{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T05:09:49Z","timestamp":1791522589285,"version":"4.3.4"},"reference-count":42,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,2,18]],"date-time":"2021-02-18T00:00:00Z","timestamp":1613606400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,2,18]],"date-time":"2021-02-18T00:00:00Z","timestamp":1613606400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Biofluorescence has been detected in several nocturnal-crepuscular organisms from invertebrates to birds and mammals. Biofluorescence in mammals has been detected across the phylogeny, including the monotreme duck-billed platypus (<jats:italic>Ornithorhyncus anatinus<\/jats:italic>), marsupial opossums (Didelphidae), and New World placental flying squirrels (<jats:italic>Gluacomys<\/jats:italic> spp.). Here, we document vivid biofluorescence of springhare (Pedetidae) in both museum specimens and captive individuals\u2014the first documented biofluorescence of an Old World placental mammal. We explore the variation in biofluorescence across our sample and characterize its physical and chemical properties. The striking visual patterning and intensity of color shift was unique relative to biofluorescence found in other mammals. We establish that biofluorescence in springhare likely originates within the cuticle of the hair fiber and emanates, at least partially, from several fluorescent porphyrins and potentially one unassigned molecule absent from our standard porphyrin mixture. This discovery further supports the hypothesis that biofluorescence may be ecologically important for nocturnal-crepuscular mammals and suggests that it may be more broadly distributed throughout Mammalia than previously thought.<\/jats:p>","DOI":"10.1038\/s41598-021-83588-0","type":"journal-article","created":{"date-parts":[[2021,2,20]],"date-time":"2021-02-20T09:14:59Z","timestamp":1613812499000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":38,"title":["Vivid biofluorescence discovered in the nocturnal Springhare (Pedetidae)"],"prefix":"10.1038","volume":"11","author":[{"given":"Erik R.","family":"Olson","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michaela R.","family":"Carlson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"V. M. Sadagopa","family":"Ramanujam","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lindsay","family":"Sears","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sharon E.","family":"Anthony","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paula Spaeth","family":"Anich","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leigh","family":"Ramon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alissa","family":"Hulstrand","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michaela","family":"Jurewicz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Adam S.","family":"Gunnelson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Allison M.","family":"Kohler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan G.","family":"Martin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,2,18]]},"reference":[{"key":"83588_CR1","doi-asserted-by":"publisher","first-page":"8980","DOI":"10.1073\/pnas.0813201106","volume":"106","author":"H Zhao","year":"2009","unstructured":"Zhao, H. et al. The evolution of color vision in nocturnal mammals. PNAS 106, 8980\u20138985 (2009).","journal-title":"PNAS"},{"key":"83588_CR2","doi-asserted-by":"publisher","first-page":"1471","DOI":"10.1098\/rspb.2013.2995","volume":"281","author":"RH Douglas","year":"2014","unstructured":"Douglas, R. H. & Jeffery, G. The spectral transmission of ocular media suggests ultraviolet sensitivity is widespread among mammals. Proc. R. Soc. B. 281, 1471\u20132954. https:\/\/doi.org\/10.1098\/rspb.2013.2995 (2014).","journal-title":"Proc. R. Soc. B."},{"key":"83588_CR3","doi-asserted-by":"publisher","first-page":"2273","DOI":"10.1098\/rspb.2001.1813","volume":"268","author":"SM Pearn","year":"2001","unstructured":"Pearn, S. M., Bennett, A. T. & Cuthill, I. C. Ultraviolet vision, fluorescence and mate choice in a parrot, the budgerigar Melopsittacus undulates. Proc. R. Soc. B. 268, 2273\u20132279. https:\/\/doi.org\/10.1098\/rspb.2001.1813 (2001).","journal-title":"Proc. R. Soc. B."},{"key":"83588_CR4","doi-asserted-by":"publisher","first-page":"e10798","DOI":"10.1371\/journal.pone.0010798","volume":"5","author":"M Olofsson","year":"2010","unstructured":"Olofsson, M., Vallin, A., Jakobsson, S. & Wiklund, C. Marginal eyespots on butterfly wings deflect bird attacks under low light intensities with UV wavelengths. PLoS ONE 5, e10798. https:\/\/doi.org\/10.1371\/journal.pone.0010798 (2010).","journal-title":"PLoS ONE"},{"key":"83588_CR5","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1034\/j.1600-0706.2002.980315.x","volume":"98","author":"J Honkavaara","year":"2008","unstructured":"Honkavaara, J., Koivula, M., Korpimaki, E., Siitari, H. & Viitala, J. Ultraviolet vision and foraging in terrestrial vertebrates. Oikos 98, 505\u2013511. https:\/\/doi.org\/10.1034\/j.1600-0706.2002.980315.x (2008).","journal-title":"Oikos"},{"key":"83588_CR6","doi-asserted-by":"publisher","DOI":"10.1093\/jmammal\/gyaa083","author":"B McDonald","year":"2020","unstructured":"McDonald, B., Geiger, B. & Vrla, S. Ultraviolet vision in Ord\u2019s kangaroo rat (Dipodomys ordii). J. Mammal. https:\/\/doi.org\/10.1093\/jmammal\/gyaa083 (2020).","journal-title":"J. Mammal."},{"key":"83588_CR7","doi-asserted-by":"publisher","first-page":"2941","DOI":"10.1098\/rstb.2009.0044","volume":"364","author":"DM Hunt","year":"2009","unstructured":"Hunt, D. M., Carvalho, L. S., Cowing, J. A. & Davies, W. L. Evolution and spectral tuning of visual pigments in birds and mammals. Phil. Trans. R. Soc. B. 364, 2941\u20132955. https:\/\/doi.org\/10.1098\/rstb.2009.0044 (2009).","journal-title":"Phil. Trans. R. Soc. B."},{"key":"83588_CR8","doi-asserted-by":"publisher","first-page":"R161","DOI":"10.1016\/j.cub.2007.01.037","volume":"17","author":"WL Davies","year":"2007","unstructured":"Davies, W. L. et al. Visual pigments of the platypus: a novel route to mammalian colour vision. Curr. Biol. 17, R161\u2013R163. https:\/\/doi.org\/10.1016\/j.cub.2007.01.037 (2007).","journal-title":"Curr. Biol."},{"key":"83588_CR9","doi-asserted-by":"publisher","unstructured":"Jeng, M.-L. Biofluorescence in terrestrial animals, with emphasis on fireflies: A review and field observation. In Bioluminescence \u2013 analytical applications and basic biology (ed. Suzuki, H.) Ch. 6, https:\/\/doi.org\/10.5772\/intechopen.86029 (IntechOpen, 2019).","DOI":"10.5772\/intechopen.86029"},{"key":"83588_CR10","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0083259","author":"JS Sparks","year":"2014","unstructured":"Sparks, J. S. et al. The covert world of fish biofluorescence: A phylogenetically widespread and phenotypically variable phenomenon. PLoS ONE https:\/\/doi.org\/10.1371\/journal.pone.0083259 (2014).","journal-title":"PLoS ONE"},{"key":"83588_CR11","doi-asserted-by":"publisher","first-page":"1277","DOI":"10.1016\/j.isci.2019.07.019","volume":"19","author":"HB Park","year":"2019","unstructured":"Park, H. B. et al. Bright green biofluorescence in sharks derives from Bromo-kynurenine metabolism. iScience 19, 1277\u20131286. https:\/\/doi.org\/10.1016\/j.isci.2019.07.019 (2019).","journal-title":"iScience"},{"key":"83588_CR12","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1206\/3845.1","volume":"3845","author":"DF Gruber","year":"2015","unstructured":"Gruber, D. F. & Sparks, J. S. First observation of fluorescence in marine turtles. Am. Mus. Novit. 3845, 1\u20138. https:\/\/doi.org\/10.1206\/3845.1 (2015).","journal-title":"Am. Mus. Novit."},{"key":"83588_CR13","doi-asserted-by":"publisher","first-page":"3672","DOI":"10.1073\/pnas.1701053114","volume":"114","author":"C Taboada","year":"2017","unstructured":"Taboada, C. et al. Naturally occurring fluorescence in frogs. PNAS 114, 3672\u20133677. https:\/\/doi.org\/10.1073\/pnas.1701053114 (2017).","journal-title":"PNAS"},{"key":"83588_CR14","doi-asserted-by":"publisher","first-page":"698","DOI":"10.1038\/s41598-017-19070-7","volume":"8","author":"D Pr\u00f6tzel","year":"2018","unstructured":"Pr\u00f6tzel, D. et al. Widespread bone-based fluorescence in chameleons. Sci. Rep. 8, 698. https:\/\/doi.org\/10.1038\/s41598-017-19070-7 (2018).","journal-title":"Sci. Rep."},{"key":"83588_CR15","doi-asserted-by":"publisher","first-page":"5388","DOI":"10.1038\/s41598-019-41959-8","volume":"9","author":"S Goutte","year":"2019","unstructured":"Goutte, S. et al. Intense bone fluorescence reveals hidden patterns in pumpkin toadlets. Sci. Rep. 9, 5388. https:\/\/doi.org\/10.1038\/s41598-019-41959-8 (2019).","journal-title":"Sci. Rep."},{"key":"83588_CR16","doi-asserted-by":"publisher","first-page":"2821","DOI":"10.1038\/s41598-020-58528-9","volume":"10","author":"JY Lamb","year":"2020","unstructured":"Lamb, J. Y. & Davis, M. P. Salamanders and other amphibians are aglow with biofluorescence. Sci. Rep. 10, 2821. https:\/\/doi.org\/10.1038\/s41598-020-58528-9 (2020).","journal-title":"Sci. Rep."},{"key":"83588_CR17","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1676\/09-125.1","volume":"123","author":"CS Weidensaul","year":"2011","unstructured":"Weidensaul, C. S., Colvin, B. A., Brinker, D. F. & Huy, J. S. Use of ultraviolet light as an aid in age classification of owls. Wilson J Ornithol. 123, 373\u2013377. https:\/\/doi.org\/10.1676\/09-125.1 (2011).","journal-title":"Wilson J Ornithol."},{"key":"83588_CR18","doi-asserted-by":"publisher","first-page":"19115","DOI":"10.1038\/s41598-019-55522-y","volume":"9","author":"C Camacho","year":"2019","unstructured":"Camacho, C., Negro, J. J., Redondo, I., Palacios, S. & S\u00e1ez-G\u00f3mez, P. Correlates of individual variation in the porphyrin-based fluorescence of red-necked nightjars. Sci. Rep. 9, 19115. https:\/\/doi.org\/10.1038\/s41598-019-55522-y (2019).","journal-title":"Sci. Rep."},{"key":"83588_CR19","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1093\/jmammal\/gyy177","volume":"100","author":"AM Kohler","year":"2019","unstructured":"Kohler, A. M., Olson, E. R., Martin, J. G. & Anich, P. S. Ultraviolet fluorescence discovered in New World flying squirrels (Glaucomys). J. Mammal. 100, 21\u201330. https:\/\/doi.org\/10.1093\/jmammal\/gyy177 (2019).","journal-title":"J. Mammal."},{"key":"83588_CR20","first-page":"4","volume":"83","author":"DH Meisner","year":"1983","unstructured":"Meisner, D. H. Psychedelic opossums: fluorescence of the skin and fur of Didelphis virginiana Kerr. Ohio J. Sci. 83, 4 (1983).","journal-title":"Ohio J. Sci."},{"key":"83588_CR21","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1515\/mamm.1985.49.2.249","volume":"49","author":"RH Pine","year":"1985","unstructured":"Pine, R. H., Rice, J. E., Bucher, J. E., Tank, D. J. Jr. & Greenhall, A. M. Labile pigments and fluorescent pelage in Didelphid marsupials. Mammalia 49, 249\u2013256 (1985).","journal-title":"Mammalia"},{"key":"83588_CR22","doi-asserted-by":"publisher","DOI":"10.1515\/mammalia-2020-0027","author":"PS Anich","year":"2020","unstructured":"Anich, P. S. et al. Biofluorescence in the platypus (Orinthorhynchus anatinus). Mammalia https:\/\/doi.org\/10.1515\/mammalia-2020-0027 (2020).","journal-title":"Mammalia"},{"key":"83588_CR23","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1023\/A:1027331727034","volume":"4","author":"CA Matthee","year":"1997","unstructured":"Matthee, C. A. & Robinson, T. J. Mitochondrial DNA phylogeography and comparative cytogenetics of the springhare, Pedetes capensis (Mammalia: Reodentia). J. Mammal. Evol. 4, 53\u201373. https:\/\/doi.org\/10.1023\/A:1027331727034 (1997).","journal-title":"J. Mammal. Evol."},{"key":"83588_CR24","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1111\/j.1365-2028.1995.tb01044.x","volume":"33","author":"DJ Augustine","year":"1995","unstructured":"Augustine, D. J., Manzon, A., Klopp, C. & Elter, J. Habitat selection and group foraging of the springhare, Pedetes capensis larvalis Hollister, East Africa. Afr. J. Ecol. 33, 347\u2013357 (1995).","journal-title":"Afr. J. Ecol."},{"issue":"2","key":"83588_CR25","first-page":"103","volume":"36","author":"DM Peinke","year":"2006","unstructured":"Peinke, D. M. & Brown, C. R. Habitat use by the southern springhare (Pedetes capensis) in the Eastern Cape Province, South Africa. S. Afr. J. Wildl. Res. 36(2), 103\u2013111 (2006).","journal-title":"S. Afr. J. Wildl. Res."},{"key":"83588_CR26","doi-asserted-by":"publisher","first-page":"663","DOI":"10.1017\/S002531540002693X","volume":"33","author":"GY Kennedy","year":"1954","unstructured":"Kennedy, G. Y. & Vevers, H. G. The occurrence of porphyrins in certain marine invertebrates. J. Mar. Biol. Ass. UK 33, 663\u2013576 (1954).","journal-title":"J. Mar. Biol. Ass. UK"},{"key":"83588_CR27","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1111\/j.1469-185X.1951.tb01358.x","volume":"26","author":"A Comfort","year":"1951","unstructured":"Comfort, A. The pigmentation of molluscan shells. Biol. Rev. 26, 285\u2013301. https:\/\/doi.org\/10.1111\/j.1469-185X.1951.tb01358.x (1951).","journal-title":"Biol. Rev."},{"key":"83588_CR28","doi-asserted-by":"publisher","first-page":"20121065","DOI":"10.1098\/rsif.2012.1065","volume":"10","author":"DB Thomas","year":"2012","unstructured":"Thomas, D. B., McGoverin, C. M., McGraw, K. J., James, H. F. & Madden, O. Vibrational spectroscopic analyses of unique yellow feather pigments (spheniscins) in penguins. J. R. Soc. Interface 10, 20121065. https:\/\/doi.org\/10.1098\/rsif.2012.1065 (2012).","journal-title":"J. R. Soc. Interface"},{"key":"83588_CR29","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1016\/0020-711X(78)90066-6","volume":"9","author":"TK With","year":"1978","unstructured":"With, T. K. On porphyrins in feathers of owls and bustards. Int. J. Biochem. 9, 893\u2013895 (1978).","journal-title":"Int. J. Biochem."},{"key":"83588_CR30","doi-asserted-by":"publisher","first-page":"398","DOI":"10.3109\/00365515709079995","volume":"9","author":"TK With","year":"1957","unstructured":"With, T. K. Pure unequivocal uroporphyrin III simplified method of preparation from turaco feathers. J. Clin. Lab Invest. 9, 398\u2013401 (1957).","journal-title":"J. Clin. Lab Invest."},{"key":"83588_CR31","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1080\/02724634.2012.639422","volume":"32","author":"AC Dooley Jr","year":"2012","unstructured":"Dooley, A. C. Jr. & Moncrief, N. D. Fluorescence provides evidence of congenital erythropoietic porphyria in 7000-year-old specimens of the eastern fox squirrel (Sciurus niger) from the Devil\u2019s Den. J. Vert. Paleontol. 32, 495\u2013497 (2012).","journal-title":"J. Vert. Paleontol."},{"key":"83588_CR32","doi-asserted-by":"publisher","first-page":"723","DOI":"10.1016\/j.bbamcr.2006.05.005","volume":"1763","author":"RS Ajioka","year":"2006","unstructured":"Ajioka, R. S., Phillips, J. D. & Kushner, J. P. Biosynthesis of heme in mammals. Biochem. Biophys. Acta. 1763, 723\u2013736. https:\/\/doi.org\/10.1016\/j.bbamcr.2006.05.005 (2006).","journal-title":"Biochem. Biophys. Acta."},{"key":"83588_CR33","doi-asserted-by":"publisher","DOI":"10.1117\/12.811913","author":"I Seo","year":"2009","unstructured":"Seo, I., Tseng, S. H., Cula, G. O., Bargo, P. R. & Kollias, N. Fluorescence spectroscopy for endogenous porphyrins in human facial skin. Proc. SPIE. https:\/\/doi.org\/10.1117\/12.811913 (2009).","journal-title":"Proc. SPIE."},{"key":"83588_CR34","doi-asserted-by":"publisher","DOI":"10.1117\/12.2527105","author":"C Heckl","year":"2019","unstructured":"Heckl, C. et al. Rapid spectrophotometric quantification of urinary porphyrins and porphobilinogen as screening tool for attacks of acute porphyria. Proc. SPIE. https:\/\/doi.org\/10.1117\/12.2527105 (2019).","journal-title":"Proc. SPIE."},{"key":"83588_CR35","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1172\/JCI107178","volume":"52","author":"EY Levin","year":"1973","unstructured":"Levin, E. Y. & Flyger, V. Erythropoietic Porphyria of Fox Squirrel Sciurus niger. J. Clin. Invest. 52, 96\u2013105 (1973).","journal-title":"J. Clin. Invest."},{"key":"83588_CR36","doi-asserted-by":"publisher","first-page":"519","DOI":"10.1016\/S0021-9258(18)74508-0","volume":"118","author":"WJ Turner","year":"1937","unstructured":"Turner, W. J. Studies on porphyria. I. Observations on the fox squirrel, Sciurus niger. J. Biol. Chem. 118, 519\u2013530 (1937).","journal-title":"J. Biol. Chem."},{"key":"83588_CR37","doi-asserted-by":"publisher","first-page":"2008","DOI":"10.1111\/j.1749-4877.2008.00088.x","volume":"216\u2013218","author":"DF Rivera","year":"2008","unstructured":"Rivera, D. F. & Leung, L.K.-P. A rare autosomal recessive condition, congenital erythropoietic porphyria, found in canefield rat Rattus sordidus Gould 1858. Integative Zool. 216\u2013218, 2008. https:\/\/doi.org\/10.1111\/j.1749-4877.2008.00088.x (2008).","journal-title":"Integative Zool."},{"issue":"1","key":"83588_CR38","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1111\/1523-1747.ep12485121","volume":"68","author":"DR Bickers","year":"1977","unstructured":"Bickers, D. R., Keogh, L., Rifkind, A. B., Harber, L. C. & Kappas, A. Studies in porphyria VI. Biosynthesis of porphyrins in mammalian skin and in the skin of porphyric patients. J. Invest. Dermatol. 68(1), 5\u20139. https:\/\/doi.org\/10.1111\/1523-1747.ep12485121 (1977).","journal-title":"J. Invest. Dermatol."},{"key":"83588_CR39","doi-asserted-by":"publisher","first-page":"14","DOI":"10.2307\/1379253","volume":"55","author":"RL Yolton","year":"1974","unstructured":"Yolton, R. L., Yolton, D. P., Renz, J. & Jacobs, G. H. Preretinal absorbance in sciurid eyes. J. Mammal. 55, 14\u201320 (1974).","journal-title":"J. Mammal."},{"key":"83588_CR40","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1016\/0003-2697(84)90115-5","volume":"137","author":"HC Friedmann","year":"1984","unstructured":"Friedmann, H. C. & Baldwin, E. T. Reverse-phase purification and silica gel thin-layer chromatography of porphyrin carboxylic acids. Anal. Biochem 137, 473\u2013480 (1984).","journal-title":"Anal. Biochem"},{"key":"83588_CR41","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1016\/0009-8981(84)90192-X","volume":"139","author":"CK Lim","year":"1984","unstructured":"Lim, C. K. & Peters, T. J. Urine and faecal porphyrin profiles by reversed-phase high performance liquid chromatography in the porphyrias. Clin. Chim. Acta. 139, 55\u201363 (1984).","journal-title":"Clin. Chim. Acta."},{"key":"83588_CR42","first-page":"251","volume":"33","author":"J To-Figueras","year":"2003","unstructured":"To-Figueras, J., Ozalla, D. & Mateu, C. H. Long-standing changes in the urinary profile of porphyrin isomers after clinical remission of porphyria cutanea tarda. Ann. Clin. Lab. Sci. 33, 251\u2013256 (2003).","journal-title":"Ann. Clin. Lab. Sci."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-83588-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-83588-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-83588-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,3]],"date-time":"2022-12-03T03:51:58Z","timestamp":1670039518000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-83588-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,2,18]]},"references-count":42,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["83588"],"URL":"https:\/\/doi.org\/10.1038\/s41598-021-83588-0","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,2,18]]},"assertion":[{"value":"27 October 2020","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 January 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 February 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"4125"}}