Chinese Journal of Animal Nutrition >

2024 , Vol. 36 >Issue 7: 4499 - 4507

DOI: https://doi.org/10.12418/CJAN2024.386

RESEARCH PAPER

Effects of Dietary Clostridium butyricum on Digestion, Metabolism and Serum Biochemical Indexes of Lambs

  • DONG Yingrui ,
  • HAO Xiaoyan ,
  • ZHANG Xuanzi ,
  • ZHANG Jianxin , *
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  • College of Animal Science, Shanxi Agricultural University, Taigu 030801, China
*E-mail:

Received date: 2024-01-25

  Online published: 2024-07-09

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Abstract

This experiment was conducted to investigate the effects of dietary Clostridium butyricum (CB) on digestion, metabolism and serum biochemical indexes of lambs. Forty-eight 3-month-old Dolper×small-tail Han crossbred male lambs with average body weight of (22.82±1.67) kg were randomly divided into 4 groups with 12 replicates per group and 1 lamb per replicate. The basal diets of four groups were supplemented with 0 (control group), 5×107 (LCB group), 5×108 (MCB group) and 5×109 CFU/kg (HCB group) CB (viable count≥5×109 CFU/g), respectively. The pre-experimental period lasted for 15 days, and experimental period lasted for 60 days. After the end of the formal period, the digestion and metabolism experiment was carried out, the adaptation period was 5 days, and the sampling period was 5 days. The results showed as follows: 1) compared with the control group, the neutral detergent fiber (NDF) apparent digestibility of LCB, MCB and HCB groups was significantly increased (P<0.05), and the acid detergent fiber (ADF) apparent digestibility of HCB group was significantly increased (P<0.05). 2) Compared with the control group, the digestible energy of MCB and HCB groups was significantly increased (P<0.05), and the digestibility of gross energy of HCB group was significantly increased (P<0.05). 3) Compared with the control group, the nitrogen apparent digestibility of MCB and HCB groups was significantly increased (P<0.05), and the nitrogen deposition of HCB group was significantly increased (P<0.05). 4) Compared with the control group, the serum total cholesterol (TC) content of MCB and HCB groups was significantly decreased (P<0.05), and the serum low density lipoprotein (LDL) content of HCB group was significantly decreased (P<0.05). In conclusion, dietary appropriate level of CB can improve digestion, metabolism and serum lipid metabolism indexes of lambs. In this experiment, the dietary appropriate CB supplemental level is 5×109 CFU/kg.

Key words： Clostridium butyricum; lambs; growth performance; digestion and metabolism; serum biochemical indexes

Cite this article

DONG Yingrui , HAO Xiaoyan , ZHANG Xuanzi , ZHANG Jianxin . Effects of Dietary Clostridium butyricum on Digestion, Metabolism and Serum Biochemical Indexes of Lambs[J]. Chinese Journal of Animal Nutrition, 2024 , 36(7) : 4499 -4507 . DOI: 10.12418/CJAN2024.386

随着我国肉羊产业的发展,集约化舍饲养殖逐渐成为肉羊育肥的主要生产模式,通过营养调控和外源补充饲料添加剂来优化肉羊的生长性能,以节约成本、提升产品品质、增加生产效益,成为肉羊产业的重要发展趋势。丁酸梭菌(Clostridium butyricum,CB)是从健康动物的肠道中分离出的一种革兰氏芽孢杆菌,因其具有耐高温、耐酸、耐抗生素等特性,已被广泛应用于食品、制药及饲料领域[1-2]。CB能在肠道中定植,产生丁酸等有益物质,调节肠道pH,修复肠黏膜[3];同时,CB能通过调节肠道菌群并分泌多种消化酶来增强动物的消化能力,改善肠道健康[4];CB还具有激活吞噬细胞、提高抗炎抗应激的能力[5]。饲粮中添加CB可提高肉鸡的生长性能和肉品质[6],平衡肠道菌群并促进免疫反应[7]。饲粮中添加CB能改善断奶仔猪小肠形态、肠道微生物区系和免疫功能,减少断奶后的腹泻[8-9]。在反刍动物上的研究发现,CB对提高犊牛生长、缓解腹泻、增强机体免疫功能方面发挥着积极的作用[10]。此外,CB还能促进羔羊骨骼肌发育并改善肉品质[11],改善山羊瘤胃发酵并提高营养物质消化率[12],增强山羊缓解急性免疫应激的能力[13],改善生长性能。
本课题组前期关于CB对羔羊生长性能、屠宰性能及肉品质影响的研究结果显示,饲粮中添加适宜水平的CB能提高羔羊生长性能、屠宰性能,改善肉品质[14]。在此基础上,本试验旨在探讨饲粮中添加CB对羔羊消化代谢和血清生化指标的影响,为提供绿色环保、科学增效的饲料添加剂应用于羔羊饲养中提供科学依据。

1 材料与方法

1.1 试验设计

本试验在2022年7—10月完成,开展于山西右玉肉羊科技小院(宏宇牧业有限责任公司)。试验选用3月龄、平均体重(22.82±1.67) kg的杜泊×小尾寒羊杂交公羔48只,采用单因素试验设计,随机分为4组,每组12个重复,每个重复1只羊。4个组在基础饲粮中分别添加0(对照组)、5×107(LCB组)、5×108(MCB组)、5×109 CFU/kg(HCB组)的CB(活菌数≥5×109 CFU/g)。预试期15 d,正试期60 d。
参照《肉羊营养需要量》(NY/T 816—2021)配制精粗比为60∶40的基础饲粮,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 the basal diet (air-dry basis)%

项目 Items 含量 Content
原料 Ingredients
玉米 Corn 31.42
棉籽粕 Cottonseed meal 3.00
喷浆玉米皮 Sprayed corn husk 6.00
豆粕 Soybean meal 6.00
玉米胚芽粕 Corn germ meal 8.40
石粉 Limestone 0.18
预混料 Premix1) 5.00
燕麦草 Oat grass 30.00
玉米秸秆 Corn straw 5.00
柠条 Caragana korshinskii 5.00
合计 Total 100.00
营养水平 Nutrient levels2)
干物质 DM 91.81
粗灰分 Ash 9.63
粗脂肪 EE 2.34
粗蛋白质 CP 12.18
中性洗涤纤维 NDF 42.16
酸性洗涤纤维 ADF 18.02
代谢能 ME/(MJ/kg) 8.76

1)预混料为每千克饲粮提供 The premix provided the following per kg of the diet:VA 20 000 IU,VD 4 000 IU,VE 40 IU,Fe 55 mg,Se 0.3 mg,Cu 15 mg,Mn 40 mg,I 0.5 mg,Zn 25 mg,Co 0.2 mg。

2)代谢能为计算值[15],其余为实测值。ME was a calculated value[15], while the others were measured values.

1.2 饲养管理

试验前清扫、消毒羊舍。预试期内完成试验羊的分组、编号、内驱和接种疫苗相关工作。晨饲喂料前,将CB与少量饲粮混合,确保试验羊全部摄入后再将剩余饲粮倒入料槽内,通过日采食量对CB添加量进行调整。试验期间,所有试验羊饲养于单栏(1 m×2 m)内,每日07:00和17:00各喂料1次,期间自由采食和饮水。

1.3 样品采集与指标测定

1.3.1 饲粮中营养成分含量的测定

饲粮中干物质(DM)含量参照GB/T 6435—2014测定,粗蛋白质(CP)含量参照GB/T 6432—2018测定,粗脂肪(EE)含量参照GB/T 6433—2006测定,粗灰分(Ash)含量参照GB/T 6438—2007测定,中性洗涤纤维(NDF)含量参照GB/T 20806—2022测定,酸性洗涤纤维(ADF)含量参照NY/T 1459—2022测定。

1.3.2 粪样、尿样收集与处理

正试期结束后对所有试验羊称重,各组随机选取6只试验羊,放置于代谢笼里进行消化代谢试验,并记录每只羊的体重,消化代谢试验结束后再次称重;其中24只试验羊的消化代谢平均初重为40.45 kg,平均末重为42.42 kg。适应期5 d,取样期5 d。适应期结束后,采取全收粪收尿法,称量、收集试验羊5 d的粪便和尿液,并记录每天的采食量。其中,粪样被代谢笼下装置的铁筛网收集,按粪重的10%取样并在每100 g粪样中加入10 mL 10%硫酸;尿液被筛网下的带孔漏板收集到装有100 mL 10%硫酸的塑料桶内,收集的尿液经4层纱布过滤后按1/10取样,将所收集的粪样、尿样分别混匀,放入-20 ℃冰箱保存。

1.3.3 消化代谢相关指标测定

将混匀的粪样称重后置于65 ℃烘箱内烘48 h,回潮12 h后再次称重。将烘干的样品粉碎,过0.4 mm筛,装袋,根据1.3.1中的国标方法测定粪样中的Ash、CP、NDF、ADF含量,计算有机物(OM)含量(OM=1-Ash);总能、粪能、尿能通过氧弹量热仪(IKA C 6000 Isoperibol Package,德国)检测;甲烷能依据赵一广[16]建立的肉用绵羊甲烷估测模型进行计算。相关指标计算公式如下:
某营养物质表观消化率=[(食入该营养物质含量-粪中该营养物质含量)/食入该营养物质含量]×100;
消化能=总能-粪能;
总能表观消化率=(消化能/总能)×100;
代谢能=消化能-尿能-甲烷能;
消化能代谢率=(代谢能/消化能)×100;
总能代谢率=(代谢能/总能)×100;
氮的表观消化率=[(摄入氮-粪氮)/摄入氮]×100;
沉积氮=摄入氮-粪氮-尿氮。

1.3.4 血清生化指标测定

正试期第60天,晨饲前对48只试验羊空腹进行颈静脉采血,采集3管5 mL血液,将采血管倾斜30°放置于37 ℃水浴锅中静置30 min后,在3 000 r/min的转速下离心10 min,分离上清,装在3个1.5 mL离心管内,-20 ℃保存。
血清总蛋白(TP)、尿素氮(UN)、白蛋白(ALB)、总胆固醇(TC)、葡萄糖(GLU)、甘油三酯(TG)、高密度脂蛋白(HDL)、低密度脂蛋白(LDL)含量及谷丙转氨酶(ALT)、谷草转氨酶(AST)、碱性磷酸酶(ALP)活性采用BS200生化分析仪测定,试剂盒购自上海酶联生物科技有限公司。

1.4 数据处理与统计分析

试验数据采用SPSS 26.0软件进行单因素方差分析,Duncan氏法进行多重比较,并进行线性和二次回归分析。结果用平均值和均值标准误(SEM)表示,P<0.05为差异显著。

2 结果

2.1 饲粮中添加CB对羔羊营养物质表观消化率的影响

由表2可知,与对照组相比,LCB、MCB和HCB组的NDF表观消化率显著升高(P<0.05),HCB组的ADF表观消化率显著升高(P<0.05);随着饲粮中CB添加水平增加,NDF和ADF表观消化率呈线性和二次变化(P<0.05),DM表观消化率呈线性升高(P<0.05)。各组之间DM和OM表观消化率无显著差异(P>0.05)。
表2 饲粮中添加CB对羔羊营养物质表观消化率的影响

Table 2 Effects of dietary CB on nutrient apparent digestibility of lambs%

项目
Items		 组别 Groups SEM P值 P-value
对照
Control		 LCB MCB HCB 处理
Treatment		 线性
Linear		 二次
Quadratic
干物质 DM 62.59 64.64 64.10 65.32 0.42 0.120 0.039 0.111
有机物 OM 65.62 67.77 66.83 68.53 0.47 0.136 0.060 0.173
中性洗涤纤维 NDF 43.60b 47.31a 47.58a 48.47a 0.68 0.038 0.009 0.018
酸性洗涤纤维 ADF 24.48b 25.77b 26.27ab 29.71a 0.70 0.040 0.006 0.018

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

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

2.2 饲粮中添加CB对羔羊能量代谢的影响

由表3可知,与对照组相比,MCB和HCB组的消化能显著升高(P<0.05),HCB组的总能消化率显著升高(P<0.05);随着饲粮中CB添加水平增加,消化能和总能消化率呈线性和二次变化(P<0.05),甲烷能、代谢能和总能代谢率呈线性升高(P<0.05)。各组之间摄入总能、粪能、尿能、甲烷能、代谢能、消化能代谢率和总能代谢率无显著差异(P>0.05)。
表3 饲粮中添加CB对羔羊能量代谢的影响

Table 3 Effects of dietary CB on energy metabolism of lambs

项目
Items		 组别 Groups SEM P值 P-value
对照
Control		 LCB MCB HCB 处理
Treatment		 线性
Linear		 二次
Quadratic
摄入总能 GE intake/(MJ/d) 31.98 33.75 33.75 34.11 0.52 0.489 0.172 0.319
粪能 FE/(MJ/d) 11.58 11.76 11.52 11.23 0.22 0.886 0.533 0.728
尿能 UE/(MJ/d) 0.78 0.79 0.93 0.87 0.03 0.396 0.199 0.404
甲烷能 Eg/(MJ/d) 3.05 3.30 3.39 3.45 0.07 0.134 0.023 0.058
消化能 DE/(MJ/d) 20.40b 21.99ab 22.23a 22.88a 0.34 0.048 0.007 0.022
代谢能 ME/(MJ/d) 16.57 17.90 17.91 18.55 0.29 0.092 0.017 0.052
消化能代谢率
Metabolizability of DE/%		 81.15 81.41 80.08 81.08 0.24 0.693 0.637 0.871
总能消化率
Digestibility of GE/%		 63.72b 65.25ab 65.92ab 67.06a 0.43 0.032 0.003 0.012
总能代谢率
Metabolizability of GE/%		 51.72 53.11 53.12 54.38 0.42 0.163 0.028 0.096

2.3 饲粮中添加CB对羔羊氮代谢的影响

由表4可知,与对照组相比,MCB和HCB组的氮表观消化率显著升高(P<0.05),HCB组的沉积氮显著升高(P<0.05);随着饲粮中CB添加水平增加,氮表观消化率和沉积氮呈线性和二次变化(P<0.05),可消化氮呈线性升高(P<0.05)。各组之间摄入氮、粪氮、尿氮和可消化氮无显著差异(P>0.05)。
表4 饲粮中添加CB对羔羊氮代谢的影响

Table 4 Effects of dietary CB on nitrogen metabolism of lambs

项目
Items		 组别 Groups SEM P值 P-value
对照
Control		 LCB MCB HCB 处理
Treatment		 线性
Linear		 二次
Quadratic
摄入氮 N intake/(g/d) 38.28 40.40 40.40 40.82 0.62 0.490 0.173 0.320
粪氮 Fecal N/(g/d) 10.42 10.69 10.03 10.07 0.16 0.429 0.238 0.475
尿氮 Urinary N/(g/d) 13.51 12.96 12.84 12.13 0.50 0.835 0.355 0.658
可消化氮 Digestible N/(g/d) 27.86 29.70 30.37 30.76 0.54 0.241 0.049 0.117
氮表观消化率
Apparent digestibility of N/%		 72.78b 73.49ab 74.98a 75.31a 0.36 0.023 0.002 0.010
沉积氮 Retained N/(g/d) 14.35b 16.75ab 17.46ab 18.63a 0.57 0.041 0.004 0.016

2.4 饲粮中添加CB对羔羊血清生化指标的影响

由表5可知,与对照组相比,MCB和HCB组的血清TC含量显著降低(P<0.05),HCB组的血清LDL含量显著降低(P<0.05);随着饲粮中CB添加水平增加,血清TC和LDL含量呈线性和二次变化(P<0.05),血清UN含量呈线性降低(P<0.05)。各组之间血清TP、ALB、TG、HDL、UN、GLU含量及ALT、AST、ALP活性无显著差异(P>0.05)。
表5 饲粮中添加CB对羔羊血清生化指标的影响

Table 5 Effects of dietary CB on serum biochemical indexes of lambs

项目
Items		 组别 Groups SEM P值 P-value
对照
Control		 LCB MCB HCB 处理
Treatment		 线性
Linear		 二次
Quadratic
总蛋白 TP/(g/L) 70.72 75.97 77.07 75.43 1.68 0.572 0.317 0.363
白蛋白 ALB/(g/L) 45.98 47.42 46.53 45.24 1.09 0.919 0.757 0.792
总胆固醇 TC/(mmol/L) 4.53a 4.36ab 4.08b 3.88b 0.09 0.049 0.005 0.019
甘油三酯 TG/(mmol/L) 1.14 0.91 1.19 1.11 0.05 0.291 0.723 0.751
高密度脂蛋白 HDL/(mmol/L) 1.50 1.61 1.62 1.59 0.06 0.861 0.547 0.685
低密度脂蛋白 LDL/(mmol/L) 2.80a 2.70a 2.38ab 2.14b 0.09 0.030 0.003 0.011
尿素氮 UN/(mmol/L) 5.84 5.13 5.24 4.52 0.18 0.083 0.017 0.061
葡萄糖 GLU/(mmol/L) 6.92 7.12 7.23 7.14 0.25 0.979 0.738 0.909
谷丙转氨酶 ALT/(U/L) 25.38 27.70 26.40 21.63 1.65 0.615 0.402 0.401
谷草转氨酶 AST/(U/L) 24.29 23.22 28.40 29.90 1.54 0.364 0.110 0.260
碱性磷酸酶 ALP/(U/L) 110.28 118.59 118.44 116.26 3.27 0.801 0.551 0.615

3 讨论

3.1 饲粮中添加CB对羔羊营养物质表观消化率的影响

饲粮中营养物质表观消化率对反刍动物的生长发育尤为重要,能反映动物对营养物质的消化吸收能力。益生菌在体内定植可增强胃肠道消化酶活性,调节胃肠道菌群,促进机体消化吸收,提高营养物质的利用率[17]。研究表明,益生菌对犊牛营养物质表观消化率具有积极的影响,其中酵母菌和乳酸菌对NDF、ADF表观消化率的提高优于其他益生菌[18]。给育肥牛添加复合益生菌制剂后,DM、CP、NDF表观消化率均有所提高,ADF表观消化率也呈上升趋势[19]。CB通过增加体内乙酸和丁酸等有机酸的含量,降低胃肠道pH,促进乳酸菌、双歧杆菌等有益菌的增殖,促进胃肠道发育,提高机体对营养物质的消化吸收[20]。Cai等[21]研究发现,饲粮中添加适量CB后,热应激山羊的瘤胃纤维降解酶活性显著提高,同时增加了瘤胃内乙酸、丙酸、丁酸和总挥发性脂肪酸含量,并有效提高了DM、NDF、ADF表观消化率,从而促进生长。此外,小肠是营养物质吸收的主要场所,饲粮中添加CB可通过改善肉兔小肠形态、调节盲肠菌群和影响肠道代谢物组成,提高营养物质的吸收利用率,从而达到促生长的作用[22]。本试验中,羔羊NDF和ADF表观消化率随着饲粮中CB添加水平的增加呈线性和二次升高。结合生长性能数据[14],表明CB通过促进营养物质消化吸收,增加机体对饲粮的利用程度,提高羔羊日增重,进一步促进羔羊生长。

3.2 饲粮中添加CB对羔羊能量代谢的影响

反刍动物摄入的总能伴随着营养物质在体内的消化代谢过程,大部分被机体和瘤胃微生物所利用,一部分以粪能、尿能、甲烷能等的形式损耗[23]。反刍动物维持生长发育所需的能量,来源于饲粮中的碳水化合物、蛋白质和脂肪[24]。其中,饲粮中的碳水化合物含量较高,经采食后主要通过瘤胃微生物降解为挥发性脂肪酸为机体供能,挥发性脂肪酸供能占动物摄入消化能的70%~80%[25],是反刍动物的重要能量来源。脂肪被分解为甘油和脂肪酸等,蛋白质被分解为氨基酸提供能量。研究发现,CB在厌氧发酵过程中可产生大量短链脂肪酸,改善瘤胃发酵,提高能量的利用效率[26]。通过对南美白对虾的肠道微生物代谢分析发现,与对照组相比,3个添加CB试验组的碳水化合物、聚合物、氨基酸等碳源的相对利用率均有所增加[27],这也进一步说明CB能增强肠道微生物的代谢活性,从而提高机体的消化代谢。在本试验中,饲粮中添加不同水平的CB后,消化能、代谢能均有所提高,总能表观消化率较对照组相比分别提高了2.53%、3.58%和5.50%,说明添加适量的CB能加速能量的利用。结合生长性能数据[14],表明CB通过提高羔羊的消化能、代谢能和总能表观消化率,增加机体对能量的利用,进而提高羔羊日增重并促进羔羊生长。

3.3 饲粮中添加CB对羔羊氮代谢的影响

反刍动物摄入的饲粮经瘤胃微生物降解利用合成微生物蛋白,部分未被降解的蛋白质进入小肠。氮代谢可以直观反映反刍动物对饲粮蛋白质的消化利用程度[28],其中沉积氮可反映出反刍动物对饲粮蛋白质的吸收。张环宇[29]研究发现,饲粮中添加30 mg/kg CB显著提高豫东山羊对CP的表观消化率。刘燕娜等[30]研究发现,饲粮中添加添加CB提高了仔猪DM、CP、EE、Ash的表观消化率,促进了营养物质的消化吸收,从而达到提高饲料利用率的效果。溶纤维丁酸弧菌、嗜淀粉瘤胃杆菌、栖瘤胃普雷沃菌是瘤胃中主要的蛋白质降解菌[31],饲粮中添加CB显著提高了荷斯坦小母牛瘤胃黄色瘤胃球菌、白色瘤胃球菌、溶纤维丁酸弧菌、嗜淀粉瘤胃杆菌、牛链球菌的数量[26],其中溶纤维丁酸弧菌、嗜淀粉瘤胃杆菌数量的增加对促进饲粮中氮的降解和合成微生物蛋白具有积极的影响。在本试验研究中,饲粮中添加CB提高了氮表观消化率和沉积氮,这可能与添加CB影响瘤胃微生物的数量及促进蛋白质的消化吸收有关。而添加适量的CB能增强断奶仔猪十二指肠和空肠的脂肪酶活性及胰腺胰蛋白酶活性,从而加速小肠中的蛋白质在肠腔内的消化,提高机体对氮的消化利用率[20]。饲料中添加CB能改善南美白对虾的肠道微生物组成,提高消化相关基因的相对表达水平[27]。这也进一步说明CB可通过改善肠道菌群和提高消化酶基因的表达来增强肠道的消化能力,提高对蛋白质等营养物质的利用。

3.4 饲粮中添加CB对羔羊血清生化指标的影响

血清TP、ALB和UN含量反映了机体内蛋白质的吸收和代谢情况。血清UN含量可直观反映机体的营养水平和蛋白质代谢状况,血清UN含量的降低表明UN在机体中的沉积增多,更有利于机体合成蛋白质,提高动物的生长发育。饲粮中添加含CB的复合益生菌极显著提高了塔里木鸽血清TP含量,并随着复合益生菌添加水平的增加,血清UN含量有下降趋势[32]。对断奶仔猪的研究发现,当饲粮中CB添加量达到500 mg/kg时,血清TP含量显著提高;当饲粮中CB添加量达到2 000 mg/kg时,血清ALB含量显著升高[33]。而饲粮中添加30 mg/kg CB可显著降低豫东山羊血清UN和TC含量[29]。在本试验中,饲粮中添加CB虽不影响血清TP、ALB含量,但3个CB添加组的血清TP含量较对照组分别提高了7.42%、8.98%和6.66%,并有降低血清UN含量的趋势,这与饲粮中添加CB后提高氮表观消化率和沉积氮的结果一致,进一步说明CB能提高羔羊对蛋白质的利用。
血清生化指标TG、TC、HDL和LDL用于评估血脂的代谢及心血管健康。HDL和LDL是胆固醇的运输载体,HDL有助于清除血液中胆固醇,LDL含量的增加可能会导致胆固醇沉积,导致动脉粥样硬化[34]。侯冉等[34]研究发现,饲粮中添加5 g/d CB(活菌数5×108 CFU/g)显著降低了小尾寒羊血清TG、TC和LDL含量。在不同精粗比饲粮的条件下,添加1×108 CFU/kg CB均有降低血清TC含量的趋势[35]。本试验中,饲粮中添加CB降低了血清TC和LDL含量,与前人的研究结果一致。

4 结论

饲粮中添加CB能提高羔羊对NDF、ADF表观消化率,提高消化能、总能消化率及氮表观消化率、沉积氮,从而促进羔羊生长,并改善血清脂质代谢指标。本试验条件下,饲粮中CB的适宜添加水平为5×109 CFU/kg。

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