Biology of Japanese encephalitis virus

 
Principal Investigator :  Sudhanshu Vrati

Project Associates/Assistants
Kaushik Bharati
Mohan Babu
Shivali Duggal
VG Anoop Kumar

Ph D Students
Surender Vashist
Manu Anantapadma

Japanese encephalitis virus (JEV) is a member of the Flaviviridae family of animal viruses that contains several other medically important viruses such as Dengue and Yellow fever. JEV is a major cause of human encephalitis and is responsible for considerable mortality and morbidity in India. Frequent epidemics of JEV are being reported from various parts of India and the virus has become endemic in several parts of the country. Different aspects of JEV biology are being studied in our laboratory, some of which are outlined below.

A.    Development of a plasmid DNA-based vaccine for JEV

The objective of this work is to develop plasmid DNA-based immunization procedures for vaccination against JEV. To this end, we intend to make recombinant plasmids carrying various structural and non-structural protein genes of JEV under the control of a strong eukaryotic promoter. These plasmid constructs will be evaluated for their potential to generate protective immunity in experimental animals against JEV.

In order to improve the efficacy of the plasmid DNA immunization against JEV, methods need to be employed that would enhance neutralizing antibody titers. In order to improve the uptake of the DNA in animal, we have adsorbed the plasmid DNA on cationic microparticles and have compared its immunogenicity with the naked plasmid DNA. We used plasmid pMEa for these studies. As seen in the ELISA, the microparticle-adsorbed DNA induced higher titers of anti-JEV antibodies when compared to those induced by the naked DNA. No difference, however, was seen in JEV neutralization titers. The microparticle-adsorbed DNA induced mixed Th1-Th2 kind of immune responses as opposed to Th1 type of immune responses elicited by the naked DNA.

B.    Development of a tissue culture-derived vaccine against JEV

A mouse brain derived vaccine using the Japanese Nakayama strain of JEV is currently being produced in limited amounts at the Central Research Institute, Kasauli. However, the cost of this vaccine precludes its large-scale usage for mass vaccination as may be required for the effective control of the virus activity in the field. In order to bring down the cost of vaccine production, efforts are being made to grow the virus to high titers in tissue cultured cells.

The virus grown in vero cells cultured on microcarriers in a spinner flask was formalin-inactivated and its immunogenicity and protective efficacy in mice were tested in comparison with the commercially available vaccine. Our studies indicated that formalin-inactivated, vero cell-cultured JEV induced high levels of protective immunity in mice. Virus inactivation with formalin at 22°C, which required shorter incubation period, was found to be as good or better to virus inactivation at 4°C for generating high titers of anti-JEV antibodies. Similarly, the 22°C-inactivated virus generated JEV neutralizing antibody titers as good or higher than those induced by the 4°C-inactivated virus. Thus for the vaccine production, inactivation of JEV with formalin at 22°C would be a preferred method as it is faster and does not require cold room storage.

C.    Interaction of cellular proteins with JEV RNA

JEV genome is a plus-sense single-stranded RNA of ~11 kilo bases. A minus-sense RNA template is generated during virus replication, which is then copied to produce a large number of plus-sense genomic RNA molecules. Based on the amino acid sequence homologies with other replicases, a couple of viral proteins have been speculated to be involved in replication of the JEV genome. However, we do not know if any of the cellular proteins also are needed for viral replication. We are, therefore, studying cellular proteins that interact with JEV genome sequences, which are likely to be involved in viral replication.

One of the proteins interacting with the 3'-NCR of JEV was identified to be the La protein of ~50 kDa. Out of the 408 amino acids of the protein, the N-terminal segment of 208 amino acids was found to bind the the 3'-NCR of JEV.

D.    Development of an oral JEV vaccine

Human adenoviruses cause mild, benign infections and in recent years these viruses are increasingly being used for human gene therapy and receombinant vaccine development. From the vaccine point of view, the added advantage that adenoviruses present is that they can be delivered orally. We are developing adenovirus recombinats that synthesise JEV E protein as potential vaccine candidate for immunization against Japanese encephalitis. We will also be developing transgenic plants that synthesize JEV E protein and study their potential in animals for oral immunization against JEV.

Replication-defective recombinant adenoviruses (RAds) were constructed that synthesized the pre-membrane and envelope (E) proteins of Japanese encephalitis virus. Recombinant virus RAdEa synthesized Ea, the membrane-anchored E protein, and RAdEs synthesized Es, the secretory E protein. Compared with RAdEs, RAdEa replicated poorly in HEK 293A cells and synthesized lower amounts of E protein. Oral immunization of mice with RAds generated low titers of anti-JEV antibodies that had only little JEV neutralizing activity. Intra-muscular (IM) immunization of mice with both RAds generated high titers of anti-JEV antibodies. Interestingly, RAdEa induced only low titers of JEV neutralizing antibodies, however, these were significantly higher in case of RAdEs immunization. Splenocytes from mice immunized IM with RAds secreted large amounts of interferon-g and moderate amounts of interleukin-5 in presence of JEV and showed cytotoxic activity against JEV-infected cells. Mice immunized IM with RAdEs showed complete protection against lethal dose of JEV given intra-cerebral.

Transgenic tobacco plants have been developed that contain the cDNA encoding JEV E protein in its full-length membrane-anchored or the truncated secretory forms under the control of the CaMV promoter. The tobacco plants are being used as model system to test the integrity of our gene constructs and their ability to express protein in plants. Subsequently, we plan to develop transgenic plants that could be consumed uncooked.

E.    Development of DNA molecules to inhibit JEV replication

JEV is a RNA virus which actively replicates in the host brain leading to the development of encephalitis. There is no specific treatment available to deal with JEV infection as no drugs are known that specifically inhibit JEV replication. We will attempt to design molecules that could be used for inhibiting JEV replication.

DNAzymes are short oligodeoxynucleotides that have the capability to cleave RNA molecules at defined site in a sequence-specific manner. The 3'- and the 5'-NCRs of JEV genome play crucial role in JEV replication. We have designed DNAzymes that can specifically cleave JEV RNA within the 3’- and the 5'-NCR. These oligonucleotides were found to inhibit JEV replication in cultured macrophage cells.

Publications

Original peer-reviewed articles

1.     Kaur R, Rauthan M and Vrati S (2004) Immunogenicity in mice of a cationic microparticle-adsorbed plasmid DNA encoding Japanese encephalitis virus envelope protein. Vaccine (in press).

2.     Appaiahgari MB and Vrati S (2004) Immunogenicity and protective efficacy in mice of a formaldehyde-inactivated Indian strain of Japanese encephalitis virus grown in vero cells. Vaccine (in press).

3.     Viswanathan P, Venkaiah B, Kumar MS, Rasheedi S, Vrati S, Bashyam MD and Hasnain SE (2003) The    homologous region sequence (hr1) of Autographa californica multinucleocapsid polyhedrosis virus can    enhance transcription from non-baculoviral promoters in mammalian cells. J Biol Chem 278:52564-52571.

4.     *Rath A., Batra D, Kaur R, Vrati S and Gupta SK (2003) Characterization of immune response in mice to    plasmid DNA encoding dog zona pellucida glycoprotein-3. Vaccine 21:1913-1923. (*in press last year, since published)

Reviews/Proceedings

1.     Kaur R and Vrati S (2003) Development of recombinant vaccine against Japanese encephalitis. J Neurovirol 9:421-431.

Patents

1.     Vrati S and Appaiahgari A (2003) Vaccine for Japanese encephalitis and its process thereof. Indian patent application # 1130/DDEL/2003.

2.      Vrati S (2003) Recombinant adenovirus-based Japanese encephalitis virus vaccine. Indian patent application # 1516/DEL/2003.