<?xml version="1.0" encoding="UTF-8"?><!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.2 20190208//EN" "http://jats.nlm.nih.gov/publishing/1.2/JATS-journalpublishing1.dtd"><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article" dtd-version="1.2" xml:lang="en">
    <front>
        <journal-meta>
            <journal-id journal-id-type="pmc">VeriXiv</journal-id>
            <journal-title-group>
                <journal-title>VeriXiv</journal-title>
            </journal-title-group>
            <issn pub-type="epub">3029-0988</issn>
            <publisher>
                <publisher-name>F1000 Research Limited</publisher-name>
                <publisher-loc>London, UK</publisher-loc>
            </publisher>
        </journal-meta>
        <article-meta>
            <article-id pub-id-type="doi">10.12688/verixiv.1186.1</article-id>
            <article-categories>
                <subj-group subj-group-type="heading">
                    <subject>Research Article</subject>
                </subj-group>
                <subj-group>
                    <subject>Articles</subject>
                </subj-group>
            </article-categories>
            <title-group>
                <article-title>Estimating the basic reproduction number of measles in low- and middle-income settings using 172 serostudies: a modelling approach</article-title>
                <fn-group content-type="pub-status">
                    <fn>
                        <p>[version 1]</p>
                    </fn>
                </fn-group>
            </title-group>
            <contrib-group>
                <contrib contrib-type="author" corresp="yes">
                    <name>
                        <surname>Fu</surname>
                        <given-names>Han</given-names>
                    </name>
                    <role content-type="http://credit.niso.org/">Conceptualization</role>
                    <role content-type="http://credit.niso.org/">Data Curation</role>
                    <role content-type="http://credit.niso.org/">Formal Analysis</role>
                    <role content-type="http://credit.niso.org/">Methodology</role>
                    <role content-type="http://credit.niso.org/">Software</role>
                    <role content-type="http://credit.niso.org/">Validation</role>
                    <role content-type="http://credit.niso.org/">Visualization</role>
                    <role content-type="http://credit.niso.org/">Writing &#x2013; Original Draft Preparation</role>
                    <uri content-type="orcid">https://orcid.org/0000-0002-8934-7347</uri>
                    <xref ref-type="corresp" rid="c1">a</xref>
                    <xref ref-type="aff" rid="a1">1</xref>
                </contrib>
                <contrib contrib-type="author" corresp="no">
                    <name>
                        <surname>Sbarra</surname>
                        <given-names>Alyssa N.</given-names>
                    </name>
                    <role content-type="http://credit.niso.org/">Data Curation</role>
                    <role content-type="http://credit.niso.org/">Validation</role>
                    <role content-type="http://credit.niso.org/">Writing &#x2013; Review &amp; Editing</role>
                    <xref ref-type="aff" rid="a1">1</xref>
                    <xref ref-type="aff" rid="a2">2</xref>
                </contrib>
                <contrib contrib-type="author" corresp="no">
                    <name>
                        <surname>Russell</surname>
                        <given-names>Timothy</given-names>
                    </name>
                    <role content-type="http://credit.niso.org/">Methodology</role>
                    <role content-type="http://credit.niso.org/">Visualization</role>
                    <role content-type="http://credit.niso.org/">Writing &#x2013; Review &amp; Editing</role>
                    <xref ref-type="aff" rid="a1">1</xref>
                </contrib>
                <contrib contrib-type="author" corresp="no">
                    <name>
                        <surname>Abbas</surname>
                        <given-names>Kaja</given-names>
                    </name>
                    <role content-type="http://credit.niso.org/">Conceptualization</role>
                    <role content-type="http://credit.niso.org/">Validation</role>
                    <role content-type="http://credit.niso.org/">Writing &#x2013; Review &amp; Editing</role>
                    <uri content-type="orcid">https://orcid.org/0000-0003-0563-1576</uri>
                    <xref ref-type="aff" rid="a1">1</xref>
                    <xref ref-type="aff" rid="a3">3</xref>
                    <xref ref-type="aff" rid="a4">4</xref>
                    <xref ref-type="aff" rid="a5">5</xref>
                </contrib>
                <contrib contrib-type="author" corresp="no">
                    <name>
                        <surname>Auzenbergs</surname>
                        <given-names>Megan</given-names>
                    </name>
                    <role content-type="http://credit.niso.org/">Writing &#x2013; Review &amp; Editing</role>
                    <xref ref-type="aff" rid="a1">1</xref>
                </contrib>
                <contrib contrib-type="author" corresp="no">
                    <name>
                        <surname>Jit</surname>
                        <given-names>Mark</given-names>
                    </name>
                    <role content-type="http://credit.niso.org/">Conceptualization</role>
                    <role content-type="http://credit.niso.org/">Funding Acquisition</role>
                    <role content-type="http://credit.niso.org/">Investigation</role>
                    <role content-type="http://credit.niso.org/">Supervision</role>
                    <role content-type="http://credit.niso.org/">Validation</role>
                    <role content-type="http://credit.niso.org/">Writing &#x2013; Review &amp; Editing</role>
                    <xref ref-type="aff" rid="a1">1</xref>
                    <xref ref-type="aff" rid="a6">6</xref>
                    <xref ref-type="aff" rid="a7">7</xref>
                    <xref ref-type="aff" rid="a8">8</xref>
                </contrib>
                <aff id="a1">
                    <label>1</label>Department of Infectious Disease Epidemiology &amp; Dynamics, London School of Hygiene and Tropical Medicine, London, England, WC1E 7HT, UK</aff>
                <aff id="a2">
                    <label>2</label>Department of Epidemiology, Johns Hopkins University, Baltimore, MD, 21205, USA</aff>
                <aff id="a3">
                    <label>3</label>School of Tropical Medicine and Global Health, Nagasaki University, Nagasaki, 852-8523, Japan</aff>
                <aff id="a4">
                    <label>4</label>Institute of Tropical Medicine, Nagasaki University, Nagasaki, 852-8523, Japan</aff>
                <aff id="a5">
                    <label>5</label>Public Health Foundation of India, New Delhi, 110030, India</aff>
                <aff id="a6">
                    <label>6</label>School of Global Public Health, New York University, New York, New York, 10003, USA</aff>
                <aff id="a7">
                    <label>7</label>Saw Swee Hock School of Public Health, National University of Singapore, Singapore, 117549, Singapore</aff>
                <aff id="a8">
                    <label>8</label>School of Public Health, The University of Hong Kong, Hong Kong, Hong Kong</aff>
            </contrib-group>
            <author-notes>
                <corresp id="c1">
                    <label>a</label>
                    <email xlink:href="mailto:Han.Fu@lshtm.ac.uk">Han.Fu@lshtm.ac.uk</email>
                </corresp>
                <fn fn-type="conflict">
                    <p>No competing interests were disclosed.</p>
                </fn>
            </author-notes>
            <pub-date pub-type="epub">
                <day>26</day>
                <month>6</month>
                <year>2025</year>
            </pub-date>
            <pub-date pub-type="collection">
                <year>2025</year>
            </pub-date>
            <volume>2</volume>
            <elocation-id>154</elocation-id>
            <history>
                <date date-type="accepted">
                    <day>29</day>
                    <month>5</month>
                    <year>2025</year>
                </date>
            </history>
            <permissions>
                <copyright-statement>Copyright: &#x00a9; 2025 Fu H et al.</copyright-statement>
                <copyright-year>2025</copyright-year>
                <license xlink:href="https://creativecommons.org/licenses/by/4.0/">
                    <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution Licence, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.</license-p>
                </license>
            </permissions>
            <self-uri content-type="pdf" xlink:href="https://verixiv.org/articles/2-154/pdf"/>
            <abstract>
                <p>The basic reproduction number, R
                    <sub>0</sub>, is an epidemiological measure to describe the transmissibility of infectious diseases and evaluate the potential effect of interventions. R
                    <sub>0</sub> for measles has historically been considered to be between 12&#x2013;18 but rarely estimated using data across multiple countries to address contextual factors contributing to its heterogeneity. Our study aims to estimate measles R
                    <sub>0</sub> in low- and middle-income countries using a standardised database of 172 serostudies extracted from a recent systematic review. Fitting an age-structured compartmental model of measles transmission dynamics and vaccination to the age-specific seroprevalence data using Markov Chain Monte Carlo, we estimated setting-specific posterior distributions of R
                    <sub>0</sub> in the included serostudies with unique survey years and locations. We pooled bootstrapped samples of posterior R
                    <sub>0</sub> estimates by serostudy characteristics, including survey period, geography, and overall bias in sampling, measurement, and reporting serological results. Measles R
                    <sub>0</sub> estimates varied substantially across studies, ranging from 0.94 (95% credible interval: 0.73&#x2013;1.00) to 154 (69.8&#x2013;212), with fewer than 20% of studies having median R
                    <sub>0</sub> values in the range of 12&#x2013;18. Pooled R
                    <sub>0</sub> estimates showed smaller medians and variation in serostudies conducted after 2000 or including the adult population, while no distinguishable variation was identified across the geographical regions. We also found an inverse association between the human development index and R
                    <sub>0</sub>. Our revised estimates demonstrated the wide range of measles R
                    <sub>0</sub> across low- and middle-income settings. Considering the contextual heterogeneity in measles transmissibility is crucial when modelling epidemics and planning vaccination strategies and control interventions.</p>
            </abstract>
            <kwd-group kwd-group-type="author">
                <kwd>Measles</kwd>
                <kwd>basic reproduction number</kwd>
                <kwd>seroprevalence</kwd>
                <kwd>vaccination</kwd>
                <kwd>modelling</kwd>
            </kwd-group>
            <funding-group>
                <award-group id="fund-1" xlink:href="https://doi.org/10.13039/100009619">
                    <funding-source>Japan Agency for Medical Research and Development</funding-source>
                    <award-id>JP223fa627004</award-id>
                </award-group>
                <award-group id="fund-2" xlink:href="https://doi.org/10.13039/100000865">
                    <funding-source>Gates Foundation</funding-source>
                    <award-id>INV-034281</award-id>
                    <award-id>OPP1157270/INV-009125</award-id>
                </award-group>
                <funding-statement>This work was supported, in whole or in part, by the Gates Foundation, via the Vaccine Impact Modelling Consortium (Grant Number INV-034281), previously (OPP1157270 / INV-009125) and Gavi, the Vaccine Alliance. Under the grant conditions of the Foundation, a Creative Commons Attribution 4.0 Generic License has already been assigned to the Author Accepted Manuscript version that might arise from this submission.
KA is supported by the Japan Agency for Medical Research and Development (JP223fa627004). 
</funding-statement>
                <funding-statement>
                    <italic>The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.</italic>
                </funding-statement>
            </funding-group>
        </article-meta>
    </front>
    <body>
        <sec id="sec1" sec-type="intro">
            <title>Introduction</title>
            <p>Measles is a contagious disease prone to epidemics and has resulted in a large health burden worldwide.
                <sup>
                    <xref ref-type="bibr" rid="ref1">1</xref>,
                    <xref ref-type="bibr" rid="ref2">2</xref>
                </sup> The basic reproduction number (R
                <sub>0</sub>) for measles is generally recognised as one of the highest amongst common vaccine-preventable diseases, with suggestions that each primary measles case can generate 12&#x2013;18 secondary infections when the population is completely susceptible.
                <sup>
                    <xref ref-type="bibr" rid="ref3">3</xref>
                </sup> Due to this characteristic of high transmissibility, measles requires a high level of two-dose vaccine coverage for disease control. However, a systematic review of R
                <sub>0</sub> estimates reveals that measles transmissibility varied widely from 1.43 to 770 by country settings.
                <sup>
                    <xref ref-type="bibr" rid="ref4">4</xref>
                </sup> This variation in measles transmissibility can affect the estimation of measles burden and vaccine impact in individual settings. However, it was unclear the extent to which this variation stemmed from the diversity in data sources or estimation methods across studies, as opposed to true heterogeneity.</p>
            <p>Serosurveys examine the presence of specific antibodies to understand the immunity profile against infectious diseases within a population over a defined period of time. Because presence of these measles-specific antibodies is understood to be a correlate of protection from clinical disease, seroprevalence data stratified by age, vaccination history, and other selected characteristics can be used to inform susceptibility gaps in the population and understand the heterogeneity of transmissibility across settings. With appropriate sampling and data collection methods, serosurveys provide a less biased snapshot of the population-level immunity profiles in the community, compared to notification data that are subject to under-ascertainment and underreporting issues in the surveillance systems.
                <sup>
                    <xref ref-type="bibr" rid="ref5">5</xref>
                </sup>
            </p>
            <p>To address the heterogeneity of measles transmissibility across low- and middle-income countries (LMICs) that contribute to the majority of global measles burden,
                <sup>
                    <xref ref-type="bibr" rid="ref2">2</xref>
                </sup> we extracted age-specific seroprevalence data from each study in a recently conducted systematic review of serosurveys. We then inferred the setting-specific R
                <sub>0</sub> using each seroprevalence dataset by fitting it using the Dynamic Measles Immunization Calculation Engine (DynaMICE), a dynamic model of measles transmission and vaccination. DynaMICE incorporates fine-scale age structure and country-specific inputs, including demography, routine and campaign vaccination coverage, and social contact patterns.
                <sup>
                    <xref ref-type="bibr" rid="ref6">6</xref>&#x2013;
                    <xref ref-type="bibr" rid="ref8">8</xref>
                </sup> The combined analysis of R
                <sub>0</sub> model-based estimates and setting-specific characteristics will enhance the understanding of the key factors that contribute to measles transmissibility across settings.</p>
        </sec>
        <sec id="sec2" sec-type="results">
            <title>Results</title>
            <sec id="sec3">
                <title>Posterior R
                    <sub>0</sub> estimates</title>
                <p>Fitting the DynaMICE model to the 172 seroprevalence studies using Markov Chain Monte Carlo (MCMC), we obtained 500 sets of posterior parameters from the converged chains. The posterior model estimates of age-specific seroprevalence show a good fit using the root mean square error (RMSE) measure, with two-thirds of the studies having an average difference less than 10% between the model results and observed data. There was a positive correlation between median R
                    <sub>0</sub> and vaccine effectiveness estimates (Spearman&#x2019;s correlation coefficient (R)=0.444, p-value (p) &lt;0.001. However, the correlation became weaker after excluding studies with a severe or critical bias in study population selection, serological measurement and result reporting (R=0.380, p=0.003). No significant correlation between R
                    <sub>0</sub> and duration of maternal immunity was found (R=0.032, p=0.680).</p>
                <p>
                    <xref ref-type="fig" rid="f1">Figure 1</xref> presents the median and 95% credible intervals (CrIs) of the R
                    <sub>0</sub> posterior distributions obtained from fitting the DynaMICE model to age-specific seroprevalence data in 172 individual studies. We observed a right skewed distribution of R
                    <sub>0</sub> estimates, ranging widely across studies from 0.94 (95% CrI: 0.73&#x2013;1.00) to 154 (95%CrI: 69.8&#x2013;212). Only 18% (n=31) of the median estimates were in the commonly cited range of 12&#x2013;18,
                    <sup>
                        <xref ref-type="bibr" rid="ref3">3</xref>
                    </sup> while 70% (n=121) of them presented a value below 12 and 12% (n=20) above 18. In 21% of the included studies (n=35), posterior R
                    <sub>0</sub> showed a broad range of 95% CrIs, with the difference between lower and upper bounds exceeding 10. These R
                    <sub>0</sub> estimates with a broad range tended to result from serosurveys with a smaller sample size, fewer categories of age groups, or conducted in the pre-vaccination era. Geographically, R
                    <sub>0</sub> estimates concentrated at a range of less than 15 based on serosurveys conducted in the World Health Organization (WHO) Eastern Mediterranean Region, while in other regions, estimates of &#x2265;30. The variation of R
                    <sub>0</sub> estimates remained large between studies in the same country, as seen in China, India, Brazil, and T&#x00fc;rkiye, where median R
                    <sub>0</sub> estimates disperse over a range of 20 in each country.</p>
                <fig fig-type="figure" id="f1" orientation="portrait" position="float">
                    <label>Figure 1. </label>
                    <caption>
                        <title>Posterior measles R
                            <sub>0</sub> obtained by fitting to age-specific seroprevalence data.</title>
                        <p>The median posterior R
                            <sub>0</sub> estimates are presented with 95% credical intervals, ordered from highest to lowest in each WHO region. Arrows indicate the upper R
                            <sub>0</sub> estimates exceed 40. Two studies (JAM-1986 &amp; IND-1983) with the whole intervals above 40 are not shown.</p>
                    </caption>
                    <graphic id="gr1" orientation="portrait" position="float" xlink:href="https://verixiv-files.f1000.com/manuscripts/1213/18ac17ed-152f-4652-98fd-ff2ebb6d233f_figure1.gif"/>
                </fig>
            </sec>
            <sec id="sec4">
                <title>Factors associated with R
                    <sub>0</sub>
                </title>
                <p>Across different study periods, age groups, and bias levels of serosurveys, pooled R
                    <sub>0</sub> estimates mostly presented a right-skewed distribution and often peaked at a value &lt;10 (
                    <xref ref-type="fig" rid="f2">Figure 2</xref>). However, R
                    <sub>0</sub> estimates based on the same study feature could still have wide variation, as seen in the multimodal curves of pooled R
                    <sub>0</sub> estimates in studies with a larger number of seroprevalence data points and low bias in sampling, measuring, and reporting measles serology. Our findings suggested that the survey period and age of populations may be associated with the peaks and shapes of pooled R
                    <sub>0</sub> distributions. The median R
                    <sub>0</sub> estimates were 8.23 (interquartile range (IQR)=10.2) in the serostudies conducted prior to 1990, 13.0 (IQR=13.0) in 1991&#x2013;2000, 5.50 (IQR=5.76) in 2001&#x2013;2010, and 6.39 (IQR=5.42) in 2011&#x2013;2019, showing a smaller value and narrower variability in the estimates based on more recent studies. A wider variation and a greater value were also found in R
                    <sub>0</sub> estimates based on the studies involving only children (median=13.0, IQR=12.1) compared to those with both children and adults (median=6.43, IQR=7.24).</p>
                <fig fig-type="figure" id="f2" orientation="portrait" position="float">
                    <label>Figure 2. </label>
                    <caption>
                        <title>Pooled measles R
                            <sub>0</sub> by (A) survey period, (B) number of age-specific data points per survey, (C) age groups of survey population, (D) bias in sampling study population, (E) bias in measuring serology, and (F) bias in reporting serological results.</title>
                        <p>Density of 10,000 bootstrapped R
                            <sub>0</sub> estimates by each selected feature is presented.</p>
                    </caption>
                    <graphic id="gr2" orientation="portrait" position="float" xlink:href="https://verixiv-files.f1000.com/manuscripts/1213/18ac17ed-152f-4652-98fd-ff2ebb6d233f_figure2.gif"/>
                </fig>
                <p>In exploring other country- and year-specific factors in the broader context, we identified a weak negative correlation (R=&#x2013;0.1917, p=0.0117) between the posterior median R
                    <sub>0</sub> and Human Development Index, suggesting that measles transmissibility decreased with improvement in health and socioeconomics. Nonetheless, no significant correlation was found between median R
                    <sub>0</sub> and average household size (R=0.1472, p=0.0577) or population density (R=&#x2013;0.0181, p=0.8139). Based on the univariate log-linear regression analysis on R
                    <sub>0</sub>, Human Development Index (HDI) was a significant predictor but with limited explanation of the variance (R-squared=2.3%, 
                    <xref ref-type="fig" rid="f3">Figure 3</xref>).</p>
                <fig fig-type="figure" id="f3" orientation="portrait" position="float">
                    <label>Figure 3. </label>
                    <caption>
                        <title>Median R
                            <sub>0</sub> in relation to (A) Human Development Index, (B) household size and (C) population density.</title>
                        <p>Regression coefficients and R squared were derived based on univarate log-linear models. Where available, data were matched to the year each serosurvey was conducted.</p>
                    </caption>
                    <graphic id="gr3" orientation="portrait" position="float" xlink:href="https://verixiv-files.f1000.com/manuscripts/1213/18ac17ed-152f-4652-98fd-ff2ebb6d233f_figure3.gif"/>
                </fig>
            </sec>
            <sec id="sec5">
                <title>Estimated measles cases</title>
                <p>WHO data on reported measles cases was not available especially in the early years when case surveillance and notification systems had not yet been fully operated. Among 155 studies included for comparison, 60% of our median model-based estimates (n=93) show greater numbers than the reported cases. Assuming these model-based estimates to reflect the actual magnitude of measles incidence, we found a proportion of 1.27% measles cases were reported to WHO (25 
                    <sup>th</sup>&#x2013;75
                    <sup>th</sup> percentiles: 0.56%&#x2013;2.94%). However, in 32 studies, we estimated fewer than 100 cases with upper bound model estimates, while the WHO data reported more than 1000 cases. We also noted that the majority of serostudies where model-estimated measles cases were lower than the WHO-reported cases were conducted in China (n=18) and having severe bias in measuring and reporting serological results.</p>
                <p>The comparison with the Global Burden of Disease (GBD) 2021 Study was available for 134 studies (78%) conducted between 1988&#x2013;2019. Our model-based case estimates did not show consistent prediction trends with the GBD incidence estimates, with 60% of included studies having no overlapping between the two interval estimates.</p>
            </sec>
            <sec id="sec6">
                <title>Sensitivity analysis</title>
                <p>Fitting the DynaMICE model to the age-specific seroprevalence data, we presented a median RMSE of 6.93% (25
                    <sup>th</sup>&#x2013;75
                    <sup>th</sup> percentiles: 3.37%&#x2013;12.2%). The sensitivity analysis was restricted to the top 80% best-fit studies, where RMSEs are less than 14.5%. Compared to the main analysis with a full set of studies, pooled R
                    <sub>0</sub> estimates of the restricted studies showed consistent magnitude and wide intervals for WHO regions, study period, age groups, and serological measurement and reporting biases (
                    <xref ref-type="fig" rid="f4">Figure 4</xref>).</p>
                <fig fig-type="figure" id="f4" orientation="portrait" position="float">
                    <label>Figure 4. </label>
                    <caption>
                        <title>Pooled measles R
                            <sub>0</sub> estimates in the full set of studies and restricted set of studies with better model fitness.</title>
                        <p>Median and 95% credible intervals of pooled R
                            <sub>0</sub> estimates from studies with each selected feature are presented. Arrows indicate the upper estimates exceed 50.</p>
                    </caption>
                    <graphic id="gr4" orientation="portrait" position="float" xlink:href="https://verixiv-files.f1000.com/manuscripts/1213/18ac17ed-152f-4652-98fd-ff2ebb6d233f_figure4.gif"/>
                </fig>
            </sec>
        </sec>
        <sec id="sec7" sec-type="discussion">
            <title>Discussion</title>
            <p>This is the first analysis to systematically estimate measles R
                <sub>0</sub> across LMICs using age-specific serological data. Based on dynamic model simulation of measles transmission and vaccination, we identified great variability of R
                <sub>0</sub> across 172 study settings. Our model-based measles R
                <sub>0</sub> estimates were mostly below the commonly cited range between 12&#x2013;18 and demonstrated large heterogeneity even within the same countries or WHO regions. The value of R
                <sub>0</sub> tended to be higher in serosurveys conducted in earlier years prior to 2000 or involved with the adult population, and it was also found to be negatively associated with Human Development Index. Furthermore, the serology-informed R
                <sub>0</sub> estimates we applied to quantify the measles incidence suggested a low reporting of disease burden in the surveillance system.</p>
            <p>The great variability of measles R
                <sub>0</sub> identified in our analysis was also reported in a comprehensive review by Guerra et al.,
                <sup>
                    <xref ref-type="bibr" rid="ref4">4</xref>
                </sup> where the extracted estimates range from 1.43 to 770. Both our analysis and Guerra&#x2019;s review found that most R
                <sub>0</sub> estimates to be outside of the commonly cited range of 12&#x2013;18, which was derived from a single study taking early data in England and New York.
                <sup>
                    <xref ref-type="bibr" rid="ref3">3</xref>
                </sup> However, Guerra et al.
                <sup>
                    <xref ref-type="bibr" rid="ref4">4</xref>
                </sup> summarised estimates from independent modelling studies based on surveillance or/and serology data sources, and it was therefore not able to separate to what extent the R
                <sub>0</sub> variability could be attributed to the methodological differences. Another multi-country analysis based on the same model structure reported more than five times greater measles transmission rates in the United Kingdom than in Ethiopia, but only two LMICs were included and the homogenous mixing assumption limited further inspection of the heterogeneity of transmissibility.
                <sup>
                    <xref ref-type="bibr" rid="ref9">9</xref>
                </sup> Instead, our analysis utilised all the available serological data in LMICs conducted prior to 2019 and the DynaMICE model incorporated with fine age structure and country-specific demographics and contact patterns contributing to measles immunity. With a systematic framework for serosurvey selection and model parameterisation, we improved the quantification of measles R
                <sub>0</sub> heterogeneity and assessment of potential contributing factors.</p>
            <p>Advancements in healthcare, education and living standards could partly explain the variability of measles R
                <sub>0</sub>, as suggested in previous studies and our regression analysis with HDI.
                <sup>
                    <xref ref-type="bibr" rid="ref4">4</xref>
                </sup> However, we did not find a correlation between R
                <sub>0</sub> and average household size and population density. In DynaMICE, the effect of household structure and crowdedness on age-related mixing across countries may have been partially adjusted through the incorporation of synthetic social contact matrices, which were constructed with contact diary surveys and demography data.
                <sup>
                    <xref ref-type="bibr" rid="ref7">7</xref>,
                    <xref ref-type="bibr" rid="ref10">10</xref>
                </sup> In such cases, other indicators like clustering of susceptibility or proportion of large-size households could be more informative to explain the complicated mechanism of household transmission.
                <sup>
                    <xref ref-type="bibr" rid="ref11">11</xref>,
                    <xref ref-type="bibr" rid="ref12">12</xref>
                </sup> In addition to selecting appropriate indicators, addressing the interactions between them is essential to disentangle the heterogeneity in measles transmissibility. An advanced understanding of the R
                <sub>0</sub> heterogeneity will help characterise measles transmission in settings where serodata are unavailable
                <sup>
                    <xref ref-type="bibr" rid="ref11">11</xref>
                </sup> and update disease burden under evolving settings.</p>
            <p>We estimated annual numbers of measles cases by leveraging the strength of serological data in being able to capture disease incidence regardless of symptom occurrence and access to healthcare.
                <sup>
                    <xref ref-type="bibr" rid="ref13">13</xref>
                </sup> The comparison between our serology-based burden estimates and WHO notification data suggested that fewer than 5% of cases were reported in most study settings. The low level of reporting was also identified in a previous study.
                <sup>
                    <xref ref-type="bibr" rid="ref14">14</xref>
                </sup> We also compared our measles case numbers with GBD 2021, the only publicly available incidence estimates at the country level. We found limited alignment in the magnitude of disease burden between the two studies, largely owing to methodological differences&#x2014;the GBD estimates were derived from a mixed-effects linear regression model with routine and campaign MCV coverage as covariates and fitted to the WHO notification data.
                <sup>
                    <xref ref-type="bibr" rid="ref15">15</xref>
                </sup>
            </p>
            <p>Our study has several limitations. First, we assumed that each serosurvey was nationally representative and used country-level demography and vaccine coverage inputs for model fitting. However, some serosurveys were conducted in geographically restricted populations due to time, cost, and personnel constraints, especially in LMICs.
                <sup>
                    <xref ref-type="bibr" rid="ref16">16</xref>
                </sup> A mismatch of population representativeness between serodata and model inputs could lead to inaccurate estimation of measles R
                <sub>0</sub> and case burden. Whilst high-spatial-resolution estimates of routine MCV coverage are available for addressing subnational heterogeneity,
                <sup>
                    <xref ref-type="bibr" rid="ref17">17</xref>
                </sup> local data on campaign coverage, contact patterns, and population structure are still limited in LMICs. In many serostudies, reported information was insufficient in defining the geographic areas of the catchment population.
                <sup>
                    <xref ref-type="bibr" rid="ref18">18</xref>
                </sup>
            </p>
            <p>Second, we assumed that R
                <sub>0</sub> and social contact rates were constant over time, even though the different R
                <sub>0</sub> values in each serosurvey for the same country suggested a potential temporal trend. Even though we addressed some of the temporal effects in DynaMICE through yearly inputs of population age structure, social and environmental factors can still contribute to the temporal variation of measles transmissibility. For example, factors such as urbanisation, education, infrastructure improvement, and major public health events like COVID-19 may alter measles transmission and contact behaviour in particular settings.
                <sup>
                    <xref ref-type="bibr" rid="ref4">4</xref>,
                    <xref ref-type="bibr" rid="ref19">19</xref>
                </sup> The temporal trend of R
                <sub>0</sub>is likely to be non-linear, sensitive to occasional events, and jointly driven by multiple time-varying components. Repeated cross-sectional serosurveys in the same population, supported by longitudinal data for model inputs, could facilitate the characterisation of the temporal trends in measles transmissibility.</p>
            <p>Third, the serostudies included in our analysis used different laboratory methods and antibody levels to define seropositivity, in part likely due to weak evidence for an antibody threshold correlating to protection against measles infection.
                <sup>
                    <xref ref-type="bibr" rid="ref20">20</xref>
                </sup> Although we excluded studies with a critical bias in measuring serology,
                <sup>
                    <xref ref-type="bibr" rid="ref18">18</xref>
                </sup> we were not able to consider the sensitivity and specificity of the laboratory methods because of limited information on whether laboratory protocols were specifically followed, or on sample quality at time of testing. We also did not perform sensitivity analyses across studies that used different thresholds to define seroprotection. In addition to the performance of serological tools, serosurveys with sufficient age-specific data points and broad age range (e.g. both children and adults) will help address the changing vaccination strategies over time and reduce bias and uncertainty in estimating measles R
                <sub>0</sub>.</p>
            <p>In conclusion, we have comprehensively explored measles R
                <sub>0</sub> across LMICs by fitting to all the available age-specific seroprevalence data extracted from a systematic review, accounting for country-specific demographic characteristics, contact patterns, and immunisation programmes using the DynaMICE model. We have illustrated the significant variability of measles R
                <sub>0</sub> estimates across different settings. We demonstrated that the variation in R
                <sub>0</sub> is not purely driven by differences in model structure, but also dependent on serosurvey features and country-specific demographic and developmental characteristics. Heterogeneity in measles is a crucial consideration when modelling epidemics and planning intervention and vaccination strategies in LMICs. Further research to improve the quality, granularity, and completeness of data collection for serostudies and other model inputs will enhance the estimation of measles R
                <sub>0</sub> and understanding of its key determinants.</p>
        </sec>
        <sec id="sec8">
            <title>Materials &amp; methods</title>
            <sec id="sec9">
                <title>Serological data</title>
                <p>We obtained 221 articles from a 2024 systematic review by Sbarra et al. that summarised the scope and bias in population-level measles serosurveys.
                    <sup>
                        <xref ref-type="bibr" rid="ref18">18</xref>
                    </sup> Surveys that did not provide information on the age distribution of the sampled population (n=16) were excluded. We also excluded survey data from non-representative populations, such as immunocompromised patients, pre-term babies, and refugees or border populations (n=21). Based on the bias assessment in the previous review,
                    <sup>
                        <xref ref-type="bibr" rid="ref18">18</xref>
                    </sup> we excluded surveys having a critical bias in the representativeness of the study populations and in the validity of the tools used to measure serology (n=19). We did not exclude serosurveys that were rated lower quality because they had insufficient information when reporting serological results, because a standard reporting style was not widely used, especially in early studies, and because reporting completeness would not necessarily reflect the implementation quality of serosurveys.</p>
                <p>From the 165 included articles, a total of 172 unique studies were defined with a unique combination of country, survey year, and first author. Where only the publication years were available, we approximated the survey years using the average duration between the survey and publication years in studies with complete information (3.7 years). In mother-newborn studies examining both the samples from mother-newborn pairs, we extracted the data only from mothers. Most such studies reported very high correlation in the antibody concentration between mother and newborn. The age-specific seroprevalence estimates were derived from surveys conducted between 1963 and 2019 in 57 LMICs, with the majority found in China, India, T&#x00fc;rkiye, and Brazil. Most of the included studies (n=132, 76.7%) consisted of seroprevalence estimates across multiple age groups, and only a small proportion did not cover children (n=21, 12.2%).</p>
            </sec>
            <sec id="sec10">
                <title>Model fitting</title>
                <p>We fitted DynaMICE to each unique set of age-specific seroprevalence data per country-time period using MCMC with component-wise Metropolis algorithm and random order for updating parameters in each blocked iteration.
                    <sup>
                        <xref ref-type="bibr" rid="ref21">21</xref>
                    </sup> Three parameters&#x2014;R
                    <sub>0</sub>, duration of maternal immunity protection, and vaccine effectiveness for the first dose of measles-containing vaccine (MCV)&#x2014;were varied. These parameters were assumed to be constant over time within the same seroprevalence study and were drawn from prior distributions based on previous literature (
                    <xref ref-type="table" rid="T1">
Table 1</xref>). For 69 serosurveys conducted prior to country-specific MCV introduction, we removed the parameter for vaccine effectiveness and fit the seroprevalence data with a two-parameter model. We constructed the likelihood by assuming the seroprevalence in each age group to be independently binomially distributed. We ran two chains with different starting parameter values and conducted diagnostic checks for convergence of chains and goodness of fit. We obtained 500 posterior samples for each study and calculated their medians and 95% CrI. All analysis and visualisation were conducted in R 4.2.3, and the computer codes can be found at 
                    <ext-link ext-link-type="uri" xlink:href="https://zenodo.org/records/15424461">https://zenodo.org/records/15424461</ext-link>.</p>
                <table-wrap id="T1" orientation="portrait" position="float">
                    <label>
Table 1. </label>
                    <caption>
                        <title>Prior distribution of parameters for model fitting.</title>
                    </caption>
                    <table content-type="article-table" frame="hsides">
                        <thead>
                            <tr>
                                <th align="left" colspan="1" rowspan="1" valign="top">
Estimated parameter</th>
                                <th align="left" colspan="1" rowspan="1" valign="top">
Prior distribution, median (5
                                    <sup>th</sup>&#x2013;95
                                    <sup>th</sup> percentiles)</th>
                                <th align="left" colspan="1" rowspan="1" valign="top">
Source and note</th>
                            </tr>
                        </thead>
                        <tbody>
                            <tr>
                                <td align="left" colspan="1" rowspan="1" valign="top">Basic reproduction number (R
                                    <sub>0</sub>)</td>
                                <td align="left" colspan="1" rowspan="1" valign="top">Log-normal, 13.1 (5.44&#x2013;31.8)</td>
                                <td align="left" colspan="1" rowspan="1" valign="top">Guerra et al.
                                    <sup>
                                        <xref ref-type="bibr" rid="ref27">27</xref>
                                    </sup>
                                </td>
                            </tr>
                            <tr>
                                <td align="left" colspan="1" rowspan="1" valign="top">Duration of maternal immunity, months</td>
                                <td align="left" colspan="1" rowspan="1" valign="top">Truncated log-normal, 2.84 (1.24&#x2013;6.45) minimum: 1, maximum: 12</td>
                                <td align="left" colspan="1" rowspan="1" valign="top">Leuridan et al.
                                    <sup>
                                        <xref ref-type="bibr" rid="ref28">28</xref>
                                    </sup>
                                    <break/>Fit to the longitudinal study</td>
                            </tr>
                            <tr>
                                <td align="left" colspan="1" rowspan="1" valign="top">Vaccine effectiveness at 9 months old (only for surveys conducted in the post-vaccination era)</td>
                                <td align="left" colspan="1" rowspan="1" valign="top">Truncated normal, 77.5% (49.3%-100%) minimum: 26%, maximum: 100%</td>
                                <td align="left" colspan="1" rowspan="1" valign="top">Hughes et al.
                                    <sup>
                                        <xref ref-type="bibr" rid="ref29">29</xref>
                                    </sup>
                                    <break/>Fit to the reported range and further capped by two-dose effectiveness 98%</td>
                            </tr>
                        </tbody>
                    </table>
                </table-wrap>
                <p>In our model-based outputs, we added up the non-susceptible populations&#x2014;maternally immune, infectious, and recovered sub-populations (compartments)
                    <sup>
                        <xref ref-type="bibr" rid="ref7">7</xref>
                    </sup> to calculate the seroprevalence in the corresponding age groups and calendar years. We included all the non-susceptible states because IgG antibodies, as the detection target in serosurveys, cannot distinguish between maternally acquired, infection-acquired and vaccine-induced immunity. For a given parameter set, models were first simulated using the country-specific demographics in 1980
                    <sup>
                        <xref ref-type="bibr" rid="ref22">22</xref>
                    </sup> and assumed no implementation of vaccination until they reached equilibrium, and then begun with annual inputs of demographics and national-level MCV coverage estimates
                    <sup>
                        <xref ref-type="bibr" rid="ref1">1</xref>
                    </sup> from 1980, when the coverage data is earliest available. For serosurveys conducted before 1980, we extracted the model estimates in 1980, assuming that seroprevalence was likely similar during earlier years given low coverage of vaccination and other control measures for measles.</p>
            </sec>
            <sec id="sec11">
                <title>Factors associated with R
                    <sub>0</sub>
                </title>
                <p>Measles R
                    <sub>0</sub> is influenced by demographic, socioeconomic, and environmental factors, which change contact behaviours and transmission risk per contact in the population.
                    <sup>
                        <xref ref-type="bibr" rid="ref4">4</xref>
                    </sup> Although the DynaMICE model has accounted for country-specific age structure and age-dependent contact patterns, other factors could further contribute to the temporal and geographical variation of measles R
                    <sub>0</sub>. In addition, the survey quality and available details of seroprevalence data also affect the estimation of R
                    <sub>0</sub> through fitting to seroprevalence data. To understand these dependencies, we evaluated the association between R
                    <sub>0</sub> estimates and different study features, including survey period, age of the survey population, number of age-specific data points, and biases in sampling, measurement and reporting serology.
                    <sup>
                        <xref ref-type="bibr" rid="ref18">18</xref>
                    </sup> By drawing 10,000 bootstrapped samples with replacement, we pooled R
                    <sub>0</sub> estimates in studies with the same selected categories and compared their distributions.</p>
                <p>We assessed the association of median R
                    <sub>0</sub> for each study with average household size,
                    <sup>
                        <xref ref-type="bibr" rid="ref23">23</xref>
                    </sup> population density,
                    <sup>
                        <xref ref-type="bibr" rid="ref22">22</xref>
                    </sup> and HDI.
                    <sup>
                        <xref ref-type="bibr" rid="ref24">24</xref>
                    </sup> HDI is a country-specific and time-dependent summary measure combining the dimensions of health and longevity, education and living standard. When these data were missing at specific survey years, we performed linear interpolation for household size and imputed the HDI value at the closest year on a country basis.</p>
            </sec>
            <sec id="sec12">
                <title>Estimated measles cases</title>
                <p>Using the posterior parameters of R
                    <sub>0</sub>, duration of maternal immunity, and vaccine effectiveness, we estimated the annual numbers of measles cases during the survey year and the two years before and after. We calculated the mean annual cases over the five years to reduce the influence of year-to-year fluctuations in the time series. We compared the model-based case estimates with the WHO country-specific reported cases
                    <sup>
                        <xref ref-type="bibr" rid="ref25">25</xref>
                    </sup> to assess their consistency in measuring measles burden and the potential issue of underreporting. We also compared our model-based estimates to measles incidence estimates in the GBD Study 2021,
                    <sup>
                        <xref ref-type="bibr" rid="ref26">26</xref>
                    </sup> as it is the key publicly available resource for health decision makers.</p>
            </sec>
            <sec id="sec13">
                <title>Sensitivity analysis</title>
                <p>We investigated the goodness-of-fit of the two-parameter and three-parameter models by calculating the RMSE between the observed seroprevalence data and median posterior predictions at specified age groups in each study. To assess how model fit affects the estimation of R
                    <sub>0</sub>, we ranked the RMSE and conducted a sensitivity analysis on the pooled R
                    <sub>0</sub> results after removing 20% of the studies with the highest error size.</p>
            </sec>
        </sec>
    </body>
    <back>
        <sec id="sec16" sec-type="data-availability">
            <title>Data availability statement</title>
            <p>Zenodo: dynamice_seroR0, 
                <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.5281/zenodo.15424461">https://doi.org/10.5281/zenodo.15424461</ext-link>.
                <sup>
                    <xref ref-type="bibr" rid="ref30">30</xref>
                </sup> This repository contains the curated datasets of measles vaccine coverage and seroprevalence estimates, and the computer codes of this manuscript.</p>
            <p>Data are available under the terms of the 
                <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution 4.0 International license</ext-link> (CC-BY 4.0).</p>
        </sec>
        <ack>
            <title>Acknowledgement</title>
            <p>This work was carried out as part of the Vaccine Impact Modelling Consortium (
                <ext-link ext-link-type="uri" xlink:href="http://www.vaccineimpact.org">www.vaccineimpact.org</ext-link>), but the views expressed are those of the authors and not necessarily those of the Consortium or its funders. The funders were given the opportunity to review this paper prior to publication, but the final decision on the content of the publication was taken by the authors.</p>
        </ack>
        <ref-list>
            <title>References</title>
            <ref id="ref1">
                <label>1</label>
                <mixed-citation publication-type="other">
                    <collab>World Health Organization</collab>:
                    <article-title>Immunization data.</article-title>
                </mixed-citation>
            </ref>
            <ref id="ref2">
                <label>2</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Minta</surname>
                            <given-names>AA</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Progress Toward Measles Elimination - Worldwide, 2000-2023.</article-title>
                    <source>

                        <italic toggle="yes">MMWR Morb. Mortal Wkly. Rep.</italic>
</source>
                    <year>2024</year>;<volume>73</volume>:<fpage>1036</fpage>&#x2013;<lpage>1042</lpage>.
                    <pub-id pub-id-type="pmid">39541251</pub-id>
                    <pub-id pub-id-type="doi">10.15585/mmwr.mm7345a4</pub-id>
                    <pub-id pub-id-type="pmcid">PMC11576049</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref3">
                <label>3</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Anderson</surname>
                            <given-names>RM</given-names>
                        </name>

                        <name name-style="western">
                            <surname>May</surname>
                            <given-names>RM</given-names>
                        </name>
</person-group>:
                    <article-title>Directly transmitted infections diseases: control by vaccination.</article-title>
                    <source>

                        <italic toggle="yes">Science.</italic>
</source>
                    <year>1982</year>;<volume>215</volume>:<fpage>1053</fpage>&#x2013;<lpage>1060</lpage>.
                    <pub-id pub-id-type="doi">10.1126/science.7063839</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref4">
                <label>4</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Guerra</surname>
                            <given-names>FM</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>The basic reproduction number (R0) of measles: a systematic review.</article-title>
                    <source>

                        <italic toggle="yes">Lancet Infect. Dis.</italic>
</source>
                    <year>2017</year>;<volume>17</volume>:<fpage>e420</fpage>&#x2013;<lpage>e428</lpage>.
                    <pub-id pub-id-type="pmid">28757186</pub-id>
                    <pub-id pub-id-type="doi">10.1016/S1473-3099(17)30307-9</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref5">
                <label>5</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Gibbons</surname>
                            <given-names>CL</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Measuring underreporting and under-ascertainment in infectious disease datasets: a comparison of methods.</article-title>
                    <source>

                        <italic toggle="yes">BMC Public Health.</italic>
</source>
                    <year>2014</year>;<volume>14</volume>:<fpage>147</fpage>.
                    <pub-id pub-id-type="pmid">24517715</pub-id>
                    <pub-id pub-id-type="doi">10.1186/1471-2458-14-147</pub-id>
                    <pub-id pub-id-type="pmcid">PMC4015559</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref6">
                <label>6</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Auzenbergs</surname>
                            <given-names>M</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Health effects of routine measles vaccination and supplementary immunisation activities in 14 high-burden countries: a Dynamic Measles Immunization Calculation Engine (DynaMICE) modelling study.</article-title>
                    <source>

                        <italic toggle="yes">Lancet Glob. Health.</italic>
</source>
                    <year>2023</year>;<volume>11</volume>:<fpage>e1194</fpage>&#x2013;<lpage>e1204</lpage>.
                    <pub-id pub-id-type="pmid">37474227</pub-id>
                    <pub-id pub-id-type="doi">10.1016/S2214-109X(23)00220-6</pub-id>
                    <pub-id pub-id-type="pmcid">PMC10369016</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref7">
                <label>7</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Fu</surname>
                            <given-names>H</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Effect of evidence updates on key determinants of measles vaccination impact: a DynaMICE modelling study in ten high-burden countries.</article-title>
                    <source>

                        <italic toggle="yes">BMC Med.</italic>
</source>
                    <year>2021</year>;<volume>19</volume>:<fpage>281</fpage>.
                    <pub-id pub-id-type="pmid">34784922</pub-id>
                    <pub-id pub-id-type="doi">10.1186/s12916-021-02157-4</pub-id>
                    <pub-id pub-id-type="pmcid">PMC8594955</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref8">
                <label>8</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Verguet</surname>
                            <given-names>S</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Controlling measles using supplemental immunization activities: a mathematical model to inform optimal policy.</article-title>
                    <source>

                        <italic toggle="yes">Vaccine.</italic>
</source>
                    <year>2015</year>;<volume>33</volume>:<fpage>1291</fpage>&#x2013;<lpage>1296</lpage>.
                    <pub-id pub-id-type="pmid">25541214</pub-id>
                    <pub-id pub-id-type="doi">10.1016/j.vaccine.2014.11.050</pub-id>
                    <pub-id pub-id-type="pmcid">PMC4336184</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref9">
                <label>9</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Trentini</surname>
                            <given-names>F</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Poletti</surname>
                            <given-names>P</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Merler</surname>
                            <given-names>S</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Measles immunity gaps and the progress towards elimination: a multi-country modelling analysis.</article-title>
                    <source>

                        <italic toggle="yes">Lancet Infect. Dis.</italic>
</source>
                    <year>2017</year>;<volume>17</volume>:<fpage>1089</fpage>&#x2013;<lpage>1097</lpage>.
                    <pub-id pub-id-type="pmid">28807627</pub-id>
                    <pub-id pub-id-type="doi">10.1016/S1473-3099(17)30421-8</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref10">
                <label>10</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Prem</surname>
                            <given-names>K</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Cook</surname>
                            <given-names>AR</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Jit</surname>
                            <given-names>M</given-names>
                        </name>
</person-group>:
                    <article-title>Projecting social contact matrices in 152 countries using contact surveys and demographic data.</article-title>
                    <source>

                        <italic toggle="yes">PLoS Comput. Biol.</italic>
</source>
                    <year>2017</year>;<volume>13</volume>:<fpage>e1005697</fpage>.
                    <pub-id pub-id-type="pmid">28898249</pub-id>
                    <pub-id pub-id-type="doi">10.1371/journal.pcbi.1005697</pub-id>
                    <pub-id pub-id-type="pmcid">PMC5609774</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref11">
                <label>11</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Papadopoulos</surname>
                            <given-names>T</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Vynnycky</surname>
                            <given-names>E</given-names>
                        </name>
</person-group>:
                    <article-title>Estimates of the basic reproduction number for rubella using seroprevalence data and indicator-based approaches.</article-title>
                    <source>

                        <italic toggle="yes">PLoS Comput. Biol.</italic>
</source>
                    <year>2022</year>;<volume>18</volume>:<fpage>e1008858</fpage>.
                    <pub-id pub-id-type="pmid">35239641</pub-id>
                    <pub-id pub-id-type="doi">10.1371/journal.pcbi.1008858</pub-id>
                    <pub-id pub-id-type="pmcid">PMC8893344</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref12">
                <label>12</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Kuylen</surname>
                            <given-names>E</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Willem</surname>
                            <given-names>L</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Broeckhove</surname>
                            <given-names>J</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Clustering of susceptible individuals within households can drive measles outbreaks: an individual-based model exploration.</article-title>
                    <source>

                        <italic toggle="yes">Sci. Rep.</italic>
</source>
                    <year>2020</year>;<volume>10</volume>:<fpage>19645</fpage>.
                    <pub-id pub-id-type="pmid">33184409</pub-id>
                    <pub-id pub-id-type="doi">10.1038/s41598-020-76746-3</pub-id>
                    <pub-id pub-id-type="pmcid">PMC7665185</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref13">
                <label>13</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Haselbeck</surname>
                            <given-names>AH</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Serology as a Tool to Assess Infectious Disease Landscapes and Guide Public Health Policy.</article-title>
                    <source>

                        <italic toggle="yes">Pathogens.</italic>
</source>
                    <year>2022</year>;<volume>11</volume>.
                    <pub-id pub-id-type="pmid">35889978</pub-id>
                    <pub-id pub-id-type="doi">10.3390/pathogens11070732</pub-id>
                    <pub-id pub-id-type="pmcid">PMC9323579</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref14">
                <label>14</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Eilertson</surname>
                            <given-names>KE</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Fricks</surname>
                            <given-names>J</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Ferrari</surname>
                            <given-names>MJ</given-names>
                        </name>
</person-group>:
                    <article-title>Estimation and prediction for a mechanistic model of measles transmission using particle filtering and maximum likelihood estimation.</article-title>
                    <source>

                        <italic toggle="yes">Stat. Med.</italic>
</source>
                    <year>2019</year>;<volume>38</volume>:<fpage>4146</fpage>&#x2013;<lpage>4158</lpage>.
                    <pub-id pub-id-type="pmid">31290184</pub-id>
                    <pub-id pub-id-type="doi">10.1002/sim.8290</pub-id>
                    <pub-id pub-id-type="pmcid">PMC6771900</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref15">
                <label>15</label>
                <mixed-citation publication-type="other">
                    <collab>Institute for Health Metrics and Evaluation</collab>:
                    <article-title>Global Burden of Disease (GBD) 2021 Methods Appendices: Measles.</article-title>
                    <year>2024</year>.</mixed-citation>
            </ref>
            <ref id="ref16">
                <label>16</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Thompson</surname>
                            <given-names>KM</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Odahowski</surname>
                            <given-names>CL</given-names>
                        </name>
</person-group>:
                    <article-title>Systematic Review of Measles and Rubella Serology Studies.</article-title>
                    <source>

                        <italic toggle="yes">Risk Anal.</italic>
</source>
                    <year>2016</year>;<volume>36</volume>:<fpage>1459</fpage>&#x2013;<lpage>1486</lpage>.
                    <pub-id pub-id-type="pmid">26077609</pub-id>
                    <pub-id pub-id-type="doi">10.1111/risa.12430</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref17">
                <label>17</label>
                <mixed-citation publication-type="journal">
                    <collab>Local Burden of Disease Vaccine Coverage Collaborators</collab>:
                    <article-title>Mapping routine measles vaccination in low- and middle-income countries.</article-title>
                    <source>

                        <italic toggle="yes">Nature.</italic>
</source>
                    <year>2020</year>;<volume>589</volume>:<fpage>415</fpage>&#x2013;<lpage>419</lpage>.
                    <pub-id pub-id-type="pmid">33328634</pub-id>
                    <pub-id pub-id-type="doi">10.1038/s41586-020-03043-4</pub-id>
                    <pub-id pub-id-type="pmcid">PMC7739806</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref18">
                <label>18</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Sbarra</surname>
                            <given-names>AN</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Evaluating Scope and Bias of Population-Level Measles Serosurveys: A Systematized Review and Bias Assessment.</article-title>
                    <source>

                        <italic toggle="yes">Vaccines (Basel).</italic>
</source>
                    <year>2024</year>;<volume>12</volume>.
                    <pub-id pub-id-type="doi">10.3390/vaccines12060585</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref19">
                <label>19</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Delamater</surname>
                            <given-names>PL</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Street</surname>
                            <given-names>EJ</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Leslie</surname>
                            <given-names>TF</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Complexity of the Basic Reproduction Number (R(0)).</article-title>
                    <source>

                        <italic toggle="yes">Emerg. Infect. Dis.</italic>
</source>
                    <year>2019</year>;<volume>25</volume>:<fpage>1</fpage>&#x2013;<lpage>4</lpage>.
                    <pub-id pub-id-type="pmid">30560777</pub-id>
                    <pub-id pub-id-type="doi">10.3201/eid2501.171901</pub-id>
                    <pub-id pub-id-type="pmcid">PMC6302597</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref20">
                <label>20</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Bolotin</surname>
                            <given-names>S</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>What Is the Evidence to Support a Correlate of Protection for Measles? A Systematic Review.</article-title>
                    <source>

                        <italic toggle="yes">J. Infect. Dis.</italic>
</source>
                    <year>2020</year>;<volume>221</volume>:<fpage>1576</fpage>&#x2013;<lpage>1583</lpage>.
                    <pub-id pub-id-type="pmid">31674648</pub-id>
                    <pub-id pub-id-type="doi">10.1093/infdis/jiz380</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref21">
                <label>21</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Metropolis</surname>
                            <given-names>N</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Rosenbluth</surname>
                            <given-names>AW</given-names>
                        </name>

                        <name name-style="western">
                            <surname>Rosenbluth</surname>
                            <given-names>MN</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Equation of State Calculations by Fast Computing Machines.</article-title>
                    <source>

                        <italic toggle="yes">J. Chem. Phys.</italic>
</source>
                    <year>1953</year>;<volume>21</volume>:<fpage>1087</fpage>&#x2013;<lpage>1092</lpage>.
                    <pub-id pub-id-type="doi">10.1063/1.1699114</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref22">
                <label>22</label>
                <mixed-citation publication-type="other">
                    <collab>United Nations Population Division</collab>:
                    <article-title>World Population Prospects.</article-title>
                </mixed-citation>
            </ref>
            <ref id="ref23">
                <label>23</label>
                <mixed-citation publication-type="other">
                    <article-title>United Nations Population Division, Household Size and Composition.</article-title>
                    <ext-link ext-link-type="uri" xlink:href="https://www.un.org/development/desa/pd/sites/www.un.org.development.desa.pd/files/undesa_pd_2022_hh-size-composition.xlsx">Reference Source</ext-link>
                </mixed-citation>
            </ref>
            <ref id="ref24">
                <label>24</label>
                <mixed-citation publication-type="other">
                    <collab>United Nations Development Programme</collab>:
                    <article-title>Human Development Index.</article-title>
                    <year>2024</year>.</mixed-citation>
            </ref>
            <ref id="ref25">
                <label>25</label>
                <mixed-citation publication-type="other">
                    <collab>World Health Organization</collab>:
                    <article-title>Provisional monthly measles and rubella data. in 1980-2022.</article-title>
                </mixed-citation>
            </ref>
            <ref id="ref26">
                <label>26</label>
                <mixed-citation publication-type="book">
                    <collab>Global Burden of Disease Collaborative Network</collab>:
                    <source>

                        <italic toggle="yes">Global Burden of Disease Study 2021 (GBD 2021) Results.</italic>
</source>
                    <publisher-loc>Seattle</publisher-loc>:
                    <publisher-name>Institute for Health Metrics and Evaluation (IHME)</publisher-name>;<year>2022</year>.</mixed-citation>
            </ref>
            <ref id="ref27">
                <label>27</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Guerra</surname>
                            <given-names>FM</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Waning of measles maternal antibody in infants in measles elimination settings - A systematic literature review.</article-title>
                    <source>

                        <italic toggle="yes">Vaccine.</italic>
</source>
                    <year>2018</year>;<volume>36</volume>:<fpage>1248</fpage>&#x2013;<lpage>1255</lpage>.
                    <pub-id pub-id-type="pmid">29398276</pub-id>
                    <pub-id pub-id-type="doi">10.1016/j.vaccine.2018.01.002</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref28">
                <label>28</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Leuridan</surname>
                            <given-names>E</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>Early waning of maternal measles antibodies in era of measles elimination: longitudinal study.</article-title>
                    <source>

                        <italic toggle="yes">BMJ.</italic>
</source>
                    <year>2010</year>;<volume>340</volume>:<fpage>c1626</fpage>.
                    <pub-id pub-id-type="pmid">20483946</pub-id>
                    <pub-id pub-id-type="doi">10.1136/bmj.c1626</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref29">
                <label>29</label>
                <mixed-citation publication-type="journal">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Hughes</surname>
                            <given-names>SL</given-names>
                        </name>

                        <etal/>
</person-group>:
                    <article-title>The effect of time since measles vaccination and age at first dose on measles vaccine effectiveness - A systematic review.</article-title>
                    <source>

                        <italic toggle="yes">Vaccine.</italic>
</source>
                    <year>2020</year>;<volume>38</volume>:<fpage>460</fpage>&#x2013;<lpage>469</lpage>.
                    <pub-id pub-id-type="pmid">31732326</pub-id>
                    <pub-id pub-id-type="doi">10.1016/j.vaccine.2019.10.090</pub-id>
                    <pub-id pub-id-type="pmcid">PMC6970218</pub-id>
                </mixed-citation>
            </ref>
            <ref id="ref30">
                <label>30</label>
                <mixed-citation publication-type="other">
                    <person-group person-group-type="author">

                        <name name-style="western">
                            <surname>Fu</surname>
                            <given-names>H</given-names>
                        </name>
</person-group>:
                    <article-title>dynamice_seroR0.</article-title>
                    <source>

                        <italic toggle="yes">Zenodo.</italic>
</source>
                    <year>2025</year>.
                    <pub-id pub-id-type="doi">10.5281/zenodo.15424461</pub-id>
                </mixed-citation>
            </ref>
        </ref-list>
    </back>
</article>
