Mercury Transport in Bacteria

Mercuric ions (Hg2+) and methylmercury are major, human-generated, toxic contaminants present in fish and our waterways. Bacteria provide a means of bioremediation by taking up these compounds and reducing them to volatile, non-toxic, elemental mercury (Hg°). Three types of mercury/methylmercury transporters have previously been identified: MerC, MerF and MerT. Each of these sets of homologues has distinct topologies. MerF proteins are characterized by a 2-transmembrane α-helical segment (TMS) topology; most MerTs have three TMSs, and MerCs have four TMSs. This report shows that MerT and MerF proteins are related by common descent and are similar in sequence throughout their first two TMSs. One of the MerF proteins is internally duplicated, generating a protein with four TMSs, while several MerT homologues bear a C-terminal extracytoplasmic Hg2+-binding MerP domain. MerPs are homologous to heavy metal-binding domains present in copper chaperone proteins, at the N-termini of mercuric reductases and in from one to six copies in heavy metal transporting P-type ATPases. Phylogenetic analyses reveal that mercuric ion transporters have been horizontally transferred with high frequency between bacteria. Some MerTs function with MerP receptors while others do not, and the MerP-dependent MerTs cluster separately from the MerP-independent MerTs on a phylogenetic tree. MerTs possessing a MerP appear to have co-evolved with their cognate receptors. Conserved sequence and motif analyses serve to define the mercuric transporter family fingerprints and allow prediction of specific subfunctions. This report provides the first detailed bioinformatic description of two apparently unrelated families of Hg2+ uptake transporters. We propose that all members of these two families function by a simple channel-type mechanism to allow influx of Hg2+ in response to the membrane potential in preparation for reduction and detoxification. This information should facilitate the exploitation of these transporters for purposes of microbial and phytobioremediation.