Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 6: 4651 - 4662

DOI: https://doi.org/10.12418/CJAN2026.372

RESEARCH PAPER

Effects of Different Mixed Silage Ratios and Additives on Fermentation Quality of Alfalfa and Sunflower Mixed Silage

  • YAN Yuting , 1 ,
  • ZHAO Muqier 1 ,
  • BAO Jian 2 ,
  • ZHU La 1 ,
  • WANG Wei 1 ,
  • GE Gentu , 1, * ,
  • JIA Yushan 1 ,
  • WANG Zhijun 1
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  • 1 Key Laboratory of Grassland Resources of the Ministry of Education, Key Laboratory of Forage Cultivation and the Processing and Highly Efficient Utilization of the Ministry of Agriculture and Rural Affairs, College of Grassland Science, Inner Mongolia Agricultural University, Hohhot 010019, China
  • 2 Inner Mongolia Academy of Agricultural and Animal Husbandry Sciences, Hohhot 010031, China
* professor, E-mail:

Received date: 2025-11-13

  Online published: 2026-06-13

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Abstract

This experiment was conducted to study the effects of different mixed silage ratios and additives on the fermentation quality of alfalfa and sunflower mixed silage. Using a two-factor experimental design, three mass ratios of alfalfa and sunflower were set at 3∶7, 5∶5 and 7∶3, respectively, and four treatments were set for each ratio, namely the Lactobacillus plantarum treatment (LP treatment), the Lactobacillus buchneri treatment (LB treatment), the sucrose treatment (SUG treatment) and the treatment without additives (CK treatment, control), with 3 replicates in each treatment. Silage was carried out at room temperature for 60 days to determine the nutrient contents, fermentation quality and microbiota. The results showed as follows: 1) under each mixed silage ratio, the contents of dry matter and water-soluble carbohydrates in SUG treatment were higher, and the ether extract content in LP treatment was higher. Compared with the other mixed silage ratios, when the mixed silage ratio was 3∶7, the crude protein content was lower and the neutral detergent fiber content was higher. 2) Under each mixed silage ratio, LP treatment had a lower pH, a higher lactic acid content, and a lower ammonia nitrogen (NH3-N) content. When the mixed silage ratio was 7∶3, the pH in each treatment was higher than that of the other mixed silage ratios. Meanwhile, when the mixed silage ratio was 5∶5, the lactic acid content was higher and the NH3-N content was lower. In addition, when the mixed silage ratio was 5∶5, the acetic acid content in LP treatment was significantly higher than that in the other treatments (P<0.05), and also significantly higher than that in the other mixed silage ratios (P<0.05); the propionic acid content in SUG treatment was significantly higher than that in other treatments (P<0.05). 3) The results of microbiota analysis indicated that at a 5∶5 mixed silage ratio, Lactiplantibacillus was enriched in LP and SUG treatments, Lentilactobacillus was enriched in LB treatment, and Lactococcus, Weissella and Enterobacter were enriched in CK treatment. In conclusion, when the mixed silage ratio of alfalfa and sunflower is 5∶5, the fermentation effect is better. Adding Lactobacillus plantarum can improve the fermentation quality, while adding sucrose is beneficial for nutrient preservation.

Key words： alfalfa; sunflower; mixed silage; fermentation quality; microbiota

Cite this article

YAN Yuting , ZHAO Muqier , BAO Jian , ZHU La , WANG Wei , GE Gentu , JIA Yushan , WANG Zhijun . Effects of Different Mixed Silage Ratios and Additives on Fermentation Quality of Alfalfa and Sunflower Mixed Silage[J]. Chinese Journal of Animal Nutrition, 2026 , 38(6) : 4651 -4662 . DOI: 10.12418/CJAN2026.372

近年来,全球变暖加剧导致气温持续攀升,极端天气事件频发[1]。气候变异性引发的降水模式变化导致多个地区出现前所未有的降雨异常现象[2],这些气候扰动加剧了人们对农业不稳定性的担忧,因为不可预测的天气条件严重威胁着农作物产量。以内蒙古为例,2024年平均降水量达到458.8 mm,比历史平均水平高出41.6%,且伴随强风暴和空间分布不均的现象[3-4]。这种异常降水对农作物和牧草造成双重打击,表现为生长迟缓、普遍倒伏以及腐烂风险增加,从而加剧了该地区的农业困境。正是在这种因气候异常导致原料质量不稳定的背景下,探索并优化青贮调制技术就显得尤为迫切。
常规青贮技术对原料固有特性有较高依赖性,且在发酵过程、营养保存和操作管理方面存在多种导致养分损失和品质下降的风险[5-6]。混合青贮作为一种优化策略,通过整合不同青贮原料的理化特性优势,采用科学配比将2种或多种原料进行混合青贮,可显著提升青贮饲料的发酵品质和营养价值[7]。该方法不仅具有成本效益优势,还能在青绿饲料季节性供应不足时有效拓展饲料资源。例如:碳水化合物含量较低的象草单独青贮很难达到优质青贮水平,而将象草与玉米秸秆进行混合青贮后发现饲料的可溶性糖含量显著增加,缓冲能值显著降低,青贮品质得到有效提升[8];张利媛等[9]将紫花苜蓿与小麦秸秆进行混合青贮,显著提高了青贮饲料的品质。
由于苜蓿本身具有高缓冲能力且水溶性碳水化合物(WSC)含量较低[10],其青贮加工面临巨大挑战,通常需要添加外源添加剂才能达到最佳发酵品质。相比之下,菊科一年生植物向日葵因能量密度和粗蛋白质(CP)含量相对较高[11]而展现出较高的饲用价值,但其在青贮生产中的实际应用受限于干物质(DM)含量低和细胞壁比例偏高等因素,这些因素可能影响发酵效率[12]。基于苜蓿和向日葵的特点及优势,本试验通过将向日葵与优质牧草苜蓿按不同比例、不同添加剂处理进行混合青贮,以期通过平衡基质组成来提升这2种饲草资源的青贮效果,为缓解草食动物饲料季节性供应不足的问题提供参考。

1 材料与方法

1.1 试验地点

本试验地点位于内蒙古自治区呼和浩特市土默特左旗海流图村内蒙古农业大学科技园区内(东经111°23'46″,北纬40°31'17″,海拔1 018 m),该地属温带大陆性季风气候,年日照时数达2 876 h,无霜期为130 d,年平均气温为6.3 ℃,年平均降水量为400 mm。

1.2 试验材料

青贮原料:本试验于2024年9月1日分别在苜蓿开花期和向日葵灌浆期对作物进行收割,将刈割后的青贮饲料原料经自然晾晒至含水量为70%左右,使用铡草机进行切割处理(长度2~3 cm)。青贮原料主要营养成分含量见表1。
表1 青贮原料主要营养成分含量

Table 1 Main nutrient contents of silage raw materials

项目
Items		 干物质
DM/% FW		 粗蛋白质
CP/% DM		 粗脂肪
EE/% DM		 水溶性碳水化合物
WSC/% DM		 中性洗涤纤维
NDF/% DM		 酸性洗涤纤维
ADF/% DM
苜蓿 Alfalfa 36.12±0.84 17.79±0.93 3.19±0.08 3.54±0.36 43.23±1.67 38.88±2.31
向日葵 Sunflower 32.71±1.12 12.76±0.76 9.23±0.69 2.47±0.44 45.11±0.92 40.02±2.77
青贮添加剂:青贮添加剂中的乳酸菌制剂为市购产品,其中植物乳杆菌活菌数为2×1011 CFU/g,布氏乳杆菌活菌数为2×1011 CFU/g;蔗糖为市购产品,其中蔗糖含量为500 mg/g。

1.3 试验设计

试验采用双因素设计,将苜蓿与向日葵分别按质量比为3∶7、5∶5和7∶3设置3种调制比例,每种比例均设置4个处理,各处理分别为添加植物乳杆菌处理(LP处理)、添加布氏乳杆菌处理(LB处理)、添加蔗糖处理(SUG处理)和无添加剂处理(CK处理,对照),每个处理3个重复。将添加剂(乳酸菌0.001 g,蔗糖4 g)溶解于20 mL蒸馏水中,并均匀喷洒在600 g按既定质量比例的苜蓿与向日葵混合样品上,然后装入30 cm×40 cm聚乙烯袋中,每袋填装200 g,采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、发酵品质以及微生物群落等指标。

1.4 测定指标及方法

1.4.1 营养成分含量

参照GB/T 6435—2014中方法测定DM含量;参照GB/T 6432—2018中方法测定CP含量;采用Ankom 2000型纤维分析仪(美国)按照所附说明书测定中性洗涤纤维(NDF)和酸性洗涤纤维(ADF)含量;采用VELP SER148/6脂肪测定仪(意大利)按照所附说明书测定粗脂肪(EE)含量;采用蒽酮-硫酸比色法[13]测定WSC含量。

1.4.2 发酵品质

混贮开袋后取10 g样品,加入90 mL蒸馏水,采用均质拍打仪匀速拍打2 min,过滤得到青贮饲料浸提液。采用酸度计测量pH;采用Agilent 1100高效液相色谱仪(美国)测定乳酸(lactic acid,LA)、乙酸(acetic acid,AA)、丙酸(propionic acid,PA)和丁酸(butyric acid,BA)含量;采用苯酚-次氯酸钠比色法[14]测定氨态氮(ammonia nitrogen,NH3-N)含量。

1.4.3 微生物群落

对筛选出的青贮营养成分含量及发酵品质最优的混贮比例(5∶5)组合进行微生物群落检测。首先提取微生物总DNA,然后采用引物338F(5'-ACTCCTACGGGGAGGCAGCAG-3')和806R(5'-GGACTACHVGGGTWTCTAAT-3')对16S rRNA基因V3~V4可变区进行扩增;PCR产物采用AxyPrep DNA凝胶提取试剂盒进行纯化,经Tris-HCl洗脱液洗脱后,采用2%琼脂糖凝胶电泳进行检测;采用QuantiFluorTM-ST荧光计用于定量检测,经纯化的扩增片段按Illumina MiSeq平台标准操作规程构建PE 2×300文库;采用Illumina MiSeq PE300平台(上海美吉生物医药科技有限公司)进行测序。

1.5 数据统计分析

采用Excel 2021对试验数据进行预处理,然后采用SAS 9.2软件对数据进行双因素方差分析(two-way ANOVA),并采用Duncan氏法进行多重比较检验,结果采用“平均值±标准差”形式表示,P<0.05表示差异显著;采用Origin 2024软件进行绘图。

2 结果与分析

2.1 不同添加剂对苜蓿与向日葵混贮营养成分含量的影响

由表2可知,添加剂处理对苜蓿与向日葵混贮DM、CP、EE、WSC和ADF含量有显著影响(P<0.05),混贮比例对CP、EE、WSC、NDF和ADF含量有显著影响(P<0.05),添加剂处理与混贮比例对CP、EE、WSC和ADF含量有显著交互作用(P<0.05)。在各混贮比例下,SUG处理DM和WSC含量均较高,LB处理DM含量较低,CK处理WSC含量较低。与其他混贮比例相比,当混贮比例为3∶7时,苜蓿与向日葵混贮CP含量较低,NDF含量较高。当混贮比例为3∶7和7∶3时,SUG处理CP含量较高;但当混贮比例为5∶5时,SUG处理CP含量显著低于其他处理(P<0.05)。在各混贮比例下,LP处理EE含量较高。
表2 不同添加剂对苜蓿与向日葵混贮营养成分含量的影响

Table 2 Effects of different additives on nutrient contents of alfalfa and sunflower mixed silage

混贮比例
(苜蓿∶向日葵)
Mixed silage ratios
(alfalfa∶sunflower)		 处理
Treatments		 干物质
DM/% FW		 粗蛋白质
CP/% DM		 粗脂肪
EE/% DM		 水溶性碳水
化合物
WSC/% DM		 中性洗
涤纤维
NDF/% DM		 酸性洗
涤纤维
ADF/% DM



3∶7		 LP 32.07±1.38AB 12.43±0.68Bc 4.30±0.16Ac 2.96±0.04Ba 36.49±3.64a 34.68±1.16ABa
LB 30.57±0.43B 13.21±0.95Bb 4.29±0.08Aa 1.58±0.06Cb 44.94±4.05a 38.01±0.52Aa
SUG 34.22±0.55A 17.15±0.71Ab 4.26±0.06Ab 4.06±0.04Ab 30.60±2.60a 29.08±0.67Cab
CK 31.77±0.49AB 13.17±0.90Bb 3.81±0.15Bb 1.23±0.01Db 37.41±0.70a 32.05±2.17BCa



5∶5		 LP 31.61±0.71A 19.03±0.42Aa 4.93±0.06Ab 1.71±0.19Bb 37.51±2.06a 33.13±1.20Aa
LB 30.74±0.59A 16.86±0.39Aa 4.56±0.13Ba 1.77±0.02Ba 33.73±1.62b 28.69±2.79ABa
SUG 33.93±1.04A 10.69±1.15Bc 5.14±0.11Aa 2.63±0.02Ac 33.92±3.24a 26.47±1.37Bb
CK 33.09±1.33A 18.61±0.69Aa 4.45±0.10Ba 1.17±0.04Cb 33.55±1.82a 29.14±0.65ABa



7∶3		 LP 32.18±0.45B 13.33±0.68Cc 6.17±0.01Aa 1.60±0.05Bb 36.08±2.12a 31.54±0.09Aa
LB 30.92±0.21B 16.19±1.03Ba 4.59±0.21Ba 1.59±0.03Bb 36.62±2.14ab 27.72±0.62Ba
SUG 34.65±0.44A 20.40±0.88Aa 2.84±0.03Cc 5.46±0.12Aa 30.45±2.85a 32.02±0.84Aa
CK 31.61±0.45B 18.82±0.87ABa 4.85±0.24Ba 1.50±0.06Ba 34.10±4.07a 32.58±0.91Aa
P值 P-value
处理 Treatment <0.001 <0.001 <0.001 <0.001 0.242 0.015
混贮比例 Mixed silage ratio 0.933 0.042 <0.001 <0.001 0.035 0.001
处理×混贮比例
Treatment×mixed silage ratio		 0.832 <0.001 <0.001 <0.001 0.245 0.002

LP:植物乳杆菌;LB:布氏乳杆菌;SUG:蔗糖;CK:无添加剂。同列数据肩标不同大写字母表示同一混贮比例不同处理间差异显著(P<0.05),不同小写字母表示同一处理不同混贮比例间差异显著(P<0.05),相同字母或无字母表示差异不显著(P>0.05)。表3同。

LP: Lactobacillus plantarum; LB: Lactobacillus buchneri; SUG: sucrose; CK: no additive. In the same column, values with different capital letter superscripts indicated significant differences among different treatments for the same mixed silage ratio (P<0.05), and with different small letter superscripts indicated significant differences among different mixed silage ratios for the same treatment (P<0.05), while with the same letter or no letter superscripts indicated no significant differences (P>0.05). The same as Table 3.

2.2 不同添加剂对苜蓿与向日葵混贮发酵品质的影响

由表3可知,添加剂处理对苜蓿与向日葵混贮pH以及乳酸、乙酸、丙酸和NH3-N含量有显著影响(P<0.05),混贮比例对pH以及乳酸、乙酸和NH3-N含量有显著影响(P<0.05),添加剂处理与混贮比例对pH以及乳酸、乙酸、丙酸和NH3-N含量有显著交互作用(P<0.05)。在各混贮比例下,LP处理pH均较低;同时,当混贮比例为7∶3时,各处理pH均高于其他混贮比例。在各混贮比例下,LP处理乳酸含量均较高,CK处理乳酸含量较低;同时,当混贮比例为5∶5时,乳酸含量较高。当混贮比例为5∶5时,LP处理乙酸含量显著高于其他处理(P<0.05),同时也显著高于其他混贮比例(P<0.05);SUG处理丙酸含量显著高于其他处理(P<0.05)。在各混贮比例下,LP处理NH3-N含量较低,SUG处理NH3-N含量较高;同时,当混贮比例为5∶5时,NH3-N含量较低。
表3 不同添加剂对苜蓿与向日葵混贮发酵品质的影响

Table 3 Effects of different additives on fermentation quality of alfalfa and sunflower mixed silage

混贮比例
(苜蓿∶向日葵)
Mixed silage ratios
(alfalfa∶sunflower)		 处理
Treatments		 pH 乳酸
LA/(g/kg
DM)		 乙酸
AA/(g/kg
DM)		 丙酸
PA/(g/kg
DM)		 丁酸
BA/(g/kg
DM)		 氨态氮
NH3-N/%
FW



3∶7		 LP 4.03±0.04Cc 4.12±0.23Ab 0.70±0.03BCc 0.28±0.28A 0.00±0.00 0.20±0.00Ca
LB 4.23±0.02Bb 3.11±0.14Ba 0.50±0.02Cb 0.29±0.29A 0.00±0.00 0.21±0.00Bb
SUG 4.31±0.02Bb 2.99±0.09Ba 1.15±0.12Aa 0.00±0.00A 0.01±0.01 0.27±0.00Ab
CK 4.47±0.03Ab 2.36±0.07Cc 0.86±0.06Ba 0.00±0.00A 0.01±0.01 0.20±0.00Cc



5∶5		 LP 4.31±0.01Bb 6.97±0.07Aa 1.44±0.06Aa 0.25±0.00D 0.00±0.00 0.19±0.00Dc
LB 4.37±0.07Bb 2.97±0.05Ca 0.90±0.01Ba 0.35±0.00C 0.00±0.00 0.21±0.00Cb
SUG 4.39±0.02Ba 3.11±0.36Ca 0.77±0.05BCb 1.05±0.04A 0.00±0.00 0.24±0.00Bc
CK 4.97±0.03Ab 4.77±0.17Ba 0.67±0.03Cb 0.80±0.00B 0.00±0.00 0.25±0.00Aa



7∶3		 LP 4.41±0.02Ca 3.55±0.10Ac 0.94±0.06Ab 0.28±0.28A 0.00±0.00 0.20±0.00Db
LB 4.63±0.04Ba 3.21±0.06Aa 0.92±0.02Aa 0.00±0.00A 0.00±0.00 0.22±0.00Ba
SUG 4.61±0.02Ba 3.54±0.13Aa 0.85±0.00Ab 0.00±0.00A 0.00±0.00 0.30±0.00Aa
CK 5.23±0.05Aa 2.81±0.10Bb 0.64±0.01Bb 0.30±0.30A 0.01±0.01 0.21±0.00Cb
P值 P-value
处理 Treatment <0.001 <0.001 0.001 <0.001 0.479 <0.001
混贮比例 Mixed silage ratio <0.001 <0.001 <0.001 0.545 0.227 <0.001
处理×混贮比例
Treatment×mixed silage ratio		 <0.001 <0.001 <0.001 0.005 0.822 <0.001

2.3 不同添加剂对苜蓿与向日葵混贮微生物群落的影响

综合苜蓿与向日葵混贮营养成分含量和发酵品质,选择发酵效果较好的混贮比例(苜蓿∶向日葵为5∶5)组合进行微生物群落分析。如图1所示,4个处理共检测到112个操作分类单元(OTU),其中各处理共有23个OTU,占OTU总数的20.54%;LB处理独有OTU最多,为30个,占OTU总数的26.79%;SUG处理独有OTU最少,为8个,占OTU总数的7.14%。
图1 苜蓿与向日葵混贮微生物群落操作分类单元韦恩图

LP:植物乳杆菌;LB:布氏乳杆菌;SUG:蔗糖;CK:无添加剂。图2同。

Fig.1 Venn diagram of OTU in microbiota of alfalfa and sunflower mixed silage

LP: Lactobacillus plantarum; LB: Lactobacillus buchneri; SUG: sucrose; CK: no additive. The same as Fig.2.

Circos图通常用于展示微生物群落类群的组成和分布情况,如图2所示,在相对丰度排名前10的菌属中,乳植杆菌属(Lactiplantibacillus)是相对丰度最高的菌属,大量存在于LP处理(31%)和SUG处理(30%);其次为慢生乳杆菌属(Lentilactobacillus),其大量存在于LB处理(57%),而在CK组中几乎不存在;乳球菌属(Lactococcus)绝大部分存在于CK处理(98%),在LP组中不存在;魏斯氏菌属(Weissella)大量存在于CK处理(53%)和SUG处理(40%)中,仅少量存在于LB处理(5%)和LP处理(2%);肠杆菌属(Enterobacter)大部分存在于CK处理(68%),部分存在于SUG处理(30%),极少部分存在于LB处理(2%),而LP处理不存在;肠球菌属(Enterococcus)和促生乳杆菌属(Levilactobacillus)也大量存在于CK处理,与肠杆菌属不同的是它们在其余3个处理中均少量存在。
图2 苜蓿与向日葵混贮微生物群落属水平Circos图

Lactiplantibacillus:乳植杆菌属;Lentilactobacillus:慢生乳杆菌属;Lactococcus:乳球菌属;Unclassified_f_Enterobacteriaceae:未分类肠杆菌科;Weissella:魏斯氏菌属;Enterobacter:肠杆菌属;Enterococcus:肠球菌属;Levilactobacillus:促生乳杆菌属;Unclassified_o_Enterobacterales:未分类肠杆菌目;Pantoea:泛菌属;Others:其他。图3同 the same as Fig.3。

Fig.2 Circos plot of microbiota at genus level of alfalfa and sunflower mixed silage

本试验进一步将苜蓿与向日葵混贮微生物群落在属水平上相对丰度与营养成分含量和发酵品质进行相关性分析,如图3所示,WSC和EE含量均与肠球菌属和乳球菌属相对丰度呈极显著负相关(P<0.01),DM和NH3-N含量以及pH与慢生乳杆菌属相对丰度呈显著负相关(P<0.05),pH以及丙酸和NH3-N含量与魏斯氏菌属和肠杆菌属相对丰度呈显著或极显著正相关(P<0.05或P<0.01),乙酸含量与魏斯氏菌属和肠杆菌属相对丰度呈极显著负相关(P<0.01),乳酸和CP含量与促生乳杆菌属相对丰度呈显著或极显著正相关(P<0.05或P<0.01)。
图3 苜蓿与向日葵混贮微生物群落(属水平)与营养成分含量和发酵品质相关性热图

*表示显著相关(P<0.05),**(P<0.01)和***(P<0.001)表示极显著相关。

Fig.3 Heatmap of correlation between microbiota (at genus level) and nutrient contents and fermentation quality of alfalfa and sunflower mixed silage

* indicated significant correlation (P<0.05), and ** (P<0.01) and *** (P<0.001) indicated extremely significant correlation.

3 讨论

3.1 不同添加剂对苜蓿与向日葵混贮营养成分含量的影响

评价饲料品质优劣最直观的指标就是营养成分含量。本试验中,苜蓿与向日葵混贮后DM含量较原料并无大幅度下降,这是由于向日葵秸秆的DM含量通常较高,而苜蓿的含水量较高,两者混合后可以平衡整体水分含量,减少发酵过程中的水分流失或渗出,从而维持DM含量的稳定性[15]。随着混贮中向日葵比例的提高,CP含量相应降低,这是因为全株向日葵(含花盘、籽壳等)CP含量较苜蓿低,其比例提高影响了混合体系的总氮含量[16]。在各混贮比例下,SUG处理WSC含量较高,其原因为蔗糖既直接提供了外源可溶性糖,又通过“先供糖-后促酸-再水解”的三步链式反应,使青贮体系的可溶性碳水化合物总量得到提升[17-18]。LP处理EE含量较高,其可能的原因包括:植物乳杆菌在发酵早期快速产酸降低了pH,有效抑制了脂肪酶活性及好氧腐败菌(如酵母、霉菌)生长,从而减缓了脂肪的分解和氧化[19];同时,植物乳杆菌在生长过程中可能合成部分胞内脂质,直接提高了EE含量[20]。由NDF和ADF含量可以看出,添加剂处理和混贮比例的改变对其的影响呈“非线性”规律,这可能与多因素交互作用有关,过多的变量导致其变化趋势混乱[21],但青贮后NDF和ADF含量较原料有一定程度的降低。

3.2 不同添加剂对苜蓿与向日葵混贮发酵品质的影响

一般情况下,pH可以反映青贮饲料品质的好坏,当pH低于4.2时通常代表青贮发酵较为成功,而当pH高于5.0时表明发酵失败而产生了较劣质的饲料[16,22]。本试验中,混贮pH随向日葵添加比例的升高而降低,这归因于向日葵秸秆具有比苜蓿更低的缓冲能力,使得等量乳酸能更有效地降低体系pH[11]。不同添加剂下乳酸菌处理较蔗糖处理的混贮pH更低,原因是蔗糖必须先被乳酸菌分泌的转化酶水解为葡萄糖和果糖才能被利用,这一过程延缓了乳酸的启动和积累,导致pH下降速度慢于可直接利用单糖的乳酸菌制剂[23]。在5∶5混贮比例下,乳酸含量较高,这可能与底物和微生物的“双平衡”效应有关,苜蓿蛋白质含量高、缓冲能高,但WSC含量低;而向日葵秸秆WSC含量相对高、缓冲能低,当二者以5∶5比例进行混合时,WSC足以支持乳酸菌的快速增殖,而缓冲能被“稀释”到乳酸菌可快速克服的阈值,从而有利于乳酸积累[15]。植物乳杆菌属于同型发酵乳酸菌,理论上每1 mol葡萄糖可生成2 mol乳酸,碳转化效率>90%[24],因此LP处理乳酸含量高于其他处理。青贮中乙酸主要来源于异型发酵乳酸菌(如布氏乳杆菌)的代谢。当混贮比例改变时,原料WSC含量与缓冲能比值随之变化,直接影响异型乳酸菌的活性[25]。混贮比例本身对乙酸含量的影响无固定方向,乙酸含量主要取决于糖源/缓冲能的实时阈值变化。丁酸作为青贮发酵中有害菌分解转化青贮饲料中乙酸和糖的产物,在本研究中仅少量存在于个别处理中,表明发酵较为成功[26-28]。此外,SUG处理NH3-N含量较高,这是因为当原料中乳酸菌数量不足或pH下降缓慢时,蔗糖可被肠杆菌、芽孢杆菌和丁酸菌等杂菌优先利用,这些菌在利用蔗糖产酸的同时,也分泌蛋白酶,将植物蛋白质分解为氨基酸,再进一步脱氨生成NH3-N,导致NH3-N含量升高[29-30]。综合来看,苜蓿与向日葵在5∶5比例下混贮品质更佳。

3.3 不同添加剂对苜蓿与向日葵混贮微生物群落的影响

布氏乳杆菌的异型发酵特性为混贮微生物群落多样性提供了物质基础。该菌通过异型发酵途径生成2类代谢物乙酸和1,2-丙二醇并分别构建“选择性营养筛选”环境,其中乙酸作为高亲和力碳源,为耐酸型醋酸杆菌属(Acetobacter)和兼性厌氧型芽孢杆菌属(Bacillus)提供专属碳源,而1,2-丙二醇则可被特定酵母[如假丝酵母属(Candida)]通过氧化还原途径利用,形成次级菌群的增殖优势[31];此外,乙酸介导的低pH环境有助于构建“生长抑制屏障”,对乳酸同化型酵母[如酵母属(Saccharomyces)]和腐败菌[如梭菌属(Clostridium)]的细胞膜完整性造成破坏,显著降低其比生长速率(μ<0.1 h-1),打破了单一乳酸菌的竞争优势[32],从而维持较高的Shannon和Simpson指数,从而使LB处理微生物群落在韦恩图中表现为更多独有OTU。SUG处理OTU数量较少的本质是“碳源利用单一化→优势菌群垄断→环境筛选单向化”的连锁效应,而植物乳杆菌和布氏乳杆菌的接种通过“代谢产物多样化→菌群功能互补→环境调控多元化”维持了更高的物种丰富度[33-34]。
Circos图能够直观呈现每个处理中优势类群的组成情况,并反向揭示同一类群在不同处理间的相对丰度差异。乳植杆菌属因其能高效利用多种糖源产酸,抑制杂菌增殖[35-36],成为绝对优势菌属,在添加了该菌剂的LP处理中相对丰度最高。慢生乳杆菌属主要富集于LB处理,这与其作为添加剂的来源一致,研究表明其接种后占据主导地位的时间可能较晚,但能在较长时间窗口内维持高相对丰度[37]。CK处理因缺乏外源乳酸菌,发酵初期pH下降缓慢,这为耐中性环境的乳球菌属、魏斯氏菌属和肠杆菌属提供了生长窗口,故这些菌主要存在于CK处理。值得注意的是,SUG处理也出现了较高相对丰度的肠杆菌属,这表明蔗糖在促进乳酸菌的同时,也可能直接支持了肠杆菌的生长[38-39]。可发酵碳水化合物添加剂(如蔗糖、糖蜜)的发酵效果高度依赖于原料本身的缓冲能力和“土著”微生物群落,如果原料缓冲能力高或有害菌基数大,单纯加糖可能无法确保乳酸菌快速取胜,反而可能在初期促进有害菌生长[40]。尽管肠球菌属和促生乳杆菌属均属乳酸菌成员,但其最适pH范围高于植物乳杆菌,各添加剂处理通过快速酸化与资源竞争成功抑制了它们的生长和存活,而CK处理因无快速酸化压力和底物竞争,两者得以大量定殖[41]。
当青贮体系WSC充足时,同型发酵乳酸菌(如植物乳杆菌)迅速增殖,乳酸积累使pH下降,而魏斯氏菌属适宜pH为5.5~6.0,缺乏F0F1-ATP酶(F0F1-ATPase)高表达耐酸系统,当乳酸含量≥ 90 g/kg FM时胞内氢离子(H+)无法外排,导致酸致死;肠球菌属虽具有耐酸基因(aga、atpC),但当pH<4.0时能量收益骤降,逐渐被同型乳酸菌群取代,其相对丰度随WSC含量降低及乳酸含量升高而显著下降[42-43]。EE含量升高往往伴随发酵产酸旺盛阶段,高酸环境会抑制脂肪酶活性,减少脂肪水解,同时也会抑制依赖中性pH的魏斯氏菌属和肠球菌属的脂肪分解代谢[44],因此WSC和EE含量均与肠球菌属相对丰度呈极显著负相关。在青贮发酵过程中,pH越低(酸化越快),乳植杆菌属和慢生乳杆菌属越占优势,这2种乳酸菌均具备出色的耐酸能力,在pH降至4.0以下的环境中仍能保持较高活性;同时,其他杂菌如肠杆菌、霉菌等在此酸度下生长受到抑制,使得乳酸菌能持续利用可溶性碳水化合物繁殖,进一步扩大菌群占比,形成“产酸-降pH-更占优势”的正向循环。在杂交构树青贮的研究中发现,接种植物乳杆菌后,随着pH的下降乳杆菌属成为绝对优势菌群[45]。相反,pH越高(酸化滞后),魏斯氏菌属和肠杆菌属等越繁盛,因此其相对丰度与pH呈显著或极显著正相关[11,46]。乳植杆菌属可将可溶性碳水化合物高效转化为乳酸和乙酸,因此其相对丰度与两者含量呈相关,这在苜蓿与黄花刺茄混贮及茅台酒糟发酵的研究中均有体现[44,47]。促生乳杆菌属能以高底物亲和力利用WSC,通过糖酵解途径高效合成乳酸,产酸速率可达0.2~0.5 g/(kg·h)(DM基础),这一方面抑制梭菌、肠杆菌等产氨和蛋白质分解菌的生长,减少CP向NH3-N转化;另一方面降低植物内源蛋白酶(如羧肽酶、肽酶)活性,减少CP水解为游离氨基酸的损失,使CP保存率提升3%~8%,因此其相对丰度与乳酸和CP含量呈显著或极显著正相关[48-49]。当乙酸含量升高(>1.5% DM),pH迅速降至<4.2,对耐中性pH的魏斯氏菌属和肠杆菌属形成化学胁迫,两者适宜pH为5.5~6.5,在酸性环境中生长速率显著下降,导致其相对丰度与乙酸含量呈极显著负相关[50]。丙酸本身并非优势酸,但其在腐败菌群(如梭菌属、肠杆菌属)活跃时会大量积累,同时腐败菌群分泌的金属蛋白酶可高效降解蛋白质,且脱氨基作用增强,推动丙酸和NH3-N协同累积,形成“菌群增殖-产酸-蛋白质降解-脱氨基”的正反馈循环[51]。因此,丙酸和NH3-N含量同步升高,且与肠杆菌属相对丰度呈显著正相关,表明腐败发酵占主导地位[52]。

4 结论

本试验表明,混贮比例和添加剂对苜蓿与向日葵混贮发酵品质有显著影响,综合来看,苜蓿∶向日葵比例为5∶5时发酵效果较好。在添加剂选择上,植物乳杆菌在改善发酵品质方面效果较为突出,能快速建立酸性环境,促进有益乳酸菌成为优势菌群,并有效抑制肠杆菌等有害微生物的增殖,从而提高发酵品质;蔗糖则表现出更好的营养保存效果。因此,为提高苜蓿与向日葵混贮饲料的综合品质,建议采用5∶5的混合比例,并优先选用植物乳杆菌和蔗糖作为添加剂。

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