动物营养学报 >

2026 , Vol. 38 >Issue 6: 4663 - 4675

DOI: https://doi.org/10.12418/CJAN2026.373

研究论文

乳酸菌添加剂对全株玉米与向日葵混合青贮发酵品质和微生物群落的影响

  • 珠拉 , 1 ,
  • 赵牧其尔 1 ,
  • 闫雨婷 1 ,
  • 孙林 2 ,
  • 格根图 , 1, *
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  • 1 内蒙古农业大学草业学院, 农业农村部饲草栽培、加工与高效利用重点实验室, 呼和浩特 010011
  • 2 内蒙古自治区农牧业科学院, 呼和浩特 010031
* 格根图,教授,博士生导师,E-mail:

珠 拉(1999—),女,蒙古族,内蒙古阿拉善人,博士研究生,从事牧草加工与利用研究。E-mail:

Office editor: 田艳明

收稿日期: 2025-11-27

  网络出版日期: 2026-06-13

基金资助

中央支持地方高校改革发展项目——“一区两基地”建设项目(DC2500000240)

收起

Effects of Lactic Acid Bacteria Additives on Fermentation Quality and Microbiota of Whole Plant Corn and Sunflower Mixed Silage

  • ZHU La , 1 ,
  • ZHAO Muqier 1 ,
  • YAN Yuting 1 ,
  • SUN Lin 2 ,
  • GE Gentu , 1, *
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  • 1 Key Laboratory of Forage Cultivation, Processing and Highly Efficient Utilization of Ministry of Agriculture and Rural Affairs, College of Grassland Science, Inner Mongolia Agricultural University, Hohhot 010011, China
  • 2 Inner Mongolia Academy of Agricultural and Animal Husbandry Sciences, Hohhot 010031, China
* professor, E-mail:

Received date: 2025-11-27

  Online published: 2026-06-13

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摘要

本试验旨在探究添加乳酸菌对全株玉米与向日葵混合青贮发酵品质和微生物群落的影响。试验选择全株玉米与全株向日葵以5∶5比例进行混合青贮,采用单因素设计设置3个组,分别为对照组(CK组,不添加添加剂)、布氏乳杆菌组(LB组,添加布氏乳杆菌)和戊糖片球菌组(PP组,添加戊糖片球菌),每组3个重复。室温密封发酵60 d后,测定营养成分含量、发酵品质和微生物群落。结果表明:1)LB组粗蛋白质和水溶性碳水化合物含量显著低于CK组和PP组(P<0.05),PP组酸性洗涤纤维含量显著低于CK组和LB组(P<0.05)。2)LB组和PP组pH和乙酸含量显著低于CK组(P<0.05),乳酸含量显著高于CK组(P<0.05);同时,PP组乳酸含量显著高于LB组(P<0.05)。3)PP组V-Score评分最高,LB组次之,CK组最低。4)与CK组相比,LB组和PP组微生物群落ACE指数和Chao1指数显著降低(P<0.05),Shannon指数显著提高(P<0.05)。在门水平上,与CK组相比,PP组厚壁菌门(Firmicutes)相对丰度提高,变形菌门(Proteobacteria)相对丰度降低;在属水平上,与CK组相比,LB组和PP组慢生乳杆菌属(Lentilactobacillus)相对丰度提高。综上所述,添加布氏乳杆菌和戊糖片球菌均能通过优化全株玉米与向日葵混合青贮微生物群落结构提高其发酵品质,在本试验条件下,添加戊糖片球菌青贮效果较好。

关键词： 玉米; 向日葵; 青贮; 发酵品质; 微生物群落

本文引用格式

珠拉 , 赵牧其尔 , 闫雨婷 , 孙林 , 格根图 . 乳酸菌添加剂对全株玉米与向日葵混合青贮发酵品质和微生物群落的影响[J]. 动物营养学报, 2026 , 38(6) : 4663 -4675 . DOI: 10.12418/CJAN2026.373

Abstract

This experiment was conducted to explore the effects of adding lactic acid bacteria on the fermentation quality and microbiota of whole plant corn and sunflower mixed silage. The whole plant corn and whole plant sunflower were selected for mixed ensiling in a ratio of 5∶5 in this experiment. A single-factor design was adopted to set up three groups, namely the control group (CK group, no additives were added), the Lactobacillus buchneri group (LB group, Lactobacillus buchneri was added), and the Pediococcus pentosaceus group (PP group, Pediococcus pentosaceus was added), with 3 replicates in each group. After 60 days of room-temperature sealed fermentation, the nutrient contents, fermentation quality and microbiota were determined. The results showed as follows: 1) the contents of crude protein and water-soluble carbohydrate in LB group were significantly lower than those in CK group and PP group (P<0.05), and the acid detergent fiber content in PP group was significantly lower than that in CK group and LB group (P<0.05). 2) The pH and acetic acid content in LB group and PP group were significantly lower than those in CK group (P<0.05), while the lactic acid content was significantly higher than that in CK group (P<0.05). Meanwhile, the lactic acid content in PP group was significantly higher than that in LB group (P<0.05). 3) The V-Score score was the highest in PP group, followed by LB group, and the lowest in CK group. 4) Compared with CK group, the ACE index and Chao1 index of microbiota in LB group and PP group were significantly decreased (P<0.05), and the Shannon index was significantly increased (P<0.05). At the phylum level, compared with CK group, the Firmicutes relative abundance in PP group was increased, while the Proteobacteria relative abundance was decreased; at the genus level, compared with CK group, the Lentilactobacillus relative abundance in LB group and PP group was increased. In conclusion, the addition of Lactobacillus buchneri and Pediococcus pentosaceus can both improve the fermentation quality of whole plant corn and sunflower mixed silage by optimizing the microbiota structure. Under the conditions of this experiment, the silage effect of adding Pediococcus pentosaceus is better.

Key words： corn; sunflower; silage; fermentation quality; microbiota

青贮饲料是现代畜牧业可持续发展的重要基础,其品质直接影响动物生产性能、健康状况及养殖经济效益[1]。玉米因其水溶性碳水化合物(water-soluble carbohydrate,WSC)含量高、青贮技术成熟,被普遍视为优质青贮原料[2-3]。尽管玉米干物质(dry matter,DM)含量高、WSC丰富,有利于获得优良青贮品质,但单一玉米青贮也存在蛋白质含量较低、营养结构较为单一的问题[4-5]。因此,积极探索和开发非常规、适应当地环境的饲草资源,以实现饲料供应体系多元化并增强产业韧性,已成为草地科学领域的重要研究方向[6]。向日葵(Helianthus annuus L.)具有耐旱、耐盐碱的特点,生物产量高且富含纤维资源,是一种很有潜力的青贮替代原料,青贮的向日葵花盘还可以用作牛、羊等牲畜的良好饲料[7-8]。巴西等地已普遍采用全株向日葵制作青贮饲料,而国内对其营养成分、加工技术及饲用价值等方面的研究却十分匮乏[9]。不过,全株向日葵粗蛋白质(crude protein,CP)含量较高[10]、纤维成分多[11],单独青贮时往往难以启动有效的乳酸(lactic acid,LA)发酵,导致pH下降缓慢,梭菌等有害微生物活动加剧,最终造成养分损失较大、营养品质下降,并可能产生大量丁酸(butyric acid,BA)等不良发酵产物,这限制了其在实际生产中的应用[12]。研究表明,开花期全株向日葵的可消化蛋白质含量为6.9%,养分含量达58.5%,且与禾本科牧草混合青贮可获得较优品质[13]。因此,将全株玉米与全株向日葵进行混合青贮,理论上可实现2种作物在营养成分上的互补,从而优化青贮饲料的营养结构,提升其综合应用价值。此外,除了利用原料特性进行混合青贮外,添加外源添加剂也是提高发酵品质的重要措施,其中布氏乳杆菌和戊糖片球菌在提高青贮发酵品质方面应用较广。因此,本试验选择全株玉米与全株向日葵按进行混合青贮,并通过添加乳酸菌探究其对营养成分含量、发酵品质和微生物群落的影响,以期为开发高品质玉米与向日葵青贮饲料提供参考。

1 材料与方法

1.1 试验材料

本试验采用的青贮原料为全株玉米和全株向日葵,种植于呼和浩特市土默特左旗内蒙古农业大学海流图实验教学基地,于2024年9月1日[全株玉米(1/2乳线期),全株向日葵(灌浆期)]进行刈割。全株玉米与全株向日葵混合比例为5∶5。青贮原料营养成分含量见表1。
表1 青贮原料营养成分含量(实测值)

Table 1 Nutrient contents of silage raw materials (measured values)

项目
Items		 全株玉米
Whole plant corn		 全株向日葵
Whole plant sunflower
干物质 DM/% FW 27.86 31.56
粗蛋白质 CP/% DM 7.98 15.63
水溶性碳水化合物 WSC/% DM 7.54 2.91
中性洗涤纤维 NDF/% DM 50.11 45.86
酸性洗涤纤维 ADF/% DM 32.55 39.75
粗脂肪 EE/% DM 2.22 9.83
乳酸菌添加剂均为市售产品,其中布氏乳杆菌活菌数为2×1010 CFU/g,戊糖片球菌活菌数为2×1010 CFU/g。

1.2 试验设计

试验采用单因素设计,设置3个组,分别为对照组(CK组,不添加添加剂)、布氏乳杆菌组(LB组,添加布氏乳杆菌)和戊糖片球菌组(PP组,添加戊糖片球菌),每组3个重复。将添加剂溶解在20 mL蒸馏水中,并均匀地喷洒在600 g样品上,装入30 cm×40 cm聚乙烯袋中,每袋填装200 g,使用真空包装机抽真空密封,室温条件下青贮60 d后,开袋取样分析营养成分含量、发酵品质和微生物群落等指标。

1.3 测定指标及方法

1.3.1 营养成分含量

将原料和青贮样品于105 ℃烘箱中处理15 min以实现酶失活,随后转入65 ℃烘箱持续烘干48 h至恒重,最后进行粉碎。DM含量参照GB/T 6435—2014[14]中方法测定;CP含量参照GB/T 6432—2018[15]采用凯氏定氮法测定;中性洗涤纤维(neutral detergent fiber,NDF)和酸性洗涤纤维(acid detergent fiber,ADF)含量分别参照GB/T 20806—2022[16]和NY/T 1459—2022[17]采用ANKOM A2000i全自动纤维分析仪(美国)测定;WSC含量通过蒽酮-硫酸比色法[18]测定;粗脂肪(ether extract,EE)含量参照GB/T 6433—2006[19]采用ANKOM XT15i全自动脂肪分析仪(美国)进行测定。

1.3.2 发酵品质

青贮开袋后,称取10 g样品,加入90 mL蒸馏水,经均质拍打仪匀质2 min,过滤后获得青贮浸提液。采用酸度计测定浸提液pH;乳酸、乙酸(acetic acid,AA)、丙酸(propionic acid,PA)和丁酸含量采用Agilent 1100高效液相色谱仪(美国)进行测定;氨态氮(ammonia nitrogen,NH3-N)含量采用苯酚-次氯酸钠比色法[20]进行测定。

1.3.3 青贮饲料V-Score评分

采用V-Score体系[21]对青贮饲料进行评分,该体系通过计算NH3-N/总氮(total nitrogen,TN)、乙酸+丙酸及丁酸含量3项指标的得分总和来评定青贮饲料质量。依据最终得分,青贮饲料质量等级被界定为:大于80分为良好,60~80分为尚可,低于60分为不良。V-Score评分标准见表2。
表2 V-Score评分标准

Table 2 V-Score scoring criteria

氨态氮/总氮 NH3-N/TN 乙酸+丙酸 AA+PA 丁酸 BA V-Score得分
V-Score score
含量
Content/%		 计算式
Formula		 含量
Content/(g/kg FM)		 计算式
Formula		 含量
Content/(g/kg FM)		 计算式
Formula
≤5 y1=50 ≤0.2 y2=10 ≤0.5 y3=(40-80x3)


y=y1+y2+y3
5~10 y1=60-2x1 0.2~1.5 y2=(150-100x2)/13 >0.5 0
10~20 y1=80-4x1 >1.5 y2=0
>20 y1=0

x1、x2和x3分别表示氨态氮/总氮以及乙酸+丙酸、丁酸含量,y1、y2和y3分别表示氨态氮/总氮以及乙酸+丙酸、丁酸含量对应的得分,y表示得分总和。

x1, x2 and x3 represented NH3-N/TN and the contents of AA+PA and BA, y1, y2 and y3 represented scores corresponding to NH3-N/TN and the contents of AA+PA and BA, respectively, and y represented the total score.

1.3.4 微生物群落

将青贮样品送至上海美吉生物医药科技有限公司进行微生物群落分析。采用FastDNA® Spin Kit for Soil试剂盒提取样品总DNA,通过1%琼脂糖凝胶电泳检测其完整性,并采用NanoDrop 2000分光光度计(Thermo Scientific,美国)测定其浓度和纯度。以引物27F(5'-AGRGTTTGATYNTGGCTCAG-3')和1492R(5'-TASGGHTACCTTGTTASGACTT-3')对细菌16S rRNA基因全长进行PCR扩增。扩增程序为:95 ℃预变性3 min;95 ℃ 30 s,55 ℃ 30 s,72 ℃ 30 s,27个循环;72 ℃终延伸10 min。将同一样本的PCR产物合并,经2%琼脂糖凝胶电泳检测后,进行磁珠纯化,并使用QuantusTM Fluorometer荧光计(美国)对纯化产物进行定量。随后按测序量要求将样本按比例混合,使用SMRTbell® Express Template Prep Kit 2.0构建文库,最后在PacBio Sequel Ⅱ System平台上完成测序。

1.4 数据统计与分析

试验数据采用Excel 2016进行初步整理后,采用SPSS 27.0软件对数据进行单因素方差分析(one-way ANOVA),并采用Duncan氏法进行多重比较;结果数据采用“平均值±标准差”形式表示,P<0.05表示差异显著。

2 结果与分析

2.1 乳酸菌添加剂对全株玉米与向日葵混合青贮营养成分含量的影响

由表3可知,乳酸菌添加剂对全株玉米与向日葵混合青贮CP、WSC和ADF含量有显著影响(P<0.05),对DM、NDF和EE含量无显著影响(P>0.05)。LB组CP含量显著低于CK组和PP组(P<0.05),CK组与PP组间CP含量无显著差异(P>0.05);LB组WSC含量显著低于CK组和PP组(P<0.05),同时PP组WSC含量显著低于CK组(P<0.05);PP组ADF含量显著低于CK组和LB组(P<0.05),CK组与LB组间ADF含量无显著差异(P>0.05)。
表3 乳酸菌添加剂对全株玉米与向日葵混合青贮营养成分含量的影响

Table 3 Effects of lactic acid bacteria additives on nutrient contents of whole plant corn and sunflower mixed silage

项目
Items		 组别 Groups P值
P-value
CK LB PP
干物质 DM/% FW 38.02±1.24 33.91±0.54 35.12±0.90 0.051
粗蛋白质 CP/% DM 15.98±1.34a 8.82±0.53b 19.31±0.99a 0.001
水溶性碳水化合物 WSC/% DM 5.10±0.03a 1.70±0.14c 3.83±0.42b <0.001
中性洗涤纤维 NDF/% DM 43.41±1.27 42.10±0.64 41.35±0.37 0.877
酸性洗涤纤维 ADF/% DM 34.33±1.06a 32.71±1.04a 28.52±0.67b 0.012
粗脂肪 EE/% DM 4.56±0.28 5.15±0.25 4.64±0.16 0.232

同行数据肩标不同字母表示差异显著(P<0.05),相同字母或无字母表示差异不显著(P>0.05)。表4同。

In the same row, values with different letter superscripts indicated significant difference (P<0.05), while with the same letter or no letter superscripts indicated no significant difference (P>0.05). The same as Table 4.

2.2 乳酸菌添加剂对全株玉米与向日葵混合青贮发酵品质的影响

由表4可知,乳酸菌添加剂对全株玉米与向日葵混合青贮pH以及乳酸、乙酸、丙酸和NH3-N含量有显著影响(P<0.05),各组均未检测到丁酸。LB组和PP组pH和乙酸含量显著低于CK组(P<0.05),且LB组与PP组间pH和乙酸含量无显著差异(P>0.05);LB组和PP组乳酸含量显著高于CK组(P<0.05),同时PP组乳酸含量显著高于LB组(P<0.05);LB组和PP组未检测到丙酸;LB组NH3-N含量显著低于PP组(P<0.05),有低于CK组的趋势,但差异不显著(P>0.05)。
表4 乳酸菌添加剂对全株玉米与向日葵混合青贮发酵品质的影响

Table 4 Effects of lactic acid bacteria additives on fermentation quality of whole plant corn and sunflower mixed silage

项目
Items		 组别 Groups P值
P-value
CK LB PP
pH 4.07±0.02a 4.00±0.02b 3.95±0.02b 0.006
乳酸 LA/(g/kg FM) 2.55±0.16c 4.21±0.19b 6.46±0.06a <0.001
乙酸 AA/(g/kg FM) 1.24±0.11a 0.85±0.00b 0.65±0.02b 0.002
丙酸 PA/(g/kg FM) 0.26±0.00 ND ND <0.001
丁酸 BA/(g/kg FM) ND ND ND
氨态氮/总氮 NH3-N/TN/% 0.77±0.16ab 0.57±0.15b 1.02±0.28a 0.037

ND:未检测到

not detected。

2.3 全株玉米与向日葵混合青贮V-Score评分

由表5可知,各组全株玉米与向日葵混合青贮NH3-N/TN和丁酸得分均一样,青贮品质均界定为良好。在不同乳酸菌添加剂处理下,PP组V-Score得分最高,排名第1;LB组次之,排名第2;CK组排名最后。
表5 全株玉米与向日葵混合青贮V-Score评分

Table 5 V-Score scoring of whole plant corn and sunflower mixed silage

组别
Groups		 氨态氮/总氮得分
NH3-N/TN score		 乙酸+丙酸得分
AA+PA score		 丁酸得分
BA score		 V-Score得分
V-Score score		 等级
Grade		 排名
Rank
CK 50 0 40 90 良好 3
LB 50 5 40 95 良好 2
PP 50 7 40 97 良好 1

2.4 乳酸菌添加剂对全株玉米与向日葵混合青贮微生物群落的影响

2.4.1 α多样性分析

如图1所示,各组全株玉米与向日葵混合青贮微生物群落Goods_coverage指数均超过99%,表明测序深度足以进行有效群落鉴定;LB组和PP组ACE指数和Chao1指数显著低于CK组(P<0.05),同时PP组ACE指数和Chao1指数显著低于LB组(P<0.05);LB组和PP组Shannon指数显著高于CK组(P<0.05),同时LB组Shannon指数显著高于PP组(P<0.05)。
图1 全株玉米与向日葵混合青贮微生物群落α多样性分析

*表示P<0.05,**表示P<0.01,***表示P<0.001。

Fig.1 Analysis of α diversity of microbiota of whole plant corn and sunflower mixed silage

* indicated P<0.05, ** indicated P<0.01, and *** indicated P<0.001.

2.4.2 β多样性分析

为了更清楚地了解混合青贮中微生物群落结构是否发生了变化,基于加权UniFrac距离的主坐标分析(PCoA)结果(图2-A)显示,LB组与PP组样本距离相近,微生物群落结构差异较小;同时,LB组和PP组样本离CK组均较远,微生物群落结构差异较大;不过,各组样本独立分为一个类群,表明不同组间差异明显。扩增子序列变异(ASV)韦恩图(图2-B)显示,3组微生物群落共有ASV数为32个;CK组与LB组、CK组与PP组以及LB组与PP组共有ASV数分别为15、4和4个;CK组特有ASV数为8个,LB组特有ASV数为5个,PP组特有ASV数为1个。
图2 全株玉米与向日葵混合青贮微生物群落主坐标分析(A)和扩增子序列变异韦恩图(B)

Fig.2 PCoA (A) and Venn diagram (B) of microbiota of whole plant corn and sunflower mixed silage

2.4.3 微生物群落组成及差异菌群分析

如图3-A所示,在门水平上,各组全株玉米与向日葵混合青贮微生物群落组成相似,均以厚壁菌门(Firmicutes)相对丰度最高,其次为变形菌门(Proteobacteria);同时,与CK组相比,PP组厚壁菌门相对丰度提高,变形菌门相对丰度降低。如图3-B所示,在属水平上,各组微生物群落组成存在差异,其中CK组相对丰度较高的菌属为乳植杆菌属(Lactiplantibacillus)(80.6%)和乳球菌属(Lactococcus)(6.4%);LB组的主要优势菌属为慢生乳杆菌属(Lentilactobacillus)(61.1%)和乳植杆菌属(14.4%);PP组的主要优势菌属为慢生乳杆菌属(80.2%)和乳植杆菌属(12.6%)。如3-C所示,线性判别分析(LDA)效应大小(LEfSe)分析结果(|LDA|>4)表明,PP组慢生乳杆菌属和片球菌属(Pediococcus)等显著富集(P<0.05);LB组乳球菌属和拉乌尔菌属(Raoultella)等显著富集(P<0.05)。
图3 全株玉米与向日葵混合青贮微生物群落在门(A)和属(B)水平上组成及差异菌群LEfSe分析

CK_1、CK_2和CK_3为对照组样本,LB_1、LB_2和LB_3为布氏乳杆菌组样本,PP_1、PP_2和PP_3为戊糖片球菌组样本。图4同。

Fig.3 Composition of microbiota at phylum (A) and genus (B) levels and LEfSe analysis of differential microbiota of whole plant corn and sunflower mixed silage

CK_1, CK_2 and CK_3 were samples in control group, LB_1, LB_2 and LB_3 were samples in Lactobacillus buchneri group, and PP_1, PP_2 and PP_3 were samples in Pediococcus pentosaceus group. The same as Fig.4.

2.4.4 相似性百分比(SIMPER)分析

SIMPER分析是一种用于比较不同组间物种组成差异的多变量分析方法,其主要目的是计算每个物种的贡献值。如图4所示,与CK组相比,LB组慢生乳杆菌属贡献值较高,其次是乳球菌属;PP组慢生乳杆菌属贡献值最高,其次是片球菌属。
图4 全株玉米与向日葵混合青贮微生物群落SIMPER分析

Fig.4 SIMPER analysis of microbiota of whole plant corn and sunflower mixed silage

3 讨论

3.1 乳酸菌添加剂对全株玉米与向日葵混合青贮发酵品质的影响

在青贮过程中,乳酸菌的群落结构与功能对发酵品质及营养价值具有决定性影响。尽管乳酸菌通过代谢WSC生成乳酸,从而抑制有害微生物生长的机制已较为明确[22],但其具体作用仍受菌种类型、添加方式及原料特性等因素调控。营养成分含量是评价青贮饲料品质最直接的指标之一[23],其中DM含量是反映青贮饲料保存状态和营养损失程度的重要参数[24]。本研究中,CK组DM含量最高,而LB组最低。这可能是因为外源乳酸菌迅速利用原料中的WSC进行发酵,产生乳酸及其他有机酸,在此过程中部分营养物质转化为挥发性组分而损失,从而导致DM含量下降[25-26]。这也解释了PP组WSC含量和pH较低而乳酸含量较高的原因。值得注意的是,LB组WSC含量显著低于其他2组,该组添加的布氏乳杆菌属于异型发酵乳酸菌,其代谢途径会同时产生乳酸、二氧化碳等产物,导致更多的底物消耗。林秀蔚等[27]报道,异型发酵乳酸菌组WSC含量低于对照组,与本试验结果一致。Carvalho等[28]和Reich等[29]也认为,异型发酵乳酸菌会增加DM和WSC的损失。此外,有研究报道异型发酵乳酸菌会导致CP含量降低[30]。本研究也得到了同样的结论,LB组CP含量显著低于其他组。相比之下,PP组CP含量最高,该结果与王昊然[31]的研究结果一致,表明戊糖片球菌在提高CP含量方面具有积极作用。研究表明,全株玉米与向日葵秸秆进行混合青贮后ADF含量显著降低[32]。本研究中,PP组ADF含量为28.52%,且显著低于其他2组,表明该组青贮饲料具有较好的可采食性和消化特性[33]。此外,优质玉米青贮的NDF含量通常应低于55%[34-35],本研究各组均符合该标准,达到优质青贮水平。各组间EE含量无显著差异,与多数相关研究结果[36-38]一致。
pH是评价青贮饲料质量的关键参数之一,较低的pH环境(通常低于4.60)可有效抑制不良微生物活动及蛋白质水解酶的活性,有利于饲料保存[39]。本研究显示,各乳酸菌添加组pH均显著低于CK组,这主要得益于外源添加的乳酸菌迅速启动了发酵过程。其中,戊糖片球菌作为同型发酵乳酸菌,能够高效利用WSC生成大量乳酸,直接推动pH快速下降。同时,添加剂的引入改变了青贮体系的微生物群落结构,使乳酸菌在竞争中占据优势,从而主导了以产酸为主的发酵方向。在发酵过程中,微生物通过代谢消耗部分DM和WSC,将其转化为乳酸等有机酸及其他代谢产物[40]。因此,本研究中LB组和PP组DM和WSC含量均低于CK组。相应地,PP组乳酸含量达到6.46 g/kg FM,显著高于其他组,进一步印证了其产酸能力的优势。在青贮期间,蛋白质在植物酶和微生物酶的作用下逐步降解为NH3-N、游离氨基酸及多肽等非蛋白氮组分[41],该过程反映了蛋白质的分解程度[20]。本研究中,各组NH3-N含量均维持在较低水平,表明蛋白质分解受到一定抑制。尤其是PP组在保持较高CP含量的同时,NH3-N较低,说明该处理有利于青贮饲料蛋白质的保存。该组较低的pH环境有效抑制了梭菌等有害微生物的生长和代谢,从而减少了不良发酵产物的生成。丁酸常作为评估青贮中梭菌活动的指标,其含量较高通常反映有害微生物活跃,不利于青贮品质[42]。本研究在所有组中均未检测到丁酸,进一步说明发酵过程处于较好的微生物调控状态,未出现明显的发酵异常。
本研究采用日本草地畜产种子协会制定的V-Score评分系统对青贮品质进行评估,该系统是青贮研究中广泛应用的评估工具[43]。本试验中,鉴于全株玉米与全株向日葵的原料配比本身为乳酸发酵奠定了良好基础,因此CK组取得了较高的V-Score评分;而添加乳酸菌制剂(尤其是戊糖片球菌)进一步优化了微生物群落,推动了乳酸发酵,提升了乳酸含量,同时降低了pH和NH3-N生成。最终,PP组V-Score评分高达97分,属于优质青贮的范畴。

3.2 乳酸菌添加剂对全株玉米与向日葵混合青贮微生物群落的影响

青贮饲料的微生物群落结构与其发酵品质密切相关[44]。在青贮过程中,原料表面附着的微生物在厌氧环境下发生复杂演替,其种类和数量对最终青贮品质具有重要影响[45]。乳酸菌作为青贮体系中最关键的有益菌群,其相对丰度及可利用发酵底物的充足程度是影响青贮成功与否的重要因素[46]。本研究采用16S rRNA基因高通量测序技术对青贮过程中的细菌群落结构进行解析[47],测序覆盖度均高于99%,表明数据量充分,能够可靠反映菌群组成。同时,通过ACE指数、Chao1指数和Shannon指数对微生物群落α多样性进行评估,其中ACE指数和Chao1指数用于反映物种丰富度,数值越低表明丰富度越低[48]。本研究结果显示,PP组ACE指数和Chao1指数均显著低于CK组,说明接种戊糖片球菌降低了微生物群落的丰富度,该结果与Li等[49]的研究结果一致。Zhao等[50]研究表明,在玉米秸秆与大豆残渣混合青贮中,试验组Shannon指数提高,表明微生物菌群多样性和丰富度提高;何家俊等[51]试验结果显示,与对照组相比,试验组Shannon指数提高,这均与本研究结果一致。PCoA结果显示,各组间微生物群落结构呈现明显分离,说明各处理形成了具有差异的微生物群落组成。进一步分析表明,在门水平上,厚壁菌门为各组的优势菌门,这与前人研究结果[52]一致。厚壁菌门(以革兰氏阳性菌为主)具有降解纤维素等结构物质的能力[53],并能在发酵后期推动乳酸、乙酸等有机酸的合成[54];而变形菌门中部分类群可能包含潜在致病菌,其与乳酸菌竞争底物,通常对发酵品质产生不利影响[55]。在属水平上,各组主要由乳植杆菌属和慢生乳杆菌属等组成,与前人研究结果[56]相似。综合LEfSE分析结果,LB组与PP组微生物群落存在差异,PP组片球菌属显著富集,这正是戊糖片球菌的添加所致。Zhang等[57]研究发现,接种植物乳杆菌和布氏乳杆菌可以有效改善青贮品质;Xiao等[58]研究表明,植物乳杆菌和布氏乳杆菌可显著提高青贮饲料乳酸含量,二者均可改善青贮饲料品质,这与本研究结果相一致。此外,PP组在DM、CP、WSC和乳酸含量等多项指标上均优于LB组,表明不同乳酸菌在促进发酵和保存养分方面存在功能差异。

4 结论

添加布氏乳杆菌和戊糖片球菌均能优化全株玉米与向日葵混合青贮微生物群落结构,从而提高其发酵品质,在本试验条件下,添加戊糖片球菌青贮效果较好。

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