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Hesperetin Protects Crayfish Procambarus Clarkii Against White

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Hesperetin Protects Crayfish Procambarus Clarkii Against White Fish and Shellfish Immunology 93 (2019) 116–123 Contents lists available at ScienceDirect Fish and Shellfish Immunology journal homepage: www.elsevier.com/locate/fsi Full length article Hesperetin protects crayfish Procambarus clarkii against white spot syndrome virus infection T Xiyi Qian, Fei Zhu* Zhejiang Provincial Engineering Laboratory for Animal Health Inspection and Internet Technology, College of Animal Science and Technology, Zhejiang Agriculture and Forestry University, Hangzhou, 311300, China ARTICLE INFO ABSTRACT Keywords: Hesperetin is a natural flavanone compound, which mainly exists in lemons and oranges, and has potential Hesperetin antiviral and anticancer activities. In this study, hesperetin was used in a crayfish pathogen challenge to discover WSSV its effects on the innate immune system of invertebrates. The crayfish Procambarus clarkii was used as an ex- Innate immunity perimental model and challenged with white spot syndrome virus (WSSV). Pathogen challenge experiments Procambarus clarkii showed that hesperetin treatment significantly reduced the mortality caused by WSSV infection, while the VP28 copies of WSSV were also reduced. Quantitative reverse transcriptase polymerase chain reaction revealed that hesperetin increased the expression of several innate immune-related genes, including NF-kappaB and C-type lectin. Further analysis showed that hesperetin treatment plays a positive effects on three immune parameters like total hemocyte count, phenoloxidase and superoxide dismutase activity. Nevertheless, whether or not in- fected with WSSV, hesperetin treatment would significantly increase the hemocyte apoptosis rates in crayfish. These results indicated that hesperetin could regulate the innate immunity of crayfish, and delaying and re- ducing the mortality after WSSV challenge. Therefore, the present study provided novel insights into the po- tential therapeutic or preventive functions associated with hesperetin to regulate crayfish immunity and protect crayfish against WSSV infection, provide certain theoretical basis for production practice. 1. Introduction the genus Whispovirus of the Nimaviridae family [10,11]. All the major species of cultivated penaeid shrimp can be naturally infected by this Procambarus clarkii, commonly known as crayfish, is a species of virus [6–9]. In addition to penaeid species, both natural and experi- freshwater crayfish, native to northern Mexico, and southern and mental infections have also been reported in caridean shrimp (Macro- southeastern United States [1]. This species had been introduced into brachium rosenbergii), crayfish (Orconectes punctimanus), wild crabs many areas of China and had become an important economic species of (Calappa lophos, Portunus sanguinolentus, P. pelagicus, Charybdis sp., freshwater aquaculture [2]. With the rapid development of crayfish Helice tridens and Scylla serrata), wild lobsters (Panulirus sp.), palae- pond culture and rice shrimp comprehensive breeding industry, the monidian pest shrimp, krill (Acetes sp.), planktonic copepods and pupae white spot disease of crayfish has become increasingly prominent. of an ephydridian insect [12–14]. WSSV replicates in the host cells after White spot disease is one of the most disastrous in shrimp culture, infecting the host. Virus could not be eliminated by conventional drugs having reduced shrimp production extensively worldwide. Mortality without damaging the host cells, there is no specific treatment for the rates are usually very high and cumulative mortality can reach 100% viral disease. During the breeding process, disease prevention and within 3–10 days from the onset of visible gross signs [3,4]. The clinical control measures are mostly based on prevention. Once the white spot symptoms of white spot syndrome include loss of appetite, lethargy, disease breaks out, the crayfish's mortality rate is extremely high, which and the appearance of white spots on the exoskeleton [5]. White spot leads to the farmers' blood loss [5]. Therefore, there is an urgent need to disease had been reported from several Asian countries since the early develop a therapeutic drug that is less harmful to crayfish cells and 1990s, and then from South America and from the south-eastern coast could effectively reduce the mortality rate after infected WSSV. of the USA [6–9]. By consulting a large amount of literature, we found that hesperetin White spot syndrome virus (WSSV) is a rod-shaped crustacean virus has multiple effects, such as anti-inflammatory, anti-cancer, anti-virus with a wide geographic distribution and host range, which belongs to and so on. Hesperetin is a type of natural flavonoid that occurs mainly * Corresponding author. E-mail addresses: [email protected], [email protected] (F. Zhu). https://doi.org/10.1016/j.fsi.2019.07.029 Received 21 May 2019; Received in revised form 5 July 2019; Accepted 11 July 2019 Available online 11 July 2019 1050-4648/ © 2019 Elsevier Ltd. All rights reserved. X. Qian and F. Zhu Fish and Shellfish Immunology 93 (2019) 116–123 in the form of glycoside hesperidin, which is the main flavonoid in crayfish were taken for virus testing, and then the infected crayfish lemons and oranges. It is known that hesperetin has an anti-oxidation were stored in an ultra-low temperature freezer. In the virus challenge effect. It was selected as a biological antioxidant to protect osteoblasts experiment mentioned in this article, we took out the infected crayfish and was used as an auxiliary preparation for melanin in the treatment from the ultra-low temperature freezer, thawed it, took its muscles, and of type I diabetic bone disease [15]. Additional, hesperetin has an anti- made the meat batter. In manner of feeding attack, each crayfish was inflammatory effect and can reduce the production of sebum, thereby fed with minced meat, which was about the size of rice grains. effectively inhibiting the growth of acne. Hesperetin can also be used to protect the cardiovascular, protect the nerves, and reduce triglyceride 2.2. Hesperetin treatment and pathogen challenge content in the liver [16–19]. Hesperetin has an anti-cancer effect, and citrus flavonoid hesperetin and naringin have obvious inhibitory effects Crayfish were randomly transferred into separate tanks as different on the proliferation of human breast cancer cells in vitro [20]. Most groups. In the hesperetin treatment experiment, crayfish were either fed importantly, hesperetin has antiviral properties, and has significant with hesperetin additive feed every 24 h as the hesperetin group or antiviral activity against the Chikungunya virus (CHIKV), reducing commercial pellet feed as the control group. To determine the influence CHIKV replication efficiency and down-regulating the production of of hesperetin on crayfish innate immune signal pathways, the hemo- viral proteins involved in replication [21]. Hesperetin can reduce in- lymph of each group was collected 24 h post feeding to analyze gene tracellular replication of herpes simplex virus type 1 (HSV-1), polio- expression. To determine the influence of hesperetin in the pathogen virus type 1, parainfluenza virus type 3 (Pf-3), and respiratory syncytial challenge process, after 72 h of hesperetin treatment, crayfish in each virus (RSV) [22]. Hesperetin showed its inhibitory activity on Sindbis group were fed with minced meat of WSSV-infected crayfish. After the virus infection in vitro and were not cytotoxic on Baby Hamster cells 21 feeding, crayfish fed commercial pellet feed only, 40 mg/kg hesperetin clone 15 (BHK-21) [23]. The intake of hesperetin in laying hens diet additive feed only, 50 mg/kg hesperetin additive feed only, 60 mg/kg may exhibit positive effects and increased the ratio of yolk weight/egg hesperetin additive feed only, commercial pellet feed for cray- weight and the blood serum superoxide dismutase (SOD) activity [24]. fish + minced meat, 40 mg/kg hesperetin additive feed + minced Based on this, we would explore whether hesperetin could effectively meat, 50 mg/kg hesperetin additive feed + minced meat, 60 mg/kg inhibit viral replication and reduce mortality in crayfish. hesperetin additive feed + minced meat were placed back into sepa- To elucidate whether hesperetin could be used as an effective an- rated tanks for further research. Crayfish fed with commercial pellet tiviral additive for crayfish, we detecting the immunological indicators feed alone were defined as the control group and those fed with 40, 50, before and after treatment with hesperetin, as well as the im- or 60 mg/kg hesperetin additive feed alone were defined as the 40 mg/ munological activity, VP28 copies of WSSV, mortality and apoptosis in kg hesperetin, 50 mg/kg hesperetin, or 60 mg/kg hesperetin groups, crayfish after WSSV infection. In the present study, the results indicated respectively. Crayfish fed with commercial pellet feed and minced meat that hesperetin could regulate the innate immunity, inhibit viral re- were defined as the WSSV group. Crayfish fed with either 40, 50, or plication and reducing mortality of crayfish. 60 mg/kg hesperetin additive feed and minced meat were defined as the WSSV +40 mg/kg hesperetin, WSSV +50 mg/kg hesperetin, or the 2. Materials and methods WSSV +60 mg/kg hesperetin groups, respectively. Based on the re- quirements of different analysis, crayfish samples were collected at 2.1. Crayfish, hesperetin and pathogens preparation different times after hesperetin or pathogen treatment. To analyze mortality, eight groups of crayfish were kept in the tanks Healthy crayfish (approximately 15 g and 10 cm each) were pur- for 12 days and the populations were counted every 24 h. Dead crayfish chased from a crayfish breeding case in Wenzhou, China. In order to were removed immediately, and aquatic water was replaced with clean adapt the crayfish to the new environment, the crayfish were bred for a water every day. To ensure the statistical accuracy in mortality analysis, week before the experiment, they were kept in tanks with fresh water every group contained 20 individuals. The mortality data was arranged and fed with commercial pellet feed at 5% of body weight per day, the and analyzed in Microsoft GraphPad 5.0. In the analysis of WSSV co- room temperature and water temperature were set at 25 °C constantly.
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