PMC:7115396 / 7673-19199 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/7115396","sourcedb":"PMC","sourceid":"7115396","source_url":"https://www.ncbi.nlm.nih.gov/pmc/7115396","text":"2 Materials and methods\n\n2.1 Vaccine and viruses\nThe H120 vaccine strain is currently a widely used vaccine strain, created from 120 serial passages of the H strain isolated in 1956. The ck/CH/IBYZ/2011 strain (referred to as IBYZ, GenBank KF663561.1) used in this study is classified as a QX-like IBV isolated from a flock presenting with IB symptoms by our research group in 2011 in Jiangsu Province, China. Both of these strains were recovered from a full-length clone using reverse genetics, as previously described [35], [36].\n\n2.2 Eggs and animals\nFertilized specific pathogen free (SPF) chicken embryos were obtained from Beijing Merial Vital Laboratory Animal Technology Co., Ltd. (Beijing, China) and hatched in our lab (Lab of Chicken Infectious Disease Protection and Control, Poultry Institute, Chinese Academic of Agriculture Sciences, Yangzhou, China). For hatching, one-day-old SPF chickens from the SPF chicken embryos as described previously were incubated at 37 °C in a relative humidity of 55–65%. All chickens were maintained in isolators under negative pressure, and food and water were provided ad libitum.\n\n2.3 Construction of recombinant rH120-S1/YZ\nThe rIBV rH120-S1/YZ strain used in this study is described in the schematic illustration presented in Fig. 1 . The genome RNA was synthesized in vitro by T7 RNA polymerase and transfected into BHK-21 cells, as previously described [35], [36]. The cell supernatants were harvested 48 h following transfection and propagated in 10-day-old SPF chicken embryos. The allantoic fluid was harvested for RT-PCR, whole genome sequencing, and was then stored at −80 °C until further use.\nFig. 1 Schematic diagram for the construction of the chimeric S gene and production of a full-length cDNA of rH120-S1/YZ. (A) Replacement of the H120 S1 fragment by the corresponding sequence of IBYZ for construction of pYZS1H120S2. The plasmids pIBYZS and pH120S contained the S gene of IBYZ strain and H120 strain, respectively, were constructed during the establishment of reverse genetic system. Primers PS1F and PS1R were used to amplify the S1 fragment of IBYZ and vector fragment, while primers PS2F and PS2R were used to amplify the S2 fragment of H120 strain. By overlapping PCR, the pYZS1H120S2 was constructed which contained a chimeric S gene. (B) Strategy for the construction of full-length cDNA clones of rH120-S1/YZ. Ten cDNA fragments covering the entire genome of H120 strain was amplified by RT-PCR. Unique Bsa I sites were inserted at the junctions between each clone, a unique T7 start site was inserted at the 5′ end of clone TM1, and a 28-nucleotide T tail was inserted at the 3′ end of clone TM10. The S fragment in the pMDTM8 plasmid of H120 strain was replaced by chimeric S gene. By using appropriate ligation strategy, the genomic cDNA of rH120-S1/YZ was assembled by in vitro ligation using appropriate restriction sites as indicated.\n\n2.4 Chicken embryo dwarf assay\nTo determine the recombinants’ pathogenicity in chicken embryos, 0.2 mL of rH120, rIBYZ, or rH120-S1/YZ virus (diluted to 1:100 in normal saline) was inoculated into the allantoic cavities of five 10-old-day embryonated SPF chicken eggs, respectively. Chick embryo lesions were examined for lesions at 144 h post-inoculation at 37 °C.\n\n2.5 Growth kinetics in chicken embryos\nTo examine the viral growth ability in chicken embryos, a real-time reverse transcription quantitative polymerase chain reaction (RT-qPCR) method was established. According to the sequences of IBV from GenBank, primers were designed based on conservative area in the 5′-UTR. The upstream primer was 5′-CCGTTGCTTGGGCTACCTAGT-3′, and the downstream primer was 5′-CGCCTACCGCTAGATGAACC-3′. The amplification product was cloned to pMD18-T vector (Takara) as a positive plasmid and its concentration was measured. A gradient dilution of 5 × 102–5 × 108 copies/μL of the plasmid was used as template for quantitation test. By plotting the cycle threshold (CT) values against the copies of the plasmid, the standard curve was generated.\n18 10-day-old embryonated SPF chicken eggs (six eggs/group) were inoculated with rH120, rIBYZ, and rH120-S1/YZ at a dose of 107 viral RNA copies/100 μL, and used for growth curve experiments. The allantoic fluids were collected separately by syringe from the six inoculated embryonated eggs of each group at 12, 18, 24, 36, 48, 72, 96 h per inoculation. RNA was extracted from the allantoic fluids using TRIzol Reagent (Invitrogen, Carlsbad, CA, USA) following the manufacturer’s instructions. cDNA was obtained by reverse transcription using a PrimeScript RT Master Mix Perfect Real Time Kit (TaKaRa, Otsu, Shiga, Japan). The viral copies were measured by absolute quantitative method of Real-time PCR, which was performed using SYBR® Premix Ex Taq™ II (TaKaRa, Otsu, Shiga, Japan) on an Applied Biosystems 7500 Fast Real-time PCR System. The standard curve was plotted against the log of the template copy number. All of the assays were run in triplicate and the copy number of each virus was calculated according to the standard curve.\n\n2.6 Titration of rIBVs in embryos\nSerial 10-fold dilutions from 105 copies to 109 copies per 0.1 mL of virus of the recombinant strain, rH120-S1/YZ, and its parental strains (rH120 and rIBYZ) were inoculated into the allantoic cavity of 10-day-old embryonated SPF chicken eggs. For each dilution, 0.1 mL of the virus solution was injected into each egg and 10 eggs were used for each dilution. PBS was used as a negative control. After a 24 h incubation, the dead embryos were considered nonspecific deaths and discarded. The embryos were examined for the presence of specific lesions caused by the virus after a 144-h incubation. Dead and live embryos that displayed IBV infectious signs (e.g., dwarfing, curling, and stunting) were considered positive samples. The 50% embryo infectious dose (EID50) of these rIBVs was calculated using the Reed-Muench method [37].\n\n2.7 Safety testing\n\n2.7.1 Assessment of pathogenicity in one-day-old SPF chickens\nThe chickens were separately housed in isolators under consistent conditions, and food and water was provided ad libitum. A total of 120 one-day-old SPF chickens were randomly divided into four groups (n = 30 per group). The chickens in the experimental groups were intranasally inoculated with 200 μL allantoic fluid per chick containing 105EID50 of the rH120, rIBYZ, and rH120-S1/YZ strains. The control group (n = 30) was inoculated with 200 μL sterile PBS via the same route. The morbidity and mortality was followed-up for 14 days. All experimental groups were monitored daily for clinical signs related to IB infection for 14 days post-infection (dpi), including coughing, sneezing, and tracheal rales. Dead chickens were examined for gross tracheal, lung, and kidney lesions.\n\n2.7.2 Tissue tropism of early viral infection in vivo\nA total of 100 1-day-old SPF chickens were assigned to four groups (n = 25 per group). The birds were inoculated with 200 μL allantoic fluid containing 105 EID50 of the rH120, rIBYZ, or rH120-S1/YZ strains, respectively via eye drop and the intranasal route and 200 μL PBS per chick was administered to the control group (n = 25) via the same route. The tracheas, lungs, kidneys, and bursa from the 5 inoculated birds per group were harvested at 1, 3, 5, and 7 dpi, weighed, and collected into 1 mL PBS per sample and frozen at −80 °C. After grinding the samples, the viral RNA was extracted using TRIzol, and the cDNA was obtained by reverse transcription using a PrimeScript RT Master Mix Perfect Real Time Kit (TaKaRa, Otsu, Shiga, Japan). The viral RNA copies from each of the different samples were detected by real-time PCR as described above. All assays were run in triplicate and the copy number for each virus was calculated according to the standard curve.\n\n2.7.3 Histopathology\nAfter inoculation with 105 EID50 dose of the rH120, rIBYZ, and rH120-S1/YZ strains, three infected chickens per group were randomly selected to be euthanized by bleeding, and the tissue pathology was examined in the different groups at 3, 5, 7, 10 dpi. The tracheas, lungs, kidneys, and bursas of three dead or other randomly-selected chickens from the experimental and control groups (inoculated with PBS) were collected at 5 or 7 dpi and further processed for histopathology. The samples were fixed in formalin, embedded in paraffin, cut into 5 μm sections, and stained with hematoxylin and eosin. The slides were examined under light microscopy for the presence of lesions.\n\n2.7.4 Viral shedding\nTo determine the level of viral shedding of the different recombinants, 10 birds per group were inoculated with 200 μL allantoic fluid containing 105 EID50 of the rH120, rIBYZ, or rH120-S1/YZ strains, respectively. 200 μL PBS per chick was administered to the control group via eye drop and the intranasal route. . Oral and cloacal swabs were collected from each bird into 1 mL PBS at 7, 14, 18, 22, 26, and 30 dpi and stored at −80 °C. After freezing and thawing three times and centrifuging at 8,000 × g for 5 min at 4 °C, relative amount of virus present in 200 μL of supernatants of each sample was quantified by RNA extraction, reverse transcription, and real-time PCR as described previously.\n\n2.8 Efficacy of rH120-S1/YZ vaccination\n\n2.8.1 Viral challenge\nIn total, 30 one-day-old SPF chickens were divided into three groups of 10 chickens. The three groups were intranasally vaccinated with the rH120 and rH120-S1/YZ strains at a dose of 104 EID50/200 μL, and the control group received PBS. Two weeks post-vaccination, the chickens in each group were challenged with the QX-like strain, rIBYZ, at 106 EID50/200 μL via eye drop and the intranasal route. Any clinical signs, as well as the percentage of morbidity and mortality were recorded for 14 days. The dead chickens were also examined for gross tracheal, lung, and kidney lesions.\n\n2.8.2 Antibody response measured by an enzyme-linked immunosorbent assay (ELISA)\n45 1-day-old SPF chickens were divided into three groups (n = 15 per group). After vaccinating with the rH120 and rH120-S1/YZ strains at a dose of 104 EID50/200 μL, the sera of 10 birds from each group were randomly collected and antibodies were tested using an ELISA created in our lab at 7, 14, 21, and 28 days post-vaccination (dpv). ELISA plates were coated with antigen of culture supernatants of inactivated IBV rMJ , a vero cells adaption strain domesticated from its parental strain of IBYZ in our lab. Individual chicken sera collected from different groups diluted to 1:200 were loaded and the plates were incubated at 37 °C for 60 min. IBV-specific IgG was detected with anti-chicken IgY (IgG) (whole molecule)-peroxidase antibodies produced in rabbits (Sigma-Aldrich, Germany). After incubated with 3,3′,5,5′-Tetramethylbenzidine(TMB) substrate for 15 min and terminated with 2 mol/L H2SO4 solution, the absorbance at 490 nm was measured using a Model 680 microplate reader (Bio-Rad, USA).\n\n2.9 Statistical analysis\nGraphPad Prism 7 software (GraphPad Software Inc., La Jolla, CA, USA) was used for the statistical analyses. For the antibody test, growth kinetics, tissue tropism, and viral shedding test, the collected data were analyzed using a two-way ANOVA to determine whether there was a significant difference between the different groups. The significance was considered as follows: significant at P ≤ 0.05 (*); highly significant at P ≤ 0.01 (**); and extremely significant at P ≤ 0.001 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