饲草附生乳酸菌对碳源的选择性

陈鑫珠; 董朝霞; 张建国

饲草附生乳酸菌对碳源的选择性

陈鑫珠^{1, 2,},
董朝霞¹,
张建国^1, ,

1.
华南农业大学南方草业研究中心，广东　广州　510642
2.
福建省农业科学院畜牧兽医研究所/福建省畜禽遗传育种重点实验室，福建　福州　350013

基金项目: 国家自然科学基金项目（31971764、32302791）；福建省农业高质量发展超越“5511”协同创新工程项目（XTCXGC2021019)；福建省科技计划公益类专项（2020R1021002）

详细信息

作者简介:
陈鑫珠（1985 —），女，博士，副研究员，主要从事动物营养、非常规饲料资源的开发利用和饲草加工与贮藏研究， E-mail：010622051@163.com

通讯作者:
张建国（1968 —），男，博士生导师，教授，主要从事饲草料加工利用、生物活性成分研究，E-mail：zhangjg@scau.edu.cn

中图分类号: S816.53
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出版历程
- 收稿日期: 2023-11-28
- 修回日期: 2024-02-18
- 网络出版日期: 2024-06-26

Silage Carbon Sources Preferred by Epiphytic Lactic Acid Bacteria

CHEN Xinzhu^{1, 2
,},
DONG Zhaoxia¹,
ZHANG Jianguo^{1
, ,}

1.
Department of Grassland Science, South China Agricultural University, Guangzhou, Guangdong　510642, China
2.
Institute of Animal Husbandry and Veterinary Medicine/Fujian Key Laboratory of Animal Genetics and Breeding, Fujian Academy of Agricultural Sciences, Fuzhou, Fujian　350013, China

摘要

摘要: 目的探究饲草源乳酸菌对碳源的选择性，为不同草种筛选适宜的青贮菌种。方法选取饲草中分离获得的16株代表性乳酸菌作为研究对象，在MRS液体培养基中以葡萄糖、蔗糖、果糖、木糖、棉籽糖以及5种糖混合为碳源，进行不同菌株发酵培养。每株乳酸菌设3个重复，每个糖处理设空白对照（不接菌）。采用高效液相色谱仪测定各处理发酵24 h后液体培养基中剩余碳源含量和乳酸产量。结果（1）蔗糖是大部分菌株发酵首选碳源，在棉籽糖、葡萄糖、果糖和木糖的选择上不同菌株差异较大，发酵乳杆菌（Lactobacillus fermentum，LbF-WM）首选棉籽糖和木糖，假肠膜明串珠菌（Leuconostioc pseudomesenteroides，LeP-IR）首选木糖。（2）罗伊氏乳杆菌（Lactobacillus reuteri，LbR-EG）和柠檬明串珠菌（Leuconostoc citreum，LeC-IR）对蔗糖，LbF-WM和LbR-EG对果糖， LbF-WM 、LbR-EG和LeP-IR对棉籽糖的利用率均达100%，屎肠球菌（Enterococcus faecium，EF-KG）、短乳杆菌（Lactobacillus brevis，LbB-KG）、戊糖乳杆菌（Lactobacillus pentosus，LbPe-EG）、乳酸乳球菌（Lactococcus lactis，LcL-SC）对葡萄糖的利用率高于其他碳源。（3）灰黄色肠球菌（Enterococcus durans，EG-IR）利用葡萄糖和混合糖，LbPe-EG利用果糖，肠膜明串珠菌（Leuconostoc mesenteroides，LeM-IR）利用棉籽糖和混合糖，LbPe-EG利用蔗糖，LbF-WM利用木糖，格氏乳球菌（Lactococcus garvieae，LcG-IR）利用混合糖发酵乳酸产量最高。（4）耐久肠球菌（Enterococcus durans，ED-IR）利用葡萄糖，植物乳杆菌（Lactobacillus plantarum，LbPl-IR）、LbPe-EG利用果糖， LbPl-IR和LeM-IR利用棉籽糖， LbPe-EG、LbPl-IR和ED-IR利用蔗糖， LeP- IR利用木糖，LcG-IR和EG-IR利用混合糖产乳酸效率较高。LbPl-IR在5种单糖中产酸效率均达到80%以上。结论不同菌株对同一碳源的利用率、产酸量和产酸效率均不同；同一株菌株对不同碳源的利用率、产酸量和产酸效率也不相同。其中，蔗糖为大部分菌株的首选碳源，蔗糖和葡萄糖的利用率和产乳酸量较高，葡萄糖的产酸效率较高；青贮调制时，可针对饲草中分布乳酸菌种类选择最适碳源。
- 饲草附生乳酸菌 /
- 碳源 /
- 蔗糖 /
- 葡萄糖 /
- 选择性
Abstract: Objective Preferential carbon sources of epiphytic lactic acid bacteria (LAB) were studied in laboratory to aid turning native grasses into silage. Method Sixteen representative LAB were isolated from forage and cultured on MRS broth media using glucose, sucrose, fructose, xylose, raffinose, or a mixture of the 5 sugars as carbon source. Three replicates for each LAB strain and a blank control without LAB inoculation for each sugar treatment were included. After incubation for 24h, contents of residual sugars and generated lactic acid in the medium were determined by high-performance liquid chromatography. Results (1) Sucrose appeared to be the preferential carbon source by most LAB. Among the other sugars, raffinose and xylose were preferred by Lactobacillus fermentum (LbF-WM), and xylose by Leuconostoc pseudomesenteroides (LeP-IR). (2) An 100% utilization on sucrose by Lb. reuteri (LbR-EG) and Leuconostoc citreum (LeC-IR), on fructose by LbF-WM and LbR-EG, as well as on raffinose by LbF-WM, LbR-EG, and LeP-IR was observed. Enterococcus faecium (EF-KG), Lb. brevis (LbB-KG), Lactobacillus pentosus (LbPe-EG), and Lactococcus lactis (LcL-SC) could more efficiently utilize glucose than the other sugars. (3) The greatest amounts of lactic acid were produced from the fermentation of glucose or the mixed sugars by E. durans (EG-IR), from that of fructose by LbPe-EG, from that of raffinose or the sugar mixture by Leuconostoc mesenteroides (LeM-IR), from that of sucrose by LbPe-EG, from that of xylose by LbF-WM, and from that of the mixed sugars by Lactococcus garvieae (LcG-IR). (4) The lactic acid production was more efficient by E. durans (ED-IR) on glucose, by Lb. plantarum (LbPl-IR), LeM-IR, or LbPe-EG on fructose, by Lb. plantarum (LbPl-IR) or LeM-IR on raffinose, by LbPe-EG, LbPl-IR, or ED-IR on sucrose, Le. pseudomesenteroides (LeP-IR) on xylose, and LcG-IR or EG-IR on mixed sugars. A lactic acid production efficiency greater than 80% was reached by LbPl-IR grown on the medium with the mixed sugars. Conclusion The sugar utilization and lactic acid production of the LAB fermentations on native grasses varied significantly. For efficient and effective application of LAB to obtain extended shelf-life silage, it was imperative that careful selection of microbe strains and carbon source be exercised.
- Epiphytic lactic acid bacteria /
- carbon source /
- glucose /
- sucrose /
- selectivity

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图 1 16株乳酸菌培养24 h后在5种糖混合培养基中各种糖的剩余含量

1）图中柱状数据表示混合糖培养基中剩余各糖的百分含量。2）CK，无接菌的混合糖培养基；

Figure 1. Contents of residual sugars in mixed sugar media of 16 different LAB cultured for 24 h

1) Data on a column are contents of residual individual sugars in medium of mixed sugars; 2) CK without inoculation on medium of mixed sugars; L.M.: Le. mesenteroides; L.P.: Le. pseudomesenteroides; W.P.: W. paramesenteroides. Same for below.

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表 1 16株乳酸菌的名称及来源

Table 1. Names and sources of 16 LAB strains

序号 No.	命名 Name	种名 Species	来源 Source	序号 No.	命名 Name	种名 Species	来源 Source
1	EF-KG	屎肠球菌 E. faecium	热研四号王草	9	LcG-IR	格氏乳球菌 Lc. garvieae	意大利黑麦草
2	ED-IR	耐久肠球菌 E. durans	意大利黑麦草	10	LcL-SC	乳酸乳球菌 Lc. lactis	甜玉米
3	EG-IR	灰黄色肠球菌 E. gilvus	意大利黑麦草	11	LeC-IR	柠檬明串珠菌 Le. citreum	意大利黑麦草
4	LbB-KG	短乳杆菌 Lb. brevis	热研四号王草	12	LeM-IR	肠膜明串珠菌 Le. mesenteroides	意大利黑麦草
5	LbF-WM	发酵乳杆菌 Lb. fermentum	糯玉米	13	LeP-IR	假肠膜明串珠菌 Le. pseudomesenteroides	意大利黑麦草
6	LbPe-EG	戊糖乳杆菌 Lb. pentosus	摩特矮象草	14	WCo-KG	融合乳杆菌 W. confusa	热研四号王草
7	LbPl-IR	植物乳杆菌 Lb. plantarum	意大利黑麦草	15	WCi-MT	魏斯特菌 W. cibaria	MT-1杂交象草
8	LbR-EG	罗伊氏乳杆菌 Lb. reuteri	摩特矮象草	16	WP-IR	类肠膜魏斯氏菌 W. paramesenteroides	意大利黑麦草

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表 2 牧草原料中糖分组成

Table 2. Sugars in native grass material (%)

项目 Item	葡萄糖 Glucose	蔗糖 Sucrose	果糖 Fructose	棉籽糖 Raffinose	木糖 Xylose
热研四号王草 Pennisetum purpureum × P. americanum cv. Reyan No.4	0.07±0.01c	1.91±0.37a	1.34±0.23b	1.86±0.06a	0.00±0.00c
MT-1象草 P. purpureum cv. MT-1	0.10±0.01c	1.82±0.08a	1.41±0.25b	1.28±0.27b	0.03±0.00c
摩特矮象草 P. purputeum cv. Mott	0.12±0.01c	1.75±0.29ab	2.11±0.13a	1.58±0.31b	0.09±0.09c
甜玉米 Zea mays L. saccharata Sturt.	1.84±0.26b	6.00±0.96a	1.12±0.11b	0.00±0.00c	0.00±0.00c
桂牧一号杂交象草 (P. americanum × P. Purpureum) × (P. Purpureum cv. Mott) Guimu No.1	0.03±0.02d	1.87±0.14a	0.21±0.07c	1.24±0.17b	0.00±0.00d
糯玉米 Zea mays L. Sinensisa Kulesh	1.81±0.36b	5.06±0.37a	0.93±0.02c	0.00±0.00d	0.00±0.00d
意大利黑麦草 Lolium multiflorum	0.70±0.39ab	0.94±0.36a	0.34±0.07b	0.08±0.08c	0.10±0.04c
同行数据后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。 Data with different lowercase letters on same row represent significant difference at P＜0.05; those with same or no letter, no significant difference at P＞0.05. Same for below.

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表 3 16株乳酸菌对6种糖源培养液发酵后的碳源利用率

Table 3. Sugar utilization by 16 different LAB in 6 treatments (单位：%)

菌株编号 Strain code	葡萄糖MRS培养基 MRS-P	果糖MRS培养基 MRS-G	棉籽糖MRS培养基 MRS-MZ	蔗糖MRS培养基 MRS-Z	木糖MRS培养基 MRS-M	混合糖MRS培养基 MRS- MIX
EF-KG	94.68±1.01a	29.25±3.43e	0.73±0.53f	89.99±1.72b	59.95±1.48c	50.27±0.54d
ED-IR	44.76±2.10b	60.48±1.52a	63.67±0.91a	69.87±1.16a	5.22±0.18c	39.83±2.34b
EG-IR	71.44±3.35ab	58.97±3.27bc	49.20±1.22cd	81.23±1.54a	0.00±0.00e	39.56±0.89d
LbB-KG	72.13±1.05a	18.10±0.53c	7.80±4.49c	50.41±0.10b	46.74±0.10b	38.82±2.37b
LbF-WM	87.50±1.43b	100.00±0.00a	100.00±0.00a	89.79±3.33b	79.15±0.84b	33.33±4.13c
LbPe-EG	82.97±0.73a	60.31±0.62b	37.42±3.02c	76.75±1.05ab	1.10±0.19d	38.56±2.88c
LbPl-IR	69.56±1.06a	58.45±1.72a	60.34±1.81a	72.80±2.56a	0c	39.57±2.86b
LbR-EG	82.07±2.63b	100.00±0.00a	100.00±0.00a	100.00±0.00a	0d	31.01±5.99c
LcG-IR	78.09±0.84ab	60.59±1.02b	74.69±0.71ab	81.48±1.64a	1.92±0.42d	38.11±3.78c
LcL-SC	93.58±1.01a	0e	18.52±0.52d	90.29±0.82a	37.25±0.12c	52.81±5.62b
LeC-IR	83.17±3.13b	79.15±0.61b	10.49±0.03d	100.00±0.00a	15.54±0.16d	44.51±10.98c
LeM-IR	73.89±1.05a	55.36±4.22b	70.94±0.01a	74.02±0.65a	0d	24.70±4.80c
LeP-IR	80.19±1.64b	90.59±0.10a	100.00±0.00a	52.23±6.10c	24.86±20.14c	33.81±2.39c
WCo-KG	76.66±1.64b	9.93±5.08e	17.11±1.42d	96.40±1.00a	48.27±0.10c	53.27±6.55c
WCi-MT	67.93±3.46a	56.89±0.82a	68.83±1.55a	71.00±0.86a	8.80±0.18b	27.70±7.52b
WP-IR	79.14±1.24a	73.84±0.01a	79.16±2.21a	79.09±1.13a	0c	41.12±7.76b

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表 4 16株乳酸菌在不同碳源培养基发酵中的乳酸产量

Table 4. Lactic acid production by 16 different LAB on different media (单位：%)

菌株名称 Name	葡萄糖MRS培养基 MRS-P	果糖MRS培养基 MRS-G	棉籽糖MRS培养基 MRS-MZ	蔗糖MRS培养基 MRS-Z	木糖MRS培养基 MRS-M	混合糖MRS培养基 MRS- MIX
EF-KG	0.69±0.01a	0.32±0.02b	0. 01±0.00c	0.62±0.01a	0.58±0.03a	0.55±0.02a
ED-IR	0.89±0.02a	0.85±0.05a	0.21±0.03b	1.14±0.01a	0.02±0.00c	0.86±0.06a
EG-IR	1.25±0.05a	0.73±0.05b	0.66±0.03b	0.87±0.00b	0.01±0.00c	1.00±0.03a
LbB-KG	0.71±0.00a	0.29±0.03c	0.23±0.00c	0.56±0.06b	0.63±0.04b	0.61±0.04b
LbF-WM	0.67±0.01b	0.36±0.04c	0.71±0.01b	0.73±0.00b	0.94±0.00a	0.67±0.05b
LbPe-EG	1.08±0.04b	1.10±0.06a	0.43±0.04c	1.31±0.00a	0.01±0.00c	0.98±0.02b
LbPl-IR	1.14±0.01a	1.07±0.05a	1.01±0.02a	1.23±0.03a	0.61±0.02b	0.77±0.00b
LbR-EG	0.73±0.02ab	0.69±0.04ab	0.86±0.02a	0.55±0.03b	0.63±0.04ab	0.67±0.00 ab
LcG-IR	1.18±0.04a	0.88±0.00b	1.02±0.04a	1.15±0.01b a	0.31±0.00c	1.08±0.05a
LcL-SC	0.69±0.02a	0.45±0.06b	0.15±0.00c	0.52±0.05ab	0.50±0.02ab	0.58±0.00a
LeC-IR	0.71±0.02a	0.67±0.02a	0.14±0.05c	0.42±0.02b	0.15±0.00c	0.45±0.01b
LeM-IR	1.13±0.04a	1.01±0.05a	1.16±0.00a	1.13±0.00a	0.71±0.01b	1.06±0.02a
LeP-IR	0.79±0.00a	0.54±0.02c	0.76±0.02a	0.65±0.01bc	0.78±0.00a	0.61±0.01bc
WCo-KG	0.60±0.02a	0.43±0.00b	0.13±0.00c	0.62±0.06a	0.55±0.02a	0.50±0.02ab
WCi-MT	1.11±0.05a	0.54±0.00b	0.36±0.02c	0.67±0.05b	0.10±0.01d	0.69±0.02b
WP-IR	0.58±0.00a	0.52±0.00a	0.08±0.01c	0.32±0.00b	0.64±0.02a	0.31±0.00b

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表 5 16株乳酸菌的产酸效率

Table 5. Acid production efficiency of 16 different LAB (单位：%)

菌株名称 Name	葡萄糖MRS培养基 MRS-P	果糖MRS培养基 MRS-G	棉籽糖MRS培养基 MRS-MZ	蔗糖MRS培养基 MRS-Z	木糖MRS培养基 MRS-M	混合糖MRS培养基 MRS- MIX
EF-KG	36.44±0.01c	54.69±0.06a	34.26±1.17d	34.45±0.01d	48.38±0.06b	54.71±0.59a
ED-IR	99.41±0.22a	70.27±0.02d	16.49±0.00f	81.58±0.07b	19.16±0.66e	71.47±0.00c
EG-IR	87.49±0.06a	61.90±0.02d	67.07±0.02c	53.55±0.03e	50.00±0.00f	82.72±0.00b
LbB-KG	49.21±0.04b	80.12±0.15ab	81.69±0.20a	55.55±0.11b	67.40±0.15ab	49.85±0.00b
LbF-WM	38.29±0.01d	18.00±0.00f	35.50±0.00e	40.65±0.01c	59.38±0.03a	49.12±0.00b
LbPe-EG	65.08±0.02d	91.20±0.03a	57.46±0.03e	85.34±0.06b	23.10±3.98f	79.76±0.00c
LbPl-IR	81.94±0.06d	91.53±0.03a	83.69±0.01c	84.70±0.45b	81.33±0.00e	63.71±0.00f
LbR-EG	44.48±0.02c	34.50±0.00e	43.00±0.00d	27.50±0.00f	45.99±0.00b	48.56±0.00a
LcG-IR	75.55±0.04b	72.61±0.02c	68.28±0.01e	70.57±0.03d	41.77±0.00f	87.25±0.00a
LcL-SC	36.87±0.01d	66.18±0.00b	40.49±0.05e	28.80±0.01f	67.11±0.22a	61.45±0.00c
LeC-IR	42.69±0.02c	42.32±0.01c	66.75±0.14a	21.00±0.00e	48.28±0.50b	40.55±0.00d
LeM-IR	76.47±0.05d	91.22±0.03a	81.76±0.01b	76.33±0.05e	79.78±0.00c	70.38±0.00f
LeP-IR	49.26±0.03c	29.80±0.01f	38.00±0.00e	62.23±0.12b	86.66±0.00a	46.08±0.00d
WCo-KG	39.13±0.02d	43.10±0.00c	37.98±0.05e	32.16±0.00f	56.97±0.12a	53.50±0.00b
WCi-MT	81.70±0.07a	47.46±0.02c	26.15±0.00d	47.18±0.04c	56.85±1.12b	47.72±0.00c
WP-IR	36.65±0.02a	35.21±0.01a	5.05±0.00b	20.23±0.01ab	15.80±27.37ab	26.33±0.00ab

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参考文献(27)

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