ε-聚赖氨酸(ε-poly-l-lysine, ε-PL)是抑菌谱广泛的天然抑菌剂,由通过α-羧基与ε-氨基连接的25-35个赖氨酸聚合而成。ε-PL主要由白色链霉菌发酵生产所得,比化学生产更加高效和环保。ε-PL具有水溶性好、耐热和对环境无污染等特点,具有良好的应用前景。本文从发酵生产入手,着重综述了ε-PL对各种微生物抑菌性能、抑菌机制及抑菌机制模型的研究进展。推测ε-PL是通过对细胞膜的破坏而改变细胞的通透性,或者作用到细胞内引起活性氧(reactive oxygen species, ROS)胁迫而影响调节基因的表达,从而起到抑菌作用。根据这2种抑菌方式分别建立了相应的抑菌模型,即毡毯模型和ROS诱导细胞凋亡模型。本文可为ε-PL对微生物抑制性能的深入研究提供依据,同时也提出了ε-PL抑菌机制的新模型,为扩展ε-PL应用领域提供了一定的参考。
ε-poly-l-lysine (ε-PL) is a natural antibacterial agent with a wide range of bacteriostatic spectrum. It is synthesized by the polymerization of 25 to 35 lysines linked by α-carboxyl group and ε-amino group. ε-PL is mainly produced by the fermentation of
ε-聚赖氨酸是一种生物聚合物,主要由25−35个l-赖氨酸通过α-羧基和ε-氨基形成的酰胺键连接而成,固体ε-聚赖氨酸主要为浅黄色粉末,吸湿性较强,易溶于水,微溶于乙醇,不溶于结构复杂的有机溶剂,具有良好的热稳定性,但与金属离子共同作用会影响其活性[
目前,ε-聚赖氨酸在日本已经有了工业化生产,主要用作食品添加剂,我国也在2014年将ε-聚赖氨酸列为食品添加剂新品种[
为了能够全面、深入地了解ε-聚赖氨酸的抑菌作用,本文综述ε-聚赖氨酸对各种微生物抑菌性能、机制及抑菌模型的研究进展。在现有的抑菌模型基础上,我们提出了一种可能的新型抑制模型。此外,为了解ε-聚赖氨酸分子特点与其抑菌性能之间的关系,本文也简要综述了微生物发酵生产ε-聚赖氨酸状况,为研究具有不同分子大小的ε-聚赖氨酸发酵生产及其抑菌性能之间的联系提供参考,将来可为ε-聚赖氨酸的特定应用提供定制化生产。
ε-聚赖氨酸是1977年由学者[
由于过去缺乏高效的手段,无法大量筛选ε-聚赖氨酸产生菌,导致无法大规模培养ε-聚赖氨酸产生菌。随着科技的发展,研究人员利用ε-聚赖氨酸是聚阳离子多肽且呈酸性的特点找到了一个快速筛选该菌种的方法,即基础酸性染料法,将所有菌种培养在同样带正电荷的琼脂中,并用碱性染料(如甲基蓝),利用静电斥力在ε-聚赖氨酸产生菌周围形成一个透明圈,从而达到分离的目的[
由于从自然界中筛选的ε-聚赖氨酸产生菌产量较少且培养周期长,很难满足应用需求,因此,研究人员对ε-聚赖氨酸产生菌进行了大量的育种工作,希望可以提高ε-聚赖氨酸的产量。目前,对于菌种分泌ε-聚赖氨酸的原理尚在研究之中,所以育种方式大多采用诱变育种[
目前,欧美一些国家及韩国、日本等通过发酵培养已经实现了ε-聚赖氨酸的商业化应用。我国仅有几家公司对ε-聚赖氨酸实现了工业化生产。对于ε-聚赖氨酸的发酵生产,目前所做的研究主要包括利用废弃物制作培养基[
总而言之,生物聚合物的分子量与其实际应用密切相关[
如引言所述,ε-聚赖氨酸产生菌通过二氨基庚二酸途径合成前体l-赖氨酸,具体的合成途径如
ε-聚赖氨酸的合成途径
The synthetic route of ε-poly-l-lysine.
对于ε-聚赖氨酸合成机理的研究仍在深入。Kawai等[
ε-聚赖氨酸作为食品加工工艺中应用较为广泛的防腐剂,其优势在于对人体无害,摄入后在人体体内即可转化成赖氨酸并且具有广泛的抑菌谱,少量即可达到抑菌效果。魏奇等[
由于ε-聚赖氨酸被广泛应用于食品防腐剂中,所以关于其灭菌机理主要集中在食物中常见的细菌,如李斯特菌和金黄色葡萄球菌等。ε-聚赖氨酸针对真菌、革兰氏阳性菌和革兰氏阴性菌有着不同的抑菌效果[
由于酿酒酵母可以作为真菌中具有代表性的菌种,所以薄涛等[
ε-聚赖氨酸不仅对酿酒酵母细胞的细胞膜产生影响,对细胞壁也有相同的作用。另外,ε-聚赖氨酸对酿酒酵母的抑制作用可能与细胞内活性氧(reactive oxygen species, ROS)水平上升有关。Tan等[
此外,相关研究[
在抑制植物病原真菌方面,目前发现ε-聚赖氨酸在对白叶枯病菌作用时效果最佳。Liu等[
除了ε-聚赖氨酸直接对真菌的细胞膜和细胞内部的DNA进行攻击达到抑制细菌的效果外,ε-聚赖氨酸同样可以与被细菌附着在表面的宿主共同作用来抑制真菌病原体。Dou等[
对于革兰氏阴性菌菌种的选取,通常选用具有代表性的大肠杆菌。与ε-聚赖氨酸对酵母的抑菌作用相同,在0–150 μg/mL的浓度范围内ε-聚赖氨酸的浓度越高,则抑菌效果越好。付萍[
目前,对于ε-聚赖氨酸与其他抑菌素联合抑菌的作用依然是十分热门的话题。Ning等[
对于革兰氏阳性菌,主要选取了几个具有代表性的菌种,分别为枯草芽孢杆菌(
Hyldgaard等[
在复合材料做食品包装材料保鲜方面,利用ε-聚赖氨酸制作的纳米纤维对李斯特菌依旧有很好的抑菌效果。Lin等[
与上述2种革兰氏阳性菌的情况不同,单独的ε-聚赖氨酸对金黄色葡萄球菌的抑菌效果较好。有学者通过研究ε-聚赖氨酸对巴氏杀菌奶中金黄色葡萄球菌的抑制作用得出结论,400 μg/mL的ε-聚赖氨酸即可使89.4%的细胞膜破损,通过电镜观察发现细胞壁与细胞膜同样被严重破坏[
关于ε-聚赖氨酸对革兰氏阳性菌的抑菌机理研究较少,目前还有待进一步研究。
通过对各菌种抑菌情况分析,ε-聚赖氨酸的抑菌效果与自身浓度和目标菌种的结构有关。在对酿酒酵母作用时,500 μg/mL的ε-聚赖氨酸可使酵母细胞死亡;而在对大肠杆菌作用时,150 μg/mL的ε-聚赖氨酸即可使大肠杆菌内外膜发生破损,细胞完整性被破坏[
虽然ε-聚赖氨酸对革兰氏阳性菌和革兰氏阴性菌的抑菌效果有明显的区别,但近年来的研究依然致力于寻找可以使ε-聚赖氨酸对阳性菌及阴性菌均产生抑制作用的方法。Bastarrachea等[
ε-聚赖氨酸对不同菌种的抑菌情况
Inhibitory effect of ε-poly-l-lysine on different strains
微生物 |
最佳抑菌浓度 |
机制 |
应用 |
参考文献 |
300 | 破坏细胞膜 |
延长食品保质期 |
[ |
|
500 | 破坏细胞膜,ROS胁迫共同作用 |
作为食品防腐剂 |
[ |
|
512 | ROS胁迫 |
作为临床抗真菌药物 |
[ |
|
500 | 破坏细胞膜,ROS胁迫共同作用 |
用于防止水果采后损失 |
[ |
|
100 | 破坏细胞膜,ROS胁迫共同作用 |
农业上用作杀菌剂 |
[ |
|
600 | 破坏细胞膜,并且激活宿主细胞中防御相关酶活性 |
用作食品防腐剂 |
[ |
|
150 | 破坏细胞膜 |
用作抗生素 |
[ |
|
200 | 与溶解素Lysqdvp001共同作用引起ROS胁迫 |
用于食品工业中的抑菌剂 |
[ |
|
No data | ε-聚赖氨酸对其并无太大影响 |
[ |
||
400 | 降低跨膜电位导致K+外泄引起ROS胁迫 |
用于乳制品的杀菌剂 |
[ |
由于ε-聚赖氨酸是通过破坏细胞膜结构实现抑菌效果,王梓源等[
毡毯模型
Model of carpet.
为了进一步研究毡毯模型的抑菌过程及现象,薄涛等[
综合以上研究人员对ε-聚赖氨酸抑菌机理的研究,我们猜测ε-聚赖氨酸首先以静电形式铺满整个细胞膜,随后,类似强力胶,将细胞膜撕扯使其破损,造成疏水集团以及疏水蛋白外露,最终造成细胞死亡。
在研究ε-聚赖氨酸抑菌机制时,Ye等[
ROS诱导细胞凋亡模型
Model of ROS induces apoptosis.
随着研究人员对抗菌肽的不断研究,发现有多种抗菌肽的抑菌过程可以用ROS诱导细胞凋亡模型进行解释,如PAF26 (一种合成的杀菌六肽)、coprisin (一种多肽)和melittin (蜂毒肽)等,其中Lee等[
以上2种模型均对ε-聚赖氨酸的抑菌作用给出了合理的解释,但无论是毡毯模型还是ROS诱导细胞凋亡模型都并不完整,不能够解释ε-聚赖氨酸对所有细菌的抑制机理。因此,本文在原有模型的基础上提出一个新的模型,称其为复合抑菌模型(
复合抑菌模型
Model of compound bacteriostatic.
ε-聚赖氨酸用途广泛,作为食品中的防腐剂、工业生产中的化妆品、药物载体及基因工程中的载体,具有十分广阔的商业前景。本文综述了ε-聚赖氨酸对不同菌种的抑菌作用,并对ε-聚赖氨酸的抑菌模型进行了概括,同时在原有的模型基础上提出了一个新抑菌模型。根据本文综述内容及目前的研究情况,提出如下几点展望:
1) ε-聚赖氨酸作为防腐剂广泛应用于食品工业,但应用在防止水果采后变质、对农作物致病微生物抑制等相关内容较少,因此,对ε-聚赖氨酸应用在农业方面的研究还应加强。现代生物技术已显著提升了ε-聚赖氨酸在工业生产中的效率,并扩大了工业化应用。
2) ε-聚赖氨酸虽然有良好的抑菌性和较为广泛的抑菌谱,但单一物质的抑菌能力依旧有其局限性,未来的关注点可以适当放在与其他抑菌物质联合使用增强其抑制效果,或与高分子材料结合进行环境污染处理或医疗材料(如创可贴、医用材料)等。目前,较明确的研究方向在于,与某些高分子材料结合,利用ε-聚赖氨酸的抑菌性使感染创面快速愈合[
3) ε-聚赖氨酸在初次使用后会迅速损耗,其活性也会逐渐降低。因此,如何延长ε-聚赖氨酸的抑菌能力也是值得研究人员关注的方面。
今后在探究上述问题时,利用高通量基因测序技术或许会找到上述问题的答案。最终实现对ε-聚赖氨酸更全面的应用。
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