2001 Annual Science Report

Marine Biological Laboratory Reporting  |  JUL 2000 – JUN 2001

Eukaryotic Origins and the Evolution of Cellular Complexity -- Evolution of Tubulins

Project Summary
4 Institutions
3 Teams
0 Publications
0 Field Sites
Field Sites

Project Progress

Evolution of Tubulins (dm)

Jakobids are free-living, heterotrophic flagellates that might represent early-diverging mitochondriate protists. They share ultrastructural similarities with other eukaryotes that occupy basal positions in molecular phylogenies. Their mitochondria genome architecture is also eubacteria-like, suggesting a close affinity with the ancestral alpha proteobacterial symbiont that gave rise to mitochondria and hydrogenosomes. Tubulin gene phylogenies are in general agreement with mitochondrial gene phylogenies and ultrastructural data, indicating that the “jakobids” may be polyphyletic. Relationships to the putatively deep-branching amitochondriate diplomonads remain uncertain.

Since the 1980s, diplomonads and their relatives retortamonads have been proposed as primitive eukaryotes that diverged before the acquisition of mitochondria. On this basis they became key organisms in attempts to understand the evolution of eukaryotic cells. Recent findings of mitochondrial-like genes in Giardia suggest that diplomonads may have lost mitochondria. However, the mitochondrial origins of some of these genes are highly questionable, and premitochondrial diplomonad scenarios are still advanced. In this phylogenetic examination we focus on a series of mostly poorly-known eukaryotes argued to be related to diplomonads on morphological grounds; the excavate taxa. Phylogenies of SSUrRNA genes, alpha tubulin, beta tubulin and concatenated alpha + beta tubulin all scatter the various excavate taxa across the diversity of eukaryotes. However all phylogenies place the excavate taxon Carpediemonas as a close relative of diplomonads (and with SSUrRNA, retortamonads), to the exclusion of all other eukaryotes. This novel relationship is recovered across phylogenetic methods and various taxon-deletion experiments and under different SSUrRNA alignments. Statistical support is strongest under maximum likelihood (when among-site rate variation is modelled) and when the most divergent diplomonad sequences are omitted, suggesting a true relationship rather than a long-branch attraction artefact. As Carpediemonas contains double membrane-bound organelles that are reminiscent of mitochondria, we speculate on the possible occurrence of mitochondria in the ancestors of diplomonads. The phylogenetic position of Carpediemonas suggests that it will be valuable in interpreting the evolutionary significance of many molecular and cellular biological peculiarities of diplomonads.

  • PROJECT INVESTIGATORS:
  • PROJECT MEMBERS:
    Mitchell Sogin
    Project Investigator

    Virginia Edgcomb
    Co-Investigator

    Andrew Roger
    Co-Investigator

    David Kysela
    Research Staff

    Alastair Simpson
    Doctoral Student

  • RELATED OBJECTIVES:
    Objective 2.0
    Develop and test plausible pathways by which ancient counterparts of membrane systems, proteins and nucleic acids were synthesized from simpler precursors and assembled into protocells.

    Objective 4.0
    Expand and interpret the genomic database of a select group of key microorganisms in order to reveal the history and dynamics of evolution.