Emergence of SARS-CoV-2 Omicron lineages BA.4 and BA.5 in South Africa
Open accessPublished version of a preprint
Abstract
Three lineages (BA.1, BA.2 and BA.3) of the severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) Omicron variant of concern predominantly drove South Africa’s fourth Coronavirus Disease 2019 (COVID-19) wave. We have now identified two new lineages, BA.4 and BA.5, responsible for a fifth wave of infections. The spike proteins of BA.4 and BA.5 are identical, and similar to BA.2 except for the addition of 69–70 deletion (present in the Alpha variant and the BA.1 lineage), L452R (present in the Delta variant), F486V and the wild-type amino acid at Q493. The two lineages differ only outside of the spike region. The 69–70 deletion in spike allows these lineages to be identified by the proxy marker of S-gene target failure, on the background of variants not possessing this feature. BA.4 and BA.5 have rapidly replaced BA.2, reaching more than 50% of sequenced cases in South Africa by the first week of April 2022. Using a multinomial logistic regression model, we estimated growth advantages for BA.4 and BA.5 of 0.08 (95% confidence interval (CI): 0.08–0.09) and 0.10 (95% CI: 0.09–0.11) per day, respectively, over BA.2 in South Africa. The continued discovery of genetically diverse Omicron lineages points to the hypothesis that a discrete reservoir, such as human chronic infections and/or animal hosts, is potentially contributing to further evolution and dispersal of the virus.
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Written from the paper’s full text.
TL;DR
Two new SARS-CoV-2 Omicron lineages, BA.4 and BA.5, were identified in South Africa and drove a fifth infection wave after rapidly replacing BA.2. They share an identical spike protein and showed estimated daily growth advantages over BA.2 of 0.08 for BA.4 and 0.10 for BA.5.
Key findings
- BA.4 and BA.5 have identical spike proteins, most similar to BA.2 but with additional spike mutations 69–70 deletion, L452R, F486V and wild-type amino acid at Q493; they differ outside spike. They reached more than 50% of sequenced cases in South Africa by the first week of April 2022 and approximately 90% of sequenced cases in Gauteng and KwaZulu-Natal by the week starting 18 April 2022.
- Estimated growth advantages over BA.2 in South Africa were 0.08 per day (95% CI: 0.08–0.09) for BA.4 and 0.10 per day (95% CI: 0.09–0.11) for BA.5.
- Among 198 samples with paired valid qPCR and sequence, 77 had S-gene target failure (SGTF); of these, 66 were BA.4 or BA.5, nine were BA.1 and two were BA.2. No BA.4 or BA.5 genomes were S-gene target positive on qPCR.
- The BA.4/BA.5 wave peaked at ~24% test positivity, lower than the BA.1 wave (~34%), with much lower hospital admissions and deaths than earlier waves in South Africa.
Methods
- Analyzed 15,225 SARS-CoV-2 sequences from South Africa collected 1 November 2021 to 19 May 2022 through the Network for Genomics Surveillance in South Africa, using phylogenetic, phylogeographic and lineage classification methods.
- Estimated lineage growth advantage with a multinomial logistic regression model; monitored SGTF using the TaqPath COVID-19 qPCR assay and validated SGTF against genome lineage assignment in 296 unselected samples, of which 198 had paired valid qPCR and sequence results.
- Used RASCL and natural selection analysis pipelines to assess selection on BA.4 and BA.5 genes; no statistical method was used to predetermine sample size.
Limitations
- The paper states that estimated growth advantage could be biased by stochastic effects such as superspreading in a low-incidence setting at the start of a wave, potentially overestimating the advantage.
- The paper states that reliable estimates of population immunity against BA.1 in South Africa were not yet available, making precise estimates of transmissibility or immune evasion difficult; origin uncertainty remains, phylogenetic inference is limited by low sampling coverage (<1% of confirmed cases), and lack of an ancestor complicates placements.
- Evident from the text: SGTF as a proxy for BA.4/BA.5 may be less useful where BA.1 prevalence is high, and the phenotypic effects of the spike mutations are inferred from prior literature rather than measured directly in this study.
Model-written summary. Check it against the paper before relying on it.
Integrity signals
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Cites
Statistics
Claims the paper makes
0 of 20 cited sources checked · 0 supported · 0 flagged
- Findingnone given
The displacement of BA.1 by BA.2 in South Africa was not associated with a substantial resurgence in cases, hospital admissions or deaths.
“Despite being associated with a modest prolongation of the fourth wave, the displacement of BA.1 by BA.2 in South Africa was not associated with a substantial resurgence in cases, hospital admissions or deaths.”
Quote in the paperNot a substantial claim - Findingown data
BA.5 is estimated to have originated in early January 2022 (95% HPD: 10 December 2021 to 6 February 2022).
“BA.4 and BA.5 are estimated to have originated in mid-December 2021 (95% highest posterior density (HPD): 25 November 2021 to 1 January 2022) and early January 2022 (HPD: 10 December 2021 to 6 February 2022), respectively (Fig. 1a).”
Quote in the paperSubstantial claim - Findingown data
The most recent common ancestor of BA.4 and BA.5 is estimated to have originated in mid-November 2021 (95% HPD: 29 September 2021 to 6 December 2021).
“The most recent common ancestor of BA.4 and BA.5 is estimated to have originated in mid-November 2021 (HPD: 29 September 2021 to 6 December 2021) (Fig. 1a), coinciding with the emergence of the other lineages—for example, BA.2 in early November 2021 (HPD: 9 October 2021 to 29 November 2021).”
Quote in the paperSubstantial claim - Findingown data
Before the emergence of BA.4 and BA.5, F486V in the receptor-binding domain (RBD) of spike had been observed in only 54 of 10 million publicly available genome sequences in GISAID.
“Before the emergence of BA.4 and BA.5, F486V in the receptor-binding domain (RBD) of spike had been observed in only 54 of 10 million publicly available genome sequences in GISAID (https://cov-spectrum.org/explore/World/AllSamples/AllTimes/variants?aaMutations=S%3AF486V&).”
Quote in the paperSubstantial claim - Findingnone given
Relative to BA.2, BA.5 has additional reversions at ORF6:D61 and nucleotide positions 26,858 and 27,259.
“Relative to BA.2, BA.5 has additional reversions at ORF6:D61 and nucleotide positions 26,858 and 27,259.”
Quote in the paperNot a substantial claim
- Findingown data
Relative to BA.2, the spike proteins of BA.4 and BA.5 have additional mutations 69-70 deletion, L452R, F486V and wild-type amino acid at position Q493.
“similar to BA.2 except for the addition of 69–70 deletion (present in the Alpha variant and the BA.1 lineage), L452R (present in the Delta variant), F486V and the wild-type amino acid at Q493.”
- Also stated in: § Main · plain
“Relative to BA.2, BA.4 and BA.5 have the additional spike mutations 69–70 deletion, L452R, F486V and wild-type amino acid at position Q493”
Quote in the paperSubstantial claim - Also stated in: § Main · plain
- Findingown data
Using the RASCL pipeline, no compelling evidence was found of imbalances between ratios of synonymous and non-synonymous substitutions indicative of positive or negative selection acting on any of the genes of viruses in either the BA.4 or BA.5 lineages.
“Using the RASCL pipeline7 (which employs a battery of tests that analyze ratios of synonymous and non-synonymous substitutions both at individual codon sites and entire protein-coding regions), we found no compelling evidence of imbalances between ratios of synonymous and non-synonymous substitutions such as would be indicative of positive”
- Also stated in: § Main · plain
“we found no compelling evidence of imbalances between ratios of synonymous and non-synonymous substitutions such as would be indicative of positive selection (that is, favoring amino acid changes) or negative selection (that is, disfavoring amino acid changes) acting on any of the genes of viruses in either the BA.4 or BA.5 lineages.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [7] Lucaci, A. G. et al. RASCL: rapid assessment of SARS-CoV-2 clades through molecular sequence analysis. Preprint at https://www.biorxiv.org/content/10.1101/2022.01.15.476448v1 (2022). source
Not checked in the free run.
- Also stated in: § Main · plain
- Findingown data
Phylogeographic analysis suggests early dispersal of BA.5 from Gauteng to KwaZulu-Natal, with more limited onward spread to other provinces.
“and early dispersal of BA.5 from Gauteng to KwaZulu-Natal, with more limited onward spread to other provinces (Fig. 1c).”
Quote in the paperSubstantial claim - Findingown data
BA.4 is estimated to have originated in mid-December 2021 (95% HPD: 25 November 2021 to 1 January 2022).
“BA.4 and BA.5 are estimated to have originated in mid-December 2021 (95% highest posterior density (HPD): 25 November 2021 to 1 January 2022) and early January 2022 (HPD: 10 December 2021 to 6 February 2022), respectively (Fig. 1a).”
Quote in the paperSubstantial claim - Findingown data
Phylogeographic analysis suggests early dispersal of BA.4 from Limpopo to Gauteng, with later spread to other provinces.
“Phylogeographic analysis suggests early dispersal of BA.4 from Limpopo to Gauteng, with later spread to other provinces (Fig. 1b)”
Quote in the paperSubstantial claim - Findingown data
The spike proteins of BA.4 and BA.5 are identical.
“The spike proteins of BA.4 and BA.5 are identical”
- Also stated in: § Main · plain
“BA.4 and BA.5 have identical spike proteins”
Quote in the paperSubstantial claim - Also stated in: § Main · plain
- Findingown data
BA.4 and BA.5 rapidly replaced BA.2, reaching more than 50% of sequenced cases in South Africa by the first week of April 2022.
“BA.4 and BA.5 have rapidly replaced BA.2, reaching more than 50% of sequenced cases in South Africa by the first week of April 2022.”
Quote in the paperSubstantial claim - Findingown data
Bayesian phylogenetic methods revealed that BA.4 and BA.5 are distinct from the other Omicron lineages.
“Bayesian phylogenetic methods revealed that BA.4 and BA.5 are distinct from the other Omicron lineages (molecular clock signal: correlation coefficient = 0.6, R2 = 0.4; Extended Data Fig. 1).”
Quote in the paperSubstantial claim - Findingown data
Since December 2020, S:486 has been evolving under strong negative selection favoring the F state at this site.
“Selection analyses focusing on ratios of non-synonymous and synonymous substitution rates at individual codons have indicated that, since December 2020, S:486 has been evolving under strong negative selection favoring the F state at this site (that is, the amino acid that is found in Wuhan-Hu-1) (Extended Data Fig. 3).”
Quote in the paperSubstantial claim - Findingown data
Outside of spike, BA.5 has the M:D3N mutation.
“Outside of spike, BA.4 has additional mutations at ORF7b:L11F and N:P151S and a triple amino acid deletion in NSP1:141–143 deletion, whereas BA.5 has the M:D3N mutation.”
Quote in the paperNot a substantial claim - Findingnone given
As the fourth wave receded in mid-January 2022, BA.2 became the dominant South African lineage.
“However, as that wave receded in mid-January 2022, BA.2 became the dominant South African lineage.”
Quote in the paperNot a substantial claim - Findingnone given
BA.4 and BA.5 differ only outside of the spike region.
“The two lineages differ only outside of the spike region.”
Quote in the paperNot a substantial claim - Findingown data
The estimated daily growth advantage of BA.4 over BA.2 in South Africa was 0.08 (95% CI: 0.08-0.09).
“we estimated growth advantages for BA.4 and BA.5 of 0.08 (95% confidence interval (CI): 0.08–0.09) and 0.10 (95% CI: 0.09–0.11) per day, respectively, over BA.2 in South Africa.”
- Also stated in: § Main · hedged
“We estimated that Omicron BA.4 and BA.5 had a daily growth advantage of 0.08 (95% CI: 0.08–0.09) and 0.10 (95% CI: 0.09–0.11), respectively, relative to BA.2 in South Africa in May 2022”
Quote in the paperSubstantial claim - Also stated in: § Main · hedged
- Findingown data
The estimated daily growth advantage of BA.5 over BA.2 in South Africa was 0.10 (95% CI: 0.09-0.11).
“we estimated growth advantages for BA.4 and BA.5 of 0.08 (95% confidence interval (CI): 0.08–0.09) and 0.10 (95% CI: 0.09–0.11) per day, respectively, over BA.2 in South Africa.”
- Also stated in: § Main · hedged
“We estimated that Omicron BA.4 and BA.5 had a daily growth advantage of 0.08 (95% CI: 0.08–0.09) and 0.10 (95% CI: 0.09–0.11), respectively, relative to BA.2 in South Africa in May 2022”
Quote in the paperSubstantial claim - Also stated in: § Main · hedged
- Findingown data
The BA.4/BA.5 wave was characterized by a peak in test positivity rate of ~24%, lower than during the Omicron BA.1 wave (~34%).
“This wave was characterized by a peak in test positivity rate of ~24%, lower than during the Omicron BA.1 wave (~34%), and, because of high population immunity, much lower hospital admissions and deaths than previously recorded during waves of infection in South Africa.”
Quote in the paperNot a substantial claim - Findingown data
At the time of writing, BA.4 and/or BA.5 have been confirmed in all nine provinces in South Africa in samples collected between 10 January 2022 and 19 May 2022.
“At the time of this writing, we have confirmed BA.4 and/or BA.5 in all nine provinces in South Africa (Eastern Cape, Gauteng, KwaZulu-Natal, Limpopo, Mpumalanga, North West, Northern Cape, Free State and Western Cape) in samples collected between 10 January 2022 and 19 May 2022 (Fig. 2b).”
Quote in the paperSubstantial claim - Findingreasoning
The RECoVERY pipeline resolved the variant calls.
“This pipeline fixed the observed frameshifts, confirming that they were artifacts of mis-assembly; this subsequently resolved the variant calls.”
Quote in the paperSubstantial claim - Findingown data
At the time of writing, a wave of infections caused by the BA.4 and BA.5 lineages was ending in South Africa.
“At the time of this writing, a wave of infections caused by the BA.4 and BA.5 lineages was ending in South Africa (Fig. 2d).”
Quote in the paperNot a substantial claim - Findingown data
Several of the initial Ion Torrent genomes contained several frameshifts, which caused unknown variant calls.
“Several of the initial Ion Torrent genomes contained several frameshifts, which caused unknown variant calls.”
Quote in the paperNot a substantial claim - Findingown data
The SARS-CoV-2 sequences evolved in a relatively strong clock-like manner, with a correlation coefficient of 0.6 and R2 of 0.4 in linear regression of root-to-tip genetic distances against sampling dates.
“Linear regression of root-to-tip genetic distances against sampling dates indicated that the SARS-CoV-2 sequences evolved in a relatively strong clock-like manner (correlation coefficient = 0.6, R2 = 0.4).”
Quote in the paperNot a substantial claim - Findingown data
The estimated daily growth advantage of BA.2 over BA.1 in February 2022 was 0.07 (95% CI: 0.07-0.06).
“These estimates are similar to the estimated daily growth advantage of 0.07 (95% CI: 0.07–0.06) of BA.2 over BA.1 in February 2022 (Fig. 2c).”
Quote in the paperNot a substantial claim - Findingown data
Of the 77 samples with SGTF on qPCR, 66 were BA.4 or BA.5, nine were BA.1 and two were BA.2.
“Of the 77 samples with SGTF on qPCR, 66 were BA.4 or BA.5, nine were BA.1 and two were BA.2.”
Quote in the paperSubstantial claim - Findingown data
No BA.4 and BA.5 genomes were S-gene target positive on qPCR.
“No BA.4 and BA.5 genomes were S-gene target positive on qPCR (Extended Data Table 1).”
Quote in the paperSubstantial claim - Findingown data
The initial phylogenetic analysis identified clusters corresponding to the BA.4 (n = 120) and BA.5 (n = 51) lineages.
“The initial phylogenetic analysis enabled us to identify clusters corresponding to the BA.4 (n = 120) and BA.5 (n = 51) lineages.”
Quote in the paperSubstantial claim - Findingreasoning
The RECoVERY pipeline fixed the observed frameshifts.
“This pipeline fixed the observed frameshifts, confirming that they were artifacts of mis-assembly; this subsequently resolved the variant calls.”
Quote in the paperSubstantial claim - Premisecitation
The F486L mutation has been shown to directly enhance entry into cells expressing mink or ferret ACE2.
“The F486L mutation has been shown to directly enhance entry into cells expressing mink or ferret ACE2 (ref. 12).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [12] Zhou, J. et al. Mutations that adapt SARS-CoV-2 to mink or ferret do not increase fitness in the human airway. Cell Rep. 38, 110344 (2022). source
Not checked in the free run.
- Premisecitation
Mutations at F486 are associated with a reduction in neutralizing activity by class 1 (and some class 2) neutralizing antibodies and by polyclonal sera.
“Mutations at F486 are associated with a reduction in neutralizing activity by class 1 (and some class 2) neutralizing antibodies and by polyclonal sera9–11.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [9] Greaney, A. J. et al. Mapping mutations to the SARS-CoV-2 RBD that escape binding by different classes of antibodies. Nat. Commun. 12, 4196 (2021). source
Not checked in the free run.
- cites [10] Greaney, A. J. et al. Comprehensive mapping of mutations in the SARS-CoV-2 receptor-binding domain that affect recognition by polyclonal human plasma antibodies. Cell Host Microbe 29, 463–476 (2021). source
Not checked in the free run.
- cites [11] Greaney, A. J. et al. Complete mapping of mutations to the SARS-CoV-2 spike receptor-binding domain that escape antibody recognition. Cell Host Microbe 29, 44–57.e9 (2021). source
Not checked in the free run.
- Interpretationreasoning
BA.4 and BA.5 may be even better at evading neutralizing antibody responses, including those recently elicited by BA.1 infections.
“This suggests that BA.4 and BA.5 may be even better at evading neutralizing antibody responses, including those recently elicited by BA.1 infections.”
Quote in the paperSubstantial claim - Interpretationreasoning
Combined with waning population immunity against infection from the initial Omicron/BA.1 wave, this could create the conditions for a substantial resurgence in infections.
“Combined with waning population immunity against infection from the initial Omicron/BA.1 wave, this could create the conditions for a substantial resurgence in infections.”
Quote in the paperSubstantial claim - Premisecitation
Mutations at position 452 have been associated with a reduction in neutralization by monoclonal antibodies (particularly class 2 antibodies) and polyclonal sera.
“mutations at this position have been associated with a reduction in neutralization by monoclonal antibodies (particularly class 2 antibodies) and polyclonal sera9–11.”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [9] Greaney, A. J. et al. Mapping mutations to the SARS-CoV-2 RBD that escape binding by different classes of antibodies. Nat. Commun. 12, 4196 (2021). source
Not checked in the free run.
- cites [10] Greaney, A. J. et al. Comprehensive mapping of mutations in the SARS-CoV-2 receptor-binding domain that affect recognition by polyclonal human plasma antibodies. Cell Host Microbe 29, 463–476 (2021). source
Not checked in the free run.
- cites [11] Greaney, A. J. et al. Complete mapping of mutations to the SARS-CoV-2 spike receptor-binding domain that escape antibody recognition. Cell Host Microbe 29, 44–57.e9 (2021). source
Not checked in the free run.
- Premisecitation
In some countries, BA.2 was responsible for a greater share of cases, hospitalizations and deaths during the Omicron wave.
“in some countries, BA.2 was responsible for a greater share of cases, hospitalizations and deaths during the Omicron wave2–4.”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [2] Rahimi, F. & Talebi Bezmin Abadi, A. The Omicron subvariant BA.2: birth of a new challenge during the COVID-19 pandemic. Int. J. Surg. 99, 106261 (2022). source
Not checked in the free run.
- cites [3] Fonager, J. et al. Molecular epidemiology of the SARS-CoV-2 variant Omicron BA.2 sub-lineage in Denmark, 29 November 2021 to 2 January 2022. Euro. Surveill 27, 2200181 (2022). source
Not checked in the free run.
- cites [4] Chen, L.-L. et al. Contribution of low population immunity to the severe Omicron BA.2 outbreak in Hong Kong. Nat. Commun. 13, 3618 (2022). source
Not checked in the free run.
- Premisecitation
The L452R mutation has been associated with an increased affinity for receptor binding with a resultant increased in vitro infectivity.
“The L452R mutation has been associated with an increased affinity for receptor binding with a resultant increased in vitro infectivity8.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [8] Motozono, C. et al. SARS-CoV-2 spike L452R variant evades cellular immunity and increases infectivity. Cell Host Microbe 29, 1124–1136 (2021). source
Not checked in the free run.
- Interpretationreasoning
Changes at spike amino acids 452, 486 and 493 are likely to influence human angiotensin-converting enzyme-2 (hACE2) and antibody binding.
“Changes at spike amino acids 452, 486 and 493 are likely to influence human angiotensin-converting enzyme-2 (hACE2) and antibody binding.”
Quote in the paperSubstantial claim- Rests on an assumption: The 452 residue is in immediate proximity to the interaction interface of the hACE2 receptor.
- Premisecitation
Deep mutational scanning suggests that F486 is a key site for escape of vaccine-elicited and infection-elicited RBD-targeted antibodies, including those still able to neutralize Omicron/BA.1.
“Deep mutational scanning suggests that F486 is a key site for escape of vaccine-elicited and infection-elicited RBD-targeted antibodies, including those still able to neutralize Omicron/BA.1 (https://jbloomlab.github.io/SARS2_RBD_Ab_escape_maps/escape-calc/)14.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [14] Greaney, A. J., Starr, T. N. & Bloom, J. D. An antibody-escape estimator for mutations to the SARS-CoV-2 receptor-binding domain. Virus Evol. 8, veac021 (2022). source
Not checked in the free run.
- Interpretationreasoning
The 69-70 deletion in spike allows BA.4 and BA.5 to be identified by the proxy marker of S-gene target failure (SGTF), on the background of variants not possessing this feature.
“The 69–70 deletion in spike allows these lineages to be identified by the proxy marker of S-gene target failure, on the background of variants not possessing this feature.”
Quote in the paperSubstantial claim - Interpretationreasoning
BA.4 and BA.5 may be related by a recombination event, with breakpoint between the E and M genes, before their emergence into the general population.
“This suggests that BA.4 and BA.5 may be related by a recombination event, with breakpoint between the E and M genes, before their emergence into the general population.”
Quote in the paperSubstantial claim - Interpretationown data
SGTF surveillance (where the assay is available) may, for now, be a reasonable proxy to identify BA.4 and BA.5 for countries with a low prevalence of BA.1.
“These results suggest that SGTF surveillance (where the assay is available) may, for now, be a reasonable proxy to identify BA.4 and BA.5 for countries with a low prevalence of BA.1.”
Quote in the paperSubstantial claim - Study design and conductown data
Phylogenetic inference is limited by the relatively low sampling coverage in our genomic surveillance (<1% of confirmed cases in South Africa).
“There also remains some uncertainty about the origin of the different Omicron lineages, and phylogenetic inference is limited by the relatively low sampling coverage in our genomic surveillance (<1% of confirmed cases in South Africa).”
Quote in the paperSubstantial claim - Interpretationown data
The observed frameshifts were artifacts of mis-assembly.
“This pipeline fixed the observed frameshifts, confirming that they were artifacts of mis-assembly; this subsequently resolved the variant calls.”
Quote in the paperSubstantial claim - Interpretationcitation
The ability of the BA.4 and BA.5 lineages to drive a new wave of infections can potentially be explained by their ability to evade immunity induced by the BA.1 lineage roughly 3 months after infection.
“The ability of the BA.4 and BA.5 lineages to drive a new wave of infections can potentially be explained by their ability to evade immunity induced by the BA.1 lineage roughly 3 months after infection18.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [18] Khan, K. et al. Omicron sub-lineages BA.4/BA.5 escape BA.1 infection elicited neutralizing immunity. Preprint at https://www.medrxiv.org/content/10.1101/2022.04.29.22274477v1 (2022).
Not checked in the free run.
- Interpretationreasoning
The observation that both BA.4 and BA.5 (and many lineages within them) have recently started to grow in frequency suggests that the growth advantage is recent and uniform across these lineages.
“The observation that both BA.4 and BA.5 (and many lineages within them) have recently started to grow in frequency suggests that the growth advantage is recent and uniform across these lineages.”
Quote in the paperSubstantial claim - Interpretationreasoning
Given that the transmission advantage becomes apparent approximately 4 months from the start of the Omicron wave, it is plausible that waning immunity (particularly that acquired from BA.1 infection) is an important contributory factor.
“Given that the transmission advantage becomes apparent approximately 4 months from the start of the Omicron wave, it is plausible that waning immunity (particularly that acquired from BA.1 infection) is an important contributory factor.”
Quote in the paperNot a substantial claim - Study design and conductreasoning
The lack of sampling of an ancestor of the different Omicron lineages complicates phylogenetic placements.
“Furthermore, the lack of sampling of an ancestor of the different Omicron lineages complicates phylogenetic placements.”
Quote in the paperSubstantial claim - Interpretationreasoning
The frameshifts in the initial Ion Torrent genomes were probably sequencing errors resulting in mis-assembled regions, probably due to the known error profile of Ion Torrent sequencers.
“Manual inspection revealed that these were probably sequencing errors resulting in mis-assembled regions (probably due to the known error profile of Ion Torrent sequencers).”
Quote in the paperNot a substantial claim
- Classified as probably not a substantial claim. 22
- May combine two claims. 19
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingown dataBA.4 and BA.5 have a nuc:G12160A synonymous mutation in NSP8 that was present in Epsilon (B.1.429) and has arisen in BA.2 in some locations.
“In addition, BA.4 and BA.5 have a nuc:G12160A synonymous mutation in NSP8 that was present in Epsilon (B.1.429) and has arisen in BA.2 in some locations (Extended Data Fig. 2).”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataBA.4 and BA.5 have identical mutational patterns in the 5′ genome region (from ORF1ab to Envelope) yet exhibit genetic divergence in the 3′ region (from M to the 3′ genome end).
“BA.4 and BA.5 have identical mutational patterns in the 5′ genome region (from ORF1ab to Envelope) yet exhibit genetic divergence in the 3′ region (from M to the 3′ genome end).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataOutside of spike, BA.4 has additional mutations at ORF7b:L11F and N:P151S and a triple amino acid deletion in NSP1:141-143 deletion.
“Outside of spike, BA.4 has additional mutations at ORF7b:L11F and N:P151S and a triple amino acid deletion in NSP1:141–143 deletion, whereas BA.5 has the M:D3N mutation.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataTwo new Omicron lineages, BA.4 and BA.5, were identified and were responsible for a fifth wave of SARS-CoV-2 infections in South Africa.
“We have now identified two new lineages, BA.4 and BA.5, responsible for a fifth wave of infections.”
Quote in the paperSubstantial claim- Rests on an assumption: BA.1 caused most of the infections in South Africa's fourth epidemic wave.
May combine two claims.
FindingreasoningThe BA.4/BA.5 wave had much lower hospital admissions and deaths than previously recorded during waves of infection in South Africa, because of high population immunity.
“This wave was characterized by a peak in test positivity rate of ~24%, lower than during the Omicron BA.1 wave (~34%), and, because of high population immunity, much lower hospital admissions and deaths than previously recorded during waves of infection in South Africa.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Findingnone givenBotswana's fourth wave was driven by BA.1, followed by BA.2 lasting about 3.5 months, and the country's fifth wave is occurring approximately 2 months after the fourth wave.
“To note, Botswana’s fourth wave was driven by BA.1, followed by BA.2 lasting about 3.5 months, and the country’s fifth wave is occurring approximately 2 months after the fourth wave.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataBA.4 and BA.5 also show a growth advantage against non-Omicron lineages, although these are minimally circulating in the discussed timeframe.
“The BA.4 and BA.5 lineages also show a growth advantage against non-Omicron lineages, although these are minimally circulating in the discussed timeframe (Extended Data Table 2).”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingnone givenBotswana was experiencing a rapid rise in cases driven by BA.4 and BA.5, with 19 of 24 health districts experiencing resurgence in cases.
“At the time of writing this report, Botswana was experiencing a rapid rise in cases driven by BA.4 and BA.5, with 19 of 24 health districts experiencing resurgence in cases.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataThe data suggest geographic heterogeneity in the distribution of BA.4 and BA.5, with growth predominantly of BA.4 in Gauteng and BA.5 in KwaZulu-Natal.
“The data suggest geographic heterogeneity in the distribution of these two new lineages, with growth predominantly of BA.4 in Gauteng and BA.5 in KwaZulu-Natal (Extended Data Fig. 4).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataAlthough Bayesian phylogenetic methods suggest that BA.4 and BA.5 are independent lineages that originated around the same time as BA.1-BA.3, maximum likelihood estimations suggest that they could have descended from BA.2.
“Although the Bayesian phylogenetic methods employed here suggest that BA.4 and BA.5 are independent lineages that originated around the same time as BA.1–BA.3, maximum likelihood estimations suggest that they could have descended from BA.2.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingnone givenTwo new Omicron lineages (BA.4 and BA.5) were identified, which are associated with a resurgence in infections in South Africa approximately 4 months on from the start of the Omicron wave.
“In conclusion, we have identified two new Omicron lineages (BA.4 and BA.5), which are associated with a resurgence in infections in South Africa approximately 4 months on from the start of the Omicron wave.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Findingnone givenRecorded death metrics were further decoupled from cases and hospitalizations compared to the BA.1 wave.
“It is worth noting that recorded death metrics were further decoupled from cases and hospitalizations compared to the BA.1 wave.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Findingnone givenInternationally, by the end of May 2022, BA.4 and BA.5 had also been detected and were rising in prevalence in several countries: Botswana (estimated prevalence 60%), Europe (Portugal, Spain and Austria) and the United States.
“Internationally, by the end of May 2022, BA.4 and BA.5 had also been detected and were rising in prevalence in several countries: in neighboring Botswana (estimated prevalence 60%), in Europe (Portugal, Spain and Austria) and in the United States.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataRecent data from public laboratories in South Africa suggest that the proportion of positive PCR tests with SGTF has been increasing since early March, suggesting that BA.4 and BA.5 may be responsible for a growing share of recently confirmed cases.
“Recent data from public laboratories in South Africa suggest that the proportion of positive polymerase chain reaction (PCR) tests with SGTF has been increasing since early March, suggesting that BA.4 and BA.5 may be responsible for a growing share of recently confirmed cases (Fig. 2a).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataIn Gauteng and KwaZulu-Natal, BA.4 and BA.5 rapidly replaced BA.2 and were responsible for approximately 90% of sequenced cases by the week starting 18 April 2022.
“In the two most populous provinces in South Africa—Gauteng and KwaZulu-Natal—BA.4 and BA.5 rapidly replaced BA.2 and were responsible for approximately 90% of sequenced cases by the week starting 18 April 2022 (Extended Data Fig. 4).”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Findingnone givenThe fifth wave in South Africa, driven by BA.4 and BA.5, occurred around 4 months after the fourth wave, driven by BA.1.
“The fifth wave in South Africa, driven by BA.4 and BA.5, occurred around 4 months after the fourth wave, driven by BA.1.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationSGTF was successfully used to track the early spread of BA.1, later also enabling discrimination between BA.1 and BA.2 infections, because BA.2 viruses generally lack the S:69-70 deletion.
“SGTF was successfully used to track the early spread of BA.1 (which also demonstrates SGTF), later also enabling discrimination between BA.1 and BA.2 infections, because BA.2 viruses generally lack the S:69–70 deletion15.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [15] Scott, L. et al. Track Omicron’s spread with molecular data. Science 374, 1454–1455 (2021). source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
InterpretationreasoningThe continued discovery of genetically diverse Omicron lineages points to the hypothesis that a discrete reservoir, such as human chronic infections and/or animal hosts, is potentially contributing to further evolution and dispersal of the virus.
“The continued discovery of genetically diverse Omicron lineages points to the hypothesis that a discrete reservoir, such as human chronic infections and/or animal hosts, is potentially contributing to further evolution and dispersal of the virus.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationWhen binding to hACE2, spike amino acid F486 interacts with hACE2 residues L79, M82 and Y83, which collectively comprise a hotspot for ACE2 differences between mammalian species.
“When binding to hACE2, spike amino acid F486 interacts with hACE2 residues L79, M82 and Y83, which collectively comprise a hotspot for ACE2 differences between mammalian species13.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [13] Lan, J. et al. Structure of the SARS-CoV-2 spike receptor-binding domain bound to the ACE2 receptor. Nature 581, 215–220 (2020). source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenNo statistical method was used to predetermine sample size.
“No statistical method was used to predetermine sample size.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
InterpretationreasoningThe effects of these different Omicron lineages may differ by location, depending on the immune landscape and, particularly, the patterns of exposure to BA.1 and BA.2.
“This would also suggest that the effects of these different Omicron lineages may differ by location, depending on the immune landscape and, particularly, the patterns of exposure to BA.1 and BA.2.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningThe continued discovery of genetically diverse Omicron lineages shifts the level of support for hypotheses regarding their origin, from an unsampled location to a discrete reservoir, such as human chronic infections and/or animal reservoirs, potentially contributing to further evolution and dispersal of the virus, although currently the data do not provide any definitive evidence in any direction.
“The continued discovery of genetically diverse Omicron lineages shifts the level of support for hypotheses regarding their origin, from an unsampled location to a discrete reservoir, such as human chronic infections (or even a network of chronic human infections) and/or animal reservoirs, potentially contributing to further evolution and dispersal of the virus, although, currently, the data do not”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductnone givenReliable estimates of the level of population immunity against BA.1 in South Africa are not yet available, making it difficult to precisely estimate transmissibility or immune evasion of the new lineages.
“Second, reliable estimates of the level of population immunity against BA.1 in South Africa are not yet available, making it difficult to precisely estimate transmissibility or immune evasion of the new lineages.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim. May combine two claims.
InterpretationreasoningThe growth advantage of Omicron BA.4 and BA.5 could be mediated by an increase in intrinsic transmissibility relative to other variants, an increase in capacity to infect and be transmitted from previously infected and vaccinated individuals, or both.
“The growth advantage of Omicron BA.4 and BA.5 could be mediated by (1) an increase in its intrinsic transmissibility relative to other variants; (2) an increase relative to other variants in its capacity to infect, and be transmitted from, previously infected and vaccinated individuals; or (3) both.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim.
Study design and conductcitationConvergence of MCMC chains was checked using Tracer version 1.7.1.
“Convergence of MCMC chains was checked using Tracer version 1.7.1 (ref. 30).”
- Also stated in: § Phylogeographic analysis · plain
“with convergence assessed using Tracer version 1.7.1.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [30] Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67, 901–904 (2018). source
Not checked in the free run.
- Also stated in: § Phylogeographic analysis · plain
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationTo date, the only reverse zoonoses cases reported from the African region were in African lions and a puma in a private zoo in Johannesburg, South Africa.
“To date, the only reverse zoonoses cases reported from the African region were in African lions and a puma in a private zoo in Johannesburg, South Africa19.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [19] Koeppel, K. N. et al. SARS-CoV-2 reverse zoonoses to pumas and lions, South Africa. Viruses 14, 120 (2022). source
Not checked in the free run.
Classified as probably not a substantial claim.
Study design and conductcitationAs part of the Network for Genomics Surveillance in South Africa (NGS-SA), seven sequencing hubs receive randomly selected samples for sequencing every week according to approved protocols at each site.
“As part of the Network for Genomics Surveillance in South Africa (NGS-SA)23, seven sequencing hubs receive randomly selected samples for sequencing every week according to approved protocols at each site.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [23] Msomi, N., Mlisana, K. & de Oliveira, T. & Network for Genomic Surveillance in South Africa writing group. A genomics network established to respond rapidly to public health threats in South Africa. Lancet Microbe 1, e229–e230 (2020). source
Not checked in the free run.
Classified as probably not a substantial claim.
Study design and conductnone givenThese estimates are based on unselected sampling for genomic surveillance (samples not selected based on SGTF or genotyping).
“These estimates are based on unselected sampling for genomic surveillance (samples not selected based on SGTF or genotyping).”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S4 (based).
Premisenone givenEstimation of the time of the most recent common ancestor (TMRCA) and dispersal dynamics of the sampled viruses is best achieved using Bayesian phylogenetic methods.
“Given that the estimation of time of the most recent common ancestor (TMRCA) and dispersal dynamics of the sampled viruses is best achieved using Bayesian phylogenetic methods, we then estimated time-calibrated phylogenies using the Bayesian software package BEAST version 1.10.4 (ref. 28).”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenData exclusion was not applicable to this study.
“Data exclusion, randomization and blinding to allocation during experiments and outcome assessment were not applicable to this study.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationAfter the emergence of Omicron, the World Organisation for Animal Health released a statement calling for enhanced surveillance in animals to identify the origin of new variants.
“After the emergence of Omicron, the World Organisation for Animal Health released a statement calling for enhanced surveillance in animals to identify the origin of new variants20.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [20] World Organisation for Animal Health. Statement from the Advisory Group on SARS-CoV-2 Evolution in Animals concerning the origins of Omicron variant. https://www.oie.int/en/document/statement-from-the-advisory-group-on-sars-cov-2-evolution-in-animals-concerning-the-origins-of-omicron-variant/ (2022).
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe estimated time to most recent common ancestor for both BA.4 and BA.5 (mid-November 2021, similar to that for BA.1 and BA.2) argues against the first option because that suggests both lineages would have been circulating throughout the period dominated by BA.1 and then BA.2 without exhibiting a transmission advantage.
“The estimated time to most recent common ancestor for both BA.4 and BA.5 (mid-November 2021, similar to that for BA.1 and BA.2) argues against the first option because that suggests both lineages would have been circulating throughout the period dominated by BA.1 and then BA.2 without exhibiting a transmission advantage.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenRandomization was not applicable to this study.
“Data exclusion, randomization and blinding to allocation during experiments and outcome assessment were not applicable to this study.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenBlinding to allocation during experiments and outcome assessment was not applicable to this study.
“Data exclusion, randomization and blinding to allocation during experiments and outcome assessment were not applicable to this study.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenThe estimated growth advantage of the BA.4 and BA.5 lineages could be biased due to stochastic effects (such as superspreading) in a low-incidence setting at the start of a wave, which can lead to overestimates of the growth advantage.
“First, the estimated growth advantage of the BA.4 and BA.5 lineages could be biased due to stochastic effects (such as superspreading) in a low-incidence setting at the start of a wave, which can lead to overestimates of the growth advantage.”
Quote in the paperNot a substantial claim
The paper relies on this without support.
The 452 residue is in immediate proximity to the interaction interface of the hACE2 receptor.
“The 452 residue is in immediate proximity to the interaction interface of the hACE2 receptor.”
The paper relies on this without support.
BA.1 caused most of the infections in South Africa's fourth epidemic wave.
“BA.1 caused most of the infections in South Africa’s fourth epidemic wave.”
The paper relies on this without support.
The NHLS has a single laboratory information system connecting laboratory testing platforms to a corporate data warehouse, where data can be mined in near real time.
“The NHLS has a single laboratory information system connecting laboratory testing platforms to a corporate data warehouse, where data can be mined in near real time.”
The paper relies on this without support.
The TaqPath COVID-19 assay accounts for around 20% of NHLS PCR tests performed, with around half of those performed in Gauteng.
“The TaqPath COVID-19 assay (Thermo Fisher Scientific) accounts for around 20% of NHLS PCR tests performed, with around half of those performed in Gauteng.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
BA.2 dominated in South Africa by late January and globally by March.
“Subvariant BA.2 dominated in South Africa by late January and globally by March [11,12].”
From Assessing COVID-19 vaccine effectiveness against Omicron subvariants: Report from a meeting of the World Health Organization (2023) · cites this paper as [11]
The Main section says BA.2 became the dominant South African lineage as the fourth wave receded in mid-January 2022, supporting dominance there by late January. It does not say BA.2 dominated globally by March; its worldwide discussion refers only to some countries and gives no March date. (LLM, conf=0.88)
Experimental — accuracy not yet measured
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
BA.2 growth-advantage confidence interval has reversed endpoints
MinorThe reported 95% confidence interval runs from 0.07 to 0.06: its stated lower endpoint is greater than its upper endpoint. The interval therefore cannot be interpreted as written.
“0.07 (95% CI: 0.07–0.06)”
“The estimated daily growth advantage of 0.07”
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RASCL: Rapid Assessment of Selection in CLades through molecular sequence analysis
Lucaci, A. G., Zehr, J. D., Shank, S. D., Bouvier, D., Ostrovsky, A., Han, M., Nekrutenko, A., Martin, D. P., & Pond, S. L. K. (2022) · PLoS ONE · doi:10.1371/journal.pone.0275623 · Cited by 10 · see citing works
Self-citationCoronavirus disease (COVID-19) case data - South Africa
Marivate, V. N., Arbi, R., Combrink, H., et al. (2020) · Zenodo (CERN European Organization for Nuclear Research) · doi:10.5281/zenodo.3819126 · Cited by 10 · see citing works
RASCL: Rapid Assessment Of SARS-CoV-2 Clades Through Molecular Sequence Analysis
Lucaci, A. G., Zehr, J. D., Shank, S. D., Bouvier, D., Mei, H., Nekrutenko, A., Martin, D. P., & Kosakovsky Pond, S. L. (2022) · doi:10.1101/2022.01.15.476448 · Cited by 2 · see citing works
Self-citationRapid epidemic expansion of the SARS-CoV-2 Omicron BA.2 subvariant during China’s largest outbreaksPreprint
Dai, Y. (2022) · Research Square · doi:10.21203/rs.3.rs-1516063/v3 · Cited by 1 · see citing works
Flagged first, then by citations
Research works printed but not linked: 1
- Khan, K. et al. Omicron sub-lineages BA.4/BA.5 escape BA.1 infection elicited neutralizing immunity. Preprint at https://www.medrxiv.org/content/10.1101/2022.04.29.22274477v1 (2022). · DOI
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Markov, P. V., Ghafari, M., Beer, M., Lythgoe, K., Simmonds, P., Stilianakis, N. I., & Katzourakis, A. (2023) · Nature Reviews Microbiology · doi:10.1038/s41579-023-00878-2 · Cited by 1,179 · see citing works
Bivalent Omicron BA.1–Adapted BNT162b2 Booster in Adults Older than 55 Years
Winokur, P., Gayed, J., Fitz-Patrick, D., Thomas, S. J., Diya, O., Lockhart, S., Xu, X., Zhang, Y., Bangad, V., Schwartz, H. I., et al. (2023) · New England Journal of Medicine · doi:10.1056/nejmoa2213082 · Cited by 116 · see citing works
Tseng, H. F., Ackerson, B. K., Bruxvoort, K. J., Sy, L. S., Tubert, J. E., Lee, G. S., Ku, J. H., Florea, A., Luo, Y., Qiu, S., et al. (2023) · Nature Communications · doi:10.1038/s41467-023-35815-7 · Cited by 105 · see citing works
BNT162b2 vaccine effectiveness against SARS-CoV-2 omicron BA.4 and BA.5
Tartof, S. Y., Slezak, J. M., Puzniak, L., Hong, V., Frankland, T. B., Ackerson, B. K., Takhar, H., Ogun, O. A., Simmons, S., Zamparo, J. M., et al. (2022) · The Lancet Infectious Diseases · doi:10.1016/s1473-3099(22)00692-2 · Cited by 66 · see citing works
Rapidly shifting immunologic landscape and severity of SARS-CoV-2 in the Omicron era in South Africa
Sun, K., Tempia, S., Kleynhans, J., von Gottberg, A., McMorrow, M. L., Wolter, N., Bhiman, J. N., Moyes, J., Carrim, M., Martinson, N. A., et al. (2023) · Nature Communications · doi:10.1038/s41467-022-35652-0 · Cited by 45 · see citing works
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Feikin, D. R., Higdon, M. M., Andrews, N., Collie, S., Deloria Knoll, M., Kwong, J. C., Link-Gelles, R., Pilishvili, T., & Patel, M. K. (2023) · Vaccine · doi:10.1016/j.vaccine.2023.02.020 · Cited by 36 · see citing works
“Subvariant BA.2 dominated in South Africa by late January and globally by March [11,12].”
BackgroundInfluentialKislaya, I., Casaca, P., Borges, V., Sousa, C., Ferreira, B. I., Fonte, A., Fernandes, E., Dias, C. M., Duarte, S., Almeida, J. P., et al. (2023) · Emerging Infectious Diseases · doi:10.3201/eid2903.221367 · Cited by 13 · see citing works
Association of SARS-CoV-2 BA.4/BA.5 Omicron lineages with immune escape and clinical outcome
Lewnard, J. A., Hong, V., Kim, J. S., Shaw, S. F., Lewin, B., Takhar, H., & Tartof, S. Y. (2022) · doi:10.1101/2022.07.31.22278258 · Cited by 35 · see citing works
- Authors
- Tegally, H., Moir, M., Everatt, J., Giovanetti, M., Scheepers, C., Wilkinson, E., Subramoney, K., Makatini, Z., Moyo, S., Amoako, D. G., Baxter, C., Althaus, C. L., Anyaneji, U. J., Kekana, D., Viana, R., Giandhari, J., Lessells, R. J., Maponga, T., Maruapula, D., Choga, W., Matshaba, M., Mbulawa, M. B., Msomi, N., NGS-SA consortium, Bester, A. P., Claassen, M., Doolabh, D., Mudau, I., Mbhele, N., Engelbrecht, S., Goedhals, D., Hardie, D., Hsiao, N.-Y., Iranzadeh, A., Ismail, A., Joseph, R., Maharaj, A., Mahlangu, B., Mahlakwane, K., Davis, A., Marais, G., Mlisana, K., Mnguni, A., Mohale, T., Motsatsi, G., Mwangi, P., Ntuli, N., Nyaga, M., Olubayo, L., Radibe, B., Ramphal, Y., Ramphal, U., Strasheim, W., Tebeila, N., van Wyk, S., Wilson, S., Lucaci, A. G., Weaver, S., Maharaj, A., Pillay, Y., Davids, M., Mendes, A., Mayaphi, S., Naidoo, Y., Pillay, S., Sanko, T. J., San, J. E., Scott, L., Singh, L., Magini, N. A., Smith-Lawrence, P., Stevens, W., Dor, G., Tshiabuila, D., Wolter, N., Preiser, W., Treurnicht, F. K., Venter, M., Chiloane, G., McIntyre, C., O’Toole, A., Ruis, C., Peacock, T. P., Roemer, C., Kosakovsky Pond, S. L., Williamson, C., Pybus, O. G., Bhiman, J. N., Glass, A., Martin, D. P., Jackson, B., Rambaut, A., Laguda-Akingba, O., Gaseitsiwe, S., von Gottberg, A., & de Oliveira, T.
- Affiliations
- Stellenbosch University (ZA)
- National Health Laboratory Service (ZA)
- Universidade Federal de Minas Gerais (BR)
- University of the Witwatersrand (ZA)
- Harvard University (US)
- University of Bern (CH)
- University of KwaZulu-Natal (ZA)
- Lancet Laboratories (ZA)
- Ministry of Health (BW)
- University of Cape Town (ZA)
- University of the Free State (ZA)
- Johannesburg Hospital (ZA)
- Temple University (US)
- University of Pretoria (ZA)
- University of Edinburgh (GB)
- University of Cambridge (GB)
- Imperial College London (GB)
- University of Basel (CH)
- University of Oxford (GB)
- ORCID
- Tegally, Houriiyah
- Moir, Monika
- Everatt, Josie
- Giovanetti, Marta
- Scheepers, Cathrine
- Wilkinson, Eduan
- Subramoney, Kathleen
- Makatini, Zinhle
- Moyo, Sikhulile
- Amoako, Daniel G.
- Baxter, Cheryl
- Althaus, Christian L.
- Anyaneji, Ugochukwu J.
- Kekana, Dikeledi
- Viana, Raquel
- Giandhari, Jennifer
- Lessells, Richard J.
- Maponga, Tongai
- Maruapula, Dorcas
- Choga, Wonderful
- Matshaba, Mogomotsi
- Msomi, Nokukhanya
- Claassen, Mathilda
- Mbhele, Nokuzola
- Engelbrecht, Susan
- Goedhals, Dominique
- Hardie, Diana
- Iranzadeh, Arash
- Ismail, Arshad
- Joseph, Rageema
- Maharaj, Arisha
- Mahlangu, Boitshoko
- Mahlakwane, Kamela
- Marais, Gert
- Mlisana, Koleka
- Mnguni, Anele
- Mohale, Thabo
- Mwangi, Peter
- Nyaga, Martin
- Ramphal, Upasana
- Strasheim, Wilhelmina
- Lucaci, Alexander G.
- Weaver, Steven
- Pillay, Yusasha
- Davids, Michaela
- Mendes, Adriano
- Mayaphi, Simnikiwe
- Naidoo, Yeshnee
- Pillay, Sureshnee
- Sanko, Tomasz Janusz
- San, James E.
- Scott, Lesley
- Singh, Lavanya
- Stevens, Wendy
- Dor, Graeme
- Tshiabuila, Derek
- Wolter, Nicole
- Preiser, Wolfgang
- Treurnicht, Florette K.
- Venter, Marietjie
- O’Toole, Aine
- Ruis, Christopher
- Peacock, Thomas P.
- Roemer, Cornelius
- Kosakovsky Pond, Sergei L.
- Williamson, Carolyn
- Pybus, Oliver G.
- Bhiman, Jinal N.
- Glass, Allison
- Martin, Darren P.
- Jackson, Ben
- Rambaut, Andrew
- Laguda-Akingba, Oluwakemi
- Gaseitsiwe, Simani
- von Gottberg, Anne
- de Oliveira, Tulio
- Venue
- Nature Medicine
- Published
- 2022-06-27
- Volume
- 28
- Issue
- 9
- Pages
- 1785-1790
- ISSN
- 1078-8956 (ISSN-L)
- 1546-170X
- PMC
- PMC9499863
- OpenAlex
- W4283704819
- Semantic Scholar
- e708146ef3ab87e2f7381416caf7255675d8898b
- Cited by
- 707
- References
- 48
- Citations by year
- 2026: 37 · 2025: 62 · 2024: 142 · 2023: 259 · 2022: 196 · 2018: 1 · 2017: 1 · 2015: 1 · 2014: 2 · 2013: 1
- FWCI
- 80.66
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Locations
- Nature Medicine (publishedVersion, cc-by)
- medRxiv (acceptedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- UpSpace Institutional Repository (University of Pretoria) (submittedVersion)
- Arca - Repositório Institucional da Fiocruz (submittedVersion, other-oa)
- Bern Open Repository and Information System (University of Bern) (submittedVersion, cc-by)
- TUScholarShare (Temple University) (submittedVersion, cc-by)
- Edinburgh Research Explorer (University of Edinburgh) (submittedVersion, cc-by)
- LA Referencia (Red Federada de Repositorios Institucionales de Publicaciones Científicas) (submittedVersion, other-oa)
- Funders
- U.S. Department of Health & Human Services | NIH | National Institute of Allergy and Infectious Diseases (U01 AI151698)
- U.S. Department of Health & Human Services | National Institutes of Health (U54 TW012041)
- South African Medical Research Council
- Bill and Melinda Gates Foundation (INV-018978)
- Wellcome Trust (221003/Z/20/Z)
- U.S. Department of Health & Human Services | NIH | Center for Information Technology (U54 TW012041)
- Topics
- Health Sciences › Medicine › Infectious Diseases › SARS-CoV-2 and COVID-19 Research
- Physical Sciences › Mathematics › Modeling and Simulation › COVID-19 epidemiological studies
- Health Sciences › Medicine › Infectious Diseases › COVID-19 Clinical Research Studies
- Keywords
- Omicron variant, SARS-CoV-2 lineages, spike protein mutations, S-gene target failure, L452R, growth advantage, COVID-19 waves, South Africa, viral evolution
- MeSH terms
- COVID-19, SARS-CoV-2, Amino Acids, Animals, Humans, South Africa, Spike Glycoprotein, Coronavirus
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