RU2812251C1 - Tetravalent vector vaccine and method of its use for prevention of dengue fever - Google Patents

Abstract

FIELD: biotechnology; genetic engineering; immunology.

SUBSTANCE: tetravalent vector vaccine has been obtained for the prevention of dengue fever, containing a recombinant vector based on a modified vaccinia virus strain Ankara (MVA) with an integrated nucleotide sequence encoding the d34 antigen (46.5 kDa), which consists of four DHI domains of the E-protein of dengue virus serotypes 1, 2, 3 and 4, connected by a flexible linker 2x(GGGGS).

EFFECT: method has been proposed for using a tetravalent vector vaccine to induce a specific immune response against four serotypes of the dengue virus, including administering the vaccine into the body of a mammal in an effective dosage.

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Description

Technical field

The invention relates to the field of biotechnology, genetic engineering and immunology, in particular to a quadrivalent vector vaccine for the prevention of dengue fever, as well as a method of its use.

State of the art

Dengue fever is a zooanthroponotic viral infection endemic to more than 100 countries in tropical and subtropical regions of the world (Africa, Eastern Mediterranean, North and South America, Southeast Asia, Western Pacific), but in recent years dengue fever has been identified in non-endemic areas. territories - in Russia, Germany, France, Croatia, England and other countries. According to the World Health Organization (WHO), about 390 million cases of the disease are registered annually, and about 4 billion people are at risk of infection. In 2019, WHO included dengue fever in the list of potential threats to humanity: from 2000 to 2019, the incidence of dengue fever in the world increased more than 10 times.

The causative agent of the disease is dengue virus (DENV), a single-stranded RNA virus of the Flaviviridae family. For humans, four serotypes of dengue virus (DENV-1-DENV-4) are of epidemiological significance. The dengue virus is transmitted to humans through the bites of Aedes mosquitoes (mainly A aegypti), whose range is expanding due to climate warming. Sources of infection can be infected people, monkeys, bats.

Most patients with dengue fever are asymptomatic or have mild symptoms; after the illness, serotype-specific immunity develops. However, in some cases, a severe course of the disease (hemorrhagic/shock form) may occur, which is characterized by multiple organ failure, tissue and capillary damage, plasma leakage and massive hemorrhages, with possible death. The risk of developing severe dengue fever increases when infected with a heterologous viral serotype - an antibody-dependent enhancement effect (ADE) is observed.

There are no specific treatments for dengue fever, so one of the priority technical challenges is the development of an effective preventive vaccine that induces a strong immune response against all four circulating dengue virus serotypes. Currently, only one tetravalent vaccine is licensed for the prevention of dengue fever - Dengvaxia (CYD-TDV, Sanofi Pasteur, France) - this is a live attenuated recombinant vaccine based on the vaccine strain 17D of the yellow fever virus (YFV), in which the prM and E protein genes are replaced homologous genes of each of the four dengue virus serotypes. The Dengvaxia vaccine is intended for three doses of immunization at an interval of 6 months.

However, in phase III clinical studies involving more than 35 thousand children aged 2-16 years from the Pacific region and Latin America, the effectiveness of the Dengvaxia vaccine was shown to differ depending on the serotype of the virus, as well as the age and serostatus of the patient. The protective effect of the vaccine was 54.7% against serotype 1, 43.0% against serotype 2, 71.6% against serotype 3 and 76.9% against serotype 4. In the group of children 2-9 years old, the effectiveness was lower than in the group children 9-16 years old (44.6 and 65.6%, respectively), and the overall effectiveness of the vaccine was only 60.3% [Hadinegoro S.R., Arredondo-Garcia J.L., Capeding M.R. et al. Efficacy and long-term safety of a dengue vaccine in regions of endemic disease // N. Engl. J. Med. 2015. V. 373. P. 1195-1206; Gubler D.J., Halstead S.B. Is Dengvaxia a useful vaccine for dengue endemic areas? // BMJ. 2019. V. 367. P. 15710]. In addition, a significant drawback of this technical solution was identified, namely the danger of vaccinating seronegative patients. It has been shown that during post-vaccination infection, such patients may develop antibody-dependent enhancement of infection (ADE), which leads to a more severe course of the disease, including death. Children are most susceptible to this phenomenon. Thus, in children 2-5 years of age, within three years after immunization, there was an 8-fold increase in cases of hospitalization with severe dengue fever [Shukla R., Ramasamy V., Shanmugam R.K. et al. Antibody-dependent ehancement: a challenge for developing a safe dengue vaccine // Front. Cell Infect. Microbiol. 2020. V. 10. P. 572681]. Therefore, before vaccination, a mandatory determination of serological status is required. Currently, the Dengvaxia vaccine is recommended only for seropositive patients - only in this case the vaccine is considered safe [Dengue vaccine: WHO position paper - September 2018 // Wkly Epidemiol. Rec. 2018. V. 93. P. 457-476; Sridhar S., Luedtke A., Langevin E. et al. Effect of dengue serostatus on dengue vaccine safety and efficacy //N. Engl. J. Med. 2018. V. 379. P. 327-340].

To prevent the development of ADE and severe complications, the search for new approaches to creating effective and safe preventive vaccines that ensure the development of a strong immune response against all four circulating dengue virus serotypes is still relevant.

Currently, several pharmaceutical companies and research teams are developing candidate tetravalent vaccines against dengue fever, and preclinical and clinical studies of their immunogenicity and safety are being conducted.

Known technical solutions are candidate live attenuated quadrivalent vaccines: 1) vaccine TV003/TV005 [Whitehead S.S. Development of TV003/TV005, a single dose, highly immunogenic live attenuated dengue vaccine; what makes this vaccine different from the Sanofi-Pasteur CYD™ vaccine? // Expert Rev. Vaccines. 2016. V. 15. P. 509-517], which is undergoing phase III clinical trials; 2) vaccine TAK-003 (Takeda Pharmaceutical Company) [Biswal S., Reynales H., Saez-Llorens X. et al. Efficacy of a tetravalent dengue vaccine in healthy children and adolescents // N. Engl. J. Med. 2019. V. 381. P. 2009-2019], which successfully completed phase III clinical trials and, under the name Qdenga, was approved by the European Medicines Agency (EMA) for use in persons over 4 years of age in accordance with national recommendations [Angelin M., Sjölin J ., Kahn F. et al. Qdenga® - a promising dengue fever vaccine; can it be recommended to non-immune travelers? // Travel Med. Infect. Dis. 2023. V. 54. P. 102598].

The antigen of choice in the design of tetravalent vaccines against dengue virus is the DIII domain of the surface E protein, which is involved in the recognition of receptors on target cells. Antibodies against the DIII domain of the E protein ensure the development of a pronounced serotype-specific immune response without the ADE effect during infection with heterologous dengue virus serotypes [Wahala W.M., Kraus A.A., Haymore L.B. et al. Dengue virus neutralization by human immune sera: role of envelope protein domain 111-reactive antibody // Virology. 2009. V. 392. P. 103-113].

Previously, several variants of vaccines based on the DIII domain were proposed, in which different platforms were used: 1) DNA vaccines [Slon Campos J.L., Poggianella M., Burrone O.R. Long-term stability of antibody responses identified by dengue virus envelope Dill-based DNA vaccines // J. Gen. Virol. 2018. V. 99. P. 1078-1085; Poggianella M., Slon Campos J.L., Chan K.R. et al. Dengue E protein domain Ill-based DNA immunization induces strong antibody responses to all four viral serotypes // PLoS Negl. Trop.Dis. 2015. V. 9. P. e0003947]; 2) recombinant proteins [Izquierdo A., Garcia A., Lazo L. et al. A tetravalent dengue vaccine containing a mix of domain III-P64k and domain 111-capsid proteins induces a protective response in mice //Arch. Virol. 2014. V. 159. P. 2597-2604]; 3) virus-like particles (VLP); 4) viral vectors (adenovirus, vesicular stomatitis virus, poxvirus, etc.).

A known technical solution is a vaccine based on the DSV4 immunogen expressed in Pichia pastoris [Ramasamy V., Arora U., Shukla R. et al. A tetravalent virus-like particle vaccine designed to display domain III of dengue envelope proteins induces multi-serotype neutralizing antibodies in mice and macaques which confer protection against antibody dependent enhancement in AG129 mice // PLoS Negl. Trop.Dis. 2018. V. 12. e0006191], which can be considered as a prototype. The DSV4 immunogen consists of the DIII domains of the E protein of four dengue virus serotypes and the hepatitis B surface antigen (S-antigen), connected by hexaglycol linkers, which, when co-expressed with the S-antigen, form mosaic virus-like particles (VLPs). In experiments on mice and macaques, the high immunogenicity of this vaccine was demonstrated - the formation of neutralizing antibodies against all four serotypes of the dengue virus was revealed, which minimizes the risk of an antibody-dependent increase in infection when infected with heterologous serotypes. However, it should be noted that the vaccine included an MPLA (monophosphoryl lipid A) adjuvant to further stimulate the immune response - this may indicate insufficient immunogenicity of the main component of the vaccine (VLP with DSV4 antigen) when used without the addition of an adjuvant.

One of the promising areas for developing vaccines against pathogens with intraspecific variability, such as dengue virus, is the creation of recombinant viral vaccines, in which the genetic material encoding the target antigen is delivered inside the host cell using a viral vector.

A well-known technical solution, which we have chosen as the closest prototype, is a vector vaccine based on the recombinant measles virus [Brandler S., Ruffie C, Najburg V. et al. Pediatric meats vaccine expressing a dengue tetravalent antigen detects neutralizing antibodies against all four dengue viruses. Vaccine. 2010. V. 28. P. 6730-6739]. The authors used an antigen consisting of a chain of DIII domains of surface protein E from four serotypes of dengue virus. The nucleotide sequence encoding this antigen was inserted into the genome of a weakened measles virus between the P and M genes. Three variants of the antigen were tested, the difference being the use of different linkers connecting the DIII domains to each other, and a variant without a linker. The authors assumed that linkers consisting of fragments of the M protein (1-40) from the dengue virus should increase the immunogenicity of the vaccine, but in experiments the version without a linker showed the best result. However, in this case, the titer of neutralizing antibodies was not high enough and amounted to no more than 1/20.

An attractive vector for creating vaccines is the modified vaccinia virus strain Ankara (MVA) due to its high immunogenicity and safety demonstrated in clinical studies [Orlova O.V., Glazkova D.V., Bogoslovskaya E.V. et al. Development of modified vaccinia virus ankara-based vaccines: advantages and applications // Vaccines. 2022. V. 10. P. 1516], as well as low cost and the ability to quickly produce the required number of vaccine doses for mass immunization, which can be extremely important for the active formation of collective immunity during outbreaks of dengue fever or a pandemic.

Thus, taking into account the annual increase in the prevalence of dengue fever, the increase in incidence and restrictions on the use of existing quadrivalent vaccines against dengue fever, we can conclude that it is urgent to develop new technical solutions for the prevention of this viral infection, ensuring the induction of an immune response against all four circulating serotypes of the virus dengue.

Disclosure of the invention

The required technical result is to develop a new effective and safe vaccine for the prevention of dengue fever.

This technical result is achieved by creating a quadrivalent vector vaccine for the prevention of dengue fever caused by the dengue virus of all four circulating serotypes (1, 2, 3 and 4). The claimed quadrivalent vector vaccine is a recombinant vector with the nucleotide sequence SEQ ID NO 1, derived from a modified vaccinia virus strain Ankara (MVA). The nucleotide sequence SEQ ID NO 2 was inserted into this vector, encoding the d34 antigen with the amino acid sequence SEQ ID NO 3. The antigen consists of four DIII domains of the surface E protein of dengue virus serotypes 1, 2, 3 and 4, connected by a flexible linker 2x (GGGGS).

The mechanism of action of the quadrivalent vector vaccine for the prevention of dengue fever is based on the induction of a serotype-specific immune response to all four circulating dengue virus serotypes. The effectiveness of the claimed quadrivalent vector vaccine is confirmed by the results of preclinical studies in vitro and in vivo [Mintaev R.R., Glazkova D.V., Orlova O.V. et al. Development of MVA-d34 tetravalent dengue vaccine: design and immunogenicity //Vaccines. 2023. V. 11. P. 831].

Experiments with patient blood sera demonstrated the antigen specificity of the target antigen d34 and the absence of its cross-reactive interactions with antibodies to closely related flaviviruses (tick-borne encephalitis and yellow fever viruses).

In mice immunized with the quadrivalent vector vaccine, a high titer of specific IgG was detected that interacted with antigens of all four serotypes of the dengue virus, which indicates the effective induction of the body's immune response to vaccination. A pronounced virus-neutralizing activity of blood sera from experimental mice at a dilution of 1:64 was also shown.

A method of using a quadrivalent vector vaccine for the prevention of dengue fever has also been proposed, which consists of introducing the vaccine into the body of mammals in an effective dosage.

Brief description of drawings

In fig. Figure 1 schematically shows the structure of the d34 antigen (A) and the location in the MVA genome of the 136/137 locus for inserting the expression cassette (B). Squares indicate homology arms (HA). Legend: Leader - anti-mouse Ig leader sequence, linker - flexible linker 2x(GGGGS). A light rectangle with a black rectangle is the left homological arm (Left HA), a black rectangle is a short left homological arm (Short HA), a purple rectangle is the right homological shoulder (Right HA). Gray rectangles are the p11 and mH5 promoters of the vaccinia virus, stimulating the expression of the RFP and d34 antigens, respectively. The vaccinia virus termination signal TTTTTNT (Term) was used to stop transcription.

In fig. Figure 2 shows a diagram of a shuttle vector with the included sequence encoding the d34 antigen. Legend: mouse Ig k leader - mouse Ig leader sequence; d3_1, d3_2, d3_3, d3_4 - DIII domains connected by the linker 2x(GGGGS) (nucleotide sequence encoding the d34 antigen), Left HA - left homological arm, short HA - short left homological arm, Right HA - right homological arm; p11 and mH5 are vaccinia virus promoters that stimulate the expression of RFP and d34 antigens, respectively. Term—vaccinia virus termination signal TTTTTNT (Term)—was used to stop transcription. Amp, ori, f1 ori - plasmid elements necessary for production of the plasmid in bacteria: Amp - resistance gene to the antibiotic ampicillin; ori, f1 ori - bacterial origins of plasmid replication.

In fig. Figure 3 shows an analysis of the expression of the d34 antigen in BHK-21 cells after treatment with the wild-type (wt) MVA vector or a recombinant vector by Western blotting using antibodies to 6xHis. M - marker of molecular weight of proteins. K - uninfected BHK-21 cells. The molecular weight of the antigen is 46.5 kDa. M - marker ECT™ Rainbow™.

In fig. Figure 4 presents an analysis of the interaction of the d34 antigen with patient blood sera using Western blotting. HEK293FT cells were transduced with wild-type (wt) MVA vector or a recombinant vector with an inserted nucleotide sequence encoding the target antigen (d34). Hybridization was carried out with antibodies to 6xHis (A) or with sera obtained from the following patients: a patient who had dengue fever (without specifying the serotype) (serum 1) (B), immunized with a vaccine against tick-borne encephalitis (serum 2) (C), immunized vaccine against tick-borne encephalitis and yellow fever (serum 3) (D), immunized with vaccine against yellow fever (serum 4) (E). M - molecular weight marker for PageRuler™ Plus Prestained Protein. K - control HEK293FT cells (not infected with virus). The molecular weight of the d34 antigen is 46.5 kDa.

Figure 5 shows the antibody titer in the blood sera of mice immunized with quadrivalent vector vaccine (n=5) or MVA vector (wt) without insert (n=5), determined by enzyme-linked immunosorbent assay (ELISA). The average value is shown with a thick line. * Statistically significant difference between groups (p<0.05, (Mann-Whitney /-test).

Table 1 presents the results of a study of the reactivity of the d34 antigen with the blood sera of people who have had dengue fever, as well as those vaccinated against tick-borne encephalitis and yellow fever.

Table 2 shows the titer of antibodies to four serotypes of dengue virus in the blood sera of mice immunized with a quadrivalent vector vaccine.

Table 3 presents the results of a study of the virus neutralizing activity of blood sera of mice immunized with a quadrivalent vector vaccine.

Carrying out the invention

To create a safe and effective prophylactic vaccine that induces a specific immune response against four circulating dengue virus serotypes, a modified vaccinia virus Ankara (MVA) vector was selected. The MVA vector is characterized by a high safety profile, confirmed both in clinical studies and during mass vaccination against smallpox, and has a number of advantages: 1) provides high vaccination efficiency due to the high level of expression of the target antigen and activation of both parts of the immune system - induces pronounced humoral and T-cell immune response to the pathogen antigen; 2) is highly immunogenic even when vaccinating immunocompromised people, elderly people and HIV-infected people; 3) does not cause the development of local reactions after vaccination [Orlova O.V., Glazkova D.V., Bogoslovskaya E.V. et al. Development of modified vaccinia virus ankara-based vaccines: advantages and applications // Vaccines. 2022. V. 10. P. 1516]. The MVA vector is characterized by high capacity - it can include up to 25 kb. foreign nucleotide sequences, which allows the insertion of rather long fragments encoding several antigens of the disease pathogen into it, and the expression of antigens with the correct post-translational modification.

To generate an antibody response, the DIII domain of the surface E protein was selected. It has been shown that antibodies against the DIII domain of the E protein, which is involved in the recognition of receptors on target cells, are serotype-specific, exhibit protection against homologous infection and do not cause an increase in disease symptoms during heterologous infection.

Since dengue fever is caused by four serotypes of the virus, when developing the vaccine target antigen, we combined four variants of the DIII domain of the E protein belonging to the four serotypes of the dengue virus into one polypeptide using linkers (Example 1, Fig. 1A). In contrast to previously described approaches, in this invention, a flexible linker 2x(GGGGS) is used to connect the domains, which, in our opinion, should increase the spatial mobility of the DIII domains of the E protein and improve the stability of their tertiary structure. Combining the domains into a single d34 antigen (with the sequence SEQ ID NO 3) allows you to stimulate an immune response against all four circulating dengue virus serotypes, as well as achieve a high level of expression of the target d34 antigen (single domains are known to have a lower level of expression in cells) .

To deliver the antigen, we used a vector based on a modified vaccinia virus strain Ankara (MVA). The nucleotide sequence encoding the target antigen d34 (Example 1, Fig. 1A) was inserted into the shuttle vector pShuttle-136/137, resulting in the shuttle vector pShuttle-136/137-d34 (Fig. 2). BHK21 cells were infected with the MVA viral vector (wt) and then transfected with the shuttle vector pShuttle-136/137-d34. After screening and selection procedures using a fluorescent marker and the use of real-time PCR, a purified recombinant vector with an inserted sequence encoding the d34 antigen, which is the immunogenic component of a quadrivalent vector vaccine, was obtained.

The expression of the d34 antigen was assessed by Western blotting using antibodies to 6xHis in BHK-21 cells transduced with the recombinant vector (example 3, Fig. 3). It has been shown that the target antigen d34 is produced in cells and secreted into the medium.

To assess the specificity of the d34 antigen, an analysis of its interaction with the blood sera of patients who had suffered from dengue fever or were vaccinated against tick-borne encephalitis and yellow fever was carried out using ELISA methods (example 4, table 1) and Western blotting (example 5, Fig. 4 ). It has been shown that the d34 antigen binds to IgG from the blood sera of people who have had dengue fever and does not interact with IgG antibodies against closely related viruses from the flavivirus family. The results obtained allow us to conclude about the antigenic specificity of the target d34 antigen and the absence of its cross-reactive interactions with antibodies to related flaviviruses (tick-borne encephalitis and yellow fever viruses).

The immunogenic characteristics of the quadrivalent vector vaccine were assessed using an animal model - mice immunized twice with the experimental quadrivalent vector vaccine with an interval between injections of 21 days. On the 21st day after the second immunization, blood serum samples were taken from mice from each group and the antibody titer was determined by ELISA (example 6, Fig. 5). In all experimental mice, high titer antibodies to the d34 antigen were detected in the blood sera.

A more detailed study of the immune response using indirect ELISA (example 7, Table 2) showed the formation of specific IgG in experimental animals that interact with each of the 4 serotypes of the dengue virus in limiting dilutions from 1:400 to 1:800, which indicates about the effective induction of an immune response to all four serotypes of dengue virus.

In addition, it was found that blood sera obtained from mice immunized with a quadrivalent vector vaccine have virus-neutralizing activity against the dengue virus (using the example of dengue virus serotype 2) at a dilution of 1:64 (example 8, Table 3), which is significant exceeds the virus-neutralizing activity of the prototype, in which the titer of neutralizing antibodies was no more than 1: 20 [Brandler S., Ruffle C, Najburg V., Frenkiel M.R., Bedouelle H., Despres P., Tangy F. Pediatric measles vaccine expressing a dengue tetravalent antigen detects neutralizing antibodies against all four dengue viruses. Vaccine. 2010. V. 28. P. 6730-6739].

The quadrivalent vector vaccine contains an immunologically effective amount of a recombinant vector dispersed in a pharmaceutically acceptable carrier. The term "effective dosage" as used herein means an amount of recombinant vector that, when administered to a human or mammalian animal, induces a significant humoral response against all four types of dengue virus. A specialist in the field can select such an immunologically effective dosage of a quadrivalent vector vaccine, guided by the above description of the invention, as well as theoretical knowledge in the field and his practical experience, without conducting additional experiments.

The immunogenic component of a quadrivalent vector vaccine - a recombinant vector with an integrated nucleotide sequence encoding the target d34 antigen - is dispersed in a pharmaceutically acceptable carrier.

Suitable pharmaceutically acceptable carriers well known to those skilled in the art include, but are not limited to, saline, buffer solutions (e.g., phosphate buffer), and may also contain excipients including, but not limited to, diluents, stabilizers, preservatives, pH-regulating buffers, etc.

Said quadrivalent vector vaccine may additionally contain one or more adjuvants from among the pharmaceutically acceptable adjuvants used in the art to create vector vaccines, or may not contain additional adjuvants.

The quadrivalent vector vaccine may be in a form suitable for intramuscular, intranasal, intraperitoneal, subcutaneous, or intradermal administration to a patient, as well as in a form suitable for administration by any other desired route.

To deliver the target d34 antigen, other vectors suitable for the expression of recombinant genes can be used, which may be necessary if the patient has pre-existing immunity to the smallpox virus (which was formed, for example, as a result of vaccination with the smallpox vaccine), since the neutralizing agents formed in this case antibodies to the vector can affect the formation of an immune response to the antigen, reducing the effectiveness of vaccination. Non-limiting examples of such vectors are adenovirus-based vectors [Mercado N.B., Zahn R. et al. //Nature. 2020. V. 586. P. 583-588; Logunov D.Y., Dolzhikova I.V. et al. // Lancet. 2021. V. 397. P. 671-681], vesicular stomatitis virus [Yahalom-Ronen Y., Tamir H. et al. //Nat. Commun. 2020. V. 11. P. 6402], poxvirus, adeno-associated virus, cytomegalovirus, etc.

We will consider the implementation of the invention using typical examples, which are given solely for the purpose of illustration and are not intended to limit the scope of this invention in any way. The given examples of implementation of the invention are not exhaustive. Possible other examples of implementation corresponding to the scope of the patent claims.

Example 1. Preparation of an expression cassette encoding a fusion recombinant d34 antigen and a plasmid shuttle vector

The d34 antigen sequence included four DIII domains of the dengue virus E protein (one for each of the four dengue virus serotypes) linked by a 2x(GGGGS) flexible liqueur (Fig. 1). To compile the antigen coding sequence, genome records of dengue virus serotypes 1 (record ID NP 05943), 2 (record ID NP 056776), 3 (record ID YP 001621843), 4 (record ID NP 073286) were used. At the 5' end of the open reading frame of the gene, a sequence encoding a mouse leader Ig was inserted to direct the protein along the secretory pathway [rm6], and at the 3' end there was a sequence encoding a 6xHis tag necessary for determining antigen expression.

The synthesis of the expression cassette encoding the d34 antigen and the nucleotide sequence of the functional part of the plasmid shuttle vector was carried out by TOP Gene Technologies, Inc. (Quebec, Canada). The following plasmids were obtained: pShuttlel, pShuttle2 and pUC16-d34.

Sequences from plasmids pShuttlel and pShuttle2 were cloned into plasmid pGEM-Easy (Promega, USA) to obtain the plasmid shuttle vector pShuttle-136/137. The plasmid shuttle vector pShuttle-136/137 contained the red fluorescent marker gene (rfp) under the control of the p11 MVA promoter, homologous regions of the 136/137 MVA locus necessary for insertion of the sequence into this locus, the mH5 MVA promoter, transcription termination signals, and sites Nhel and KpnI restriction.

The synthesized nucleotide sequence encoding the d34 antigen (expression cassette) was inserted at the NheI and KpnI sites into the polylinker of the plasmid shuttle vector, resulting in the plasmid pShuttle-136/137-d34 with an integrated cassette for the expression of the d34 antigen (Fig. 2). Plasmid DNA was produced in Escherichia coli cells (strain DH5a) and isolated using Plasmid Mini Kit (Qiagen, Germany) or Plasmid Maxi Kit (Qiagen, Germany) under sterile conditions. The correctness of the construct was confirmed by Sanger sequencing.

Example 2. Preparation of a recombinant vector with an integrated nucleotide sequence encoding the d34 antigen

The production and development of the recombinant vector (rMVA) was carried out in BHK-21 cells in DMEM medium containing 2% FBS, 4 mM L-glutamine, 1 mM sodium pyruvate, 10 mM HEPES. BHK-21 cells were infected with the wild-type (wt) MVA viral vector, after 90 min the growth medium was replaced with fresh one and transfected with the plasmid shuttle vector pShuttle-136/137-d34. After 48 h, the recombinant vector mixture was collected and seeded to form individual plaques. Identification of plaques was carried out using an inverted microscope in the UV spectrum. Fluorescent plaques that expressed RFP protein were marked and transferred to a tube containing 500 μl of fresh medium. The procedure was repeated until a recombinant vector free of wild-type MVA was obtained. The absence of wild-type MVA contamination in the recombinant vector (rMVA) was confirmed using real-time PCR by assessing threshold cycles (Ct) in the rMVA and total MVA amplification reactions, selecting samples with a minimum ΔCt value. After 2-3 rounds of purification, samples were obtained in which the ΔCt fell in the range from 0 to 2.

Next, testing was carried out using classical PCR, amplifying the insertion region of the shuttle vector into the MVA genome using primers: 136-137 reg forw - CTTACTCAAATGAACGCCAGAG and 137_reg_rev - CATGATTACTAACGTGGGTCA. Based on the PCR results, clones were selected whose electrophoretic mobility was at the appropriate level (5022 bp with rfp, 3903 bp without rfp) without an additional 1256 bp band, the presence of which indicates the presence of wild-type MVA admixture.

Three rounds of purification resulted in six clones being selected. rMVA DNA was isolated using the AmpliTest RIBO-prep kit (FGBU TsSP FMBA of Russia) and used as a template for PCR on the region of insertion of the shuttle vector into the MVA genome (primers 136-137 reg forw and 137_reg_rev). Two clones were selected for further development. Each subsequent amplification of the recombinant virus was assessed using PCR (primers 136-137 reg forw and 137_reg_rev).

The average titer of the purified recombinant vector (without wild-type MVA impurities) was 5×10 ⁸ PFU/ml.

Thus, an immunogenic component of a quadrivalent vector vaccine for the prevention of dengue fever has been obtained - a recombinant vector with an integrated nucleotide sequence encoding the d34 antigen.

Example 3. Analysis of d34 antigen expression by Western blotting in BHK-21 cells transduced with a recombinant vector

Expression of the d34 antigen in BHK-21 cells was assessed by Western blot analysis using antibodies to 6xHis.

BHK-21 cells were transduced with a recombinant vector expressing the d34 antigen or an MVA (wt) vector without insert at an MOI of 2 for 90 min. Cells without vector addition were used as a negative control. Then the vector medium was replaced with serum-free OptiMEM. After 24 h, cells were harvested, washed three times with PBS, resuspended in RIPA lysis buffer containing 1% NP-40, and disrupted by sonication. Cellular debris was removed by centrifugation (15,000 g, 15 min, 4°C). The culture supernatant was collected separately and concentrated using an Amicon Ultra-0.5 10 kDa centrifugal concentrator. Protein marker ECL Rainbow (Cytiva, USA) or PageRuler Plus (Thermo Scientific, USA) and samples were analyzed in 12% SDS-PaGE.

After electrophoresis under reducing conditions, the proteins were transferred to an Immun-Blot LF PVDF membrane (Bio-Rad, USA). Membranes were blocked in PBS containing 5% nonfat dry milk ECL Prime Blocking Agent (PRN418, Amersham, USA) and 0.1% Tween 20, then incubated overnight at 4°C with antibodies to the 6xHis tag (HIS.H8- HRP, Invitrogen, USA) at a dilution of 1: 20000. Then the membranes were washed 3 times in PBS with 0.1% Tween 20. Protein complexes were determined using the Clarity Western ECL Substrate reagent (Bio-Rad, USA) in accordance with the manufacturer’s recommendations and visualized using X-ray film (Fui Film, Japan).

The results of Western blotting are presented in Fig. 3. One band was detected, which corresponded to the expected size of the target d34 antigen (46.5 kDa), and the d34 antigen was detected both in cells and in the growth medium, where its amount was larger.

Thus, it was shown that the target antigen d34 is produced in BHK-21 cells transduced with the recombinant vector and is successfully secreted into the medium.

Example 4. Evaluation of the antigen specificity of the target antigen d34 by indirect ELISA using patient blood sera

Antigen specificity was studied by indirect ELISA using blood sera that were obtained from patients who had dengue fever (without specifying the serotype). This method is aimed at detecting antibodies in blood serum that can interact with the d34 antigen sorbed on a substrate. To check the absence of cross-reactivity of the antigen with antibodies against related viruses from the Flaviviridae family, blood sera obtained from patients immunized with vaccines produced by the Federal State Budgetary Institution "FNTsIRIP im. Chumakov" RAS (Russia) against tick-borne encephalitis or against yellow fever. Previously, the presence/absence of specific IgG to dengue viruses (DENV), tick-borne encephalitis (TBEV) and yellow fever (YFV) was confirmed by ELISA using kits: DENV - Anti-Dengue Virus ELISA (IgG) (EI 226b-9601G, Euroimmun AG , Germany); TBEV - VectoVKE-IgG (JSC “Vector-Best”, Russia); YFV - Qualitative Human Yellow Fever Virus Antibody IgG (YFV-IgG) ELISA Kit (MyBioSource Inc., USA).

The d34 antigen required for ELISA was obtained as follows. HEK293FT cells were transduced with the recombinant vector at a multiplicity of infection MOI of 4 for 90 min. Then the medium with the virus was replaced with serum-free OptiMEM. After 48 h, the culture supernatant was collected, clarified, and the protease inhibitor PMSF was added to a final concentration of 1 mM.

Primary purification of the target antigen d34 from the supernatant was carried out by chromatography on Ni-activated Sepharose (HisTrap, GE, USA). Then the resulting d34 antigen was further purified on the SPS-Bio-SP sorbent (Technosorbent, Russia). The fraction of the eluted d34 antigen was desalted to 1500 mM NaCl and concentrated on an Amicon Ultra-0.5 column with a pore diameter of 10 kDa (Millipore, USA).

ELISA was performed according to standard methods. The d34 antigen was adsorbed at a concentration of 2 μg/ml onto the wells of a polystyrene plate (#2592; Coming, USA). The washing solution was 0.05 M phosphate buffered saline with the addition of 0.02% Tween-20, pH 7.2 (PBS-T). After removing the blocking solution (1% sodium caseinate solution), the plate was dried in a laminar flow hood for three hours. Next, patient blood sera, previously diluted 100 times in PBS-T, were added to the wells of the plate with the sorbed d34 antigen and incubated for 1 hour at 37°C. Then, a standard procedure was carried out to detect the amount of antibodies bound to the d34 antigen using a peroxidase anti-species conjugate to Human IgG (#M-2101-HRP; Bioservice). Optical density was measured spectrophotometrically at a wavelength of 450 nm (reference wavelength - 640 nm) on a Multiskan FC instrument (Thermo Scientific, USA). The criterion for a positive result was considered to be OD ₄₅₀ >0.35.

The results of assessing antigen specificity using the indirect ELISA method are presented in Table 1. It is shown that the d34 antigen binds to IgG antibodies from the blood sera of people who have had dengue fever and does not interact with antibodies to closely related viruses of the flavivirus family - tick-borne encephalitis and yellow fever viruses .

The results obtained allow us to conclude that the target antigen d34, consisting of four DIII domains of the E protein of dengue virus of different serotypes, has antigen specificity.

Example 5. Assessment of the specificity of recognition of the d34 antigen by antibodies from patient blood sera using Western blotting

HEK293FT cells were transduced with a recombinant vector expressing the d34 antigen or an MVA (wt) vector without insert at an MOI of 2 for 90 min. Cells without vector addition were used as a negative control. Then the vector medium was replaced with serum-free OptiMEM. After 24 h, cells were collected, washed three times with PBS, resuspended in RIPA lysis buffer containing 1% NP-40, and disrupted by sonication. Cellular debris was removed by centrifugation (15,000 g, 15 min, 4°C). The culture supernatant was collected separately and concentrated using an Amicon Ultra-0.5 10 kDa centrifugal concentrator. Protein marker ECL Rainbow (Cytiva, USA) or PageRuler Plus (Thermo Scientific, USA) and samples were analyzed in 12% SDS-PAGE.

After electrophoresis under reducing conditions, the proteins were transferred to an Immun-Blot LF PVDF membrane (Bio-Rad, USA). Membranes were blocked in PBS containing 5% nonfat dry milk ECL Prime Blocking Agent (PRN418, Amersham, USA) and 0.1% Tween 20, then incubated overnight at 4°C with antibodies to the 6xHis tag (HIS.H8- HRP, Invitrogen, USA) at a dilution of 1: 20,000, or with the blood serum of a patient who has had dengue fever (without specifying the serotype) (serum 1, 1: 1000); or with the blood serum of a patient immunized with a vaccine against tick-borne encephalitis (serum 2, 1: 1000), or a vaccine against tick-borne encephalitis and yellow fever (serum 3, 1:1000), or a vaccine against yellow fever (serum 4, 1:1000) . Membranes that were hybridized with patient sera were additionally incubated with secondary antibodies Goat anti-Human IgG (H+L) HRP 1:30,000 (#31410, Invitrogen, USA) for 1 hour at room temperature. The membranes were then washed 3 times in PBS with 0.1% Tween 20. Protein complexes were determined using Clarity Western ECL Substrate reagent (Bio-Rad, USA) according to the manufacturer's recommendations and visualized using X-ray film (Fuji Film, Japan).

The results of Western blotting with patient sera are presented in Fig. 4. It has been shown that the d34 antigen binds to IgG from the blood sera of people who have had dengue fever and does not interact with IgG against closely related viruses from the flavivirus family. The results obtained allow us to conclude that the d34 antigen is antigenic specific and there is no cross-reactivity with antibodies against related flaviviruses - tick-borne encephalitis virus and yellow fever.

Example 6: In Vivo Assessment of Vaccine Immunogenicity

The experiment was carried out on sexually mature female mice (age 6-8 weeks, weight 18-20 g), first-generation hybrids of the C57 and CBA lines (Nursery of the Scientific Center for Biomedical Technologies of the Russian Academy of Sciences, Russia). Animals were immunized intramuscularly with 0.2 ml (5 × 10 ⁷ infectious particles) of the experimental quadrivalent vector vaccine or 0.2 ml of the control MVA vector (wt) without insert. Immunization was carried out twice, the interval between injections was 21 days. On the 21st day after the second immunization, blood serum samples were taken from mice from each group to determine the antibody titer using ELISA according to the standard method (see example 4); a peroxidase anti-species conjugate to IgG was used to detect the amount of antibodies bound to the d34 antigen mice (#55554, MP Biomedicals).

To determine the antibody titer, serum dilutions from 100 to 2700 were used, with a step of 3. The criterion for a positive result was OD ₄₅₀ >0.35.

The results of the experiment are presented in Fig. 5. Antibody titer values of 1: 2700 were obtained for one serum, 1: 900 for two and 1: 100 for two. At the same time, in all sera obtained from control group mice at a dilution of 1:100, the OD450 value did not exceed 0.25.

Thus, all mice (n=5) immunized with the quadrivalent vector vaccine showed an immune response to the d34 antigen.

Example 7. Immune response to different dengue virus serotypes in mice immunized with a quadrivalent vector vaccine

Antibodies to four serotypes of dengue virus from polyclonal rabbit serum (#M-2101-HRP; Bioservice, Russia) were adsorbed according to a standard protocol at a concentration of 10 μg/ml per wells of a polystyrene plate (#2592; Coming, USA). After this, antigens of each dengue virus serotype were added to the wells of the plate in a volume of 100 μl with a concentration of 10 μg/ml. Antigens of dengue serotypes were brain extracts of 4-day-old mouse sucklings infected intracranially with dengue viruses of different serotypes. Brain extracts were inactivated with β-propiolactone and clarified by centrifuge. The same volume of brain extract from mice not infected with the dengue virus was added to the control wells in a similar manner. The plate was incubated at 37°C for 60 min. After antigen sorption, the plate was washed 3 times. When performing ELISA, dilutions of mouse blood sera were pre-prepared with a dilution step of 2 from 1: 100 to 1: 3200 in PBS-T buffer. Serum dilutions were added to the wells of the plate in triplicate for each serotype and control and incubated for 60 min at 37°C. Then the wells were washed three times with PBS-T, peroxidase conjugate of goat antibodies to mouse IgG (#55554; MP Biomedicals) was added and incubated at + 37°C for 60 minutes.

When assessing the results, the following criteria were used: OD in control wells with brain antigen from an uninfected mouse should not exceed 0.2 FU. Optical density was measured spectrophotometrically at a wavelength of 450 nm (reference wavelength 640 nm) on a LisaScan instrument (Erba Mannheim). The criterion for a positive result in wells with a specific dengue antigen was an OD ₄₅₀ value of >0.35.

The titer of antibodies to different serotypes of dengue virus in the blood sera of mice immunized with the quadrivalent vector vaccine and the control MVA vector without insert (wt) is presented in Table 2. It is shown that after double immunization with the quadrivalent vector vaccine, specific IgG was formed in mice that interacted with the antigens of all four serotypes of dengue virus, which indicates the effective induction of an immune response to all four serotypes of dengue virus in experimental mice.

Example 8. Virus neutralizing activity of blood sera of mice immunized with a quadrivalent vector vaccine

The virus-neutralizing activity of the blood sera of mice immunized with a quadrivalent viral vaccine against dengue virus serotype 2 was studied in a C6/36 cell culture (Aedes albopictus mosquito cells).

The sera were inactivated at 56°C for 30 min. Serum dilutions were prepared from 1: 8 to 1: 512 with a step of 2 on Leibowitz's medium. 100 μl of each dilution was mixed with 100 μl (50 TCID ₅₀ ) of dengue virus serotype 2 and incubated for 1 h at 37°C. After this, the mixture was transferred to a monolayer of C6/36 cells (Aedes albopictus, ATCC CRL-1660) into a well of a 24-well plate and incubated at 32°C and 5% CO ₂ for 5 days. Monitoring of monolayer disruption under the influence of the virus (cytopathic effect) was carried out using a microscope. On the 5th day, the culture liquid was collected and the presence of the virus in the medium was determined using the AmpliSense Dengue-Virus-FL kit (N-2391-1, Central Research Institute of Epidemiology of Rospotrebnadzor) by RT-PCR.

Pure virus and virus mixed with blood serum from an intact mouse were used as a positive control for viral infection.

The results of the study of the virus neutralizing activity of the blood sera of mice immunized with the quadrivalent recombinant vaccine are shown in Table 3.

When infected with the virus without the addition of serum, monolayer disruption exceeding 75% was observed. The serum obtained from intact animals also did not have virus-neutralizing activity - the cytopathic effect was more than 75%, and the PCR results were positive. On the contrary, sera from mice immunized with a quadrivalent vector vaccine had virus-neutralizing activity and had a virus-neutralizing effect up to a dilution of 1:64 - in this serum dilution there was no cytopathic effect of the dengue virus serotype 2 on cells, and the PCR results were negative.

Thus, the technical task set, namely the creation of an effective and safe vaccine that ensures the specific induction of an immune response against the dengue virus of four serotypes, has been achieved, as confirmed by the examples given.

The presented examples confirm the effectiveness of the claimed quadrivalent vector vaccine, as well as industrial applicability.

List of sequences

SEQ ID 1

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tcgtaaaaaagttttatctctttctctcttcgatggtctcacaaaaatattaaacctctttctgatggagtcgtaaaaaagttttatctctttctccttcgatggtctcacaaaaatattaaacctctttctgatggagtcgtaaaaaagttttatctctttctccttcgatggtctcacaaaaatattaaacctctttctgatggagtcgtaaaaaagttttatctctttctccttcgatggtctcacaaaaatattaaacctctttctgatggagtcgtaaaaaagttttatctctttctctcttcgatggtctcacaaaaatattaaacctctttctgatggagtcgtaaaaaagttttatctctttctctcttcgatggtctcacaaaaatattaaacctctttctgatggagtcgtaaaaaagttttatctctttctccttcgatggtctcacaaaaatattaaacctctttctgatggtctctataaagcgattgatttttcttaccctctagagtttcctacggtcgttggtcacacatttttttctagacactaaataaata

SEQ ID 2

atggaaaccgacacactgctgctgtgggtcctcctgctctgggtgcctggctccacaggacatatgagctacgtgatgtgtacaggcagctttaaacttgagaaggaagtggccgagactcagcacggcacggtgctcgtgcaagtgaagtacgaagggaccgacgcaccatgtaaaattccgttttcttcccaagacgaaa agggtgtcactcagaacggcagacttatcactgccaatccgatagtcacagataaggaaaaaccagtcaatatcgaggccgaaccgccttttggagaatcatacatagtcgtcggcgccggtgagaaggccctgaagctgtcatggtttaagaagggctctcatatgaagcttggtggtggaggctctggaggaggcggcagctactccatgtgtacaggta agttcaaggtggtcaaagagatagcagaaacccagcacggaaccatcgtgattcgcgtgcagtacgaaggcgacgggagtccttgtaaaattcctttcgaaataatggatctggaaaaaaggcatgtgctgggtcgcctcatcaccgtgaatcctatcgttaccgaaaaagactctcccgtcaacatcgaagcaga accacctttcggcgatagttacattattatcggcgtggaaccaggccaactgaagctcaattggtttaaaaaagggtccaagctttctagaggtggtggagggtccggaggtggcggctcctacgccatgtgtacgaacacatttgtacttaaaaaagaagtgtccgaaactcagcacgggacaatcttgatcaaagtagaatataagggtgagg acgctccttgcaaaatacccttttccacagaggacgggcaagggaaggcccataatggccgcctgattaccgctaacccagtcgtcacaaagaaggaggagcccgtgaacattgaggctgagcctcccttcggagagtcaaacatcgtgattgggatcggcgataacgctcttaaaatcaactggtacaagaagggttcctctagacggccggga ggtggcggaagcggaggcggcggcagttatactatgtgctccggcaagttcagcatcgacaaagagatggcggagacacagcacgggacgaccgtggtcaaagtgaagtatgaaggcgcgggcgcaccctgtaaggtgccgattgagatccgcgacgtcaacaaggaaaaagtcgtgggccgcattatttcctcaac gccactggctgagaacaccaattctctgtgacaaacatcgaactcgaacctccttttggagacagttatatagtgatcggtgtgggaaattccgccctgactctgcattggtttcggaagggttctcggccgctcgagcaccatcaccatcaccac

SEQ ID 3

METDTLLLWVLLLWVPGSTGHMSYVMCTGSFKLEKEVAETQHGTVLVQVKYEGTDAPCKIPFSSQDEKGVTQNGRLITANNPIVTDKEKPVNIEAEPPFGESYIVVGAGEKALKLSWFKKGSHMKLGGGGSGGGGSYSMCTGKFKVVKEIAETQHGTIVIRVQYEGDGSPCKIPFEIMDLEKRHVLGRLITV NPIVTEKDSPVNIEAEPPFGDSYIIIGVEPGQLKLNWFKKGSKLSRGGGGSGGGGSYAMCTNTFVLKKEVSETQHGTILIKVEYKGEDAPCKIPFSTEDGQGKAHNGRLITANPVVTKKEEPVNIEAEPPFGESNIVIGIGDNALKINWYKKGSSRRPGGGGSGGGGSYTMCSGKFSIDKEMAETQHGTTVVKVKYEGAGAP CKVPIEIRDVNKEKVVGRIISSTPLAENTNSVTNIELEPPFGDSYIVIGVGNSALTLHWFRKGSRPLEHHHHHH

Claims (2)

1. Quadrivalent vector vaccine for the prevention of dengue fever, containing a recombinant vector with the nucleotide sequence SEQ ID NO 1 based on a modified vaccinia virus strain Ankara (MVA) with an integrated nucleotide sequence SEQ ID NO 2 encoding the d34 antigen (46.5 kDa) with amino acid sequence of SEQ ID NO 3, which consists of four DIII domains of the E protein of dengue virus serotypes 1, 2, 3 and 4, connected by a flexible linker 2x(GGGGS). 2. The use of a quadrivalent vector vaccine according to claim 1 for inducing a specific immune response against dengue virus serotypes 1, 2, 3 and 4, including the introduction of the vaccine into the body of a mammal in an effective dosage.