The greatest diversity of eukaryotic species is within the microbial eukaryotes, the protists, with plants and fungi/metazoa representing just two of the estimated seventy five lineages of eukaryotes. Protists are a diverse group characterized by unusual genome features and a wide range of genome sizes from 8.2？Mb in the apicomplexan parasite Babesia bovis to 112,000-220,050？Mb in the dinoflagellate Prorocentrum micans. Protists possess numerous cellular, molecular and biochemical traits not observed in “text-book” model organisms. These features challenge some of the concepts and assumptions about the regulation of gene expression in eukaryotes. Like multicellular eukaryotes, many protists encode multiple eIF4Es, but few functional studies have been undertaken except in parasitic species. An earlier phylogenetic analysis of protist eIF4Es indicated that they cannot be grouped within the three classes that describe eIF4E family members from multicellular organisms. Many more protist sequences are now available from which three clades can be recognized that are distinct from the plant/fungi/metazoan classes. Understanding of the protist eIF4Es will be facilitated as more sequences become available particularly for the under-represented opisthokonts and amoebozoa. Similarly, a better understanding of eIF4Es within each clade will develop as more functional studies of protist eIF4Es are completed. 1. Eukaryogenesis and Protein Synthesis Protein synthesis is an ancient, conserved, complex multienzyme system, involving the participation of hundreds of macromolecules in which the mRNA template is decoded into a protein sequence on the ribosome. The ribosome, a complex and dynamic nucleoprotein machine, provides the platform for amino acid polymerization in all organisms [1, 2]. This process utilizes mRNAs, aminoacyl tRNAs, and a range of protein factors, as well as the inherent peptidyl-transferase activity of the ribosome itself. The common origin of protein synthesis in all domains of life is evident in the conservation of tRNA and ribosome structure, as well as some of the additional protein factors. Although the basic molecular mechanisms are conserved across the three domains of life, the Bacteria (eubacteria), Archaea (archaebacteria), and Eukarya (eukaryotes), important divergences have taken place as eukaryotic species have evolved. The origin of the eukaryotic cell is enigmatic. Eukaryotes are thought to have evolved from a fusion of a euryarchaeon with a deep-rooted Gram-positive proteobacteria, the phylum from which mitochondria are derived [3]. It
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