.72c
P值 P-value A <0.001 <0.001 <0.001
B <0.001 <0.001 <0.001
A×B <0.001 <0.001 <0.001

2.3 不同精粗比饲粮中添加番茄红素对体外发酵CH4产量和体外发酵液中VFA浓度的影响

由表4可知,饲粮精粗比由55∶45提高到80∶20,CH4产量和乙丙比显著降低(P<0.05),总挥发性脂肪酸(TVFA)、丙酸和丁酸浓度显著升高(P<0.05),乙酸浓度无显著差异(P>0.05)。随着番茄红素添加量的增加,TVFA、丙酸、丁酸浓度均呈现先升高后降低的变化趋势;在TVFA浓度方面,B3、B4组显著高于B1、B5组(P<0.05);在丙酸浓度方面,B4组显著高于B1、B2、B5组(P<0.05);对于丁酸浓度,B4组显著高于B1组(P<0.05)。饲粮精粗比与番茄红素添加剂量的交互作用对CH4产量、各VFA浓度及乙丙比没有显著影响(P>0.05)。
表4 不同精粗比饲粮中添加番茄红素对体外发酵CH4产量和体外发酵液中VFA浓度的影响

Table 4 Effects of lycopene supplementation in diets with different concentrate-forage ratios on CH4 production of in vitro fermentation and volatile fatty acid concentrations in in vitro fermentation fluid

项目
Items		 甲烷产量
CH4
production/
mL		 总挥发性
脂肪酸
TVFA/
(mmol/L)		 乙酸
Acetate/
(mmol/L)		 丙酸
Propionate/
(mmol/L)		 丁酸
Butyrate/
(mmol/L)		 乙丙比
Acetate/
propionate
ratio
精粗比
Concentrate-
forage ratios		 番茄红素添加量
Lycopene supplemental
levels/(mg/kg)
0 (B1) 11.58 89.44 49.89 15.50 12.35 3.22
10.3 (B2) 11.54 94.51 52.77 16.75 13.02 3.15
55∶45 (A1) 20.6 (B3) 11.51 95.96 52.66 17.25 13.51 3.06
30.9 (B4) 11.57 99.22 53.55 18.38 14.21 2.91
41.2 (B5) 11.36 93.22 51.06 17.00 13.09 3.00
0 (B1) 11.21 94.05 49.67 18.12 13.73 2.74
10.3 (B2) 10.71 99.27 52.78 19.42 14.16 2.72
80∶20 (A2) 20.6 (B3) 10.49 100.24 52.83 19.86 14.27 2.66
30.9 (B4) 10.14 98.42 50.42 19.49 14.76 2.59
41.2 (B5) 10.69 95.32 49.70 18.56 14.20 2.68
均值标准误 SEM 0.299 1.237 1.140 0.315 0.395 0.070
精粗比
Concentrate-forage ratios		 55∶45 (A1) 11.51a 94.47b 51.97 16.98b 13.24b 3.07a
80∶20 (A2) 10.65b 97.46a 51.09 19.09a 14.22a 2.68b
番茄红素添加量
Lycopene supplemental
levels/(mg/kg)		 0 (B1) 11.39 91.75c 49.76 16.81d 13.04b 2.98
10.3 (B2) 11.13 96.89ab 52.78 18.08bc 13.59ab 2.94
20.6 (B3) 11.00 98.10a 52.74 18.55ab 13.89ab 2.86
30.9 (B4) 10.86 98.82a 51.99 18.93a 14.48a 2.75
41.2 (B5) 11.03 94.27bc 50.38 17.78c 13.65ab 2.84
P值 P-value A <0.001 0.003 0.249 <0.001 0.003 <0.001
B 0.489 0.001 0.074 0.001 0.049 0.063
A×B 0.497 0.197 0.582 0.095 0.840 0.775

2.4 不同精粗比饲粮中添加番茄红素对体外发酵液pH和NH3-N浓度的影响

由表5可知,饲粮精粗比由55∶45提高到80∶20,pH显著降低(P<0.05),NH3-N浓度显著升高(P<0.05)。随着番茄红素添加量的增加,pH先升高后降低,在B3组达到了最高,显著高于其他组(P<0.05);添加番茄红素的4个组(B2、B3、B4、B5组)NH3-N浓度均显著低于未添加番茄红素的B1组(P<0.05)。饲粮精粗比与番茄红素添加量的交互作用对pH和NH3-N浓度均有显著影响(P<0.05)。
表5 不同精粗比饲粮中添加番茄红素对体外发酵液pH和NH3-N浓度的影响

Table 5 Effects of lycopene supplementation in diets with different concentrate-forage ratios on pH and NH3-N concentration in in vitro fermentation fluid

项目
Items		 pH 氨态氮
NH3-N/(mg/dL)
精粗比
Concentrate-forage ratios		 番茄红素添加量
Lycopene supplemental levels/(mg/kg)
0 (B1) 6.52c 15.90cdef
10.3 (B2) 6.53c 13.39f
55∶45 (A1) 20.6 (B3) 6.55a 15.45def
30.9 (B4) 6.54b 14.56ef
41.2 (B5) 6.54b 16.29cde
0 (B1) 6.46f 24.71a
10.3 (B2) 6.49e 21.41b
80∶20 (A2) 20.6 (B3) 6.51d 16.19cde
30.9 (B4) 6.50d 17.51cd
41.2 (B5) 6.50d 18.35c
均值标准误 SEM 0.003 0.827
精粗比
Concentrate-forage ratios		 55∶45 (A1) 6.54a 15.12b
80∶20 (A2) 6.49b 19.63a
番茄红素添加量
Lycopene supplemental
levels/(mg/kg)		 0 (B1) 6.49d 20.30a
10.3 (B2) 6.51c 17.40b
20.6 (B3) 6.53a 15.82b
30.9 (B4) 6.52b 16.05b
41.2 (B5) 6.52b 17.32b
P值
P-value		 A <0.001 <0.001
B <0.001 <0.001
A×B 0.037 <0.001

2.5 不同精粗比饲粮中添加番茄红素对体外发酵液中消化酶活性的影响

由表6可知,饲粮精粗比由55∶45提高到80∶20,α-淀粉酶、蛋白酶活性显著升高(P<0.05),羧甲基纤维素酶、β-葡萄糖苷酶、木聚糖酶及果胶酶活性均无显著变化(P>0.05)。随着番茄红素添加量的增加,羧甲基纤维素酶和α-淀粉酶活性均呈现先升后降的趋势,B3组显著高于B1、B5组(P<0.05);B3、B4组的β-葡萄糖苷酶活性显著高于B1、B5组(P<0.05);番茄红素添加量对木聚糖酶、果胶酶、蛋白酶活性均无显著影响(P>0.05)。饲粮精粗比与番茄红素添加量的交互作用对各消化酶活性均无显著影响(P>0.05)。
表6 不同精粗比饲粮中添加番茄红素对体外发酵液中消化酶活性的影响

Table 6 Effects of lycopene supplementation in diets with different concentrate-forage ratios on digestive enzyme activities in in vitro fermentation fluidU/mL

项目
Items		 羧甲基
纤维素酶
Carboxymethyl
cellulase		 β-葡萄
糖苷酶
β-gluco-
sidase		 木聚糖酶
Xylanase		 果胶酶
Pectinase		 α-淀粉酶
α-amylase		 蛋白酶
Protease
精粗比
Concentrate-
forage ratios		 番茄红素添加量
Lycopene supplemental
levels/(mg/kg)
0 (B1) 0.011 0.156 0.060 0.116 0.020 0.332
10.3 (B2) 0.013 0.159 0.062 0.127 0.022 0.328
55∶45 (A1) 20.6 (B3) 0.014 0.164 0.062 0.122 0.023 0.334
30.9 (B4) 0.014 0.167 0.057 0.123 0.023 0.326
41.2 (B5) 0.012 0.153 0.055 0.119 0.022 0.328
0 (B1) 0.011 0.159 0.057 0.117 0.021 0.407
10.3 (B2) 0.012 0.163 0.060 0.124 0.029 0.406
80∶20 (A2) 20.6 (B3) 0.014 0.164 0.063 0.125 0.038 0.409
30.9 (B4) 0.014 0.162 0.059 0.127 0.025 0.412
41.2 (B5) 0.011 0.160 0.059 0.125 0.021 0.395
均值标准误 SEM 0.001 4 0.001 4 0.002 8 0.002 8 0.002 8 0.005 7
精粗比
Concentrate-forage ratios		 55∶45 (A1) 0.013 0.160 0.059 0.121 0.022b 0.329b
80∶20 (A2) 0.012 0.162 0.060 0.124 0.027a 0.406a
番茄红素添加量
Lycopene supplemental
level/(mg/kg)		 0 (B1) 0.011b 0.160b 0.063 0.121 0.025b 0.369
10.3 (B2) 0.013ab 0.164ab 0.066 0.131 0.030ab 0.367
20.6 (B3) 0.014a 0.167a 0.067 0.128 0.035a 0.372
30.9 (B4) 0.013ab 0.168a 0.063 0.130 0.029ab 0.369
41.2 (B5) 0.012b 0.160b 0.062 0.127 0.026b 0.362
P值
P-value		 A 0.120 0.222 0.795 0.302 0.020 <0.001
B 0.016 0.001 0.388 0.060 0.029 0.512
A×B 0.946 0.096 0.794 0.707 0.084 0.611

3 讨论

体外发酵产生的气体量可在很大程度上体现瘤胃微生物对发酵底物的利用情况,因而也可反映营养物质的降解率,且二者呈正相关[23]。产气量与发酵底物的营养成分及微生物活性密切相关。本研究中,与精粗比为55∶45相比,精粗比为80∶20时体外发酵24、48 h累积产气量及慢速降解部分产气量、潜在产气量和有机物消化率均显著升高,这与前人研究结果一致。田希雅等[24]研究表明,随饲粮精粗比从20∶80增至80∶20,绵羊瘤胃体外发酵总产气量及干物质降解率均随精料比例的升高而显著提升。精料中可发酵碳水化合物在瘤胃中更容易被微生物分解,从而产生更多的气体。随着番茄红素添加量的增加,24、48 h累积产气量和潜在产气量及有机物消化率、能量供应(消化能、代谢能)均先升高后降低,添加番茄红素各组的快速降解部分产气量、慢速降解部分产气量在数值也高于未添加番茄红素组,这可能是因为番茄红素通过协同调控瘤胃功能微生物如纤维分解菌、淀粉分解菌等的丰度与活性,提高了碳水化合物的降解效率。目前有关番茄红素对反刍动物瘤胃微生物的影响尚未见报道,但不少研究证明番茄红素可以增加单胃动物肠道有益菌的数量,从而调控单胃动物营养代谢[25-26]。本研究中,高剂量的番茄红素降低体外发酵产气量、有机物消化率及消化能和代谢能潜在供应量,表明高剂量番茄红素对体外瘤胃发酵有一定的抑制作用。番茄红素作为一种类胡萝卜素,具有强抗氧化和抗菌活性[27-28],高剂量的番茄红素可能通过破坏细菌细胞膜稳定性破坏细菌结构,从而抑制瘤胃发酵。因此,今后有必要深入研究番茄红素对羊瘤胃微生物区系的影响,为其在反刍动物生产中的应用提供科学支撑。
在瘤胃发酵体系中,VFA的生成与底物类型及饲粮结构密切相关[29]。其中,65%~80%的TVFA来源于碳水化合物的微生物降解,而纤维素和淀粉作为宿主动物主要的能量来源,是驱动瘤胃微生物合成VFA的关键底物。饲粮精粗比直接调控着瘤胃内VFA的构成、产量及后续代谢流向。当饲粮中粗料占比升高时,TVFA的总产量会随之降低,但瘤胃液中乙酸的相对浓度显著上升。乙酸与丁酸代谢过程中产生的氢气会被瘤胃内的产甲烷菌捕获并用于合成CH4,这一过程直接导致CH4排放量同步增加,同时也造成了部分能量的损耗。反之,若饲粮中精料占比升高时,丙酸浓度明显升高,由于丙酸在发酵过程中可直接利用体系内的氢气,一定程度上抑制了CH4生成,降低了能量以CH4形式流失的比例,从而提升了饲粮的能量利用效率[30]。本试验中,与精粗比为55∶45相比,精粗比为80∶20时体外发酵CH4产量和体外发酵液中乙丙比显著降低,TVFA、丙酸和丁酸浓度则显著升高,与Della Rosa等[31]的研究结果一致。
抗氧化剂在瘤胃中具有重要作用,能够减少氧化应激,保护瘤胃微生物和宿主健康,瘤胃中VFA浓度与抗氧化能力密切相关。李宁等[32]研究发现,与番茄红素结构相似、同属不含氧胡萝卜素的β-胡萝卜素可以改变母犏牛瘤胃微生物区系,提高拟杆菌门相对丰度,提高瘤胃液中VFA和微生物蛋白浓度,促进其对纤维物质的消化利用。本试验中,各添加番茄红素组体外发酵液中TVFA、丙酸和丁酸浓度均高于未添加番茄红素组,与岳云双等[33]的研究结果一致,说明适量的番茄红素可以促进瘤胃发酵。袁旭鹏[34]研究表明,β-胡萝卜素可以显著降低繁殖母猪肠道中甲烷杆菌科数量,但本研究中添加番茄红素未对体外发酵CH4产量产生显著影响。据此推测,本研究中番茄红素的添加量尚不足以调控产甲烷菌的活性,因此未能对CH4产量产生显著影响。
瘤胃液pH是衡量反刍动物瘤胃发酵程度及微生物活性的关键生理指标[35],pH过高或过低均会抑制微生物的正常生长与增殖,并对瘤胃发酵过程产生不利影响[36]。瘤胃液pH受饲粮结构及营养水平等因素影响,反刍动物瘤胃内pH变化范围为5.5~7.5[37],瘤胃pH小于6时纤维素分解菌的活力会大大降低[38]。本试验中pH均在正常范围内,但饲粮精粗比对体外发酵液的pH产生了显著影响,且pH随精粗比的提高而降低。这是由于精粗比提高时,非结构性碳水化合物的比例也随之上升,从而产生更多的VFA使pH降低。有研究表明,番茄红素能够提高瘤胃液pH,维持微生物群落的稳定[39]。本试验中,番茄红素的添加显著提高了发酵液pH,且在调控体外发酵液pH方面饲粮精粗比和番茄红素添加量存在显著的交互作用,在精料比例较高时番茄红素稳定pH的效果更明显。
瘤胃液中NH3-N浓度反映了饲粮中蛋白质及非蛋白氮在瘤胃中的降解程度,同时也是评估瘤胃微生物对氮利用状况的重要指标[40]。本试验中,与精粗比为55∶45相比,精粗比为80∶20时体外发酵液中NH3-N浓度显著升高,这是因为精料中快速降解蛋白质(如豆粕、玉米蛋白)比例更高,被瘤胃微生物分解产氨的速度远快于粗料中的慢速降解纤维蛋白,这与Wang等[41]的研究结果一致。已有研究证明,抗氧化剂(如维生素E、硒等)加入到反刍动物饲粮中后,能够显著影响瘤胃的氮代谢过程,使瘤胃液中的NH3-N浓度下降[42]。本试验中,在高精料饲粮中添加番茄红素后,体外发酵液中NH3-N浓度显著低于未添加组,说明番茄红素可能有减少饲粮中蛋白质被瘤胃微生物降解的作用,这对提高反刍动物氮利用效率具有重要意义。
饲粮精粗比可通过调控瘤胃微生物的种群结构与数量,进而影响瘤胃消化酶活性[43]。本试验结果显示,与精粗比为55∶45相比,精粗比为80∶20时体外发酵液中α-淀粉酶和蛋白酶活性显著升高,与Hao等[44]的研究结果一致,原因是精料富含淀粉、可溶性蛋白和氨基酸等易发酵碳源和氮源,提供了更多的底物供瘤胃微生物利用,刺激了淀粉分解菌和蛋白质分解菌增殖,同时也与饲粮精粗比增加后体外发酵液中丙酸和NH3-N浓度显著升高这一现象相符。本试验中,20.6 mg/kg番茄红素的添加使体外发酵液中羧甲基纤维素酶、β-葡萄糖苷酶、α-淀粉酶活性显著升高。羧甲基纤维素酶与β-葡萄糖苷酶是降解饲粮中纤维素、半纤维素的核心酶系,前者负责断裂纤维素主链,后者则分解纤维二糖等寡糖产物[45]。羧甲基纤维素酶、β-葡萄糖苷酶、α-淀粉酶这3种酶活性的协同升高,意味着番茄红素可能通过改善瘤胃内环境,为产酶微生物提供了更适宜的代谢条件,推测番茄红素可能通过清除瘤胃发酵过程中产生的过量活性氧,减少氧化应激对微生物细胞膜及酶蛋白结构的损伤,从而维持或增强重要功能微生物如纤维分解菌、淀粉分解菌的代谢活性。

4 结论

综上所述,番茄红素可以提高绵羊瘤胃体外发酵产气量、VFA(TVFA、丙酸、丁酸)浓度和消化酶(羧甲基纤维素酶、β-葡萄糖苷酶、α-淀粉酶)活性,降低高精料饲粮条件下NH3-N浓度并稳定pH。本试验条件下,精粗比为55∶45和80∶20的饲粮中均以番茄红素添加量为20.6 mg/kg时对绵羊体外瘤胃发酵的促进效果最佳。

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