Antibodies that specifically target surface-exposed or secreted antigens can effectively protect after natural infection and vaccination. While a multitude of antigens is recognised by antibodies following infection, most of them do not confer protection. Understanding the nature and targets of protective antibodies to Gram-negative bacteria is important to develop a new generation of vaccines against these increasingly antibiotic-resistant pathogens.
Following our identification of the outer-membrane protein (OMP) OmpD from nontyphoidal Salmonella as a protective antigen, here we report our studies on how antibody to OmpD protects and what its potential strengths and limitations as a vaccine against multiple Salmonella serovars are. Understanding of the protection afforded by antibodies requires taking into account the wider spatial context and interplay of the membrane and lipopolysaccharide (LPS) O-chain alongside the proteinateous antigens. To address this we generated the first of its kind full-atomistic molecular dynamics (FA-MD) model of the outer membrane, which revealed an unexpected complexity of antigen-antibody interaction associated with the LPS-O-chain layer.
Our data suggests that the native, trimeric OmpD creates a tunnel in the LPS-O-chain layer of sufficient size to allow epitope binding on the bacterial surface by the Fab-region of antibody, potentially explaining why antibodies to OmpD can be protective. However, surprisingly, OmpD from S. Typhimurium does not cross-protect against bacteria expressing conserved OmpD-variants that differ by a single amino-acid. Notably, loss of the O-chain restores bacterial susceptibility to killing possibly by unmasking covered epitopes.
Further analysis of structural and FA-MD modelling corroborated with in vivo and in vitro studies of immune response to Salmonella infection in mice suggests a central role of LPS O-chain in modulating the protective antigenic properties of OMP-based epitopes. Our model allows a testable a priori prediction of the antibody-binding properties of surface antigens and will be helpful in identifying protective antigens for vaccines.