Dr Maté Biro

Fields of research: Biophysics

Contact Details

Phone: +61293858020
Email: m.biro@unsw.edu.au

Tabs

Main

Visit the Biro Group website at EMBL Australia

Dr Maté Biro received his PhD with summa cum laude at the Max Planck Institute of Molecular Cell Biology and Genetics in Dresden, Germany.

Dr. Biro has previously worked at the Centenary Institute of Cancer Medicine and Cell Biology in Sydney, in the Imaging Informatics division at the Bioinformatics Institute of A*STAR in Singapore, at the MIT BioImaging Center in Cambridge, MA, USA and in the Belle collaboration at the KEK: High Energy Accelerator Research Organisation in Tsukuba, Japan.

As an EMBL Australia Group Leader he will expand his exciting studies on the way actomyosin complexes control cell shape, homeostasis and migration in cancer and immune function, focusing on the elucidation of biomechanical interaction within and between cells.

Dr. Biro is spearheading the use of complex in vitro and in vivo models for studying the cell-intrinsic actin cytoskeleton cues and dynamics that govern the invasive migration of tumour cells, the tissue scanning of T cells and their cytotoxic interaction with tumour cells. Using a multidisciplinary approach encompassing cell biology, biophysics, bioimage informatics and advanced light microscopy, he aims to unravel fundamental actin-based processes and develop new methodologies for basic cell biological research. Dr. Biro's research relies heavily on advanced microscopy and the development of image analysis platforms capable of automatically detecting and analysing the kinetics of actomyosin, cell movement and protrusions.

Dr. Biro's group is primarily concerned with the cell biology and mechanics of the actin cytoskeleton, and how immune cells (T cells) locate and kill cancer cells.

Main research topics are:

Mechanobiology of immune cell-tumour cell interactions
Single cell and population-wide immune cell migration dynamics
Advanced imaging and image analysis
Computational modelling and simulations

The group uses high-speed imaging to watch how cells move throughout complex 3-dimensional environments and also uses single molecule microscopes to observe the dynamics of actomyosin, a major and the force-generating component of the cytoskeleton.

When he is not involved with research Dr. Biro enjoys flying all things powered or unpowered and discovering the marvels of the underwater world whilst diving. He is also always happy to discuss his research and other passions over a beer...

University role:

EMBL Australia Group Leader

Research

Socio-economic Objective (SEO) tags:

Expanding Knowledge

Publications related to this Person

Book Chapters

Biro M; Maître JL, 2015, 'Dual pipette aspiration: Aunique tool for studying intercellular adhesion', in Methods in Cell Biology, pp. 255 - 267, http://dx.doi.org/10.1016/bs.mcb.2014.10.007

Journal articles

Geoghegan ND; Evelyn C; Whitehead LW; Pasternak M; McDonald P; Triglia T; Marapana DS; Kempe D; Thompson JK; Mlodzianoski MJ; Healer J; Biro M; Cowman AF; Rogers KL, 2021, '4D analysis of malaria parasite invasion offers insights into erythrocyte membrane remodeling and parasitophorous vacuole formation', Nature Communications, vol. 12, http://dx.doi.org/10.1038/s41467-021-23626-7

Zhang L; Chen C; Tay SS; Wen S; Cao C; Biro M; Jin D; Stenzel MH, 2021, 'Optimizing the Polymer Cloak for Upconverting Nanoparticles: An Evaluation of Bioactivity and Optical Performance', ACS Applied Materials and Interfaces, vol. 13, pp. 16142 - 16154, http://dx.doi.org/10.1021/acsami.1c01922

McDonald MM; Khoo WH; Ng PY; Xiao Y; Zamerli J; Thatcher P; Kyaw W; Pathmanandavel K; Grootveld AK; Moran I; Butt D; Nguyen A; Corr A; Warren S; Biro M; Butterfield NC; Guilfoyle SE; Komla-Ebri D; Dack MRG; Dewhurst HF; Logan JG; Li Y; Mohanty ST; Byrne N; Terry RL; Simic MK; Chai R; Quinn JMW; Youlten SE; Pettitt JA; Abi-Hanna D; Jain R; Weninger W; Lundberg M; Sun S; Ebetino FH; Timpson P; Lee WM; Baldock PA; Rogers MJ; Brink R; Williams GR; Bassett JHD; Kemp JP; Pavlos NJ; Croucher PI; Phan TG, 2021, 'Erratum: Osteoclasts recycle via osteomorphs during RANKL-stimulated bone resorption (Cell (2021) 184(5) (1330–1347.e13), (S0092867421001525), (10.1016/j.cell.2021.02.002))', Cell, vol. 184, pp. 1940, http://dx.doi.org/10.1016/j.cell.2021.03.010

Biro M, 2021, 'Two high-yield complementary methods to sort cell populations by their 2D or 3D migration speed (vol 31, pg 2749, 2020)', MOLECULAR BIOLOGY OF THE CELL, vol. 32, pp. 505 - 505, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000629288700007&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

McDonald MM; Khoo WH; Ng PY; Xiao Y; Zamerli J; Thatcher P; Kyaw W; Pathmanandavel K; Grootveld AK; Moran I; Butt D; Nguyen A; Warren S; Biro M; Butterfield NC; Guilfoyle SE; Komla-Ebri D; Dack MRG; Dewhurst HF; Logan JG; Li Y; Mohanty ST; Byrne N; Terry RL; Simic MK; Chai R; Quinn JMW; Youlten SE; Pettitt JA; Abi-Hanna D; Jain R; Weninger W; Lundberg M; Sun S; Ebetino FH; Timpson P; Lee WM; Baldock PA; Rogers MJ; Brink R; Williams GR; Bassett JHD; Kemp JP; Pavlos NJ; Croucher PI; Phan TG, 2021, 'Osteoclasts recycle via osteomorphs during RANKL-stimulated bone resorption', Cell, vol. 184, pp. 1330 - 1347.e13, http://dx.doi.org/10.1016/j.cell.2021.02.002

Hywood JD; Rice G; Pageon SV; Read MN; Biro M, 2021, 'Detection and characterization of chemotaxis without cell tracking', Journal of the Royal Society Interface, vol. 18, pp. 20200879, http://dx.doi.org/10.1098/rsif.2020.0879

Newman P; Osteil P; Anderson T; Sun J; Kempe D; Biro M; Shin J-W; Tam PPL; Zreiqat H, 2021, 'Chemomechanically voxelated niches for programmable histogenesis', , http://dx.doi.org/10.1101/2021.02.10.430691

Patkunarajah A; Schroeter L; Sanderson G; Biro M; Poole K, 2021, 'Disrupting ELKIN1-Dependent Mechanoelectrical Transduction Modulates Cell-Cell Interactions in Organotypic Tumour Spheroids', Biophysical Journal, vol. 120, pp. 65a - 66a, http://dx.doi.org/10.1016/j.bpj.2020.11.624

Arora A; Niño JLG; Myaing MZ; Chia S; Arasi B; Ravasio A; Huang RYJ; Dasgupta R; Biro M; Viasnoff V, 2020, 'Two high-yield complementary methods to sort cell populations by their 2D or 3D migration speed', Molecular Biology of the Cell, vol. 31, pp. 2779 - 2790, http://dx.doi.org/10.1091/mbc.E20-07-0466

Lamm N; Read MN; Nobis M; Van Ly D; Page SG; Masamsetti VP; Timpson P; Biro M; Cesare AJ, 2020, 'Nuclear F-actin counteracts nuclear deformation and promotes fork repair during replication stress', Nature Cell Biology, vol. 22, pp. 1460 - 1470, http://dx.doi.org/10.1038/s41556-020-00605-6

Niño JLG; Pageon SV; Tay SS; Colakoglu F; Kempe D; Hywood J; Mazalo JK; Cremasco J; Govendir MA; Dagley LF; Hsu K; Rizzetto S; Zieba J; Rice G; Prior V; O’neill G; Williams RJ; Nisbet DR; Kramer B; Webb AI; Luciani F; Read MN; Biro M, 2020, 'Cytotoxic T cells swarm by homotypic chemokine signalling', eLife, vol. 9, pp. 1 - 71, http://dx.doi.org/10.7554/eLife.56554

Lim HYG; Alvarez Y; Gasnier M; Wang Y; Tetlak P; Bissiere S; Wang H; Biro M; Plachta N, 2020, 'Keratins are asymmetrically inherited fate determinants in the mammalian embryo', Nature

Piper AK; Sophocleous RA; Ross SE; Evesson FJ; Saleh O; Bournazos A; Yasa J; Reed C; Woolger N; Sluyter R; Greer P; Biro M; Lemckert FA; Cooper ST, 2020, 'Loss of calpains-1 and -2 prevents repair of plasma membrane scrape injuries, but not small pores, and induces a severe muscular dystrophy', American Journal of Physiology - Cell Physiology, vol. 318, pp. C1226 - C1237, http://dx.doi.org/10.1152/ajpcell.00408.2019

Arora A; Niño JLG; Myaing MZ; Arasi B; Huang RY-J; Dasgupta R; Biro M; Viasnoff V, 2020, 'A Method to sort heterogenous cell populations based on migration in 2D and 3D environments', , http://dx.doi.org/10.1101/2020.05.06.080234

Niño JLG; Tay SS; Tearle JLE; Xie J; Govendir MA; Kempe D; Mazalo J; Drew AP; Colakoglu F; Kummerfeld SK; Proud CG; Biro M, 2020, 'The lifeact-EGFP mouse is a translationally controlled fluorescent reporter of T cell activation', Journal of Cell Science, vol. 133, http://dx.doi.org/10.1242/JCS.238014

Patkunarajah A; Stear JH; Moroni M; Schroeter L; Blaszkiewicz J; Tearle JLE; Cox CD; Fuerst C; Sánchez-Carranza O; Fernández MDÁO; Fleischer R; Eravci M; Weise C; Martinac B; Biro M; Lewin GR; Poole K, 2020, 'TMEM87a/Elkin1, a component of a novel mechanoelectrical transduction pathway, modulates melanoma adhesion and migration.', eLife, vol. 9, http://dx.doi.org/10.7554/eLife.53308

Obeidy P; Ju LA; Oehlers SH; Zulkhernain NS; Lee Q; Galeano Niño JL; Kwan RYQ; Tikoo S; Cavanagh LL; Mrass P; Cook AJL; Jackson SP; Biro M; Roediger B; Sixt M; Weninger W, 2020, 'Partial loss of actin nucleator actin-related protein 2/3 activity triggers blebbing in primary T lymphocytes', Immunology and Cell Biology, vol. 98, pp. 93 - 113, http://dx.doi.org/10.1111/imcb.12304

Meiring JCM; Bryce NS; Niño JLG; Gabriel A; Tay SS; Hardeman EC; Biro M; Gunning PW, 2019, 'Tropomyosin concentration but not formin nucleators mDia1 and mDia3 determines the level of tropomyosin incorporation into actin filaments', Scientific Reports, vol. 9, http://dx.doi.org/10.1038/s41598-019-42977-2

Mitchell CB; Black B; Sun F; Chrzanowski W; Cooper-White J; Maisonneuve B; Stringer B; Day B; Biro M; O’Neill GM, 2019, 'Tropomyosin Tpm 2.1 loss induces glioblastoma spreading in soft brain-like environments', Journal of Neuro-Oncology, vol. 141, pp. 303 - 313, http://dx.doi.org/10.1007/s11060-018-03049-z

Redpath GMI; Ecker M; Kapoor-Kaushik N; Vartoukian H; Carnell M; Kempe D; Biro M; Ariotti N; Rossy J, 2019, 'Flotillins promote T cell receptor sorting through a fast Rab5–Rab11 endocytic recycling axis', Nature Communications, vol. 10, http://dx.doi.org/10.1038/s41467-019-12352-w

Moran I; Nguyen A; Khoo WH; Butt D; Bourne K; Young C; Hermes JR; Biro M; Gracie G; Ma CS; Munier CML; Luciani F; Zaunders J; Parker A; Kelleher AD; Tangye SG; Croucher PI; Brink R; Read MN; Phan TG, 2018, 'Memory B cells are reactivated in subcapsular proliferative foci of lymph nodes', Nature Communications, vol. 9, http://dx.doi.org/10.1038/s41467-018-05772-7

Lamm N; Masamsetti P; Read M; Biro M; Cesare A, 2018, 'ATR and mTOR regulate F-actin to alter nuclear architecture and repair replication stress', Nuclear filamentous actin functions in the replication stress response, http://dx.doi.org/10.1101/451708

Pageon SV; Govendir MA; Kempe D; Biro M, 2018, 'Mechanoimmunology: Molecular-scale forces govern immune cell functions', Molecular Biology of the Cell, vol. 29, pp. 1919 - 1926, http://dx.doi.org/10.1091/mbc.E18-02-0120

Zenker J; White MD; Gasnier M; Alvarez YD; Lim HYG; Bissiere S; Biro M; Plachta N, 2018, 'Expanding Actin Rings Zipper the Mouse Embryo for Blastocyst Formation', Cell, vol. 173, pp. 776 - 791.e17, http://dx.doi.org/10.1016/j.cell.2018.02.035

Marshal AD; Bailey CG; Champ K; Vellozzi M; O'Young P; Metierre C; Feng Y; Thoeng A; Richards AM; Schmitz U; Biro M; Jayasinghe R; Ding L; Anderson L; Mardis ER; Rasko JEJ, 2017, 'CTCF genetic alterations in endometrial carcinoma are pro-tumorigenic', Oncogene, vol. 36, pp. 4100 - 4110, http://dx.doi.org/10.1038/onc.2017.25

Newman P; Galeano Niño JL; Graney P; Razal JM; Minett AI; Ribas J; Ovalle-Robles R; Biro M; Zreiqat H, 2017, 'Corrigendum: Relationship between nanotopographical alignment and stem cell fate with live imaging and shape analysis', Scientific reports, vol. 7, pp. 43073, http://dx.doi.org/10.1038/srep43073

Sedzinski J; Hannezo E; Tu F; Biro M; Wallingford JB, 2017, 'RhoA regulates actin network dynamics during apical surface emergence in multiciliated epithelial cells', Development, vol. 144, pp. e1.2 - e1.2, http://dx.doi.org/10.1242/dev.149286

Sedzinski J; Hannezo E; Tu F; Biro M; Wallingford JB, 2017, 'RhoA regulates actin network dynamics during apical surface emergence in multiciliated epithelial cells', Journal of Cell Science, vol. 130, pp. 420 - 428, http://dx.doi.org/10.1242/jcs.194704

Sedzinski J; Hannezo E; Tu F; Biro M; Wallingford JB, 2017, 'Correction: RhoA regulates actin network dynamics during apical surface emergence in multiciliated epithelial cells [J. Cell Sci. 130, (420-428)]', Journal of Cell Science, vol. 130, pp. 1017, http://dx.doi.org/10.1242/jcs.202234

Newman P; Galenano-Ninõ JL; Graney P; Razal JM; Minett AI; Ribas J; Ovalle-Robles R; Biro M; Zreiqat H, 2016, 'Relationship between nanotopographical alignment and stem cell fate with live imaging and shape analysis', Scientific Reports, vol. 6, http://dx.doi.org/10.1038/srep37909

Tay SS; Carol H; Biro M, 2016, 'TriKEs and BiKEs join CARs on the cancer immunotherapy highway', Human Vaccines and Immunotherapeutics, vol. 12, pp. 2790 - 2796, http://dx.doi.org/10.1080/21645515.2016.1198455

Galeano Niño JL; Kwan RYQ; Weninger W; Biro M, 2016, 'Antigen-specific T cells fully conserve antitumour function following cryopreservation', Immunology and Cell Biology, vol. 94, pp. 411 - 418, http://dx.doi.org/10.1038/icb.2015.105

Sedzinski J; Hannezo E; Tu F; Biro M; Wallingford JB, 2016, 'Emergence of an Apical Epithelial Cell Surface In Vivo', Developmental Cell, vol. 36, pp. 24 - 35, http://dx.doi.org/10.1016/j.devcel.2015.12.013

Newman P; Lu Z; Roohani-Esfahani SI; Church TL; Biro M; Davies B; King A; Mackenzie K; Minett AI; Zreiqat H, 2015, 'Porous and strong three-dimensional carbon nanotube coated ceramic scaffolds for tissue engineering', Journal of Materials Chemistry B, vol. 3, pp. 8337 - 8347, http://dx.doi.org/10.1039/c5tb01052g

Tong PL; Roediger B; Kolesnikoff N; Biro M; Tay SS; Jain R; Shaw LE; Grimbaldeston MA; Weninger W, 2015, 'The skin immune atlas: Three-dimensional analysis of cutaneous leukocyte subsets by multiphoton microscopy', Journal of Investigative Dermatology, vol. 135, pp. 84 - 93, http://dx.doi.org/10.1038/jid.2014.289

Bovellan M; Romeo Y; Biro M; Boden A; Chugh P; Yonis A; Vaghela M; Fritzsche M; Moulding D; Thorogate R; Jégou A; Thrasher AJ; Romet-Lemonne G; Roux PP; Paluch EK; Charras G, 2014, 'Cellular control of cortical actin nucleation', Current Biology, vol. 24, pp. 1628 - 1635, http://dx.doi.org/10.1016/j.cub.2014.05.069

Biro M; Munoz MA; Weninger W, 2014, 'Targeting Rho-GTPases in immune cell migration and inflammation', British Journal of Pharmacology, vol. 171, pp. 5491 - 5506, http://dx.doi.org/10.1111/bph.12658

Munoz MA; Biro M; Weninger W, 2014, 'T cell migration in intact lymph nodes in vivo', Current Opinion in Cell Biology, vol. 30, pp. 17 - 24, http://dx.doi.org/10.1016/j.ceb.2014.05.002

Weninger W; Biro M; Jain R, 2014, 'Leukocyte migration in the interstitial space of non-lymphoid organs', Nature Reviews Immunology, vol. 14, pp. 232 - 246, http://dx.doi.org/10.1038/nri3641

Biro M; Romeo Y; Kroschwald S; Bovellan M; Boden A; Tcherkezian J; Roux PP; Charras G; Paluch EK, 2013, 'Cell cortex composition and homeostasis resolved by integrating proteomics and quantitative imaging', Cytoskeleton, vol. 70, pp. 741 - 754, http://dx.doi.org/10.1002/cm.21142

Sedzinski J; Biro M; Oswald A; Tinevez JY; Salbreux G; Paluch E, 2011, 'Polar actomyosin contractility destabilizes the position of the cytokinetic furrow', Nature, vol. 476, pp. 462 - 468, http://dx.doi.org/10.1038/nature10286

Puah WC; Cheok LP; Biro M; Ng WT; Wasser M, 2011, 'TLM-converter: Reorganization of long time-lapse microscopy datasets for downstream image analysis', BioTechniques, vol. 51, pp. 49 - 54, http://dx.doi.org/10.2144/000113704

Biro M; Kroschwald S; Paluch E, 2010, 'Analysis of De Novo Cell Cortex Assembly in Blebs as a Novel Assay for Probing Cortical Dynamics and Regulation', BIOPHYSICAL JOURNAL, vol. 98, pp. 159A - 159A, http://dx.doi.org/10.1016/j.bpj.2009.12.853

Conference Papers

McDonald M; Khoo WH; Mohanty S; Terry R; Chai R; Quinn J; Pettitt J; Xiao Y; Baldock P; Rogers M; Croucher P; Phan T; Ng PY; Pavlos N; Biro M, 2019, 'Intravital imaging of osteoclasts in vivo reveals novel osteoclast fate which may underlie the therapeutic response to Denosumab withdrawal', in JOURNAL OF BONE AND MINERAL RESEARCH, WILEY, Orlando, FL, pp. 44 - 45, presented at Annual Meeting of the American-Society-for-Bone-and Mineral Research, Orlando, FL, 20 September 2019 - 23 September 2019, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000508356600130&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

Tong PL; Roediger B; Kolesnikoff N; Tay SS; Biro M; Shaw L; Jain R; Grimbaldeston M; Weninger W, 2014, 'The immune atlas of the skin: A topographical three-dimensional study using multiphoton microscopy and multi-parameter flow cytometry', in AUSTRALASIAN JOURNAL OF DERMATOLOGY, WILEY-BLACKWELL, pp. 20 - 21, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000335981400070&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

Sedzinski J; Biro M; Oswald A; Tinevez J-Y; Salbreux G; Paluch E, 2011, 'Mechanism of the actin cortex and shape stability during cytokinesis', in MOLECULAR BIOLOGY OF THE CELL, AMER SOC CELL BIOLOGY, Denver, CO, presented at Annual Meeting of the American-Society-for-Cell-Biology (ASCB), Denver, CO, 03 December 2011 - 07 December 2011, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000305505502499&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

Sedzinski J; Biro M; Salbreux G; Paluch E, 2011, 'Polar acto-myosin contractility and cleavage furrow stability during cytokinesis', in MOLECULAR BIOLOGY OF THE CELL, AMER SOC CELL BIOLOGY, Denver, CO, presented at Annual Meeting of the American-Society-for-Cell-Biology (ASCB), Denver, CO, 03 December 2011 - 07 December 2011, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000305505500093&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

Sedzinski J; Biro M; Oswald A; Tinevez J-Y; Salbreux G; Paluch E, 2011, 'Polar actin cortex mechanics and cell shape stability during cytokinesis', in EUROPEAN BIOPHYSICS JOURNAL WITH BIOPHYSICS LETTERS, SPRINGER, Budapest, HUNGARY, pp. 213 - 213, presented at 8th EBSA European Biophysics Congress, Budapest, HUNGARY, 23 August 2011 - 27 August 2011, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000293637300601&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

Conference Abstracts

McDonald M; Khoo WH; Mohanty S; Terry R; Chai R; Quinn J; Pettitt J; Xiao Y; Baldock P; Rogers M; Croucher P; Tri P; Ng PY; Pavlos N; Biro M, 2019, 'Intravital imaging of osteoclasts in vivo reveals novel osteoclast fate which may underlie the therapeutic response to Denosumab withdrawal', in JOURNAL OF BONE AND MINERAL RESEARCH, WILEY, Orlando, FL, Vol. 34, pp. 44 - 45, presented at Annual Meeting of the American-Society-for-Bone-and Mineral Research, Orlando, FL, 20 September 2019 - 23 September 2019, http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000508614700130&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=891bb5ab6ba270e68a29b250adbe88d1

Datasets

Weninger W; Biro M, 2015, Faculty Opinions recommendation of Actin depletion initiates events leading to granule secretion at the immunological synapse., http://dx.doi.org/10.3410/f.725506810.793507006

Weninger W; Biro M; Jain R, 2014, Faculty Opinions recommendation of Generation of compartmentalized pressure by a nuclear piston governs cell motility in a 3D matrix., http://dx.doi.org/10.3410/f.718547671.793499677