In recent decades, due to the continuous increasing demand for aquatic products, the global fishery industry has been expanding rapidly, and aquaculture output has been increasing at an unprecedented speed (Cole et al., 2009). High-density aquaculture is the current mainstream aquaculture modes worldwide. While the environmental stress induced by high rearing density results in poor resistance to disease and vulnerability to infection by pathogenic microorganisms in fish (Costas, Aragão, Mancera, Dinis, & Conceicão, 2008). For a long time, many antibiotics and chemotherapeutic agents have been used to improve non-specific immunity and to treat some bacterial infections in aquatic animals, but these agents also lead to undesirable effects, such as antibiotic resistance, pharmaceutical bioaccumulation and environmental pollution (Bilen, Bulut, & Bilen, 2011;Harikrishnan, Balasundaram, & Heo, 2011). In recent years, many harmless therapeutic techniques and