Ayala F J
Department of Ecology and Evolution, University of California, Irvine 92717, USA.
Biol Res. 1993;26(1-2):47-63.
This paper presents three results concerning the population structure of Trypanosoma cruzi, the agent of Chagas disease: (1) The mode of propagation of T. cruzi in nature is clonal; sexual reproduction is either totally absent or so rare that it leaves no traces in the population structure of the parasite. (2) The genetic diversity of the clonal lineages is large: extant T. cruzi represent lineages of descent that have evolved independently for long time spans (up to 40 million years). (3) Some genetically identical clonal lineages ("clonets") are geographically widespread ("ubiquitous"). However, most clonets are endemic, restricted in geographic distribution. These results have each in turn consequences of epidemiological significance: (1) In a sexually-reproducing organism the individual genotype is ephemeral; the entity that persists and evolves is the species ("gene pool"), and a few individuals contain most of the genetic variability of the species. In a clonally-propagating organism, the entity that persists and evolves is the clonal lineage; the genetic diversity of the species can only be captured by extensive sampling of distinct lineages. (2) The extensive genetic divergence among clonal lineages implies proportionally diverse biological characteristics, which are likely to include pathological effects, host propensity, vulnerability to drugs and vaccines, and other medically significant attributes. The extant T. cruzi lineages diverged much before human origins; hence, specific adaptation to human hosts, to whichever extent it exists, has evolved independently in separate lineages, and may not have evolved at all in some T. cruzi. (3) Epidemiological surveys and medical characterization, including search for specific vaccines and drugs, should not proceed randomly; rather, preliminary surveys must identify those clonets that are ubiquitous and target them for investigation. Review of published literature shows that Leishmania (and other parasitic protozoa) also has a clonal population structure. We advance a taxonomic and nomenclatural proposal that is appropriate for clonal organisms, yet simple.
(1)克氏锥虫在自然界中的传播方式为克隆性繁殖;有性生殖要么完全不存在,要么极为罕见,以至于在寄生虫的种群结构中未留下任何痕迹。(2)克隆谱系的遗传多样性很大:现存的克氏锥虫代表了在很长时间跨度(长达4000万年)内独立进化的谱系。(3)一些基因相同的克隆谱系(“克隆群”)在地理上广泛分布(“无处不在”)。然而,大多数克隆群是地方性的,地理分布受限。这些结果依次具有各自的流行病学意义:(1)在有性生殖的生物体中,个体基因型是短暂的;持续存在并进化的实体是物种(“基因库”),少数个体包含了物种的大部分遗传变异性。在克隆繁殖的生物体中,持续存在并进化的实体是克隆谱系;物种的遗传多样性只能通过对不同谱系的广泛采样来获取。(2)克隆谱系之间广泛的遗传差异意味着相应的生物学特征也存在差异,这可能包括病理效应、宿主倾向、对药物和疫苗的易感性以及其他医学上重要的属性。现存的克氏锥虫谱系在人类起源之前就已经分化;因此,对人类宿主的特定适应性(无论其存在程度如何)是在不同谱系中独立进化的,并且在某些克氏锥虫中可能根本没有进化。(3)流行病学调查和医学特征分析,包括寻找特定的疫苗和药物,不应随机进行;相反,初步调查必须识别出那些无处不在的克隆群,并将其作为调查目标。对已发表文献的回顾表明,利什曼原虫(以及其他寄生原生动物)也具有克隆种群结构。我们提出了一个适用于克隆生物体且简单的分类和命名建议。