“…One of the most notorious opportunistic pathogens, Vibrio harveyi , has been reported frequently since the 1990s to infect a wide range of marine fish and invertebrates [ 12 , 13 , 14 , 15 , 16 , 17 , 18 , 19 ]. Various tissues or organs, such as the brain [ 20 ], gills [ 21 ], eyeball [ 22 ], liver [ 19 ], spleen [ 23 ], head kidney [ 24 ], intestine [ 25 , 26 ] and heart [ 27 ] have been reported to be targets of V. harveyi infection. As one of the most common pathogens worldwide, the crucial virulence genes of V. harveyi are still not systematically revealed [ 28 , 29 ] since each step of its infection is usually tightly controlled by specific virulence factors [ 30 ].…”