Citation: Ávalos, I.; Lao, T.;
Rodríguez, E.M.; Zamora, Y.;
Rodríguez, A.; Ramón, A.; Lemos, G.;
Cabrales, A.; Bequet-Romero, M.;
Casillas, D.; et al. Chimeric Antigen
by the Fusion of SARS-CoV-2
Receptor Binding Domain with the
Extracellular Domain of Human
CD154: A Promising Improved
Vaccine Candidate. Vaccines 2022, 10,
897. https://doi.org/10.3390/
vaccines10060897
Academic Editor: François Meurens
Received: 16 March 2022
Accepted: 27 May 2022
Published: 3 June 2022
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Article
Chimeric Antigen by the Fusion of SARS-CoV-2 Receptor
Binding Domain with the Extracellular Domain of Human
CD154: A Promising Improved Vaccine Candidate
Ileanet Ávalos, Thailin Lao , Elsa María Rodríguez, Yasser Zamora, Alianet Rodríguez, Ailyn Ramón,
Gilda Lemos , Ania Cabrales, Monica Bequet-Romero , Dionne Casillas, Ivan Andújar , Luis Ariel Espinosa,
Luis Javier González, Yanitza Alvarez, Yamila Carpio * and Mario Pablo Estrada *
Center for Genetic Engineering and Biotechnology, CIGB, Ave. 31 E/158 y 190, Havana 10600, Cuba;
ileanet.avalos@cigb.edu.cu (I.Á.); thailin.lao@cigb.edu.cu (T.L.); elsa.rodriguez@cigb.edu.cu (E.M.R.);
yasser.zamora@cigb.edu.cu (Y.Z.); alianet.rodriguez@cigb.edu.cu (A.R.); ailyn.ramon@cigb.edu.cn (A.R.);
gilda.lemos@cigb.edu.cu (G.L.); ania.cabrales@cigb.edu.cu (A.C.); monica.bequet@cigb.edu.cu (M.B.-R.);
dionne.casillas@cigb.edu.cu (D.C.); ivan.andujar@cigb.edu.cu (I.A.); luis.espinosa@cigb.edu.cu (L.A.E.);
luis.javier@cigb.edu.cu (L.J.G.); yanitza.alvarez@cigb.edu.cu (Y.A.)
* Correspondence: yamila.carpio@cigb.edu.cu (Y.C.); mario.pablo@cigb.edu.cu(M.P.E.)
Abstract: COVID-19 is a respiratory viral disease caused by a new coronavirus called SARS-CoV-2.
This disease has spread rapidly worldwide with a high rate of morbidity and mortality. The receptor-
binding domain (RBD) of protein spike (S) mediates the attachment of the virus to the host’s cellular
receptor. The RBD domain constitutes a very attractive target for subunit vaccine development due
to its ability to induce a neutralizing antibody response against the virus. With the aim of boosting
the immunogenicity of RBD, it was fused to the extracellular domain of CD154, an immune system
modulator molecule. To obtain the chimeric protein, stable transduction of HEK-293 was carried out
with recombinant lentivirus and polyclonal populations and cell clones were obtained. RBD-CD was
purified from culture supernatant and further characterized by several techniques. RBD-CD immuno-
genicity evaluated in mice and non-human primates (NHP) indicated that recombinant protein was
able to induce a specific and high IgG response after two doses. NHP sera also neutralize SARS-CoV-2
infection of Vero E6 cells. RBD-CD could improve the current vaccines against COVID-19, based in
the enhancement of the host humoral and cellular response. Further experiments are necessary to
confirm the utility of RBD-CD as a prophylactic vaccine and/or booster purpose.
Keywords: SARS-CoV-2; RBD; HEK-293; lentivirus; vaccine
1. Introduction
The coronavirus disease (COVID-19) is a respiratory viral disease caused by a new
coronavirus called Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2). This
virus has become a severe threat to global health due to its rapid expansion around the
world with a high rate of morbidity and mortality [1]. SARS-CoV-2 belongs to the beta-
coronavirus genus of the family Coronaviridae. The Coronaviridae family comprises 4 genera:
alpha-coronavirus, beta-coronavirus, gamma-coronavirus, and delta-coronavirus. Coronaviruses
are enveloped viruses with a single-stranded, positive-sense RNA genome of 29–30 kb
in size. These viruses infect several animal species including humans [2]. The SARS-
CoV-2 genome encodes four major structural proteins: spike (S), membrane (M), envelope
(E), and nucleocapsid (N). The S protein has become the main therapeutic target due
to its critical role in viral attachment, entry, and fusion. This protein has two major
subunits: the S1 and S2. The S1 subunit contains a receptor-binding domain (RBD) of
25 kDa approximately, which is responsible for the direct binding with the angiotensin-
converting enzyme 2 (ACE2), a protein receptor on the surface of human cells that mediates
Vaccines 2022, 10, 897. https://doi.org/10.3390/vaccines10060897 https://www.mdpi.com/journal/vaccines