Model of leptospiral membrane architecture. Leptospires have two membranes, an outer membrane (OM) and a cytoplasmic or inner membrane (IM). As in Gram-positive bacteria, the peptidoglycan (PG) cell wall is closely associated with the IM. The leptospiral surface is dominated by lipopolysaccharide (LPS) carbohydrate side chains. Subsurface proteins include the cytoplasmic protein, GroEL, and the periplasmic endoflagella (EF). The IM contains lipoproteins such as LipL31 and transmembrane proteins such as signal peptidase (SP) and ImpL63. The OM contains lipoproteins including LipL41 and LipL36, and transmembrane proteins including the porin, OmpL1. Genomic sequence analysis reveals several novel types of outer membrane proteins (OMPs), including TonB-dependent OMPs involved in nutrient acquisition. BtuB is an example of a TonB-dependent OMP involved in uptake of vitamin B12. The type I efflux system is represented by TolC, which forms a complex with ATP-binding cassette (ABC) transporters to export hemolysins and other cytoplasmic components. The leptospiral genome also contains genes involved in a three-component efflux system consisting of an outer membrane factor (OMF), membrane fusion protein (MFP), and an inner membrane transporter, in this case CzcA, which is involved in heavy metal detoxification.