Remodeling of host membranes during herpesvirus assembly and egress

Many viruses,enveloped or non-enveloped,remodel host membrane structures for their replication,assembly and escape from host cells.Herpesviruses are important human pathogens and cause many diseases.As large enveloped DNA viruses,herpesviruses undergo several complex steps to complete their life cycles and produce infectious progenies.Firstly,herpesvirus assembly initiates in the nucleus,producing nucleocapsids that are too large to cross through the nuclear pores.Nascent nucleocapsids instead bud at the inner nuclear membrane to form primary enveloped virions in the perinuclear space followed by fusion of the primary envelopes with the outer nuclear membrane,to translocate the nucleocapsids into the cytoplasm.Secondly,nucleocapsids obtain a series of tegument proteins in the cytoplasm and bud into vesicles derived from host organelles to acquire viral envelopes.The vesicles are then transported to and fuse with the plasma membrane to release the mature virions to the extracellular space.Therefore,at least two budding and fusion events take place at cellular membrane structures during herpesviruses assembly and egress,which induce membrane deformations.In this review,we describe and discuss how herpesviruses exploit and remodel host membrane structures to assemble and escape from the host cell.