How Does Tremblaya princeps Get Essential Proteins from Its Nested Partner Moranella endobia in the Mealybug Planoccocus citri?

Many insects maintain intracellular mutualistic symbiosis with a wide range of bacteria which are considered essential for their survival (primary or P-endosymbiont) and typically suffer drastic genome degradation. Progressive loss of P-endosymbiont metabolic capabilities could lead to the recruitment of co-existent facultative endosymbiont (secondary or S-endosymbiont), thus adding more complexity to the symbiotic system. Planococcus citri, among other mealybug species, harbors an unconventional nested endosymbiotic system where every Tremblaya princeps cell (β-proteobacterium) harbors many Moranella endobia cells (γ-proteobacterium). In this system, T. princeps possess one of the smallest prokaryote genome known so far. This extreme genome reduction suggests the supply of many metabolites and essential gene products by M. endobia. Although sporadic cell lysis is plausible, the bacterial participation on the regulation of the predicted molecular exchange (at least to some extent) cannot be excluded. Although the comprehensive analysis of the protein translocation ability of M. endobia PCVAL rules out the existence of specific mechanisms for the exportation of proteins from M. endobia to T. princeps, immunolocation of two M. endobia proteins points towards a non-massive but controlled protein provision. We propose a sporadic pattern for the predicted protein exportation events, which could be putatively controlled by the host and/or mediated by local osmotic stress.