{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T03:46:55Z","timestamp":1790221615484,"version":"4.1.0"},"reference-count":130,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2023,8,27]],"date-time":"2023-08-27T00:00:00Z","timestamp":1693094400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>Listeria continues to be a persistent foodborne pathogen that is responsible for human cases of listeriosis when contaminated food products are consumed. Human subjects considered to be most susceptible include the elderly, immunocompromised, and pregnant women. Listeria is characterized as a saprophytic organism with the capability of responding and adapting to constantly changing environments because it possesses multiple stress response mechanisms to overcome varying temperatures, salt concentrations, and pH, among others. Primary foods and food products associated with listeriosis include dairy products and ready-to-eat meats such as turkey products. Historically, chicken eggs have not been identified as a primary source of Listeria, but the potential for contamination during egg production and processing does exist. Listeria species have been isolated from egg-processing plant equipment and are presumed to occur in egg-processing plant environments. Whether Listeria is consistently disseminated onto eggs beyond the egg-processing plant is a risk factor that remains to be determined. However, research has been conducted over the years to develop egg wash solutions that generate combinations of pH and other properties that would be considered inhibitory to Listeria. Even less is known regarding the association of Listeria with alternative egg production systems, but Listeria has been isolated from pasture flock broilers, so it is conceivable, given the nature of the outdoor environments, that layer birds under these conditions would also be exposed to Listeria and their eggs become contaminated. This review focuses on the possibility of Listeria occurring in conventional and alternative egg-laying production and processing systems.<\/jats:p>","DOI":"10.3390\/microorganisms11092164","type":"journal-article","created":{"date-parts":[[2023,8,28]],"date-time":"2023-08-28T02:23:25Z","timestamp":1693189405000},"page":"2164","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Listeria Occurrence in Conventional and Alternative Egg Production Systems"],"prefix":"10.3390","volume":"11","author":[{"given":"Steven C.","family":"Ricke","sequence":"first","affiliation":[{"name":"Meat Science and Animal Biologics Discovery Program, Department of Animal and Dairy Sciences, University of Wisconsin, Madison, WI 53706, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4738-7751","authenticated-orcid":false,"given":"Corliss A.","family":"O\u2019Bryan","sequence":"additional","affiliation":[{"name":"Food Science Department, University of Arkansas, Fayetteville, AR 72704, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"suffix":"Jr.","given":"Michael J.","family":"Rothrock","sequence":"additional","affiliation":[{"name":"U.S. National Poultry Research Center, Egg Safety & Quality Research Unit, USDA-ARS, Athens, GA 30606, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,8,27]]},"reference":[{"key":"ref_1","unstructured":"Albrecht, J.A. (2019, June 24). Listeria Monocytogenes. Available online: https:\/\/food.unl.edu\/listeria-monocytogenes."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"652","DOI":"10.4315\/0362-028X-53.8.652","article-title":"Growth, inhibition, and survival of Listeria monocytogenes as affected by acidic conditions","volume":"53","author":"Conner","year":"1990","journal-title":"J. Food Prot."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1111\/j.1365-2672.1990.tb02561.x","article-title":"Growth of Listeria monocytogenes at refrigeration temperatures","volume":"68","author":"Walker","year":"1990","journal-title":"J. Appl. Bacteriol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"308","DOI":"10.1038\/ng.3501","article-title":"Uncovering Listeria monocytogenes hypervirulence by harnessing its biodiversity","volume":"48","author":"Maury","year":"2016","journal-title":"Nat. Genet."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"125","DOI":"10.3389\/fvets.2017.00125","article-title":"Listeria occurrence in poultry flocks: Detection and potential implications","volume":"4","author":"Rothrock","year":"2017","journal-title":"Front. Vet. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1095","DOI":"10.1099\/00221287-147-5-1095","article-title":"Comparative genetic characterization of Listeria monocytogenes isolates from human and animal listeriosis cases","volume":"147","author":"Jeffers","year":"2001","journal-title":"Microbiology"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.ijmm.2010.05.002","article-title":"Listeria monocytogenes lineages: Genomics, evolution, ecology, and phenotypic characteristics","volume":"301","author":"Orsi","year":"2011","journal-title":"Int. J. Med. Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"2640","DOI":"10.4315\/0362-028X-69.11.2640","article-title":"Growth and stress resistance variation in culture broth among Listeria monocytogenes strains of various serotypes and origins","volume":"69","author":"Lianou","year":"2006","journal-title":"J. Food Protect."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1283","DOI":"10.4315\/0362-028X-72.6.1283","article-title":"Prevalence and genetic diversity of Listeria monocytogenes in vacuum-packaged ready-to-eat meat products at retail markets in Latvia","volume":"72","author":"Berzins","year":"2009","journal-title":"J. Food Protect."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.foodcont.2012.06.047","article-title":"Listeria monocytogenes prevalence and serotype diversity in various foods","volume":"30","author":"Kramarenko","year":"2013","journal-title":"Food Control"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"8115","DOI":"10.1128\/AEM.71.12.8115-8122.2005","article-title":"Relatedness of Listeria monocytogenes isolates recovered from selected ready-to-eat foods and listeriosis patients in the United States","volume":"71","author":"Gilbreth","year":"2005","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Ryser, E.T., and Marth, H.E. (2007). Listeria, Listeriosis, and Food Safety, CRC Press. [3rd ed.].","DOI":"10.1201\/9781420015188"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1881","DOI":"10.1016\/j.meegid.2011.08.001","article-title":"Recombination and positive selection contributed to the evolution of Listeria monocytogenes lineages III and IV, two distinct and well supported uncommon L. monocytogenes lineages","volume":"11","author":"Tsai","year":"2011","journal-title":"Infect. Genet. Evol."},{"key":"ref_14","unstructured":"CDC (2019, March 06). Centers for Disease Control and Prevention. Information for Health Professionals and Laboratories | Listeria | CDC, Available online: https:\/\/www.cdc.gov\/Listeria\/technical.html."},{"key":"ref_15","unstructured":"European Center for Disease Prevention and Control (2023, August 14). Listeriosis\u2014Annual Epidemiological Report for 2021. Available online: https:\/\/www.ecdc.europa.eu\/en\/publications-data\/listeriosis-annual-epidemiological-report-2021."},{"key":"ref_16","unstructured":"(2023, August 14). Healthy People 2030. Reduce Infections Caused by Listeria\u2014FS-03, Available online: https:\/\/health.gov\/healthypeople\/objectives-and-data\/browse-objectives\/foodborne-illness\/reduce-infections-caused-listeria-fs-03\/data?tab=data-table#data-table."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1128\/CMR.14.3.584-640.2001","article-title":"Listeria pathogenesis and molecular virulence determinants","volume":"14","author":"Kuhn","year":"2001","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2113","DOI":"10.3201\/eid2212.160165","article-title":"Infectious dose of Listeria monocytogenes in outbreak linked to ice cream, United States, 2015","volume":"22","author":"Pouillot","year":"2016","journal-title":"Emerg. Infect. Dis."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1016\/j.cofs.2018.03.007","article-title":"The evolution of FDA\u2019s policy on Listeria monocytogenes in ready-to-eat foods in the United States","volume":"20","author":"Archer","year":"2018","journal-title":"Curr. Opin. Food Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/S0168-1605(99)00070-7","article-title":"Inactivation of Listeria innocua in liquid whole egg by pulsed electric fields and nisin","volume":"51","author":"Swanson","year":"1999","journal-title":"Internat. J. Food Microbiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"986","DOI":"10.4315\/0362-028X-68.5.986","article-title":"Efficacy of electrolyzed water in inactivating Salmonella enteritidis and Listeria monocytogenes on shell eggs","volume":"68","author":"Park","year":"2005","journal-title":"J. Food Protect."},{"key":"ref_22","first-page":"1208","article-title":"An updated review on chicken eggs: Production, consumption, management aspects and nutritional benefits to human health","volume":"6","author":"Zaheer","year":"2015","journal-title":"Food Nutr. Sci."},{"key":"ref_23","unstructured":"(2023, June 14). ChickenFans.com. Poultry Industry Statistics (2023): Meat & Egg Production. Available online: https:\/\/www.chickenfans.com\/poultry-industry-statistics\/#Global-Egg-Production."},{"key":"ref_24","unstructured":"Bitter, A. (2023, August 16). Why Organic Eggs Are Suddenly Cheaper than Conventional Ones. Available online: https:\/\/uk.news.yahoo.com\/why-organic-eggs-suddenly-cheaper-182324110.html?guccounter=1&guce_referrer=aHR0cHM6Ly93d3cuZ29vZ2xlLmNvbS8&guce_referrer_sig=AQAAACaVG_ZMnM9qjbJuR0tGDj-siLSA_XXhgj9-rjnNSNWMz_vgc8YjdUFFA0YPvfGzc_LmvYw3Qdpb5ELGn9VTnjvaSp_UyCadJTvu4jJ4yQR2O5O4w2AmTv1xSVXglMilIX-r0cRzvV6UMZsdFloLF05PGsd8XVRMUTiaLFFdNdLF."},{"key":"ref_25","unstructured":"United Egg Producers (2023, June 14). Facts & Stats. Available online: https:\/\/unitedegg.com\/facts-stats\/."},{"key":"ref_26","unstructured":"Walmart (2023, August 14). Animal Welfare. Available online: https:\/\/corporate.walmart.com\/esgreport\/esg-issues\/animal-welfare#resources."},{"key":"ref_27","unstructured":"Sumner, D.A., Rosen-Molina, J.T., Matthews, W.A., Mench, J.A., and Richter, K.R. (2008). Economic Effects of Proposed Restrictions on Egg-Laying Hen Housing in California, University of California Agricultural Issues Center."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"241","DOI":"10.3382\/ps.2010-00822","article-title":"Economic and market issues on the sustainability of egg production in the United States: Analysis of alternative production systems","volume":"90","author":"Sumner","year":"2011","journal-title":"Poult. Sci."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"251","DOI":"10.3382\/ps.2010-00794","article-title":"The impact of different housing systems on egg safety and quality","volume":"90","author":"Holt","year":"2011","journal-title":"Poult. Sci."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"29","DOI":"10.7120\/09627286.25.1.029","article-title":"Farmer attitudes to injurious pecking in laying hens and to potential control strategies","volume":"25","author":"Palcynski","year":"2016","journal-title":"Anim. Welf."},{"key":"ref_31","unstructured":"Food and Drug Administration (FDA) (2019, June 24). Egg Safety Final Rule. 26 January 2018, Available online: https:\/\/www.fda.gov\/food\/eggs-guidance-documents-regulatory-information\/egg-safety-final-rule."},{"key":"ref_32","unstructured":"Dubois, S. (2019, June 24). Is Raw Egg in a Protein Shake Unhealthy? 27 November 2018. Available online: https:\/\/healthyeating.sfgate.com\/raw-egg-protein-shake-unhealthy-1090.html."},{"key":"ref_33","unstructured":"USDA (2019, June 24). United States Depart of Agriculture (USDA). Egg Products Inspection Act. 21 January 2016, Available online: https:\/\/www.fsis.usda.gov\/wps\/portal\/fsis\/topics\/rulemaking\/egg-products-inspection-act\/EPIA."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1017\/S0043933911000043","article-title":"Pasture-based systems for poultry production: Implications and perspectives","volume":"67","author":"Sossidou","year":"2011","journal-title":"World\u2019s Poult. Sci. J."},{"key":"ref_35","unstructured":"USDA (2023, August 14). Shell Eggs from Farm to Table, Available online: https:\/\/www.fsis.usda.gov\/food-safety\/safe-food-handling-and-preparation\/eggs\/shell-eggs-farm-table."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"33","DOI":"10.3389\/fsufs.2019.00033","article-title":"Listeria occurrence and potential control strategies in alternative and conventional poultry processing and retail","volume":"3","author":"Rothrock","year":"2019","journal-title":"Front. Sustain. Food Syst."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1093\/japr\/6.1.116","article-title":"The presence of Listeria monocytogenes in the integrated poultry industry","volume":"6","author":"Cox","year":"1997","journal-title":"J. Appl. Poult. Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/S0168-1605(00)00258-0","article-title":"Listeria spp. in broiler flocks: Recovery rates and species distribution investigated by conventional culture and the EiaFoss method","volume":"58","author":"Petersen","year":"2000","journal-title":"Int. J. Food Microbiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1292\/jvms.53.873","article-title":"Prevalence of Listeria monocytogenes in intestinal contents of healthy animals in Japan","volume":"53","author":"Iida","year":"1991","journal-title":"J. Vet. Med. Sci."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2158","DOI":"10.3382\/ps.2012-02292","article-title":"C Listeria monocytogenes and hemolytic Listeria innocua in poultry","volume":"91","author":"Milillo","year":"2012","journal-title":"Poult. Sci."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"827","DOI":"10.4315\/0362-028X-59.8.827","article-title":"Growth of Listeria monocytogenes as a biofilm on various food-processing surfaces","volume":"59","author":"Blackman","year":"1996","journal-title":"J. Food Protect."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"132","DOI":"10.3382\/japr.2012-00676","article-title":"Contamination of raw poultry meat by airborne Listeria originating from a floor drain","volume":"22","author":"Berrang","year":"2013","journal-title":"J. Appl. Poult. Res."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"791","DOI":"10.3382\/ps.2008-00396","article-title":"Listeria monocytogenes in two different poultry facilities: Manual and automatic evisceration","volume":"88","author":"Chiarini","year":"2009","journal-title":"Poult. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2537","DOI":"10.4315\/0362-028X-71.12.2537","article-title":"Comparison of antimicrobial resistance in Escherichia coli, Staphylococcus aureus, and Listeria monocytogenes strains isolated from organic and conventional poultry meat","volume":"71","author":"Miranda","year":"2008","journal-title":"J. Food Prot."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1590\/1678-457X.6808","article-title":"Prevalence of Listeria monocytogenes in poultry meat","volume":"35","author":"Elmali","year":"2015","journal-title":"Food Sci. Technol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.prevetmed.2010.11.017","article-title":"Risk factors for Listeria monocytogenes contamination in French laying hens and broiler flocks","volume":"98","author":"Aury","year":"2011","journal-title":"Prev. Veterin. Med."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1128\/jb.80.3.316-320.1960","article-title":"Survival of Listeria monocytogenes in soil","volume":"80","author":"Welshimer","year":"1960","journal-title":"J. Bacteriol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1481","DOI":"10.4315\/0362-028X.JFP-17-466","article-title":"A comparative study of the susceptibility of Listeria species to sanitizer treatments when grown under planktonic and biofilm conditions","volume":"81","author":"Fox","year":"2018","journal-title":"J. Food Protect."},{"key":"ref_49","unstructured":"Clauer, P. (2019, July 09). Modern Egg Industry. 5 July 2012. Available online: https:\/\/extension.psu.edu\/modern-egg-industry."},{"key":"ref_50","first-page":"953","article-title":"Effect of poultry housing systems on egg production","volume":"18","author":"Gerzilov","year":"2012","journal-title":"Bulg. J. Agric. Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1079\/WPS200574","article-title":"Physiological and endocrinological mechanisms associated with ovulatory cycle and induced-moulting in the domestic chicken\u2014A Review","volume":"61","author":"Oguike","year":"2005","journal-title":"World\u2019s Poult. Sci. J."},{"key":"ref_52","unstructured":"Poole, T.E. (2016). Introduction to Developing a Free-Range Poultry Enterprise, University of Maryland. [1st ed.]. Available online: https:\/\/extension.umd.edu\/sites\/default\/files\/_docs\/locations\/frederick_county\/Ag%20Pubs%20A%20Supplement%20to%20Free%20Range%20Poultry.pdf."},{"key":"ref_53","unstructured":"Pitesky, M. (2019, June 10). Free-Range vs. Pastured Poultry: What\u2019s the Difference? October 2017. Available online: https:\/\/www.hobbyfarms.com\/free-range-vs-pastured-poultry-whats-difference\/."},{"key":"ref_54","unstructured":"USDA (2019, July 10). United States Depart of Agriculture. Meat and Poultry Labeling Terms. 10 August 2015, Available online: https:\/\/www.fsis.usda.gov\/wps\/portal\/fsis\/topics\/food-safety-education\/get-answers\/food-safety-fact-sheets\/food-labeling\/meat-and-poultry-labeling-terms\/meat-and-poultry-labeling-terms."},{"key":"ref_55","unstructured":"Certified Humane (2019, July 10). \u201cFree Range\u201d and \u201cPasture Raised\u201d Officially Defined by HFAC for Certified Humane\u00ae Label. 9 October 2014. Available online: https:\/\/certifiedhumane.org\/free-range-and-pasture-raised-officially-defined-by-hfac-for-certified-humane-label\/."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"165","DOI":"10.4081\/ijas.2009.165","article-title":"Effects of two different rearing systems (organic and barn) on production performance, animal welfare traits and egg quality characteristics in laying hens","volume":"8","author":"Ferrante","year":"2009","journal-title":"Ital. J. Anim. Sci."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"434","DOI":"10.3923\/ijps.2007.434.439","article-title":"Effects of genotype and housing system on the laying performance of chickens in different seasons in the semi-humid tropics","volume":"6","author":"Yakubu","year":"2007","journal-title":"Int. J. Poult. Sci."},{"key":"ref_58","unstructured":"Castellini, C., Perella, F., Mugnai, C., and Dal Bosco, A. (2006, January 10\u201314). Welfare, productivity and qualitative traits of egg in laying hens reared under different rearing systems. Proceedings of the XII European Poultry Conference, Verona, Italy."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"M496","DOI":"10.1111\/j.1750-3841.2010.01747.x","article-title":"Prevalence and characterization of Campylobacter jejuni isolated from pasture flock poultry","volume":"75","author":"Hanning","year":"2010","journal-title":"J. Food Sci."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"S37","DOI":"10.1186\/1751-0147-43-S1-S37","article-title":"Health and welfare in organic poultry production","volume":"43","author":"Berg","year":"2001","journal-title":"Acta Vet. Scand."},{"key":"ref_61","first-page":"301","article-title":"Listeria monocytogenes infection in poultry and its public health importance with special reference to food borne zoonoses","volume":"16","author":"Dhama","year":"2013","journal-title":"PJBSBI"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"534","DOI":"10.4314\/eamj.v81i10.9237","article-title":"Sensitivity of Listeria species recovered from indigenous chickens to antibiotics and disinfectants","volume":"81","author":"Njagi","year":"2004","journal-title":"East Afr. Med. J."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"457","DOI":"10.3382\/ps.0690457","article-title":"Listeria monocytogenes colonization of broiler chickens","volume":"69","author":"Bailey","year":"1990","journal-title":"Poult. Sci."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1080\/03079450802050697","article-title":"Systemic listeriosis in caged canaries (Serinus canarius)","volume":"37","author":"Akanbi","year":"2008","journal-title":"Avian Pathol."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Gwida, M., L\u00fcth, S., El-Ashker, M., Zakaria, A., El-Gohary, F., Elsayed, M., Kleta, S., and Al Dahouk, S. (2020). Contamination pathways can be traced along the poultry processing chain by whole genome sequencing of Listeria innocua. Microorganisms, 8.","DOI":"10.3390\/microorganisms8030414"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"1996","DOI":"10.4315\/0362-028X-71.10.1996","article-title":"Prevalence of Listeria monocytogenes in poultry production in France","volume":"71","author":"Chemaly","year":"2008","journal-title":"J. Food Prot."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1016\/j.ijfoodmicro.2007.12.012","article-title":"A survey of foodborne pathogens in free-range poultry farms","volume":"123","author":"Esteban","year":"2008","journal-title":"Int. J. Food Microbiol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1111\/j.1863-2378.2008.01229.x","article-title":"Comparative analysis on antibiotic resistance characteristics of Listeria spp. and Enterococcus spp. isolated from laying hens and eggs in conventional and organic keeping systems in Bavaria, Germany","volume":"57","author":"Schwaiger","year":"2010","journal-title":"Zoonoses Public Health"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"422","DOI":"10.4315\/0362-028X.JFP-18-355","article-title":"Risk Characterization of antibiotic resistance in bacteria isolated from backyard, organic, and regular commercial eggs","volume":"82","author":"Fenollar","year":"2019","journal-title":"J. Food Prot."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.3382\/ps.2011-01795","article-title":"Prevalence of coliforms, Salmonella, Listeria, and Campylobacter associated with eggs and the environment of conventional cage and free-range egg production","volume":"91","author":"Jones","year":"2012","journal-title":"Poult. Sci."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/0963-9969(92)90156-Y","article-title":"Presence of Listeria spp. in whole eggs and wash water samples from Ontario and Quebec","volume":"25","author":"Farber","year":"1992","journal-title":"Food Res. Int."},{"key":"ref_72","first-page":"117","article-title":"Studies on the occurrence of Listeria monocytogenes in fecal samples of domestic and companion animals","volume":"198","author":"Weber","year":"1995","journal-title":"Int. J. Hyg. Environ. Med."},{"key":"ref_73","first-page":"15","article-title":"The impact of the prevalence of Listeria monocytogenes in the environment on meat and milk hygiene","volume":"8","author":"Skovgaard","year":"1990","journal-title":"Microbiol. Aliment. Nutr."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1002\/(SICI)1097-0010(199611)72:3<299::AID-JSFA656>3.0.CO;2-A","article-title":"Microbiological decontamination of poultry feed\u2014Evaluation of steam conditioners","volume":"72","author":"Blank","year":"1996","journal-title":"J. Sci. Food Agric."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"685","DOI":"10.1038\/s41598-023-27862-3","article-title":"Isolation of Listeria monocytogenes from poultry red mite (Dermanyssus gallinae) infesting a backyard chicken farm in Greece","volume":"13","author":"Sioutas","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_76","unstructured":"Sparagano, O., and Giangaspero, A. (2011). Improving the Safety and Quality of Eggs. Egg Products Egg Chemistry, Production and Consumption, Woodhead Publishing Limited."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"100338","DOI":"10.1016\/j.japr.2023.100338","article-title":"Impact of paddock area stocking density of free-range laying hens on egg and environmental microbiology","volume":"32","author":"Garcia","year":"2023","journal-title":"J. Appl. Poult. Res."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"100189","DOI":"10.1016\/j.japr.2021.100189","article-title":"Impact of organic dairy cattle manure on environmental and egg microbiology of organic free-range laying hens","volume":"30","author":"Garcia","year":"2021","journal-title":"J. Appl. Poult. Res."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Crespo, R., Garner, M.M., Hopkins, S.G., and Shah, D.H. (2013). Outbreak of Listeria monocytogenes in an urban poultry flock. BMC Vet. Res., 9.","DOI":"10.1186\/1746-6148-9-204"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"862","DOI":"10.4315\/0362-028X-55.11.862","article-title":"Survival of Listeria monocytogenes on the surface of egg shells and during frying of whole and scrambled eggs","volume":"55","author":"Brackett","year":"1992","journal-title":"J. Food Protect."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1093\/japr\/8.4.499","article-title":"Bacterial penetration of the eggshell and shell membranes of the chicken hatching egg: A review","volume":"8","author":"Berrang","year":"1999","journal-title":"J. Appl. Poult. Res."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1111\/j.1574-6968.1992.tb05684.x","article-title":"Observations on the mode of bacterial infection of hens\u2019 eggs","volume":"100","author":"Lock","year":"1992","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Board, R.G., and Fuller, R. (1994). Microbiology of the Avian Egg, Springer.","DOI":"10.1007\/978-1-4615-3060-2"},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Mead, G.C. (2005). Food Safety Control in the Poultry Industry: Bacterial Infection of Eggs, Woodhead Publishing Limited.","DOI":"10.1533\/9781845690236"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"463","DOI":"10.2307\/1591437","article-title":"Salmonella typhimurium penetration through the eggshell of hatching eggs","volume":"34","author":"Padron","year":"1990","journal-title":"Avian Dis."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"718","DOI":"10.1111\/j.1574-6976.2008.00161.x","article-title":"Mechanisms of egg contamination by Salmonella Enteritidis","volume":"33","author":"Gantois","year":"2009","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Downes, F.P., and Ito, K. (2001). Eggs and egg products, In Compendium of Methods for the Microbiological Examination of Foods, American Public Health Association. [4th ed.].","DOI":"10.2105\/9780875531755"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1080\/00071667308415999","article-title":"The cuticle: A barrier to liquid and particle penetration of the shell of hen\u2019s egg","volume":"14","author":"Board","year":"1973","journal-title":"Br. Poult. Sci."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1111\/age.12071","article-title":"Enhancing the egg\u2019s natural defense against bacterial penetration by increasing cuticle deposition","volume":"44","author":"Bain","year":"2013","journal-title":"Anim. Genet."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1079\/WPS200443","article-title":"Eggshell penetration by Salmonella: A review","volume":"61","author":"Messens","year":"2005","journal-title":"World\u2019s Poult. Sci. J."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"7571","DOI":"10.1128\/AEM.71.11.7571-7574.2005","article-title":"Factors influencing the ability of Listeria monocytogenes to pass through a membrane filter by active infiltration","volume":"71","author":"Nakazawa","year":"2005","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1590\/S1516-635X2000000200007","article-title":"Pore size distribution in chicken eggs as determined by mercury porosimetry","volume":"2","author":"Boleli","year":"2000","journal-title":"Rev. Bras. Cienc. Av\u00edcola"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1111\/j.1469-185X.1974.tb01297.x","article-title":"Non-specific antimicrobial defenses of the avian egg, embryo, and neonate","volume":"49","author":"Board","year":"1974","journal-title":"Biol. Rev."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1066","DOI":"10.1111\/mmi.13368","article-title":"The Listeria monocytogenes Fur-regulated virulence protein FrvA is an Fe(II) efflux P1B4 -type ATPase","volume":"100","author":"Pi","year":"2016","journal-title":"Mol. Microbiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"840","DOI":"10.1039\/C7MT00112F","article-title":"Ferrous iron efflux systems in bacteria","volume":"9","author":"Pi","year":"2017","journal-title":"Metallomics"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"12318","DOI":"10.1073\/pnas.0404924101","article-title":"Listeria monocytogenes regulates flagellar motility gene expression through MogR, a transcriptional repressor required for virulence","volume":"101","author":"Burrack","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_97","unstructured":"Center for Food Safety and Public Health (CFSPH) (2019, July 16). Listeriosis. Available online: http:\/\/www.cfsph.iastate.edu\/Factsheets\/pdfs\/listeriosis.pdf."},{"key":"ref_98","unstructured":"USDA (2019, June 24). United States Depart of Agriculture (USDA). Egg Products and Food Safety. 10 August 2015, Available online: https:\/\/www.fsis.usda.gov\/food-safety\/safe-food-handling-and-preparation\/eggs\/egg-products-and-food-safety."},{"key":"ref_99","unstructured":"Funk, E.M. (1935). Missouri Agriculture Experiment Station Bulletin, University of Missouri College of Agriculture, Agricultural Experiment Station. Number 350."},{"key":"ref_100","unstructured":"Anderson, K.E. (2023, July 17). Range Egg Production; Is It Better than in Cages?. Available online: https:\/\/www.wattagnet.com\/egg\/cage-free-laying-systems\/article\/15500793\/study-compares-range-cage-free-conventional-egg-production."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1093\/ps\/81.5.727","article-title":"Effects of cryogenic cooling of shell eggs on egg quality","volume":"81","author":"Jones","year":"2002","journal-title":"Poult. Sci."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1111\/j.1745-4565.2006.00048.x","article-title":"Frequency of Salmonella, Campylobacter, Listeria and Enterobacteriaceae detection in commercially cool water-washed shell eggs","volume":"26","author":"Jones","year":"2006","journal-title":"J. Food Saf."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1007\/s11259-010-9391-4","article-title":"Hot air treatment for surface decontamination of table eggs experimentally infected with Salmonella, Listeria, and Escherichia coli","volume":"34","author":"Manfreda","year":"2010","journal-title":"Vet. Res. Commun."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/0168-1605(91)90060-3","article-title":"Survival of Listeria monocytogenes in egg washwater","volume":"12","author":"Laird","year":"1991","journal-title":"Int. J. Food Microbiol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1080\/10408398.2010.507909","article-title":"A review of the ecology, genomics and stress response of Listeria innocua and Listeria monocytogenes","volume":"52","author":"Milillo","year":"2012","journal-title":"Crit. Revs. Food Sci. Nutr."},{"key":"ref_106","unstructured":"Sam Houston State University (2023, July 05). Egg Processing. Available online: https:\/\/www.google.com\/url?sa=i&rct=j&q=&esrc=s&source=web&cd=&cad=rja&uact=8&ved=0CDgQw7AJahcKEwiYjN3Kl_j_AhUAAAAAHQAAAAAQAw&url=https%3A%2F%2Fwww.shsu.edu%2Facademics%2Fagricultural-sciences-and-engineering-technology%2Fdocuments%2FEggProcessing.ppt&psig=AOvVaw0hcQngwr9OrQlVBtsPjLSK&ust=1688667711474197&opi=89978449."},{"key":"ref_107","first-page":"4","article-title":"Listeria-review of epidemiology and pathogenesis","volume":"40","author":"Ramaswamy","year":"2007","journal-title":"J. Microbiol. Immunol. Infect."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"429","DOI":"10.4315\/0362-028X.JFP-12-213","article-title":"Frequency of Salmonella and Listeria monocytogenes in five commercial brands of chicken eggs using a combined method of enrichment and nested-PCR","volume":"76","year":"2013","journal-title":"J. Food Prot."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.foodcont.2012.11.033","article-title":"Detection and isolation of Listeria spp. and Listeria monocytogenes in ready-to-eat foods with various selective culture media","volume":"32","author":"Jamali","year":"2013","journal-title":"Food Control"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"774","DOI":"10.1016\/j.foodcont.2009.09.008","article-title":"Listeria monocytogenes in raw salad vegetables sold at retail level in Malaysia","volume":"21","author":"Ponniah","year":"2010","journal-title":"Food Control"},{"key":"ref_111","first-page":"597","article-title":"Listeria monocytogenes, Yersinia enterocolitica and Salmonella enteritidis in quail eggs","volume":"28","year":"2004","journal-title":"Turk. J. Vet. Anim. Sci."},{"key":"ref_112","unstructured":"USDA (2023, July 15). Voluntary Grading of Shell Eggs. Minimum Facility and Operating Requirements for Shell Egg Grading and Packing Plants. 7 CFR 57.76(f)(11). Shell Egg Cleaning Operations, Available online: https:\/\/www.govinfo.gov\/content\/pkg\/CFR-2020-title7-vol3\/pdf\/CFR-2020-title7-vol3.pdf."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"R76","DOI":"10.1111\/j.1750-3841.2011.02053.x","article-title":"Toward better control of Salmonella contamination by taking advantage of the egg\u2019s selfdefense system: A review","volume":"76","author":"Zhang","year":"2011","journal-title":"J. Food Sci."},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Koutchma, T., Forney, L.J., and Moraru, C.I. (2009). Ultraviolet Light in Food Technology: Principles and Applications, CRC Press. [1st ed.].","DOI":"10.1201\/9781420059519"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"4433","DOI":"10.1016\/S0043-1354(02)00188-4","article-title":"Literature review\u2014Efficacy of various disinfectants against Legionella in water s ystems","volume":"36","author":"Kim","year":"2002","journal-title":"Water Res."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"6","DOI":"10.4315\/0362-028X.JFP-17-128","article-title":"Comparison of UV-C and Pulsed UV light treatments for reduction of Salmonella, Listeria monocytogenes, and Enterohemorrhagic Escherichia coli on Eggs","volume":"81","author":"Holck","year":"2018","journal-title":"J. Food Prot."},{"key":"ref_117","first-page":"25","article-title":"Agricultural electrostatic spray application: A review of significant research and development during the 20th century","volume":"51","author":"Law","year":"2001","journal-title":"J. Electrost."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1093\/ps\/82.1.158","article-title":"The effect of electrolyzed oxidative water applied using electrostatic spraying on pathogenic and indicator bacteria on the surface of eggs","volume":"82","author":"Russell","year":"2003","journal-title":"Poult. Sci."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"1864","DOI":"10.4315\/0362-028X.JFP-18-249","article-title":"Comparison of the efficacy of electrostatic versus conventional sprayer with commercial antimicrobials to inactivate Salmonella, Listeria monocytogenes, and Campylobacter jejuni for eggs and economic feasibility analysis","volume":"81","author":"Jiang","year":"2018","journal-title":"J. Food Prot."},{"key":"ref_120","unstructured":"Dreyer, J. (2023, July 07). Liquid Egg Processing Procedures, Key to Egg Market. Available online: https:\/\/www.wattagnet.com\/egg\/egg-processing\/article\/15527477\/liquid-egg-processing-procedures-key-to-egg-market-wattagnet."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"777","DOI":"10.4315\/0362-028X-52.11.777","article-title":"Listeria species in commercially broken raw liquid whole egg","volume":"52","author":"Leasor","year":"1989","journal-title":"J. Food Protect."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.ijfoodmicro.2010.01.013","article-title":"Detection of Listeria monocytogenes in raw and pasteurized liquid whole eggs and characterization by PFGE","volume":"138","author":"Rivoal","year":"2010","journal-title":"Int. J. Food Microbiol."},{"key":"ref_123","unstructured":"Stadelman, W.J., and Cotterill, O.J. (1995). Egg Science and Technology, Food Products Press, Haworth Press."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"703","DOI":"10.4315\/0362-028X-68.4.703","article-title":"Thermal resistance of Salmonella enterica serotypes, Listeria monocytogenes, and Staphylococcus aureus in high solids liquid egg mixes","volume":"68","author":"Li","year":"2005","journal-title":"J. Food Protect."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.4315\/0362-028X-58.11.1211","article-title":"Heat resistance of Listeria monocytogenes Scott A and HAL 957E1 in various liquid egg products","volume":"58","author":"Bartlett","year":"1995","journal-title":"J. Food Protect."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"100081","DOI":"10.1016\/j.jfp.2023.100081","article-title":"Ready-to-eat egg products formulated with nisin and organic acids to control Listeria monocytogenes","volume":"86","author":"Shrestha","year":"2023","journal-title":"J. Food Prot."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1111\/j.1472-765X.1989.tb00298.x","article-title":"The heat resistance of Listeria monocytogenes","volume":"9","author":"Mackey","year":"1989","journal-title":"Lett. Appl. Microbiol."},{"key":"ref_128","unstructured":"Shute, N. (2019, June 24). Recall Reveals an Egg\u2019s Long Path to the Deli Sandwich. 8 February 2012. Available online: https:\/\/www.npr.org\/sections\/thesalt\/2012\/02\/07\/146540839\/recall-reveals-an-eggs-long-path-to-the-deli-sandwich."},{"key":"ref_129","unstructured":"Powell, D. (2019, June 27). Listeria in Hard-Boiled Eggs Recall Widens to 34 States. 7 November 2012. Available online: https:\/\/www.barfblog.com\/2012\/02\/listeria-in-hard-boiled-eggs-recall-widens-to-34-states\/."},{"key":"ref_130","unstructured":"CDC (2023, July 10). Centers for Disease Control and Prevention (CDC). Outbreak of Listeria Infections Linked to Hard-Boiled Eggs, Available online: https:\/\/www.cdc.gov\/listeria\/outbreaks\/eggs-12-19\/index.html."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/11\/9\/2164\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T20:39:54Z","timestamp":1760128794000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/11\/9\/2164"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,8,27]]},"references-count":130,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2023,9]]}},"alternative-id":["microorganisms11092164"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms11092164","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,8,27]]}}}