Perspectives on the origin of microfilaments, microtubules, the relevant chaperonin system and cytoskeletal motors- a commentary on the spiro chaete origin of flagella

The origin of cytoskeleton and the origin of relevant intracelluar transportation system are big problemsfor understanding the emergence of eukaryotic cells. The present article snmmarized relevant information of evidences and molecular traces on the origin of actin, tubulin, the chaperonin system for folding them,myosins, kinesins, axonemal dyneins and cytoplasmic dyneins. On this basis the authors proposed a seriesof works, which should be done in the future, and indicated the ways for reaching the targets.These targets are mainly: 1) the reconstruction of evolutionary path from MreB protein of archaeal ancestor ofeukaryotic cells to typical actin; 2) the finding of the MreB or MreB-related proteins in crenarchaea andusing them to examine J. A. Lake''s hypothesis on the origin of eukaryote from "eocytes" (crenarchaea);3) the examinations of the existence and distribution of cytoskeleton made of MreB-related protein withincoccoid archaea, especially in amoeboid archaeon Thermoplasm acidophilnm; 4) using Thermoplasma asa model of archaeal ancestor of eukaryotic cells; 5) the searching for the homolog of ancestral dynein inpresent-day living archaea. During the writing of this article, Margulis'' famous spirochaete hypothesis onthe origin of flagella and cilia was unexpectedly involved and analyzed from aspects of tubulin.% dyneinsand spirochaetes. Actually, spirochaete cannot be reasonably ass,med as the ectosymbiotic ancestor of eukaryotic flagella and cilia, since their swing depends upon large amount of bacterial flagella beneath theflexible outer wall, but not depends upon their intracelluar tubules and the assumed dyneins. In this case,if they had "evolved" into cilia and lost their bacterial fiagella, they would immediately become immobile!In fact, tubulin and dynein-like proteins have not been found in any spirochaete.

The origin of cytoskeleton and the origin of relevant intracellular transportation system are big problems for understanding the emergence of eukaryotic cells. The present article summarized relevant information of evidences and molecular traces on the origin of actin, tubulin, the chaperonin system for folding them, myosins, kinesins, axonemal dyneins and cytoplasmic dyneins. On this basis the authors proposed a series of works, which should be done in the future, and indicated the ways for reaching the targets. These targets are mainly: 1) the reconstruction of evolutionary path from MreB protein of archaeal ancestor of eukaryotic cells to typical actin; 2) the finding of the MreB or MreB-related proteins in crenarchaea and using them to examine J. A. Lake's hypothesis on the origin of eukaryote from "eocytes" (crenarchaea); 3) the examinations of the existence and distribution of cytoskeleton made of MreB-related protein within coccoid archaea, especially in amoeboid archaeon Thermoplasm acidophilum; 4) using Thermoplasma as a model of archaeal ancestor of eukaryotic cells; 5) the searching for the homolog of ancestral dynein in present-day living archaea. During the writing of this article, Margulis' famous spirochaete hypothesis on the origin of flagella and cilia was unexpectedly involved and analyzed from aspects of tubulins, dyneins and spirochaetes. Actually, spirochaete cannot be reasonably assumed as the ectosymbiotic ancestor of eukaryotic flagella and cilia, since their swing depends upon large amount of bacterial flagella beneath the flexible outer wall, but not depends upon their intracellular tubules and the assumed dyneins. In this case, if they had "evolved" into cilia and lost their bacterial flagella, they would immediately become immobile! In fact, tubulin and dynein-like proteins have not been found in any spirochaete.