{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,12]],"date-time":"2025-10-12T04:26:23Z","timestamp":1760243183502,"version":"build-2065373602"},"reference-count":61,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2015,12,12]],"date-time":"2015-12-12T00:00:00Z","timestamp":1449878400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["Intramural"],"award-info":[{"award-number":["Intramural"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Toxins"],"abstract":"<jats:p>The US outbreak of B.anthracis infection in 2001 and subsequent cases in the US and Europe demonstrate that anthrax is a continuing risk for the developed world. While several bacterial components contribute to the pathogenesis of B. anthracis, production of lethal toxin (LT) is strongly associated with the development of hypotension and lethality. However, the mechanisms underlying the cardiovascular instability LT produces are unclear. Some evidence suggests that LT causes shock by impairing the peripheral vasculature, effects consistent with the substantial extravasation of fluid in patients dying with B. anthracis. Other data suggests that LT directly depresses myocardial function. However a clinical correlate for this latter possibility is less evident since functional studies and post-mortem examination in patients demonstrate absent or minimal cardiac changes. The purposes of this review were to first present clinical studies of cardiac functional and histologic pathology with B. anthracis infection and to then examine in vivo, in vitro, and ex vivo preclinical studies of LT\u2019s myocardial effects. Together, these data suggest that it is unclear whether that LT directly depresses cardiac function. This question is important for the clinical management and development of new therapies for anthrax and efforts should continue to be made to answer it.<\/jats:p>","DOI":"10.3390\/toxins7124891","type":"journal-article","created":{"date-parts":[[2015,12,14]],"date-time":"2015-12-14T02:57:29Z","timestamp":1450061849000},"page":"5417-5434","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["Does Bacillus anthracis Lethal Toxin Directly Depress Myocardial Function? A Review of Clinical Cases and Preclinical Studies"],"prefix":"10.3390","volume":"7","author":[{"given":"Dante","family":"Suffredini","sequence":"first","affiliation":[{"name":"Critical Care Medicine Department, Clinical Center, National Institutes of Health, Bethesda, MD 20892, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hanish","family":"Sampath-Kumar","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, Clinical Center, National Institutes of Health, Bethesda, MD 20892, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yan","family":"Li","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, Clinical Center, National Institutes of Health, Bethesda, MD 20892, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lernik","family":"Ohanjanian","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, Clinical Center, National Institutes of Health, Bethesda, MD 20892, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kenneth","family":"Remy","sequence":"additional","affiliation":[{"name":"Division of Critical Care Medicine, Department of Pediatrics, Washington University School of Medicine, St. Louis, MO 63110, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xizhong","family":"Cui","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, Clinical Center, National Institutes of Health, Bethesda, MD 20892, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Peter","family":"Eichacker","sequence":"additional","affiliation":[{"name":"Critical Care Medicine Department, Clinical Center, National Institutes of Health, Bethesda, MD 20892, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2015,12,12]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"933","DOI":"10.3201\/eid0706.010604","article-title":"Bioterrorism-related inhalational anthrax: The first 10 cases reported in the united states","volume":"7","author":"Jernigan","year":"2001","journal-title":"Emerg. Infect. Dis."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1452","DOI":"10.3201\/eid2009.131481","article-title":"Confirmed bacillus anthracis infection among persons who inject drugs, scotland, 2009\u20132010","volume":"20","author":"Booth","year":"2014","journal-title":"Emerg. Infect. Dis."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1097\/CCM.0000000000000026","article-title":"Two decades of mortality trends among patients with severe sepsis: A comparative meta-analysis","volume":"42","author":"Stevenson","year":"2014","journal-title":"Crit. Care Med."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"840","DOI":"10.1056\/NEJMra1208623","article-title":"Severe sepsis and septic shock","volume":"369","author":"Angus","year":"2013","journal-title":"N. Engl. J. Med."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1186\/1741-7015-11-217","article-title":"Anthracis associated cardiovascular dysfunction and shock: The potential contribution of both non-toxin and toxin components","volume":"11","author":"Remy","year":"2013","journal-title":"BMC Med."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1111\/j.1462-5822.2010.01438.x","article-title":"Transcriptome dysregulation by anthrax lethal toxin plays a key role in induction of human endothelial cell cytotoxicity","volume":"12","author":"Rolando","year":"2010","journal-title":"Cell. Microbiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"854","DOI":"10.1038\/nature09446","article-title":"Anthrax toxins cooperatively inhibit endocytic recycling by the Rab11\/Sec15 exocyst","volume":"467","author":"Guichard","year":"2010","journal-title":"Nature"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1278","DOI":"10.3390\/toxins3101278","article-title":"Anthrax lethal toxin-mediated disruption of endothelial VE-cadherin is attenuated by inhibition of the Rho-associated kinase pathway","volume":"3","author":"Warfel","year":"2011","journal-title":"Toxins"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1016\/j.tim.2014.02.012","article-title":"Anthrax lethal and edema toxins in anthrax pathogenesis","volume":"22","author":"Liu","year":"2014","journal-title":"Trends Microbiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1038\/nature12510","article-title":"Key tissue targets responsible for anthrax-toxin-induced lethality","volume":"501","author":"Liu","year":"2013","journal-title":"Nature"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"2291","DOI":"10.1073\/pnas.90.6.2291","article-title":"Pathology of inhalational anthrax in 42 cases from the sverdlovsk outbreak of 1979","volume":"90","author":"Abramova","year":"1993","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1038\/modpathol.3880337","article-title":"Quantitative pathology of inhalational anthrax I: Quantitative microscopic findings","volume":"14","author":"Grinberg","year":"2001","journal-title":"Mod. Pathol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e466","DOI":"10.1371\/journal.pone.0000466","article-title":"Anthrax toxins induce shock in rats by depressed cardiac ventricular function","volume":"2","author":"Watson","year":"2007","journal-title":"PLoS ONE"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Moayeri, M., Crown, D., Dorward, D.W., Gardner, D., Ward, J.M., Li, Y., Cui, X., Eichacker, P., and Leppla, S.H. (2009). The heart is an early target of anthrax lethal toxin in mice: A protective role for neuronal nitric oxide synthase (nNOS). PLoS Pathog., 5.","DOI":"10.1371\/journal.ppat.1000456"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1111\/j.1476-5381.2012.02040.x","article-title":"Toll-like receptor 4 knockout protects against anthrax lethal toxin-induced cardiac contractile dysfunction: Role of autophagy","volume":"167","author":"Kandadi","year":"2012","journal-title":"Br. J. Pharmacol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"3884","DOI":"10.1016\/j.ijcard.2013.06.050","article-title":"Anthrax lethal toxin induces acute diastolic dysfunction in rats through disruption of the phospholamban signaling network","volume":"168","author":"Golden","year":"2013","journal-title":"Int. J. Cardiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1885","DOI":"10.1086\/657408","article-title":"Anthrax lethal and edema toxins produce different patterns of cardiovascular and renal dysfunction and synergistically decrease survival in canines","volume":"202","author":"Sweeney","year":"2010","journal-title":"J. Infect. Dis."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"858","DOI":"10.1001\/jama.287.7.858","article-title":"Fatal inhalational anthrax with unknown source of exposure in a 61-year-old woman in new york city","volume":"287","author":"Mina","year":"2002","journal-title":"JAMA"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1186\/1471-2334-13-408","article-title":"Two anthrax cases with soft tissue infection, severe oedema and sepsis in danish heroin users","volume":"13","author":"Russell","year":"2013","journal-title":"BMC Infect. Dis."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"721","DOI":"10.3390\/toxins3060721","article-title":"Hemodynamic effects of anthrax toxins in the rabbit model and the cardiac pathology induced by lethal toxin","volume":"3","author":"Lawrence","year":"2011","journal-title":"Toxins"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1146\/annurev.biochem.75.103004.142728","article-title":"Anthrax toxin: Receptor binding, internalization, pore formation, and translocation","volume":"76","author":"Young","year":"2007","journal-title":"Annu. Rev. Biochem."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1038","DOI":"10.3390\/toxins2051038","article-title":"Consequences and utility of the zinc-dependent metalloprotease activity of anthrax lethal toxin","volume":"2","author":"Lee","year":"2010","journal-title":"Toxins"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1016\/j.micinf.2011.12.005","article-title":"Anthrax and the inflammasome","volume":"14","author":"Moayeri","year":"2012","journal-title":"Microbes Infect."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Rolli, J., Rosenblatt-Velin, N., Li, J., Loukili, N., Levrand, S., Pacher, P., Waeber, B., Feihl, F., Ruchat, P., and Liaudet, L. (2010). Bacterial flagellin triggers cardiac innate immune responses and acute contractile dysfunction. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0012687"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Zhang, W., Xu, X., Kao, R., Mele, T., Kvietys, P., Martin, C.M., and Rui, T. (2014). Cardiac fibroblasts contribute to myocardial dysfunction in mice with sepsis: The role of NLRP3 inflammasome activation. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0107639"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1172\/JCI113866","article-title":"Role of endotoxemia in cardiovascular dysfunction and mortality. Escherichia coli and staphylococcus aureus challenges in a canine model of human septic shock","volume":"83","author":"Natanson","year":"1989","journal-title":"J. Clin. Invest."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1016\/S0002-9440(10)63697-8","article-title":"Pathology and pathogenesis of bioterrorism-related inhalational anthrax","volume":"163","author":"Guarner","year":"2003","journal-title":"Am. J. Pathol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1607","DOI":"10.1056\/NEJMoa012948","article-title":"Index case of fatal inhalational anthrax due to bioterrorism in the united states","volume":"345","author":"Bush","year":"2001","journal-title":"N. Engl. J. Med."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2554","DOI":"10.1001\/jama.286.20.2554","article-title":"Death due to bioterrorism-related inhalational anthrax: Report of 2 patients","volume":"286","author":"Borio","year":"2001","journal-title":"JAMA"},{"key":"ref_30","first-page":"261","article-title":"Fatal case of inhalational anthrax mimicking intra-abdominal sepsis","volume":"66","author":"Quintiliani","year":"2002","journal-title":"Conn. Med."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"863","DOI":"10.1001\/jama.287.7.863","article-title":"Fatal inhalational anthrax in a 94-year-old connecticut woman","volume":"287","author":"Barakat","year":"2002","journal-title":"JAMA"},{"key":"ref_32","first-page":"457","article-title":"Human inhalation anthrax. A report of three fatal cases","volume":"36","author":"Albrink","year":"1960","journal-title":"Am. J. Pathol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"27","DOI":"10.26719\/2004.10.1-2.27","article-title":"Anthrax: Pathological aspects in autopsy cases in shiraz, islamic republic of iran, 1960\u20132001","volume":"10","author":"Tabei","year":"2004","journal-title":"East. Mediterr. Health J."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Berger, T., Kassirer, M., and Aran, A.A. (2014). Injectional anthrax\u2014New presentation of an old disease. Euro. Surveill., 19.","DOI":"10.2807\/1560-7917.ES2014.19.32.20877"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"968","DOI":"10.1086\/512372","article-title":"A case of naturally acquired inhalation anthrax: Clinical care and analyses of anti-protective antigen immunoglobulin g and lethal factor","volume":"44","author":"Walsh","year":"2007","journal-title":"Clin. Infect. Dis."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"11","DOI":"10.2807\/ese.13.51.19076-en","article-title":"Investigations and control measures following a case of inhalation anthrax in East London in a drum maker and drummer, October 2008","volume":"13","author":"Anaraki","year":"2008","journal-title":"Euro. Surveill."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"766","DOI":"10.1056\/NEJMcpc1003887","article-title":"Case records of the massachusetts general hospital. Case 25\u20132010. A 24-year-old woman with abdominal pain and shock","volume":"363","author":"Klempner","year":"2010","journal-title":"N. Engl. J. Med."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"310","DOI":"10.3201\/eid2002.130245","article-title":"Lethal factor and anti-protective antigen IGG levels associated with inhalation anthrax, minnesota, USA","volume":"20","author":"Sprenkle","year":"2014","journal-title":"Emerg. Infect. Dis."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e175","DOI":"10.1016\/j.jinf.2005.12.018","article-title":"Three rare cases of anthrax arising from the same source","volume":"53","author":"Babamahmoodi","year":"2006","journal-title":"J. Infect."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/j.jiph.2010.07.004","article-title":"A review of cutaneous anthrax and its outcome","volume":"3","author":"Doganay","year":"2010","journal-title":"J. Infect. Public Health"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"733","DOI":"10.2807\/ese.16.45.20008-en","article-title":"Two cases of infection with bacillus anthracis, Romania, October 2011","volume":"16","author":"Popescu","year":"2011","journal-title":"Euro. Surveill."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1186\/1471-2334-11-21","article-title":"A case of septicaemic anthrax in an intravenous drug user","volume":"11","author":"Powell","year":"2011","journal-title":"BMC Infect. Dis."},{"key":"ref_43","first-page":"843","article-title":"Injection anthrax\u2014A new outbreak in heroin users","volume":"109","author":"Grunow","year":"2012","journal-title":"Dtsch. Arztebl. Int."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2549","DOI":"10.1001\/jama.286.20.2549","article-title":"Clinical presentation of inhalational anthrax following bioterrorism exposure: Report of 2 surviving patients","volume":"286","author":"Mayer","year":"2001","journal-title":"JAMA"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1038\/173869a0","article-title":"Observations on experimental anthrax; demonstration of a specific lethal factor produced in vivo by bacillus anthracis","volume":"173","author":"Smith","year":"1954","journal-title":"Nature"},{"key":"ref_46","first-page":"323","article-title":"The chemical basis of the virulence of bacillus anthracis. IV. Secondary shock as the major factor in death of guinea-pigs from anthrax","volume":"36","author":"Smith","year":"1955","journal-title":"Br. J. Exp. Pathol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1164\/rccm.200608-1239CP","article-title":"Lethal and edema toxins in the pathogenesis of bacillus anthracis septic shock: Implications for therapy","volume":"175","author":"Sherer","year":"2007","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/j.mam.2009.07.003","article-title":"Cellular and systemic effects of anthrax lethal toxin and edema toxin","volume":"30","author":"Moayeri","year":"2009","journal-title":"Mol. Aspects Med."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"670","DOI":"10.1172\/JCI17991","article-title":"Bacillus anthracis lethal toxin induces TNF-alpha-independent hypoxia-mediated toxicity in mice","volume":"112","author":"Moayeri","year":"2003","journal-title":"J. Clin. Invest."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"R699","DOI":"10.1152\/ajpregu.00593.2003","article-title":"Lethality during continuous anthrax lethal toxin infusion is associated with circulatory shock but not inflammatory cytokine or nitric oxide release in rats","volume":"286","author":"Cui","year":"2004","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1016\/j.micpath.2007.12.001","article-title":"Anthrax toxin-induced shock in rats is associated with pulmonary edema and hemorrhage","volume":"44","author":"Kuo","year":"2008","journal-title":"Microb. Pathog."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"4670","DOI":"10.2741\/2416","article-title":"Lethal and edema toxins of anthrax induce distinct hemodynamic dysfunction","volume":"12","author":"Watson","year":"2007","journal-title":"Front. Biosci."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Kandadi, M.R., Hua, Y., Ma, H., Li, Q., Kuo, S.R., Frankel, A.E., and Ren, J. (2010). Anthrax lethal toxin suppresses murine cardiomyocyte contractile function and intracellular Ca2+ handling via a nadph oxidase-dependent mechanism. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0013335"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Kandadi, M.R., Yu, X., Frankel, A.E., and Ren, J. (2012). Cardiac-specific catalase overexpression rescues anthrax lethal toxin-induced cardiac contractile dysfunction: Role of oxidative stress and autophagy. BMC Med., 10.","DOI":"10.1186\/1741-7015-10-134"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2335","DOI":"10.2741\/3382","article-title":"Anthrax toxin: Pathologic effects on the cardiovascular system","volume":"14","author":"Golden","year":"2009","journal-title":"Front. Biosci (Landmark Ed.)"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"4516","DOI":"10.2741\/3544","article-title":"Pathophysiology of anthrax","volume":"14","author":"Frankel","year":"2009","journal-title":"Front. Biosci. (Landmark Ed.)"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1161\/circ.116.suppl_16.II_758-a","article-title":"Progressive left ventricle, myocyte dysfunction, and heart failure in the lethality of anthrax toxin in conscious dogs","volume":"116","author":"Cheng","year":"2007","journal-title":"Circulation"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1093\/infdis\/jir834","article-title":"Protective antigen antibody augments hemodynamic support in anthrax lethal toxin shock in canines","volume":"205","author":"Barochia","year":"2012","journal-title":"J. Infect. Dis."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"H1108","DOI":"10.1152\/ajpheart.01128.2010","article-title":"Anthrax edema toxin has camp-mediated stimulatory effects and high-dose lethal toxin has depressant effects in an isolated perfused rat heart model","volume":"300","author":"Hicks","year":"2011","journal-title":"Am. J. Physiol. Heart Circ. Physiol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"H1592","DOI":"10.1152\/ajpheart.00851.2014","article-title":"Bacillus anthracis edema but not lethal toxin challenge in rats is associated with depressed myocardial function in hearts isolated and tested in a langendorff system","volume":"308","author":"Li","year":"2015","journal-title":"Am. J. Physiol. Heart Circ. Physiol."},{"key":"ref_61","first-page":"139","article-title":"Molecular determinants for a cardiovascular collapse in anthrax","volume":"6","author":"Brojatsch","year":"2014","journal-title":"Front. Biosci. (Elite Ed.)"}],"container-title":["Toxins"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6651\/7\/12\/4891\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T20:53:58Z","timestamp":1760216038000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6651\/7\/12\/4891"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2015,12,12]]},"references-count":61,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2015,12]]}},"alternative-id":["toxins7124891"],"URL":"https:\/\/doi.org\/10.3390\/toxins7124891","relation":{},"ISSN":["2072-6651"],"issn-type":[{"type":"electronic","value":"2072-6651"}],"subject":[],"published":{"date-parts":[[2015,12,12]]}}}