Granato Elisa T, Kümmerli Rolf
Department of Plant and Microbial Biology, University of Zurich, Zurich, Switzerland.
BMC Evol Biol. 2017 Sep 11;17(1):214. doi: 10.1186/s12862-017-1060-6.
A common form of cooperation in bacteria is based on the secretion of beneficial metabolites, shareable as public good among cells within a group. Because cooperation can be exploited by "cheating" mutants, which contribute less or nothing to the public good, there has been great interest in understanding the conditions required for cooperation to remain evolutionarily stable. In contrast, much less is known about whether cheats, once fixed in the population, are able to revert back to cooperation when conditions change. Here, we tackle this question by subjecting experimentally evolved cheats of Pseudomonas aeruginosa, partly deficient for the production of the iron-scavenging public good pyoverdine, to conditions previously shown to favor cooperation.
Following approximately 200 generations of experimental evolution, we screened 720 evolved clones for changes in their pyoverdine production levels. We found no evidence for the re-evolution of full cooperation, even in environments with increased spatial structure, and reduced costs of public good production - two conditions that have previously been shown to maintain cooperation. In contrast, we observed selection for complete abolishment of pyoverdine production. The patterns of complete trait degradation were likely driven by "cheating on cheats" in unstructured, iron-limited environments where pyoverdine is important for growth, and selection against a maladaptive trait in iron-rich environments where pyoverdine is superfluous.
Our study shows that the path to re-evolve public-goods cooperation can be constrained. While a limitation of the number of mutational targets potentially leading to reversion might be one reason for the observed pattern, an alternative explanation is that the selective conditions required for revertants to spread from rarity are much more stringent than those needed to maintain cooperation.
细菌中一种常见的合作形式是基于有益代谢产物的分泌,这些代谢产物可作为公共物品在群体内的细胞间共享。由于合作可能会被“作弊”突变体利用,这些突变体对公共物品的贡献较少或没有贡献,因此人们对理解合作在进化上保持稳定所需的条件非常感兴趣。相比之下,对于作弊者一旦在种群中固定下来,在条件变化时是否能够恢复合作,人们了解得要少得多。在这里,我们通过对实验进化的铜绿假单胞菌作弊者进行研究来解决这个问题,这些作弊者在产生铁载体公共物品绿脓菌素方面部分存在缺陷,并将其置于先前已证明有利于合作的条件下。
经过大约200代的实验进化后,我们筛选了720个进化克隆,以检测它们绿脓菌素产生水平的变化。我们没有发现完全恢复合作的证据,即使是在空间结构增加和公共物品生产成本降低的环境中——这两个条件先前已被证明可以维持合作。相反,我们观察到了对完全停止产生绿脓菌素的选择。完全性状退化的模式可能是由在无结构、铁限制环境中“对作弊者作弊”驱动的,在这种环境中绿脓菌素对生长很重要,而在铁丰富环境中对一种适应不良性状的选择则是因为绿脓菌素是多余的。
我们的研究表明,重新进化公共物品合作的途径可能受到限制。虽然潜在导致回复突变的突变靶点数量有限可能是观察到这种模式的一个原因,但另一种解释是,回复突变体从稀有状态扩散所需的选择条件比维持合作所需的条件要严格得多。