维基百科

假单胞菌属

假单孢菌科的一属细菌

假单胞菌属（学名：Pseudomonas）是一类需氧的革兰氏阴性细菌，它位于假单胞菌科下，已知物种有191个。该属微生物具有极其丰富的代谢多样性，这些多样性也使得它们能够在非常广阔的生态位中生存。[1]假单胞菌属的微生物在in vitro条件下很容易培养，这使得该属成为科研的绝佳材料，比较典型的研究对象有人类致病菌绿脓杆菌、植物致病菌丁香假单胞菌、土壤细菌恋臭假单胞菌以及能促进植物生长的荧光假单胞菌。

假单胞菌属
琼脂板上的绿脓杆菌菌落
科学分类 编辑
域： 细菌域 Bacteria
门： 假單胞菌門 Pseudomonadota
纲： γ-變形菌綱 Gammaproteobacteria
目： 假單胞菌目 Pseudomonadales
科： 假单胞菌科 Pseudomonadaceae
属： 假单胞菌属 Pseudomonas
Migula, 1894
模式種
绿脓杆菌 Pseudomonas aeruginosa
(Schroeter, 1872) Migula, 1900
种

见正文

異名
  • Flavimonas Holmes et al. 1987
  • Chryseomonas Holmes et al. 1986
  • Serpens Hespell 1977

因为广泛存在于水体和诸如双子叶植物等的种子中，假单胞菌属算是较早发现的一种微生物。假单胞菌属这个名字最早在1894和1900年由Walter Migula（英语：Walter Migula）给出，当时只是一个很模糊的名词，它的意思是指“革兰氏阴性菌，杆状以及一端有鞭毛的、能够形成芽孢的细菌”。[2][3]但是后来这些被认为芽孢的东西被证明只是某些储存物质的颗粒而已。[4]现在我们已经不用这些比较模糊的描述了，而选用绿脓杆菌作为模式种来研究。[4]

分类历史

和大部分细菌属一样，假单胞菌属的最后共同祖先生活于大约几亿年前。它最早在19世纪末由Walter Migula（英语：Walter Migula）定名，但它名字的来源在当时并未给出，而是最早见于第七版的细菌命名法著作《伯杰氏系统细菌学手册》(Bergey's Manual of Systematic Microbacteriology（英语：Bergey's Manual of Systematic Microbacteriology）)：pseudo来源于希腊语的pseudes(ψευδής)，即“假”之义；-monas来源于μονάς/μονάδος，为“单个单元”之义；合起来可以把这个名字理解为“假的单元”之义。但是这个名字没有任何有价值的含义，它并不是说这种生物可能错误地会出现单个细胞的状态，因为它从来就没有以多细胞的形式存在过。一种比较可靠的猜测是，Migula这样起名可能单纯是想说这种生物是假的“Monas”，Monas是指金藻纲的一种有着微小鞭毛的原生生物。[4]不久之后，其他符合Migula这一模糊定义的微生物也陆续被发现并被加入到这一属下，但是它们当中的很多在后来通过检测保守性大分子等方法又被重新划分到其他门类下了。[5]

最近，利用16S rRNA测序分析手段，很多细菌都被重新划分分类。[6]有一些原来属于单胞菌属和黄色单胞菌属的细菌又被划分到了假单胞菌属下。[7]原来属于假单胞菌属的一些菌株现在被分到了伯克氏菌属和罗尔斯通氏菌属。[8][9]

2000年，人们获得了假单胞菌属某物种的全基因组序列，其后很多其他菌株的序列也被测出，包括：P. aeruginosa strains PAO1 (2000), P. putida KT2440 (2002), P. protegens Pf-5 (2005), P. syringae pathovar tomato DC3000 (2003), P. syringae pathovar syringae B728a (2005), P. syringae pathovar phaseolica 1448A (2005), P. fluorescens Pf0-1, and P. entomophila L48。[5]

下属物种

本属包括以下物种：

绿脓杆菌 Pseudomonas aeruginosa 组
绿叶假单胞菌Pseudomonas chlororaphis 组
荧光假单胞菌 Pseudomonas fluorescens 组
穿孔素假单胞菌Pseudomonas pertucinogena 组
恋臭假单胞菌Pseudomonas putida 组
施氏假单胞菌Pseudomonas stutzeri 组
丁香假单胞菌Pseudomonas syringae组
组的地位未定的菌种：

特征

假单胞菌属的总体特征如下：[53]

假单胞菌属的其他特征还有（部分种例外）在铁限制条件下分泌一种荧光铁载体。[54]有些假单胞菌属物种还可以分泌其他类型的铁载体，如绿脓杆菌可以分泌绿脓杆菌毒素（英语：Pyocyanin）[55]，荧光假单胞菌可以分泌一种叫做thioquinolobactin的含硫铁载体。[56]假单胞菌属细菌氧化酶测试呈阳性，这也是其一个典型特征。

假单胞菌属可能是云中形成冰晶的最重要的成核剂，它们在全世界的雨雪形成过程中都起着十分重要的作用。[57]

生物膜形成

人们以前认为，假单胞菌属中所有的种都属于专性需氧微生物。近来人们在假单胞菌形成的生物膜中发现了例外。[58]在生物膜形成过程中，许多细胞都能够产生诸如藻朊酸盐等胞外多糖（英语：Extracellular polymeric substance），这些多糖可以防止假单胞菌被白细胞吞噬。[59]胞外多糖还可以使假单胞菌在食物表面形成一片片难以去除的生物膜。腐坏食物表面的假单胞菌还会使食物闻起来有水果般的味道。

抗生素抗性

因为是革兰氏阴性细菌，大部分假单胞菌对青霉素以及很多相关β-内酰胺类抗生素都有抗性，但也有部分对哌拉西林、亚胺培南、替卡西林以及环丙沙星敏感。[59]临床治疗上的其他选择是诸如妥布霉素、庆大霉素以及阿米卡星等氨基糖苷类抗生素。

因为拥有含有孔蛋白的坚硬细胞壁，假单胞菌能够在严苛的环境下生存。它们的外排泵能够在许多抗生素發挥作用之前就把它们排出去。

绿脓杆菌是一种机会致病菌，对抗生素表现出低敏感性，较为棘手。[60]一是因为绿脓杆菌细胞壁渗透性低，二是因为其抗生素抗性基因（如mexAB-oprM、mexXY[61]）可调控多种药物外排泵协同运作。 除了内生抗性，绿脓杆菌还很容易通过基因组突变或水平基因转移来产生新的抗性。若要获得对多种药物的抗性，就需要发生不同的突变或者接受多个水平基因的转移。高频突变倾向于在绿脓杆菌中产生能够引起慢性感染的抗生素抗性，而在内含子中同时出现多个抗生素抗性基因会使菌株同时具有多种抗性。有些研究发现，有些表型抗性会随生物膜形成或小的变种菌落而出现。[5]

对镓元素的敏感性

虽然镓元素没有什么自然的生物学功能，但它能对细胞过程产生类似于三价铁的影响。当细菌错误地将镓当成铁吸收后，镓并不像铁那样能够传递电子，这会影响细胞呼吸，最终导致细胞死亡。[62][63]

分类学

在二十世纪初，依靠经典分类方法来描述和鉴定这一微生物属经历了一段颇为坎坷的过程。当人们将比对核糖体RNA的大分子组分作为一项评判标准引入后，假单胞菌的分类就豁然开朗了：根据核糖体RNA的相似性，假单胞菌属可以分为五个核糖体RNA同源组。在Migula命名假单胞菌属数十年后，被划归到该属下的微生物种数目达到了惊人的比例。目前该属的微生物种类已经缩减到了不到原来的10%，这其中有不少菌是新命名的，而原来属于假单胞菌属的微生物，其实已经没有几个目前还在该属下了。依赖除核糖体RNA外的其他保守大分子的分类方法使得我们能够控制该属不至于过于庞大。[5]

致病性

动物病原体

致病菌包括绿脓杆菌、栖稻假单胞菌栖稻假单胞菌（英语：Pseudomonas oryzihabitans）、和变形假单胞菌（英语：Pseudomonas plecoglossicida）。绿脓杆菌是医院里一个较为头疼的问题，因为它对病人来说是侵染力排第二位的致病菌（院内感染）[來源請求]。它的致病机理可能和绿脓杆菌分泌的蛋白质有一定关系。该细菌具有种类非常广泛的分泌系统，它们分泌的许多蛋白质都和临床株的致病性有关。[64]

植物病原体

丁香假单胞菌是一类植物致病菌。它具有五十多种致病变种（英语：pathovar），其中许多具有高度的宿主专一性。假单胞菌属的许多其他物种也能成为植物致病菌，但是丁香假单胞菌是研究最为透彻的一个。

尽管并非严格的植物病原体，托拉斯假单胞菌（英语：Pseudomonas tolaasii）在农业上也是一个头疼的问题，它可以造成栽培蘑菇的细菌性斑点病。[65]能够造成栽培蘑菇疾病的还有伞菌假单胞菌（英语：Pseudomonas agarici）。[66]

作为生防试剂

二十世纪八十年代中期始，假单胞菌属中的某些物种就开始被用来抑制作物致病菌的生长或定殖。这种应用通常称作生物防治。荧光假单胞菌和Pseudomonas protegens（英语：Pseudomonas protegens）的菌株（如CHAO或Pf-5）的生物防治特性是目前研究最为透彻的，尽管人们目前还不十分清楚荧光假单胞菌促进植物生长的过程是如何实现的。这其中可能的方式有：假单胞菌促进了宿主植物的系统抗性，使得植物能够更好地抵抗病原体；打败其他的土壤（致病）微生物（如分泌铁载体（：铁载体）来获取铁元素）；产生对其他土壤微生物有害的化合物，如吩嗪类抗体或氰化氢。上述所有方式均已得到了实验证实。[67]

其他具有生物防治功能的假单胞菌还有：绿叶假单胞菌（英语：Pseudomonas chlororaphis），产生吩嗪类抗生素抑制特定真菌病原体生长[68]；桔黄假单胞菌（英语：Pseudomonas aurantiaca）产生一种一直格兰仕阳性细菌的抗生素类物质di-2,4-diacetylfluoroglucylmethane。[69]

作为生物修复试剂

假单胞菌属的某些物种能够代谢环境污染物，它们可以用来进行生物修复。相关物种包括：

食物腐败

因为具有多种多样的代谢特征，能够在低温和特殊环境下生存，很多假单胞菌都能够导致食物腐败。典型的例子有：莓实假单胞菌（英语：Pseudomonas fragi）可以引起奶制品腐坏,[78]，腐臭假单胞菌（英语：Pseudomonas taetrolens）和霉味假单胞菌（英语：Pseudomonas mucidolens）能够造成鸡蛋腐臭，[79]隆德假单胞菌（英语：Pseudomonas lundensis），能够造成牛奶、奶酪、肉类和鱼类的腐败。[80]

以前属于该属的微生物种

最近，人们已经通过16s核糖体RNA测序将许多以前曾经属于假单胞菌属的微生物进行了重新分类。[6]这些被移出假单胞菌属的物种如下所示；点击相应物种原名会出现它们目前的新名字：

α 变形菌: P. abikonensis（英语：Sphingomonas abikonensis）, P. aminovorans（英语：Aminobacter aminovorans）, P. azotocolligans（英语：Sphingomonas trueperi）, P. carboxydohydrogena（英语：Bradyrhizobium）, P. carboxidovorans（英语：Oligotropha carboxidovorans）, P. compransoris（英语：Zavarzinia compransoris）, P. diminuta（英语：Brevundimonas diminuta）, P. echinoides（英语：Sphingomonas echinoides）, P. extorquens（英语：Methylobacterium extorquens）, P. lindneri（英语：Zymomonas mobilis）, P. mesophilica（英语：Methylobacterium mesophilicum）, P. paucimobilis（英语：Sphingomonas paucimobilis）, P. radiora（英语：Methylobacterium radiotolerans）, P. rhodos（英语：Methylobacterium rhodinum）, P. riboflavina（英语：Devosia riboflavina）, P. rosea（英语：Methylobacterium extorquens）, P. vesicularis（英语：Brevundimonas vesicularis）.

β 变形菌: P. acidovorans（英语：Comamonas acidovorans）, P. alliicola（英语：Burkholderia gladioli）, P. antimicrobica（英语：Burkholderia gladioli）, P. avenae（英语：Acidovorax avenae）, P. butanovorae（英语：Thauera）, P. caryophylli（英语：Burkholderia caryophylli）, P. cattleyae（英语：Acidovorax avenae）, P. cepacia（英语：Burkholderia cepacia）, P. cocovenenans（英语：Burkholderia cocovenenans）, P. delafieldii（英语：Acidovorax delafieldii）, P. facilis（英语：Acidovorax facilis）, P. flava（英语：Hydrogenophaga flava）, P. gladioli（英语：Burkholderia gladioli）, P. glathei（英语：Burkholderia glathei）, P. glumae（英语：Burkholderia glumae）, P. huttiensis（英语：Herbaspirillum huttiense）, P. indigofera（英语：Vogesella indigofera）, P. lanceolata（英语：Comamonadaceae）, P. lemoignei（英语：Paucimonas lemoignei）, P. mallei, P. mephitica（英语：Janthinobacterium lividum）, P. mixta（英语：Telluria mixta）, P. palleronii（英语：Hydrogenophaga palleronii）, P. phenazinium（英语：Burkholderia phenazinium）, P. pickettii（英语：Ralstonia pickettii）, P. plantarii（英语：Burkholderia plantarii）, P. pseudoflava（英语：Hydrogenophaga pseudoflava）, P. pseudomallei, P. pyrrocinia（英语：Burkholderia pyrrocinia）, P. rubrilineans（英语：Acidovorax avenae）, P. rubrisubalbicans（英语：Herbaspirillum rubrisubalbicans）, P. saccharophila（英语：Matsuebacter）, P. solanacearum（英语：Ralstonia solanacearum）, P. spinosa（英语：Hydrogenophaga）, P. syzygii（英语：Ralstonia syzygii）, P. taeniospiralis（英语：Hydrogenophaga taeniospiralis）, P. terrigena（英语：Comamonas terrigena）, P. testosteroni（英语：Comamonas testosteroni）.

γ-β 变形菌: P. beteli（英语：Stenotrophomonas）, P. boreopolis（英语：Xanthomonas）, P. cissicola（英语：Xanthomonas）, P. geniculata（英语：Stenotrophomonas）, P. hibiscicola（英语：Stenotrophomonas）, P. maltophilia（英语：Stenotrophomonas maltophilia）, P. pictorum（英语：Stenotrophomonas）.

γ 变形菌: P. beijerinckii（英语：Chromohalobacter）, P. diminuta（英语：Brevundimonas diminuta）, P. doudoroffii（英语：Aeromonas）, P. elongata（英语：Microbulbifer elongatus）, P. flectens（英语：Enterobacteriaceae）, P. halodurans（英语：Halomonas halodurans）, P. halophila（英语：Marinobacter）, P. iners（英语：Marinobacterium georgiense）, P. marina（英语：Halomonadaceae）, P. nautica（英语：Marinobacter hydrocarbonoclasticus）, P. nigrifaciens（英语：Pseudoalteromonas nigrifaciens）, P. pavonacea（英语：Acinetobacter）,[81]P. piscicida（英语：Pseudoalteromonas piscicida）, P. stanieri（英语：Marinobacterium stanieri）.

δ 变形菌: P. formicans（英语：Aeromonas caviae）.

噬菌体

能够侵染假单胞菌的噬菌体有很多，例如：

参考文献

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