Molecular Biology and Evolution, Vol 13, 1297-1305, Copyright © 1996 by Society for Molecular Biology and Evolution
PJ Keeling and WF Doolittle
The tubulin gene family, which includes alpha-,beta-, and gamma-tubulin
subfamilies, is composed of highly conserved proteins which are the
principle structural and functional components of eukaryotic microtubules.
We are interested in (1) establishing when in eukaryotic evolution the
duplications leading to paralogous alpha, beta, and gamma subfamilies
occurred and (2) the possible utility of tubulin sequences in
reconstructing organismal phylogeny. To broaden the taxonomic
representation of alpha-tubulins so that it roughly equals that of beta-
tubulins, alpha-tubulin genes from three Microsporidia (Encephalitozoon
hellem, Nosema locustae, and Spraguea lophii), two Parabasalia
(Monocercomonas sp. and Trichomitus batrachorum), and one Heterolobosean
(Acrasis rosea) were sequenced. With these new genes, phylogenetic trees of
alpha- and beta-tubulins were constructed and compared. Trees were
congruent with each other, but incongruent with other molecular
phylogenies. The agreement between alpha- and beta- tubulin trees could
arise by the co-adaptation of one molecule to variants of the other as a
result of their intimate steric association in microtubules. Thus, these
trees may not be providing independent support for the phylogenetic
results. However, one of these unexpected results, that microsporidia
cluster with fungi, is supported by other circumstantial evidence, and may
therefore reflect a real relationship despite the basal position usually
assigned to microsporidia. Relationships between the three tubulins were
also examined by constructing trees of all three types. These trees were
found to be of limited value for determining the position of the root
within each subfamily because of the great interfamily distances, but they
do confirm the classification of all known genes into three monophyletic
subfamilies. Divergent genes from Caenorhabditis elegans and Saccharomyces
cerevisiae that have been proposed to represent the novel classes delta-
and epsilon-tubulin were found to be specifically related to gamma-tubulins
from animals and fungi respectively, and therefore are best seen as rapidly
evolving orthologues of gamma- tubulin.
ORIGINAL ARTICLE
Alpha-tubulin from early-diverging eukaryotic lineages and the evolution of the tubulin family
Department of Biochemistry, Dalhousie University, Nova Scotia, Canada. pkeeling@is.dal.ca
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