乳酸菌发酵剂菌株的自溶特性及机理研究
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摘要
乳酸菌的自溶特性可影响发酵乳制品品质,发酵剂菌体的高度自溶不仅缩短了成熟周期、降低了生产成本,同时菌体自溶后释放的胞内物质也决定着发酵乳制品的风味和感官特性。本课题选择了发酵乳制品工业中最为常用的几种乳酸菌作为研究对象,对这些菌株的自溶特性进行研究,并将这一特性应用于酸乳发酵剂的选育工作中。主要研究结果如下:
利用碘化丙锭(PI)只能对细胞壁破损的菌体进行标记的特性,建立经PI染色联用流式细胞仪的方法(PI-FCM),可快速检测乳酸菌菌株的自溶度。将乳酸菌菌体细胞经20 mmol/L的PI-PBS染液在4℃条件下避光染色30 min后上样,流式细胞仪检测器激发波长488 nm,检测波长630 nm,每个样品收集105个细胞,得到PI染色阳性细胞数与细胞总数之间的比值,即为乳酸菌的自溶度;使用平板计数法和菌悬液吸光值法对PI-FCM测定结果的进行验证,相关性系数分别为r=0.9805和r=0.9249(p<0.01),说明PI-FCM方法可对乳酸菌自溶度进行准确测定。与传统方法相比,PI-FCM方法的灵敏度(103 cells/mL～107 cells/mL)、精密度(CV<5%)和准确度均处于较高水平,且耗时短、干扰因素少,可作为乳酸菌菌株自溶度的检测方法。
选择发酵性能好的野生型菌株进行N+离子注入诱变,离子注入能量为50 keV,注入剂量为4×2.6×1013 ion/cm2,菌体存活率在23%～33%之间,总突变率在57%～78%之间,突变幅度在-51.96%～+151.24%之间;对突变菌株进行筛选,经7次传代培养验证,得到了遗传性状稳定、自溶度改变的乳酸菌菌株。
自溶度较小的菌株细胞体积较小,易形成长链;乳酸菌自溶度因其种属不同而不同,同一种属不同菌株之间自溶度也存在差异,菌株的生物量与自溶度并无明显相关;过高或者过低的环境温度和pH值都可抑制乳酸菌的自溶,但某些菌株在低pH值条件下也能呈现较高的自溶度;悬浮介质中的Mg2+和Ca2+对菌株自溶的促进作用较为显著。相同测定条件下,乳酸乳杆菌产脂肪酶活性最高,德氏乳杆菌保加利亚亚种产蛋白酶活性最高,同一种属自溶度越大的菌株产酶活性到达峰值所需时间越短。
使用已知的自溶酶基因acmA序列设计引物,对14株乳酸乳球菌基因组中的N-乙酰基肽聚糖水解酶进行PCR扩增,在12个菌株中得到了碱基长度约为1100 bp的特异性片段。这些片段的碱基序列和氨基酸序列与其他N-乙酰基肽聚糖水解酶基因具有高度的同源性。由不同自溶度的菌株中扩增得到的特异性产物,其氨基酸序列同源性为94.91%。使用荧光定量RT-PCR技术对各菌株中N-乙酰基肽聚糖水解酶的转录水平进行分析,发现菌株自溶度与N-乙酰基肽聚糖水解酶转录水平极为相关。
在酸乳发酵过程中,自溶度较高的乳酸菌菌株产酸性能更优,凝乳时间较短;菌株的自溶度对酸乳的黏度、持水率以及风味物质的成分及含量影响不大,但对酸乳的后酸化程度有着显著影响。在模拟货架期的10℃～15℃贮藏条件下,由自溶度较高的菌株制作的酸乳后酸化程度相对较低。选育出三株自溶度较高的菌株进行复配,运用二次正交旋转组合设计复合发酵剂中各菌株比例,根据所建模型优选结果,最后确定三种菌株最优组合方案为:GS191.46%、LD3-A3 2.44%、S15-A3 6.10%。使用该复合发酵剂制作的酸乳感官性状优良,具有较好的风味和质地,后酸化程度小,可极好的满足生产需要。
Bacterial peptidoglycan hydrolases (autolysins) degrade the peptidoglycan of cell walls, causing cell autolysis. Autolysis of lactococci used as starter cultures in the manufacture of fermented milk results in the leakage of lipases, proteases and peptidases and other intracellular components, which play an important role in flavor development during ripening. Hence, autolysis properties of lactic acid bacteria are crucial for their applications as dairy starters.
To establish a rapid and sensitive method to detect autolysis rate of lactic acid bacteria, A Flow Cytometric Method (FCM) can be employed. Cell suspensions were dyed 30 min by PI-PBS (20 mmol/L) in 4℃dark environment. A 630 nm long pass filter was used to collect the red fluorescence (FL3 488 nm), and 1×105 cells were collected each sample. Data were analyzed by CellQuest online analysis system. Test time depended on different concentrations of cell suspensions, were about 40 s to 100 s. FCM enumerations were accurate down to a concentration of 104 cells/mL. Correlation test results reported that there were no significant difference between FCM test and other normal methods for the determination of autolysis rate of lactic acid bacteria. FCM by use of fluorescent probes propidium iodide (PI) is a good analytical method that allows the rapid and sensitive measurement of viability of lactic acid bacteria. This FCM method provides tools to assess the viability of different lactic acid bacteria in fermentation starters and probiotic products.
In order to obtain different autolysis rate lactic acid bacteria, Streptococcus salivarius ssp., Lactobacillus delbrueckii ssp. Bulgaricus, Lactobacillus lactis, Lactobacillus casei, and Lactococcus lactis ssp. Lactis were mutated by 50 keV N+ ions implantation. The results indicate that the survival rate curve took a saddle shape and there was a high positive mutation rate by the dose of 4×2.6×1013 ion/cm2. The survival rate was 25%～33% under this implantation condition. Strains which have different autolyisis rates were obtained among the mutated strains whose mutation rate in the range of -51.96%～+151.24%. Fermentation property of these strains were stable after 7 generation transfer inoculation. It was indicated that the ion implantation technique is a feasible method in breeding lactic acid bacteria starter strains.
Autolysis rate of different strains were determined under various conditions of incubated temperature and pH values, as these factors were closely related to cell viability and flavor development in the products. Autolysis rate increased when temperature and pH values were rasing in a certain range. Over higher or lower temperature and pH value would inhibit autolysis of strains. But there were some strains indicate high autolysis in extreme low pH. Mg2+ could affect autolysis rate of lactic acid bacteria. Strains showed higher autolysis when they were suspended in 0.1 mol/L magnesian phosphate buffer. Lipase and protease activities were closely related to the sort of strains. Time of reaching the highest activities for lipase and protease depended on the autolytic degree of strains. All Lactococcus lactis ssp. Lactis strains showed higher lipase activities, and protease activities in Lactobacillus delbrueckii ssp. Bulgaricus, were mostly higher than other strains. These properties should be considered when commercial starters are selected.
The occurrence of the acmA gene, encoding the major peptidoglycan hydrolase of lactic acid bacteria, has been determined in 14 Lactococcus lactis ssp. Lactis and Lactococcus lactis ssp. cremoris strains. On the basis of PCR amplification of the gene acmA, 12 strains exhibited a single 1,100 bp fragment highly correlated with N-acetylmuramidase of Lactococcus lactis. There was discrepancy between the high autolysis level strains and low auloysis level strains. Using fluorescent quantitative real-time PCR testified that autolysis rate was closely related with the transcript level of N-acetylmuramidase.
Coagulation time decreased significantly by using high autolysis rate lactic acid bacteria fermented yogurt, were positivly related to acid producing ability of lactic acid bacteria starters. There was no raletive correlation between autolysis rate of strains and water holdup, viscosity and flavor compounds of yogurt. However, yogurt acidities were significant infuluenced by autolysis rate of lactic acid bacteria staters, especially in the simulated shelf life of 10℃~ 15℃storage conditions. High autolysis rate starters could decrease acidification level of yogurt.
A dratic orthogonal rotary composite design is employed to study the effects of characters of yogurt fermented by high autolysis strains in mixed starters. According to regression analysis to strains GS1-A8, LD3-A3 and S15 -A3 coding value (X1, X2, X3) as independent variables, the shelf life of yogurt acidity changing degree (Δ0T) Y1 and sensory evaluation scores Y2 as the dependent variable, a regression equation were established. The optimum combinated scheme of three strains of programs was selected according the model: GS1 91.46%, LD3-A3 2.44%, S15-A3 6.10%. Yogurt fermented by the selected optimum starter has favorable sensory properties, good flavor and texture and lower acidification level.
利用碘化丙锭(PI)只能对细胞壁破损的菌体进行标记的特性,建立经PI染色联用流式细胞仪的方法(PI-FCM),可快速检测乳酸菌菌株的自溶度。将乳酸菌菌体细胞经20 mmol/L的PI-PBS染液在4℃条件下避光染色30 min后上样,流式细胞仪检测器激发波长488 nm,检测波长630 nm,每个样品收集105个细胞,得到PI染色阳性细胞数与细胞总数之间的比值,即为乳酸菌的自溶度;使用平板计数法和菌悬液吸光值法对PI-FCM测定结果的进行验证,相关性系数分别为r=0.9805和r=0.9249(p<0.01),说明PI-FCM方法可对乳酸菌自溶度进行准确测定。与传统方法相比,PI-FCM方法的灵敏度(103 cells/mL～107 cells/mL)、精密度(CV<5%)和准确度均处于较高水平,且耗时短、干扰因素少,可作为乳酸菌菌株自溶度的检测方法。
选择发酵性能好的野生型菌株进行N+离子注入诱变,离子注入能量为50 keV,注入剂量为4×2.6×1013 ion/cm2,菌体存活率在23%～33%之间,总突变率在57%～78%之间,突变幅度在-51.96%～+151.24%之间;对突变菌株进行筛选,经7次传代培养验证,得到了遗传性状稳定、自溶度改变的乳酸菌菌株。
自溶度较小的菌株细胞体积较小,易形成长链;乳酸菌自溶度因其种属不同而不同,同一种属不同菌株之间自溶度也存在差异,菌株的生物量与自溶度并无明显相关;过高或者过低的环境温度和pH值都可抑制乳酸菌的自溶,但某些菌株在低pH值条件下也能呈现较高的自溶度;悬浮介质中的Mg2+和Ca2+对菌株自溶的促进作用较为显著。相同测定条件下,乳酸乳杆菌产脂肪酶活性最高,德氏乳杆菌保加利亚亚种产蛋白酶活性最高,同一种属自溶度越大的菌株产酶活性到达峰值所需时间越短。
使用已知的自溶酶基因acmA序列设计引物,对14株乳酸乳球菌基因组中的N-乙酰基肽聚糖水解酶进行PCR扩增,在12个菌株中得到了碱基长度约为1100 bp的特异性片段。这些片段的碱基序列和氨基酸序列与其他N-乙酰基肽聚糖水解酶基因具有高度的同源性。由不同自溶度的菌株中扩增得到的特异性产物,其氨基酸序列同源性为94.91%。使用荧光定量RT-PCR技术对各菌株中N-乙酰基肽聚糖水解酶的转录水平进行分析,发现菌株自溶度与N-乙酰基肽聚糖水解酶转录水平极为相关。
在酸乳发酵过程中,自溶度较高的乳酸菌菌株产酸性能更优,凝乳时间较短;菌株的自溶度对酸乳的黏度、持水率以及风味物质的成分及含量影响不大,但对酸乳的后酸化程度有着显著影响。在模拟货架期的10℃～15℃贮藏条件下,由自溶度较高的菌株制作的酸乳后酸化程度相对较低。选育出三株自溶度较高的菌株进行复配,运用二次正交旋转组合设计复合发酵剂中各菌株比例,根据所建模型优选结果,最后确定三种菌株最优组合方案为:GS191.46%、LD3-A3 2.44%、S15-A3 6.10%。使用该复合发酵剂制作的酸乳感官性状优良,具有较好的风味和质地,后酸化程度小,可极好的满足生产需要。
Bacterial peptidoglycan hydrolases (autolysins) degrade the peptidoglycan of cell walls, causing cell autolysis. Autolysis of lactococci used as starter cultures in the manufacture of fermented milk results in the leakage of lipases, proteases and peptidases and other intracellular components, which play an important role in flavor development during ripening. Hence, autolysis properties of lactic acid bacteria are crucial for their applications as dairy starters.
To establish a rapid and sensitive method to detect autolysis rate of lactic acid bacteria, A Flow Cytometric Method (FCM) can be employed. Cell suspensions were dyed 30 min by PI-PBS (20 mmol/L) in 4℃dark environment. A 630 nm long pass filter was used to collect the red fluorescence (FL3 488 nm), and 1×105 cells were collected each sample. Data were analyzed by CellQuest online analysis system. Test time depended on different concentrations of cell suspensions, were about 40 s to 100 s. FCM enumerations were accurate down to a concentration of 104 cells/mL. Correlation test results reported that there were no significant difference between FCM test and other normal methods for the determination of autolysis rate of lactic acid bacteria. FCM by use of fluorescent probes propidium iodide (PI) is a good analytical method that allows the rapid and sensitive measurement of viability of lactic acid bacteria. This FCM method provides tools to assess the viability of different lactic acid bacteria in fermentation starters and probiotic products.
In order to obtain different autolysis rate lactic acid bacteria, Streptococcus salivarius ssp., Lactobacillus delbrueckii ssp. Bulgaricus, Lactobacillus lactis, Lactobacillus casei, and Lactococcus lactis ssp. Lactis were mutated by 50 keV N+ ions implantation. The results indicate that the survival rate curve took a saddle shape and there was a high positive mutation rate by the dose of 4×2.6×1013 ion/cm2. The survival rate was 25%～33% under this implantation condition. Strains which have different autolyisis rates were obtained among the mutated strains whose mutation rate in the range of -51.96%～+151.24%. Fermentation property of these strains were stable after 7 generation transfer inoculation. It was indicated that the ion implantation technique is a feasible method in breeding lactic acid bacteria starter strains.
Autolysis rate of different strains were determined under various conditions of incubated temperature and pH values, as these factors were closely related to cell viability and flavor development in the products. Autolysis rate increased when temperature and pH values were rasing in a certain range. Over higher or lower temperature and pH value would inhibit autolysis of strains. But there were some strains indicate high autolysis in extreme low pH. Mg2+ could affect autolysis rate of lactic acid bacteria. Strains showed higher autolysis when they were suspended in 0.1 mol/L magnesian phosphate buffer. Lipase and protease activities were closely related to the sort of strains. Time of reaching the highest activities for lipase and protease depended on the autolytic degree of strains. All Lactococcus lactis ssp. Lactis strains showed higher lipase activities, and protease activities in Lactobacillus delbrueckii ssp. Bulgaricus, were mostly higher than other strains. These properties should be considered when commercial starters are selected.
The occurrence of the acmA gene, encoding the major peptidoglycan hydrolase of lactic acid bacteria, has been determined in 14 Lactococcus lactis ssp. Lactis and Lactococcus lactis ssp. cremoris strains. On the basis of PCR amplification of the gene acmA, 12 strains exhibited a single 1,100 bp fragment highly correlated with N-acetylmuramidase of Lactococcus lactis. There was discrepancy between the high autolysis level strains and low auloysis level strains. Using fluorescent quantitative real-time PCR testified that autolysis rate was closely related with the transcript level of N-acetylmuramidase.
Coagulation time decreased significantly by using high autolysis rate lactic acid bacteria fermented yogurt, were positivly related to acid producing ability of lactic acid bacteria starters. There was no raletive correlation between autolysis rate of strains and water holdup, viscosity and flavor compounds of yogurt. However, yogurt acidities were significant infuluenced by autolysis rate of lactic acid bacteria staters, especially in the simulated shelf life of 10℃~ 15℃storage conditions. High autolysis rate starters could decrease acidification level of yogurt.
A dratic orthogonal rotary composite design is employed to study the effects of characters of yogurt fermented by high autolysis strains in mixed starters. According to regression analysis to strains GS1-A8, LD3-A3 and S15 -A3 coding value (X1, X2, X3) as independent variables, the shelf life of yogurt acidity changing degree (Δ0T) Y1 and sensory evaluation scores Y2 as the dependent variable, a regression equation were established. The optimum combinated scheme of three strains of programs was selected according the model: GS1 91.46%, LD3-A3 2.44%, S15-A3 6.10%. Yogurt fermented by the selected optimum starter has favorable sensory properties, good flavor and texture and lower acidification level.
引文
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