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 "slug": "plaque-reduction-neutralization-test",
 "title": "Plaque reduction neutralization test",
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 "excerpt": "The plaque reduction neutralization test (PRNT) is a live-virus cell-culture assay measuring neutralizing antibodies, considered the laboratory standard and the WHO gold standard for dengue.",
 "snippet": "The plaque reduction neutralization test (PRNT) is a live-virus cell-culture assay measuring neutralizing antibodies, considered the laboratory standard and the WHO gold standard for dengue.",
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 "markdown": "# Plaque reduction neutralization test\n\nThe plaque reduction neutralization test (PRNT) is a live-virus cell-culture assay that quantifies functional neutralizing antibodies in serum by measuring the dilution that reduces the number of viral plaques in a cell monolayer. The result is a titer, reported as the reciprocal of the highest serum dilution that reduces plaque counts by a stated percentage, most often 50% (PRNT50), with higher cutoffs such as 70%, 80%, and 90% (PRNT90) used less frequently.<sup>[1](https://link.springer.com/article/10.1186/1471-2334-12-233)</sup> The PRNT is considered the laboratory standard against which other neutralizing antibody assays should be compared,<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> and the WHO gold standard for dengue vaccine immunogenicity assessment.<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup>\n\n| Key fact | Detail |\n| --- | --- |\n| Output | A titer: the reciprocal of the highest serum dilution reducing plaques by 50% (PRNT50) or 90% (PRNT90) versus the virus control<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC7536930/)</sup> |\n| Reference status | Laboratory standard for neutralizing antibody assays<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup>; WHO gold standard for dengue<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> |\n| Virus challenge | 40–60 PFU per 35 mm dish (WHO dengue guidelines)<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup>; 40–120 plaques per well in a validated 24-well dengue protocol<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> |\n| Cells | Vero recommended for dengue<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup>; published dengue protocols used LLC-MK2 in 47% of articles, BHK-21 in 28%, and Vero in 19%<sup>[1](https://link.springer.com/article/10.1186/1471-2334-12-233)</sup> |\n| Dengue adaptation | Russell, Nisalak, Sukhavachana, and Vivona, The Journal of Immunology, 1967<sup>[5](https://doi.org/10.4049/jimmunol.99.2.285)</sup> |\n| SARS-CoV-2 PRNT | 5 days incubation to countable plaques<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup>; BSL-3<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> |\n| Interpretation | PRNT titer considered the best immune correlate of protection for flaviviral infections<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> |\n\n## How it works\n\nSerum is serially diluted and mixed with a standardized, constant amount of infectious virus; the mixture is plated on susceptible cells under a semi-solid overlay, so each remaining productive infection yields one countable plaque.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> [Neutralizing antibody](https://www.edgepedia.org/neutralizing-antibody) lowers the number of infectious units, and the plaque count falls in proportion. Percent inhibition is calculated against the virus-only control, and the endpoint titer is the reciprocal of the last dilution showing the desired percent reduction.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> In a validated dengue protocol, the PRNT50 is the reciprocal of the highest dilution reducing infectivity by 50% versus the challenge-virus control, calculated by four-point linear regression on \\( \\log_{10} \\)-transformed plaque counts, with a theoretical lower limit of quantitation of 10.<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> Traditional Spearman–Kärber and Reed–Muench calculations require a full 0–100% neutralization response and do not easily provide confidence intervals; curve-fitting methods give more accurate results.<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> A distinctive advantage is that the assay also registers plaque morphology, for example plaques enlarged by antibody-dependent enhancement.<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup>\n\n## How it is done\n\nSerum or plasma is heat-inactivated at 56 °C for 30 min to 1 h before testing.<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC7536930/)</sup> In a validated dengue protocol, Vero cells are seeded at \\( 4 \\times 10^{5} \\) cells per well in 24-well plates, serum is diluted 1:5 and then two-fold across 12 wells, and virus is diluted to give 40–120 plaques per well in control wells.<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> WHO dengue guidelines recommend 40–60 PFU per 35 mm dish and at least three repeat wells for challenge doses of 50 PFU per reaction or less.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> Serum and virus are incubated for 60 min at 37 °C, the mixture is adsorbed onto the monolayer for 90 min, and an overlay of 2% carboxymethylcellulose<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> or 1% agarose<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC7536930/)</sup> restricts spread so infection stays localized. Incubation runs about 4 days for dengue<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> and 5 days in a described [SARS-CoV-2](https://www.edgepedia.org/sars-cov-2) workflow.<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> Cells are then fixed; chemical fixation inactivates dengue virus, so staining and counting can proceed under BSL-1 containment instead of BSL-2 or 3.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> Plaques are visualized with non-specific dyes such as crystal violet, neutral red, or amido black, or by immunostaining with serotype-specific anti-envelope monoclonal antibodies.<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> Plaques are counted manually or with an automated imager, and back-titration plates verify the challenge dose.<sup>[7](https://www.ecdc.europa.eu/sites/default/files/documents/SARS-CoV-2-characterisation-update-standard-laboratory-protocols.pdf)</sup>\n\n## Origin\n\nPlaque neutralization was originally developed for assaying antisera to bacteriophage; before plaque methods, neutralizing activity against animal viruses was estimated by quantal methods in eggs, animals, or tissue-culture tubes.<sup>[8](https://doi.org/10.1017/s0022172400045216)</sup> [Renato Dulbecco](https://www.edgepedia.org/renato-dulbecco)'s 1952 paper in the [Proceedings of the National Academy of Sciences](https://www.edgepedia.org/proceedings-of-the-national-academy-of-sciences) produced plaques in monolayer tissue cultures from single particles of an animal virus.<sup>[9](https://doi.org/10.1073/pnas.38.8.747)</sup> Dulbecco and Marguerite Vogt extended plaque formation to poliomyelitis viruses in 1954, showing that plaque counts are reproducible and proportional to virus concentration provided plates are not crowded.<sup>[10](https://doi.org/10.1084/jem.99.2.167)</sup> Henderson and Taylor reported arthropod-borne virus plaques in agar-overlaid tube cultures in 1959.<sup>[11](https://doi.org/10.3181/00379727-101-24902)</sup> A 1956 Virology study by Dulbecco, Vogt, and A.G.R. Strickland examined the basic aspects of neutralization of [Western equine encephalitis](https://www.edgepedia.org/western-equine-encephalitis) virus and poliomyelitis virus.<sup>[12](https://doi.org/10.1016/0042-6822%2856%2990017-4)</sup> In 1959, French, Armstrong, and Nagler adopted a plaque neutralization test for poliomyelitis antisera under defined conditions, finding neutralization logarithmic with respect to time and serum dilution and titers more reproducible than by other methods then in use.<sup>[13](https://doi.org/10.1139/m59-057)</sup> The dengue adaptation, a plaque reduction test for dengue virus neutralizing antibodies, was published by Philip K. Russell, Ananda Nisalak, Pairatana Sukhavachana, and Stefano Vivona in The Journal of Immunology in 1967.<sup>[5](https://doi.org/10.4049/jimmunol.99.2.285)</sup> Roehrig, Hombach, and Barrett published guidelines for plaque-reduction neutralization testing of human antibodies to dengue viruses in 2008.<sup>[14](https://doi.org/10.1089/vim.2008.0007)</sup>\n\n## Variants\n\n**The cutoff defines the titer.** WHO dengue guidelines prefer PRNT50 for vaccinee sera because it falls on the linear portion of the titration curve and gives more accurate results, though it is more variable; the more stringent PRNT90 is more useful in dengue-endemic areas, where it reduces detection of cross-reactive flavivirus antibodies.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> A reduction of at least 80% is likewise considered useful for epidemiologic or diagnostic purposes to limit cross-reactivity.<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> Higher neutralization percentages such as 90% versus 50% trade sensitivity for specificity.<sup>[1](https://link.springer.com/article/10.1186/1471-2334-12-233)</sup> One variability analysis of dengue control assays found the evaluation point with the lowest variance lay between PRNT75 and PRNT80 rather than at the WHO-recommended PRNT50 or PRNT90, a published counterpoint to the guideline cutoffs.<sup>[15](https://journals.plos.org/plosntds/article?id=10.1371%2Fjournal.pntd.0002952)</sup>\n\n**Format variants shrink the assay.** Running the PRNT in 96-well plates is called a [microneutralization assay](https://www.edgepedia.org/microneutralization-assay), and the focus reduction neutralization test (FRNT) stains viral protein after 24–48 h instead of waiting for plaques to develop.<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC7536930/)</sup> A Zika PRNT protocol adds neutral red after 4 days and considers sera with at least 90% plaque reduction to contain neutralizing antibodies.<sup>[16](https://experiments.springernature.com/articles/10.1007/978-1-0716-0581-3_5)</sup> For dengue, an immuno-plaque/focus reduction format using infrared-conjugated antibodies cuts turnaround from six days to three with over 90% sensitivity and specificity versus PRNT, and can quantify non-plaque-forming strains such as a DENV-4 Singapore isolate.<sup>[17](https://link.springer.com/article/10.1186/s12985-024-02459-y)</sup> Automated counting has entered practice: Viridot, a free open-source counter, matched manual plaque counts with a Lin's concordance correlation coefficient of 0.99 and outputs PRNT10–PRNT90 cutoffs with 95% confidence intervals, including from camera-phone images.<sup>[18](https://doi.org/10.1371/journal.pntd.0006862)</sup>\n\n## Applications\n\nPRNTs have been developed for many viruses, including dengue, mumps, yellow fever, measles, poliovirus, Lassa, and [Japanese encephalitis](https://www.edgepedia.org/japanese-encephalitis) virus.<sup>[19](https://pmc.ncbi.nlm.nih.gov/articles/PMC10814169/)</sup> The dengue PRNT is the WHO gold standard for vaccine immunogenicity assessment,<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup> and Zika protocols use PRNT with a 90% criterion for neutralizing antibodies.<sup>[16](https://experiments.springernature.com/articles/10.1007/978-1-0716-0581-3_5)</sup> For SARS-CoV-2, PRNT quantifies neutralizing antibodies in patient serum and plasma,<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> and microneutralization and pseudotyped-virus assays used alongside it supported development and licensure of the ChAdOx1 nCoV-19 and Ad26.COV2.S vaccines.<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> The virus-neutralizing antibody titer by PRNT is considered the best immune correlate of protection for flaviviral infections, although establishing a true protective level requires vaccine-efficacy trials with fully validated assays.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> A dengue vaccine trial observed infections in vaccinated individuals despite detectable titers, so the presence of some titer does not by itself separate protected from unprotected people.<sup>[15](https://journals.plos.org/plosntds/article?id=10.1371%2Fjournal.pntd.0002952)</sup> For SARS-CoV-2, PRNT50 titers between 1:10 and 1:30 were associated with protection against symptomatic infection in convalescent patients in one meta-analysis.<sup>[20](https://www.nature.com/articles/s41598-022-07597-3)</sup>\n\n## Limitations and alternatives\n\nPRNT is labor intensive and not readily amenable to high throughput, limiting its use in large-scale surveillance and vaccine trials.<sup>[2](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)</sup> For SARS-CoV-2 it requires BSL-3 containment, and the limited number of such facilities restricts how many laboratories can run live-virus neutralization.<sup>[21](https://pmc.ncbi.nlm.nih.gov/articles/PMC8733193/)</sup> The assay also requires plaque-forming viruses, excluding many non-cytopathic agents,<sup>[17](https://link.springer.com/article/10.1186/s12985-024-02459-y)</sup> and its manual plaque counting is slow and error-prone.<sup>[22](https://www.mdpi.com/2076-0817/14/11/1129)</sup> Titers depend on the experimental setup, including the virus particles used, the strain, the cells, and the dilution scheme, so results can only be compared between laboratories with caution.<sup>[20](https://www.nature.com/articles/s41598-022-07597-3)</sup> Variability analysis of dengue PRNT found experimental factors explained less than 1% of titer variance, indicating inherent assay variability.<sup>[15](https://journals.plos.org/plosntds/article?id=10.1371%2Fjournal.pntd.0002952)</sup> [Flavivirus](https://www.edgepedia.org/flavivirus) cross-reactivity is managed with higher cutoffs at a cost of sensitivity.<sup>[3](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)</sup>\n\n**Alternatives trade fidelity for speed.** A 96-well micro-neutralization test reading absorbance after crystal violet staining correlated strongly with dengue PRNT50 (\\( r = 0.9167 \\)), is about eight times more efficient per plate, and avoids counting the small plaques of DENV-3 and DENV-4 or blurry Japanese encephalitis plaques.<sup>[23](https://www.mdpi.com/2076-0817/13/1/8)</sup> An MTT-based microneutralization assay showed only moderate correlation with PRNT50, declining further at 70% and 90% endpoints.<sup>[17](https://link.springer.com/article/10.1186/s12985-024-02459-y)</sup> Pseudovirus-based neutralization assays run at BSL-2 instead of BSL-3; a pseudotyped assay correlated with PRNT at Pearson \\( r = 0.862 \\),<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> and a lentiviral surrogate achieved 100% sensitivity and specificity against PRNT for ancestral SARS-CoV-2 with a 48-h turnaround in 96- to 384-well formats.<sup>[24](https://pmc.ncbi.nlm.nih.gov/articles/PMC9927366/)</sup> Because pseudovirus systems are single-cycle entry assays, their neutralization curves may plateau earlier and titers may run lower than PRNT at high antibody concentrations.<sup>[22](https://www.mdpi.com/2076-0817/14/11/1129)</sup> Binding ELISA detects total antibodies and cannot distinguish neutralizing from non-neutralizing antibodies, so it can underestimate functional titers.<sup>[21](https://pmc.ncbi.nlm.nih.gov/articles/PMC8733193/)</sup>\n\n**Standardization targets these gaps.** The NIBSC 1st WHO International Standard for anti-SARS-CoV-2 antibody (20/136) is used alongside neutralization assays,<sup>[6](https://www.nature.com/articles/s41596-021-00536-y)</sup> though demand exceeded availability.<sup>[25](https://epub.uni-regensburg.de/76403/1/streif-baeumner-2025-advances-in-surrogate-neutralization-tests-for-high-throughput-screening-and-the-point-of-care.pdf)</sup> An analytical [quality by design](https://www.edgepedia.org/quality-by-design) (AQbD) procedure identified 13 main risk factors among 30 potential influencing factors and reduced between-laboratory differences that had reached 4.8-fold higher and 3.3-fold lower than the standardized SOP.<sup>[26](https://www.nature.com/articles/s41392-023-01389-5)</sup> Published comparisons do not settle what specific PRNT titer threshold is protective for dengue, nor do they quantify PRNT cost in currency units.\n\n## References\n\n1. [Variation in dengue virus plaque reduction neutralization testing: systematic review and pooled analysis (BMC Infectious Diseases, 2012)](https://link.springer.com/article/10.1186/1471-2334-12-233)\n2. [Guidelines for plaque reduction neutralization testing of human antibodies to dengue viruses (WHO/IVB/07.07)](https://apps.who.int/iris/bitstream/handle/10665/69687/who_ivb_07.07_eng.pdf?isAllowed=y&sequence=1)\n3. [Optimization and Validation of a Plaque Reduction Neutralization Test for the Detection of Neutralizing Antibodies to Four Serotypes of Dengue Virus (2013)](https://pmc.ncbi.nlm.nih.gov/articles/PMC3752766/)\n4. [Viral infection neutralization tests: a focus on SARS-CoV-2 with implications for convalescent plasma therapy (2020)](https://pmc.ncbi.nlm.nih.gov/articles/PMC7536930/)\n5. [Philip K Russell and colleagues (1967). A Plaque Reduction Test for Dengue Virus Neutralizing Antibodies. The Journal of Immunology.](https://doi.org/10.4049/jimmunol.99.2.285)\n6. [Quantification of SARS-CoV-2 neutralizing antibody by wild-type plaque reduction neutralization, microneutralization and pseudotyped virus neutralization assays (Nature Protocols, 2021)](https://www.nature.com/articles/s41596-021-00536-y)\n7. [Update of standard laboratory protocols for SARS-CoV-2 characterisation (ECDC/AURORAE consortium)](https://www.ecdc.europa.eu/sites/default/files/documents/SARS-CoV-2-characterisation-update-standard-laboratory-protocols.pdf)\n8. [A comparative study of a plaque and quantal method for assaying the neutralizing activity of antisera to type 1 poliovirus](https://doi.org/10.1017/s0022172400045216)\n9. [Renato Dulbecco (1952). Production of Plaques in Monolayer Tissue Cultures by Single Particles of an Animal Virus. Proceedings of the National Academy of Sciences.](https://doi.org/10.1073/pnas.38.8.747)\n10. [R. Dulbecco, Marguerite Vogt (1954). PLAQUE FORMATION AND ISOLATION OF PURE LINES WITH POLIOMYELITIS VIRUSES. The Journal of Experimental Medicine.](https://doi.org/10.1084/jem.99.2.167)\n11. [J. R. Henderson, R. M. Taylor (1959). Arthropod-Borne Virus Plaques in Agar Overlaid Tube Cultures.. Experimental Biology and Medicine.](https://doi.org/10.3181/00379727-101-24902)\n12. [A study of the basic aspects of neutralization of two animal viruses, Western equine encephalitis virus and poliomyelitis virus (Virology, 1956)](https://doi.org/10.1016/0042-6822%2856%2990017-4)\n13. [R. C. French, R. E. Armstrong, F. P. Nagler (1959). EVALUATION OF POLIOMYELITIS ANTISERA BY PLAQUE NEUTRALIZATION TESTS. Canadian Journal of Microbiology.](https://doi.org/10.1139/m59-057)\n14. [John T. Roehrig, Joachim Hombach, Alan D.T. Barrett (2008). Guidelines for Plaque-Reduction Neutralization Testing of Human Antibodies to Dengue Viruses. Viral Immunology.](https://doi.org/10.1089/vim.2008.0007)\n15. [Variability in Dengue Titer Estimates from Plaque Reduction Neutralization Tests Poses a Challenge to Epidemiological Studies and Vaccine Development (PLOS NTD)](https://journals.plos.org/plosntds/article?id=10.1371%2Fjournal.pntd.0002952)\n16. [A Plaque Reduction Neutralization Test for the Detection of ZIKV-Specific Antibodies (Springer Nature Experiments protocol)](https://experiments.springernature.com/articles/10.1007/978-1-0716-0581-3_5)\n17. [Evaluation of three alternative methods to the plaque reduction neutralizing assay for measuring neutralizing antibodies to dengue virus serotype 2 (Virology Journal, 2024)](https://link.springer.com/article/10.1186/s12985-024-02459-y)\n18. [Leah C. Katzelnick and colleagues (2018). Viridot: An automated virus plaque (immunofocus) counter for the measurement of serological neutralizing responses with application to dengue virus. PLoS neglected tropical diseases.](https://doi.org/10.1371/journal.pntd.0006862)\n19. [Plaque Reduction Neutralization Test (PRNT) Accuracy in Evaluating Humoral Immune Response to SARS-CoV-2 (Diseases, 2024)](https://pmc.ncbi.nlm.nih.gov/articles/PMC10814169/)\n20. [Evaluation of a commercial ELISA as alternative to plaque reduction neutralization test to detect neutralizing antibodies against SARS-CoV-2](https://www.nature.com/articles/s41598-022-07597-3)\n21. [SARS-CoV-2 Antibody Neutralization Assay Platforms Based on Epitopes Sources: Live Virus, Pseudovirus, and Recombinant S Glycoprotein RBD (2021)](https://pmc.ncbi.nlm.nih.gov/articles/PMC8733193/)\n22. [Pseudovirus-Based Neutralization Assays as Customizable and Scalable Tools for Serological Surveillance and Immune Profiling (Pathogens, 2025)](https://www.mdpi.com/2076-0817/14/11/1129)\n23. [Utility of an In-Vitro Micro-Neutralizing Test in Comparison to a Plaque Reduction Neutralization Test for Dengue Virus, Japanese Encephalitis Virus, and Zika Virus Serology and Drug Screening (Pathogens, 2024)](https://www.mdpi.com/2076-0817/13/1/8)\n24. [Validation and Establishment of the SARS-CoV-2 Lentivirus Surrogate Neutralization Assay as a Prescreening Tool for the Plaque Reduction Neutralization Test](https://pmc.ncbi.nlm.nih.gov/articles/PMC9927366/)\n25. [Advances in Surrogate Neutralization Tests for High-Throughput Screening and the Point-of-Care (2025)](https://epub.uni-regensburg.de/76403/1/streif-baeumner-2025-advances-in-surrogate-neutralization-tests-for-high-throughput-screening-and-the-point-of-care.pdf)\n26. [Standardized neutralization antibody analytical procedure for clinical samples based on the AQbD concept (Signal Transduction and Targeted Therapy, 2023)](https://www.nature.com/articles/s41392-023-01389-5)\n\n---\n*Topic: Encyclopedia › Life and health › Microorganisms and fungi › Viruses and acellular agents*\n\n*Initially written Sep 29, 2026 · Edited: Oct 15, 2026*\n\n*Copyright 2026 EdgeChat AI, a subsidiary of Biostate AI.*\n\nLicense: Edgepedia Community License 1.0, https://www.edgepedia.org/edgepedia/license\n",
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 "speakable": "The plaque reduction neutralization test is a live-virus cell-culture assay measuring neutralizing antibodies, considered the laboratory standard and the WHO gold standard for dengue."
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