Making COVID-19 mRNA vaccines accessible: challenges resolved

Figures & data

Figure 1. Infectiveness and mortality from infections, including SARS-CoV-2 strains [5,6] .

Figure 2. mRNA in vitro transcription and innate immunity activation. (A) mRNA in vitro transcription. Using DNA with the antigen-encoding sequence as a template, mRNA in vitro transcription products contain single-stranded RNA (ssRNA), double-stranded RNA (dsRNA), etc. The ssRNA structure typically includes a five-prime cap (5’ cap), a five-prime untranslated region (5’ UTR), an open reading frame (ORF) region, a three-prime untranslated region (3’ UTR) and a poly (A) tail structure. (B) RNA translation and antigen presentation. Through endocytosis, mRNAs enter the cytoplasm. Some mRNAs combine with ribosomes of the host cell and translate successfully. Antigen proteins can be degraded to antigenic peptides by the proteasome in the cytoplasm and presented to cytotoxic T lymphocytes (CTLs) via the major histocompatibility complex (MHC) I pathway. Or, they can be released out of the host cell and taken up by DCs. Then, they are degraded and presented to helper T cells and B cells via the MHC-II pathway. B cells can also recognize released antigen proteins. (C) Self-adjuvant effect. Various pattern recognition receptors (PRRs) can recognize mRNA in vitro transcription products. Endosomal innate immune receptors can recognize ssRNA (e.g. Toll-like receptor 7 (TLR7), TLR8). Endosomal innate immune receptors can recognize dsRNA (e.g. TLR3) and cytoplasmic innate immune receptors (e.g. protein kinase RNA-activated (PKR), a retinoic acid-inducible gene I protein (RIG-I), and melanoma differentiation-associated protein 5 (MDA5), and 2’-5’-oligoadenylate synthase (OAS). Based on those, mRNA products can stimulate the secretion of pro-inflammatory cytokines and type I interferon (IFN), leading to activation of antigen-presenting cells (APCs) and inflammatory reaction. However, they can also activate antiviral enzymes that cause stalled mRNA translation and mRNA degradation [8]. [Under the Creative Commons Attribution 4.0 International license] .

Figure 3. Process flow of mRNA vaccine manufacturing. With permission [28] .

Table 1. The SARS-Cov-2 virus surface protein sequence and the modified sequence (shown in BOLD) translated into the three mRNA vaccines

P0DTC2 (SPIKE_SARS2) Protein Sequence

BNT162b2, mRNA-1273, CVnCov Protein Sequence

MFVFLVLLPLVSSQCVNLTTRTQLPPAYTNSFTRGVYYPDKVFRSSVLHSTQDLFLPFFSNVTWFHAIHVSGTNGTKRFDNPVLPFNDGVYFASTEKSNIIRGWIFGTTLDSKTQSLLIVNNATNVVIKVCEFQFCNDPFLGVYYHKNNKSWMESEFRVYSSANNCTFEYVSQPFLMDLEGKQGNFKNLREFVFKNIDGYFKIYSKHTPINLVRDLPQGFSALEPLVDLPIGINITRFQTLLALHRSYLTPGDSSSGWTAGAAAYYVGYLQPRTFLLKYNENGTITDAVDCALDPLSETKCTLKSFTVEKGIYQTSNFRVQPTESIVRFPNITNLCPFGEVFNATRFASVYAWNRKRISNCVADYSVLYNSASFSTFKCYGVSPTKLNDLCFTNVYADSFVIRGDEVRQIAPGQTGKIADYNYKLPDDFTGCVIAWNSNNLDSKVGGNYNYLYRLFRKSNLKPFERDISTEIYQAGSTPCNGVEGFNCYFPLQSYGFQPTNGVGYQPYRVVVLSFELLHAPATVCGPKKSTNLVKNKCVNFNFNGLTGTGVLTESNKKFLPFQQFGRDIADTTDAVRDPQTLEILDITPCSFGGVSVITPGTNTSNQVAVLYQDVNCTEVPVAIHADQLTPTWRVYSTGSNVFQTRAGCLIGAEHVNNSYECDIPIGAGICASYQTQTNSPRRARSVASQSIIAYTMSLGAENSVAYSNNSIAIPTNFTISVTTEILPVSMTKTSVDCTMYICGDSTECSNLLLQYGSFCTQLNRALTGIAVEQDKNTQEVFAQVKQIYKTPPIKDFGGFNFSQILPDPSKPSKRSFIEDLLFNKVTLADAGFIKQYGDCLGDIAARDLICAQKFNGLTVLPPLLTDEMIAQYTSALLAGTITSGWTFGAGAALQIPFAMQMAYRFNGIGVTQNVLYENQKLIANQFNSAIGKIQDSLSSTASALGKLQDVVNQNAQALNTLVKQLSSNFGAISSVLNDILSRLDKVEAEVQIDRLITGRLQSLQTYVTQQLIRAAEIRASANLAATKMSECVLGQSKRVDFCGKGYHLMSFPQSAPHGVVFLHVTYVPAQEKNFTTAPAICHDGKAHFPREGVFVSNGTHWFVTQRNFYEPQIITTDNTFVSGNCDVVIGIVNNTVYDPLQPELDSFKEELDKYFKNHTSPDVDLGDISGINASVVNIQKEIDRLNEVAKNLNESLIDLQELGKYEQYIKWPWYIWLGFIAGLIAIVMVTIMLCCMTSCCSCLKGCCSCGSCCKFDEDDSEPVLKGVKLHYT

MFVFLVLLPLVSSQCVNLTTRTQLPPAYTNSFTRGVYYPDKVFRSSVLHSTQDLFLPFFSNVTWFHAIHVSGTNGTKRFDNPVLPFNDGVYFASTEKSNIIRGWIFGTTLDSKTQSLLIVNNATNVVIKVCEFQFCNDPFLGVYYHKNNKSWMESEFRVYSSANNCTFEYVSQPFLMDLEGKQGNFKNLREFVFKNIDGYFKIYSKHTPINLVRDLPQGFSALEPLVDLPIGINITRFQTLLALHRSYLTPGDSSSGWTAGAAAYYVGYLQPRTFLLKYNENGTITDAVDCALDPLSETKCTLKSFTVEKGIYQTSNFRVQPTESIVRFPNITNLCPFGEVFNATRFASVYAWNRKRISNCVADYSVLYNSASFSTFKCYGVSPTKLNDLCFTNVYADSFVIRGDEVRQIAPGQTGKIADYNYKLPDDFTGCVIAWNSNNLDSKVGGNYNYLYRLFRKSNLKPFERDISTEIYQAGSTPCNGVEGFNCYFPLQSYGFQPTNGVGYQPYRVVVLSFELLHAPATVCGPKKSTNLVKNKCVNFNFNGLTGTGVLTESNKKFLPFQQFGRDIADTTDAVRDPQTLEILDITPCSFGGVSVITPGTNTSNQVAVLYQDVNCTEVPVAIHADQLTPTWRVYSTGSNVFQTRAGCLIGAEHVNNSYECDIPIGAGICASYQTQTNSPRRARSVASQSIIAYTMSLGAENSVAYSNNSIAIPTNFTISVTTEILPVSMTKTSVDCTMYICGDSTECSNLLLQYGSFCTQLNRALTGIAVEQDKNTQEVFAQVKQIYKTPPIKDFGGFNFSQILPDPSKPSKRSFIEDLLFNKVTLADAGFIKQYGDCLGDIAARDLICAQKFNGLTVLPPLLTDEMIAQYTSALLAGTITSGWTFGAGAALQIPFAMQMAYRFNGIGVTQNVLYENQKLIANQFNSAIGKIQDSLSSTASALGKLQDVVNQNAQALNTLVKQLSSNFGAISSVLNDILSRLDPPEAEVQIDRLITGRLQSLQTYVTQQLIRAAEIRASANLAATKMSECVLGQSKRVDFCGKGYHLMSFPQSAPHGVVFLHVTYVPAQEKNFTTAPAICHDGKAHFPREGVFVSNGTHWFVTQRNFYEPQIITTDNTFVSGNCDVVIGIVNNTVYDPLQPELDSFKEELDKYFKNHTSPDVDLGDISGINASVVNIQKEIDRLNEVAKNLNESLIDLQELGKYEQYIKWPWYIWLGFIAGLIAIVMVTIMLCCMTSCCSCLKGCCSCGSCCKFDEDDSEPVLKGVKLHYT

Table 2. BioNTech-Pfizer Vaccine Structure and Modifications (mRNA Sequence (4284) (Uridine is replaced with Ψ = 1-methyl-3’-pseudouridylyl) [26]

Element	Description	Position
cap	A modified 5’-cap1 structure (m7G+m3’-5’-ppp-5’-Am)	1–2
5’-UTR	5´-untranslated region derived from human alpha-globin RNA with an optimized Kozak sequence.	3–54
sig	S glycoprotein signal peptide (extended leader sequence), which guides translocation of the nascent polypeptide chain into the endoplasmic reticulum.	55–102
S protein_mut	Codon-optimized sequence encoding full-length SARS-CoV-2 spike (S) glycoprotein containing mutations K986P and V987P to ensure the S glycoprotein remains in an antigenically optimal pre-fusion conformation; stop codons: 3874–3879.	103–3879
3’-UTR	The 3´ untranslated region comprises two sequence elements derived from the amino-terminal enhancer of split (AES) mRNA and the mitochondrial encoded 12S ribosomal RNA to confer RNA stability and high total protein expression.	3880–4174
poly(A)	A 110-nucleotide poly(A)-tail consisting of a stretch of 30 adenosine residues, followed by a 10-nucleotide linker sequence and another 70 adenosine residues.	4175–4284

Table 3. Sequence of BioNTech-Pfizer mRNA Vaccine (Uridine is replaced with Ψ = 1-methyl-3’-pseudouridylyl) [26]

GAGAAΨAAAC	ΨAGΨAΨΨCΨΨ	CΨGGΨCCCCA	CAGACΨCAGA	GAGAACCCGC	50
CACCAΨGΨΨC	GΨGΨΨCCΨGG	ΨGCΨGCΨGCC	ΨCΨGGΨGΨCC	AGCCAGΨGΨG	100
ΨGAACCΨGAC	CACCAGAACA	CAGCΨGCCΨC	CAGCCΨACAC	CAACAGCΨΨΨ	150
ACCAGAGGCG	ΨGΨACΨACCC	CGACAAGGΨG	ΨΨCAGAΨCCA	GCGΨGCΨGCA	200
CΨCΨACCCAG	GACCΨGΨΨCC	ΨGCCΨΨΨCΨΨ	CAGCAACGΨG	ACCΨGGΨΨCC	250
ACGCCAΨCCA	CGΨGΨCCGGC	ACCAAΨGGCA	CCAAGAGAΨΨ	CGACAACCCC	300
GΨGCΨGCCCΨ	ΨCAACGACGG	GGΨGΨACΨΨΨ	GCCAGCACCG	AGAAGΨCCAA	350
CAΨCAΨCAGA	GGCΨGGAΨCΨ	ΨCGGCACCAC	ACΨGGACAGC	AAGACCCAGA	400
GCCΨGCΨGAΨ	CGΨGAACAAC	GCCACCAACG	ΨGGΨCAΨCAA	AGΨGΨGCGAG	450
ΨΨCCAGΨΨCΨ	GCAACGACCC	CΨΨCCΨGGGC	GΨCΨACΨACC	ACAAGAACAA	500
CAAGAGCΨGG	AΨGGAAAGCG	AGΨΨCCGGGΨ	GΨACAGCAGC	GCCAACAACΨ	550
GCACCΨΨCGA	GΨACGΨGΨCC	CAGCCΨΨΨCC	ΨGAΨGGACCΨ	GGAAGGCAAG	600
CAGGGCAACΨ	ΨCAAGAACCΨ	GCGCGAGΨΨC	GΨGΨΨΨAAGA	ACAΨCGACGG	650
CΨACΨΨCAAG	AΨCΨACAGCA	AGCACACCCC	ΨAΨCAACCΨC	GΨGCGGGAΨC	700
ΨGCCΨCAGGG	CΨΨCΨCΨGCΨ	CΨGGAACCCC	ΨGGΨGGAΨCΨ	GCCCAΨCGGC	750
AΨCAACAΨCA	CCCGGΨΨΨCA	GACACΨGCΨG	GCCCΨGCACA	GAAGCΨACCΨ	800
GACACCΨGGC	GAΨAGCAGCA	GCGGAΨGGAC	AGCΨGGΨGCC	GCCGCΨΨACΨ	850
AΨGΨGGGCΨA	CCΨGCAGCCΨ	AGAACCΨΨCC	ΨGCΨGAAGΨA	CAACGAGAAC	900
GGCACCAΨCA	CCGACGCCGΨ	GGAΨΨGΨGCΨ	CΨGGAΨCCΨC	ΨGAGCGAGAC	950
AAAGΨGCACC	CΨGAAGΨCCΨ	ΨCACCGΨGGA	AAAGGGCAΨC	ΨACCAGACCA	1000
GCAACΨΨCCG	GGΨGCAGCCC	ACCGAAΨCCA	ΨCGΨGCGGΨΨ	CCCCAAΨAΨC	1050
ACCAAΨCΨGΨ	GCCCCΨΨCGG	CGAGGΨGΨΨC	AAΨGCCACCA	GAΨΨCGCCΨC	1100
ΨGΨGΨACGCC	ΨGGAACCGGA	AGCGGAΨCAG	CAAΨΨGCGΨG	GCCGACΨACΨ	1150
CCGΨGCΨGΨA	CAACΨCCGCC	AGCΨΨCAGCA	CCΨΨCAAGΨG	CΨACGGCGΨG	1200
ΨCCCCΨACCA	AGCΨGAACGA	CCΨGΨGCΨΨC	ACAAACGΨGΨ	ACGCCGACAG	1250
CΨΨCGΨGAΨC	CGGGGAGAΨG	AAGΨGCGGCA	GAΨΨGCCCCΨ	GGACAGACAG	1300
GCAAGAΨCGC	CGACΨACAAC	ΨACAAGCΨGC	CCGACGACΨΨ	CACCGGCΨGΨ	1350
GΨGAΨΨGCCΨ	GGAACAGCAA	CAACCΨGGAC	ΨCCAAAGΨCG	GCGGCAACΨA	1400
CAAΨΨACCΨG	ΨACCGGCΨGΨ	ΨCCGGAAGΨC	CAAΨCΨGAAG	CCCΨΨCGAGC	1450
GGGACAΨCΨC	CACCGAGAΨC	ΨAΨCAGGCCG	GCAGCACCCC	ΨΨGΨAACGGC	1500
GΨGGAAGGCΨ	ΨCAACΨGCΨA	CΨΨCCCACΨG	CAGΨCCΨACG	GCΨΨΨCAGCC	1550
CACAAAΨGGC	GΨGGGCΨAΨC	AGCCCΨACAG	AGΨGGΨGGΨG	CΨGAGCΨΨCG	1600
AACΨGCΨGCA	ΨGCCCCΨGCC	ACAGΨGΨGCG	GCCCΨAAGAA	AAGCACCAAΨ	1650
CΨCGΨGAAGA	ACAAAΨGCGΨ	GAACΨΨCAAC	ΨΨCAACGGCC	ΨGACCGGCAC	1700
CGGCGΨGCΨG	ACAGAGAGCA	ACAAGAAGΨΨ	CCΨGCCAΨΨC	CAGCAGΨΨΨG	1750
GCCGGGAΨAΨ	CGCCGAΨACC	ACAGACGCCG	ΨΨAGAGAΨCC	CCAGACACΨG	1800
GAAAΨCCΨGG	ACAΨCACCCC	ΨΨGCAGCΨΨC	GGCGGAGΨGΨ	CΨGΨGAΨCAC	1850
CCCΨGGCACC	AACACCAGCA	AΨCAGGΨGGC	AGΨGCΨGΨAC	CAGGACGΨGA	1900
ACΨGΨACCGA	AGΨGCCCGΨG	GCCAΨΨCACG	CCGAΨCAGCΨ	GACACCΨACA	1950
ΨGGCGGGΨGΨ	ACΨCCACCGG	CAGCAAΨGΨG	ΨΨΨCAGACCA	GAGCCGGCΨG	2000
ΨCΨGAΨCGGA	GCCGAGCACG	ΨGAACAAΨAG	CΨACGAGΨGC	GACAΨCCCCA	2050
ΨCGGCGCΨGG	AAΨCΨGCGCC	AGCΨACCAGA	CACAGACAAA	CAGCCCΨCGG	2100
AGAGCCAGAA	GCGΨGGCCAG	CCAGAGCAΨC	AΨΨGCCΨACA	CAAΨGΨCΨCΨ	2150
GGGCGCCGAG	AACAGCGΨGG	CCΨACΨCCAA	CAACΨCΨAΨC	GCΨAΨCCCCA	2200
CCAACΨΨCAC	CAΨCAGCGΨG	ACCACAGAGA	ΨCCΨGCCΨGΨ	GΨCCAΨGACC	2250
AAGACCAGCG	ΨGGACΨGCAC	CAΨGΨACAΨC	ΨGCGGCGAΨΨ	CCACCGAGΨG	2300
CΨCCAACCΨG	CΨGCΨGCAGΨ	ACGGCAGCΨΨ	CΨGCACCCAG	CΨGAAΨAGAG	2350
CCCΨGACAGG	GAΨCGCCGΨG	GAACAGGACA	AGAACACCCA	AGAGGΨGΨΨC	2400
GCCCAAGΨGA	AGCAGAΨCΨA	CAAGACCCCΨ	CCΨAΨCAAGG	ACΨΨCGGCGG	2450
CΨΨCAAΨΨΨC	AGCCAGAΨΨC	ΨGCCCGAΨCC	ΨAGCAAGCCC	AGCAAGCGGA	2500
GCΨΨCAΨCGA	GGACCΨGCΨG	ΨΨCAACAAAG	ΨGACACΨGGC	CGACGCCGGC	2550
ΨΨCAΨCAAGC	AGΨAΨGGCGA	ΨΨGΨCΨGGGC	GACAΨΨGCCG	CCAGGGAΨCΨ	2600
GAΨΨΨGCGCC	CAGAAGΨΨΨA	ACGGACΨGAC	AGΨGCΨGCCΨ	CCΨCΨGCΨGA	2650
CCGAΨGAGAΨ	GAΨCGCCCAG	ΨACACAΨCΨG	CCCΨGCΨGGC	CGGCACAAΨC	2700
ACAAGCGGCΨ	GGACAΨΨΨGG	AGCAGGCGCC	GCΨCΨGCAGA	ΨCCCCΨΨΨGC	2750
ΨAΨGCAGAΨG	GCCΨACCGGΨ	ΨCAACGGCAΨ	CGGAGΨGACC	CAGAAΨGΨGC	2800
ΨGΨACGAGAA	CCAGAAGCΨG	AΨCGCCAACC	AGΨΨCAACAG	CGCCAΨCGGC	2850
AAGAΨCCAGG	ACAGCCΨGAG	CAGCACAGCA	AGCGCCCΨGG	GAAAGCΨGCA	2900
GGACGΨGGΨC	AACCAGAAΨG	CCCAGGCACΨ	GAACACCCΨG	GΨCAAGCAGC	2950
ΨGΨCCΨCCAA	CΨΨCGGCGCC	AΨCAGCΨCΨG	ΨGCΨGAACGA	ΨAΨCCΨGAGC	3000
AGACΨGGACC	CΨCCΨGAGGC	CGAGGΨGCAG	AΨCGACAGAC	ΨGAΨCACAGG	3050
CAGACΨGCAG	AGCCΨCCAGA	CAΨACGΨGAC	CCAGCAGCΨG	AΨCAGAGCCG	3100
CCGAGAΨΨAG	AGCCΨCΨGCC	AAΨCΨGGCCG	CCACCAAGAΨ	GΨCΨGAGΨGΨ	3150
GΨGCΨGGGCC	AGAGCAAGAG	AGΨGGACΨΨΨ	ΨGCGGCAAGG	GCΨACCACCΨ	3200
GAΨGAGCΨΨC	CCΨCAGΨCΨG	CCCCΨCACGG	CGΨGGΨGΨΨΨ	CΨGCACGΨGA	3250
CAΨAΨGΨGCC	CGCΨCAAGAG	AAGAAΨΨΨCA	CCACCGCΨCC	AGCCAΨCΨGC	3300
CACGACGGCA	AAGCCCACΨΨ	ΨCCΨAGAGAA	GGCGΨGΨΨCG	ΨGΨCCAACGG	3350
CACCCAΨΨGG	ΨΨCGΨGACAC	AGCGGAACΨΨ	CΨACGAGCCC	CAGAΨCAΨCA	3400
CCACCGACAA	CACCΨΨCGΨG	ΨCΨGGCAACΨ	GCGACGΨCGΨ	GAΨCGGCAΨΨ	3450
GΨGAACAAΨA	CCGΨGΨACGA	CCCΨCΨGCAG	CCCGAGCΨGG	ACAGCΨΨCAA	3500
AGAGGAACΨG	GACAAGΨACΨ	ΨΨAAGAACCA	CACAAGCCCC	GACGΨGGACC	3550
ΨGGGCGAΨAΨ	CAGCGGAAΨC	AAΨGCCAGCG	ΨCGΨGAACAΨ	CCAGAAAGAG	3600
AΨCGACCGGC	ΨGAACGAGGΨ	GGCCAAGAAΨ	CΨGAACGAGA	GCCΨGAΨCGA	3650
CCΨGCAAGAA	CΨGGGGAAGΨ	ACGAGCAGΨA	CAΨCAAGΨGG	CCCΨGGΨACA	3700
ΨCΨGGCΨGGG	CΨΨΨAΨCGCC	GGACΨGAΨΨG	CCAΨCGΨGAΨ	GGΨCACAAΨC	3750
AΨGCΨGΨGΨΨ	GCAΨGACCAG	CΨGCΨGΨAGC	ΨGCCΨGAAGG	GCΨGΨΨGΨAG	3800
CΨGΨGGCAGC	ΨGCΨGCAAGΨ	ΨCGACGAGGA	CGAΨΨCΨGAG	CCCGΨGCΨGA	3850
AGGGCGΨGAA	ACΨGCACΨAC	ACAΨGAΨGAC	ΨCGAGCΨGGΨ	ACΨGCAΨGCA	3900
CGCAAΨGCΨA	GCΨGCCCCΨΨ	ΨCCCGΨCCΨG	GGΨACCCCGA	GΨCΨCCCCCG	3950
ACCΨCGGGΨC	CCAGGΨAΨGC	ΨCCCACCΨCC	ACCΨGCCCCA	CΨCACCACCΨ	4000
CΨGCΨAGΨΨC	CAGACACCΨC	CCAAGCACGC	AGCAAΨGCAG	CΨCAAAACGC	4050
ΨΨAGCCΨAGC	CACACCCCCA	CGGGAAACAG	CAGΨGAΨΨAA	CCΨΨΨAGCAA	4100
ΨAAACGAAAG	ΨΨΨAACΨAAG	CΨAΨACΨAAC	CCCAGGGΨΨG	GΨCAAΨΨΨCG	4150
ΨGCCAGCCAC	ACCCΨGGAGC	ΨAGCAAAAAA	AAAAAAAAAA	AAAAAAAAAA	4200
AAAAGCAΨAΨ	GACΨAAAAAA	AAAAAAAAAA	AAAAAAAAAA	AAAAAAAAAA	4250
AAAAAAAAAA	AAAAAAAAAA	AAAAAAAAAA	AAAA		4284

Table 4. Lipid nanoparticle formulation of mRNA vaccines

mRNA Vaccine	Lipid Name	Lipid amount per dose [mg]
NIH-Moderna; mRNA-1273	Ionizable lipid a	1.09
	Phospholipid b	0.24
	Cholesterol	0.47
	PEG lipid c	0.13
	Total lipids	1.93
BioNTech-Pfizer; BNT162b2	Ionizable lipid d	0.43
	Phospholipid e	0.09
	Cholesterol	0.2
	PEG lipid f	0.05
	Total lipids	0.77
CureVac; CVnCoV	Ionizable lipid g	0.17
	Phospholipid h	0.04
	Cholesterol	0.08
	PEG lipid i	0.02
	Total lipids	0.31

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