Tamames Javier, Gil Rosario, Latorre Amparo, Peretó Juli, Silva Francisco J, Moya Andrés
Institut Cavanilles de Biodiversitat i Biologia Evolutiva, Universitat de València, Apartado Postal 22085, 46071 València, Spain.
BMC Evol Biol. 2007 Oct 1;7:181. doi: 10.1186/1471-2148-7-181.
Bacterial symbioses are widespread among insects. The early establishment of such symbiotic associations has probably been one of the key factors for the evolutionary success of insects, since it may have allowed access to novel ecological niches and to new imbalanced food resources, such as plant sap or blood. Several genomes of bacterial endosymbionts of different insect species have been recently sequenced, and their biology has been extensively studied. Recently, the complete genome sequence of Candidatus Carsonella ruddii, considered the primary endosymbiont of the psyllid Pachpsylla venusta, has been published. This genome consists of a circular chromosome of 159,662 bp and has been proposed as the smallest bacterial endosymbiont genome known to date.
The detailed analysis of the gene content of C. ruddii shows that the extensive degradation of the genome is not compatible with its consideration as a mutualistic endosymbiont and, even more, as a living organism. The ability to perform most essential functions for a cell to be considered alive is heavily impaired by the lack of genes involved in DNA replication, transcription and translation. Furthermore, the shortening of genes causes, in some cases, the loss of essential domains and functional residues needed to fulfill such vital functions. In addition, at least half of the pathways towards the biosynthesis of essential amino acids, its proposed symbiotic function, are completely or partially lost.
We propose that this strain of C. ruddii can be viewed as a further step towards the degeneration of the former primary endosymbiont and its transformation in a subcellular new entity between living cells and organelles. Although the transition of genes from C. ruddii to the host nucleus has been proposed, the amount of genes that should have been transferred to the germinal line of the insect would be so big that it would be more plausible to consider the implication of the mitochondrial machinery encoded in the insect nucleus. Furthermore, since most genes for the biosynthesis of essential amino acids have also been lost, it is likely that the host depends on another yet unidentified symbiont to complement its deficient diet.
细菌共生现象在昆虫中广泛存在。此类共生关系的早期建立可能是昆虫进化成功的关键因素之一,因为这可能使昆虫能够利用新的生态位和新的不均衡食物资源,如植物汁液或血液。最近,已对不同昆虫物种的几种细菌内共生体的基因组进行了测序,并对其生物学特性进行了广泛研究。最近,被认为是木虱Pachpsylla venusta主要内共生体的“Candidatus Carsonella ruddii”的完整基因组序列已发表。该基因组由一条159,662 bp的环状染色体组成,被认为是迄今为止已知的最小细菌内共生体基因组。
对“C. ruddii”基因含量的详细分析表明,基因组的广泛降解与其作为互利共生内共生体甚至作为活生物体的地位不符。由于缺乏参与DNA复制、转录和翻译的基因,细胞执行大多数基本功能的能力受到严重损害。此外,基因的缩短在某些情况下导致实现此类重要功能所需的必需结构域和功能残基的丧失。此外,其假定的共生功能——必需氨基酸生物合成的至少一半途径完全或部分丧失。
我们认为,这种“C. ruddii”菌株可被视为先前主要内共生体退化并转变为介于活细胞和细胞器之间的亚细胞新实体的又一步。尽管有人提出基因从“C. ruddii”转移到宿主细胞核,但应该转移到昆虫生殖系的基因数量如此之多,以至于认为昆虫细胞核中编码的线粒体机制起作用更合理。此外,由于必需氨基酸生物合成的大多数基因也已丧失,宿主很可能依赖另一种尚未鉴定的共生体来补充其不足的饮食。