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Polyphenol oxidase

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3645: 605:, PPO initiates the chain of browning reactions. Exposure to oxygen when sliced or pureed also leads to enzymatic browning by PPO in fruits and vegetables. Examples in which the browning reaction catalyzed by PPO may be desirable include avocados, prunes, sultana grapes, black tea, and green coffee beans. 593:
Polyphenol oxidase is an enzyme found throughout the plant and animal kingdoms, including most fruits and vegetables. PPO has importance to the food industry because it catalyzes enzymatic browning when tissue is damaged from bruising, compression or indentations, making the produce less marketable
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Rompel, Annette; Fischer, Helmut; Meiwes, Dirk; Büldt-Karentzopoulos, K.; Dillinger, Renée; Tuczek, Felix; Witzel, Herbert; Krebs, B. (1999). "Purification and spectroscopic studies on catechol oxidases from Lycopus europaeus and Populus nigra: Evidence for a dinuclear copper center of type 3 and
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Present in the chloroplasts and mitochondria of all parts of an apple, PPO is the major enzyme responsible for enzymatic browning of apples. Due to an increase in consumer demand for pre-prepared fruits and vegetables, a solution for enzymatic browning has been a targeted area of research and new
722:, PPO is used in the potato as a defense against insect predation, leading to enzymatic browning from tissue damage. Damage in the skin tissue of potato tuber causes a disruption of cell compartmentation, resulting in browning. The brown or black pigments are produced from the reaction of PPO 617:-type PPO is located in the chloroplasts of mango skin cells and its phenolic substrates in the vacuoles. Sap burn is therefore the initiating event of PPO in mango skin, as it breaks down cell compartments. PPO is located in mango skin, sap and pulp, with highest activity levels in skin. 653:
product development. As an example, pre-sliced apples are an appealing consumer product, but slicing apples induces PPO activity, leading to browning of the cut surfaces and lowering their esthetic quality. Browning also occurs in apple juices and purees when poorly handled or processed.
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A mixture of monophenol oxidase and catechol oxidase enzymes is present in nearly all plant tissues, and can also be found in bacteria, animals, and fungi. In insects, cuticular polyphenol oxidases are present and their products are responsible for
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García-Molina F, Muñoz JL, Varón R, Rodríguez-López JN, García-Cánovas F, Tudela J (November 2007). "A review on spectrophotometric methods for measuring the monophenolase and diphenolase activities of tyrosinase".
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Espín JC, Morales M, Varón R, Tudela J, García-Cánovas F (October 1995). "A continuous spectrophotometric method for determining the monophenolase and diphenolase activities of apple polyphenol oxidase".
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Haghbeen K, Wue Tan E (January 2003). "Direct spectrophotometric assay of monooxygenase and oxidase activities of mushroom tyrosinase in the presence of synthetic and natural substrates".
3032: 562:, which are the products of polyphenol oxidase-catalysed reactions, or the consumption of the substrate. Alternative spectrophotometric method that involves the coupling of 1611:
Thaler JS, Karban R, Ullman DE, Boege K, Bostock RM (April 2002). "Cross-talk between jasmonate and salicylate plant defense pathways: effects on several plant parasites".
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Several assays were developed to monitor the activity of polyphenol oxidases and to evaluate the inhibition potency of polyphenol oxidase inhibitors. In particular,
388:, a protein that can form two or more different homo-oligomers (morpheein forms), but must come apart and change shape to convert between forms. It exists as a 958: 594:
and causing economic loss. Enzymatic browning due to PPO can also lead to loss of nutritional content in fruits and vegetables, further lowering their value.
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Wuyts N, De Waele D, Swennen R (2006). "Extraction and partial characterization of polyphenol oxidase from banana (Musa acuminata Grande naine) roots".
1227:"Three recombinantly expressed apple tyrosinases suggest the amino acids responsible for mono- versus diphenolase activity in plant polyphenol oxidases" 573:
such as 3-methyl-2-benzothiazolinonehydrazone hydrochloride (MBTH) was also used. Other techniques, such as activity staining assays with the use of
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Rescigno A, Sollai F, Rinaldi AC, Soddu G, Sanjust E (March 1997). "Polyphenol oxidase activity staining in polyacrylamide electrophoresis gels".
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Duke SO, Vaughn KC (April 1982). "Lack of involvement of polyphenol oxidase in ortho-hydroxylation of phenolic compounds in mung bean seedlings".
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Li Y, Zafar A, Kilmartin PA, Reynisson J, Leung IK (November 2017). "Development and Application of an NMR-Based Assay for Polyphenol Oxidases".
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Trémolières, Michèle; Bieth, Joseph G. (1984). "Isolation and characterization of the polyphenoloxidase from senescent leaves of black poplar".
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Sugumaran M, Lipke H (May 1983). "Quinone methide formation from 4-alkylcatechols: a novel reaction catalyzed by cuticular polyphenol oxidase".
854:) since both enzymes sharing type copper active site coordination. Hemocyanin also exhibits PPO activity, but with slowed kinetics from greater 1338:
Toledo L, Aguirre C (December 2017). "Enzymatic browning in avocado (Persea americana) revisited: History, advances, and future perspectives".
2569:"Identification of the amino acid position controlling the different enzymatic activities in walnut tyrosinase isoenzymes (jrPPO1 and jrPPO2)" 181: 858:
bulk at the active site. Partial denaturation actually improves hemocyanin's PPO activity by providing greater access to the active site.
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There are two types of inhibitor of PPO, those competitive to oxygen in the copper site of the enzyme and those competitive to phenolics.
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A Sánchez-Ferrer; J N Rodríguez-López; F García-Cánovas; F García-Carmona (1995). "Tyrosinase: A Comprehensive Review of Its Mechanism".
2806:"Aurone synthase is a catechol oxidase with hydroxylase activity and provides insights into the mechanism of plant polyphenol oxidases" 2293: 690:
PPO form can spontaneously activate during the first weeks of storage, generating the active enzyme with a molecular weight of 38 kDa.
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Valero E, Garcia-Moreno M, Varon R, Garcia-Carmona F (1991). "Time-dependent inhibition of grape polyphenol oxidase by tropolone".
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Decker H, Tuczek F (August 2000). "Tyrosinase/catecholoxidase activity of hemocyanins: structural basis and molecular mechanism".
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Nicolas JJ, Richard-Forget FC, Goupy PM, Amiot MJ, Aubert SY (1994). "Enzymatic browning reactions in apple and apple products".
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enzyme with unclear synthesis and function. In functional chloroplasts, it may be involved in oxygen chemistry like mediation of
174: 1121:"High level protein-purification allows the unambiguous polypeptide determination of latent isoform PPO4 of mushroom tyrosinase" 3190: 2929: 3287: 1041: 555:-based assays are widely applied. The most common UV/Vis spectrophotometry assay involves the monitoring of the formation of 141: 1519:"Similar but Still Different: Which Amino Acid Residues Are Responsible for Varying Activities in Type-III Copper Enzymes?" 552: 2907: 3090: 625:
PPO in avocados causes rapid browning upon exposure to oxygen, a multistep process involving oxidation reactions of both
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Rocha AM, Cano MP, Galeazzi MA, Morais AM (1 August 1998). "Characterisation of 'Starking' apple polyphenoloxidase".
416: 2865:"Latent and active aurone synthase from petals of C. grandiflora: a polyphenol oxidase with unique characteristics" 1969:
Pomerantz SH (June 1964). "Tyrosine hydroxylation catalyzed by mammalian tyrosinase: an improved method of assay".
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and found in wine. This compound production is responsible for the lower level of browning in certain white wines.
3102: 3048: 135: 2720: 1178:"Latent and active abPPO4 mushroom tyrosinase cocrystallized with hexatungstotellurate(VI) in a single crystal" 117: 2393:. Novel Foods Section, Food Directorate, Health Products and Food Branch, Health Canada, Ottawa. 20 March 2015 3520: 122: 2328:
Son SM, Moon KD, Lee CY (April 2001). "Inhibitory effects of various antibrowning agents on apple slices".
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Robinson SP, Loveys BR, Chacko EK (1993). "Polyphenol Oxidase Enzymes in the Sap and Skin of Mango Fruit".
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Mallette MF, Dawson CR (August 1949). "On the nature of highly purified mushroom tyrosinase preparations".
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groups in the tuber. In potatoes, PPO genes are not only expressed in potato tubers, but also in leaves,
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In mangoes, PPO catalyzed enzymatic browning is mainly caused by sap burn which leads to skin browning.
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has also been used in recent research to eliminate the PPO activity from seedlings of higher plants.
3515: 3469: 3412: 2950: 2414:"Purification and Characterization of Latent Polyphenol Oxidase from Apricot (Prunus armeniaca L.)" 1716:
Del Signore A, Romeoa F, Giaccio M (May 1997). "Content of phenolic substances in basidiomycetes".
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McLarin, Mark-Anthony; Leung, Ivanhoe K. H. (2020). "Substrate Specificity of Polyphenol Oxidase".
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substituents at the 1, 2 positions, with no carbon between). It can also further catalyse the
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Mayer AM (November 2006). "Polyphenol oxidases in plants and fungi: going places? A review".
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is homologous to plant polyphenol oxidase, but contains certain significant modifications.
524: 405: 98: 2085:Ăśnal MĂś (2007). "Properties of polyphenol oxidase from Anamur banana (Musa cavendishii)". 8: 3665: 3474: 3208: 861: 660: 74: 2821: 2739: 2637: 2584: 2527: 1624: 1242: 1136: 1089: 37: 3407: 2971: 2889: 2864: 2840: 2805: 2701: 2601: 2568: 2544: 2511: 2487: 2462: 2438: 2413: 2030: 2005: 1675: 1644: 1593: 1553: 1518: 1503: 1363: 1308: 1283: 1259: 1226: 1202: 1177: 1153: 1120: 982: 484:(2-S glutathionyl caftaric acid) is an oxidation compound produced by action of PPO on 3117: 2782: 2747: 2645: 2512:"Conversion of walnut tyrosinase into a catechol oxidase by site directed mutagenesis" 2341: 2193: 2168: 2125:
Vámos-Vigyázó L (1981). "Polyphenol oxidase and peroxidase in fruits and vegetables".
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Molitor C, Mauracher SG, Pargan S, Mayer RL, Halbwirth H, Rompel A (September 2015).
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Plants make use of polyphenol oxidase as one in a suite of chemical defences against
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contains a single phenolic ring that may be oxidised by the action of PPOs to form
293: 201: 2256:"The Avocado Lab: An Inquiry-Driven Exploration of an Enzymatic Browning Reaction" 1351: 970: 867:
Aurone synthase catalyzes the formation of aurones. Aurone synthase purified from
3500: 3484: 3397: 3095: 1756: 1284:"A Peptide-Induced Self-Cleavage Reaction Initiates the Activation of Tyrosinase" 719: 352: 2478: 1002:
Biochimica et Biophysica Acta (BBA) - Protein Structure and Molecular Enzymology
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Proceedings of the National Academy of Sciences of the United States of America
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Vaughn KC, Duke SO (1984). "Function of polyphenol oxidase in higher plants".
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combinations constitute a promising practical anti-browning method as treated
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Derardja AE, Pretzler M, Kampatsikas I, Barkat M, Rompel A (September 2017).
2311: 2240: 2006:"Oxidation of monohydric phenol substrates by tyrosinase. An oximetric study" 1544: 1119:
Mauracher SG, Molitor C, Michael C, Kragl M, Rizzi A, Rompel A (March 2014).
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Derardja AE, Pretzler M, Kampatsikas I, Barkat M, Rompel A (December 2019).
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10.1002/(sici)1097-0010(199808)77:4<527::aid-jsfa76>3.0.co;2-e
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Mauracher SG, Molitor C, Al-Oweini R, Kortz U, Rompel A (September 2014).
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Shelby T. Peres; Kelsey A. Oonk; Kasandra J. Riley (29 October 2019).
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to suppress the expression of PPO, thus inhibiting fruit browning.
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PPO2) encoding polyphenol oxidases have been identified. The two
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Because the substrates of these PPO reactions are located in the
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Kampatsikas I, Bijelic A, Pretzler M, Rompel A (August 2017).
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Panis F, Kampatsikas I, Bijelic A, Rompel A (February 2020).
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Acta Crystallographica. Section D, Biological Crystallography
715: 703: 1383:"The Multiple Forms of Mushroom Tyrosinase. Interconversion" 1118: 3085: 3073: 3068: 3058: 3053: 3036: 3010: 2862: 1282:
Kampatsikas I, Bijelic A, Pretzler M, Rompel A (May 2019).
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spectroscopic similarities to tyrosinase and hemocyanin".
1933: 2052: 581:-based radioactive assays, oxygen consumption assay, and 458: 1715: 633:
products subsequently converted irreversibly into brown
2279: 1897: 1610: 2803: 1780:"Banana polyphenoloxidase. Preparation and properties" 3633: 2218: 1742: 667:
apples that contain a non-browning trait derived by
519:. Banana root PPO activity is strongly inhibited by 2166: 2046: 1971:
Biochemical and Biophysical Research Communications
682:as a climacteric fruit undergoes fast post-harvest 426:
differentiates between monophenol oxidase enzymes (
307:-quinones to produce black, brown or red pigments ( 2003: 663:engineered to reduce PPO activity, are a suite of 249:as substrates. The enzyme works by catalyzing the 2214: 2212: 2167:Thygesen PW, Dry IB, Robinson SP (October 1995). 1516: 881:, but the enzyme does not react with the classic 329:-quinone. Hence, PPOs may also be referred to as 235:that contains four atoms of copper per molecule. 3657: 2804:Molitor C, Mauracher SG, Rompel A (March 2016). 2623: 1709: 468: 1862: 1421: 585:(NMR)-based assay were also reported and used. 2357:Critical Reviews in Food Science and Nutrition 2282:Journal of the Science of Food and Agriculture 2209: 2162: 2160: 2158: 2156: 2127:Critical Reviews in Food Science and Nutrition 1936:Journal of Biochemical and Biophysical Methods 1575: 1569: 1462: 1340:Critical Reviews in Food Science and Nutrition 553:ultraviolet/visible (UV/Vis) spectrophotometry 3358: 2923: 2124: 2120: 2118: 2116: 2114: 2112: 2110: 2108: 1337: 2768: 2762: 1856: 1820: 1456: 1374: 2718: 2566: 2153: 2004:Naish-Byfield S, Riley PA (November 1992). 1997: 1962: 1927: 1891: 1422:Jolley RL, Robb DA, Mason HS (March 1969). 1415: 1380: 1037:"Catechol Oxidase - Structure and Activity" 1035:C Eicken; B Krebs; J C Sacchettini (1999). 1028: 949: 850:is homologous to the phenol oxidases (e.g. 3365: 3351: 2930: 2916: 2418:Journal of Agricultural and Food Chemistry 2383: 2327: 2323: 2321: 2105: 1830:Journal of Agricultural and Food Chemistry 1661: 1604: 1489: 993: 601:of plant cells damaged mainly by improper 336:Common foods producing the enzyme include 2888: 2839: 2829: 2670:Journal of Biological Inorganic Chemistry 2600: 2543: 2486: 2437: 2301: 2192: 2029: 1968: 1803: 1655: 1552: 1534: 1439: 1398: 1333: 1331: 1329: 1327: 1307: 1258: 1201: 1152: 434:-diphenol:oxygen oxidoreductase enzymes ( 379: 1517:Kampatsikas I, Rompel A (October 2020). 2318: 2080: 2078: 2076: 1790:(5). Oxford University Press: 508–513. 832:found in some invertebrates, including 714:Found in high concentrations in potato 3658: 1777: 1324: 515:. Another inhibitor of this enzyme is 3346: 2911: 2247: 1075: 1042:Current Opinion in Structural Biology 768:PPO1 shows a higher activity towards 588: 3123:24-hydroxycholesterol 7α-hydroxylase 2719:Beck G, Habicht GS (November 1996). 2712: 2084: 2073: 1069: 897:and must therefore be classified as 527:, as is banana fruit PPO by similar 1428:The Journal of Biological Chemistry 1387:The Journal of Biological Chemistry 221:polyphenol oxidase i, chloroplastic 13: 1676:10.1111/j.1399-3054.1982.tb00696.x 1504:10.1111/j.1399-3054.1984.tb04258.x 1381:Jolley RL, Mason HS (March 1965). 820: 783: 718:peel and 1–2 mm of the outer 575:polyacrylamide gel electrophoresis 14: 3682: 2748:10.1038/scientificamerican1196-60 1745:Plant Physiology and Biochemistry 417:pseudocyclic photophosphorylation 16:Enzyme involved in fruit browning 3643: 2721:"Immunity and the Invertebrates" 3103:Cholesterol 7 alpha-hydroxylase 3049:Flavin-containing monooxygenase 2856: 2797: 2660: 2617: 2567:Panis F, Rompel A (July 2020). 2560: 2503: 2454: 2405: 2348: 2273: 2261:. Rollins College, CourseSource 1771: 1736: 1682: 1510: 1483: 1145:10.1016/j.phytochem.2013.12.016 1098:10.1016/j.phytochem.2006.08.006 2771:Trends in Biochemical Sciences 2099:10.1016/j.foodchem.2005.10.048 1275: 1218: 1169: 1112: 959:Crit. Rev. Biochem. Mol. Biol. 927: 629:and polyphenols, resulting in 511:is a grape polyphenol oxidase 1: 2783:10.1016/S0968-0004(00)01602-9 2646:10.1016/s0031-9422(00)80367-2 2342:10.1016/s0308-8146(00)00274-0 1948:10.1016/S0165-022X(96)01201-8 1877:10.1016/S0003-2697(02)00408-6 1441:10.1016/S0021-9258(18)91800-4 1400:10.1016/S0021-9258(18)97603-9 1352:10.1080/10408398.2016.1175416 1055:10.1016/s0959-440x(99)00029-9 971:10.1080/10409238.2020.1768209 921: 674: 620: 498: 469:Distribution and applications 1983:10.1016/0006-291X(64)90359-6 1757:10.1016/j.plaphy.2006.06.005 1590:10.1016/0014-5793(83)80210-5 1014:10.1016/0167-4838(94)00204-t 776:PPO2 is more active towards 737: 709: 7: 3372: 3332:Deoxyhypusine monooxygenase 2479:10.1016/j.fochx.2019.100053 937:. Worthington Enzyme Manual 904: 647: 608: 10: 3687: 2593:10.1038/s41598-020-67415-6 2536:10.1038/s41598-020-57671-x 1251:10.1038/s41598-017-08097-5 828:is a modified form of the 661:genetically modified fruit 583:nuclear magnetic resonance 299:. PPO catalyses the rapid 205: 199: 3529: 3521:Michaelis–Menten kinetics 3493: 3462: 3431: 3380: 3327:Ecdysone 20-monooxygenase 3286: 3278:Stearoyl-CoA desaturase-1 3258: 3250:Dopamine beta-hydroxylase 3236: 3218:Phenylalanine hydroxylase 3199: 3167: 3131: 3031: 2958: 2881:10.1007/s00425-015-2261-0 2369:10.1080/10408399409527653 2139:10.1080/10408398109527312 1730:10.1017/S0953756296003206 1633:10.1007/s00442-002-0885-9 1194:10.1107/S1399004714013777 546: 280:molecules containing two 180: 168: 156: 151: 147: 128: 116: 104: 92: 80: 68: 56: 51: 43: 31: 26: 21: 3413:Diffusion-limited enzyme 2430:10.1021/acs.jafc.7b03210 2221:Functional Plant Biology 1465:Archives of Biochemistry 748:), two different genes ( 2831:10.1073/pnas.1523575113 2010:The Biochemical Journal 1901:Analytical Biochemistry 1865:Analytical Biochemistry 702:purees preserved their 517:potassium metabisulfite 261:molecules in which the 3312:Squalene monooxygenase 3228:Tryptophan hydroxylase 2977:HIF prolyl-hydroxylase 2067:10.1002/slct.201702144 1913:10.1006/abio.1995.1526 1536:10.1002/cbic.202000647 1300:10.1002/anie.201901332 482:Grape reaction product 380:Structure and function 292:-diphenols to produce 3506:Eadie–Hofstee diagram 3439:Allosteric regulation 3108:Methane monooxygenase 3081:Nitric oxide synthase 2682:10.1007/s007750050289 1664:Physiologia Plantarum 1492:Physiologia Plantarum 870:Coreopsis grandiflora 734:, flowers and roots. 571:nucleophilic reagents 476:desiccation tolerance 450:is controlled by the 3516:Lineweaver–Burk plot 3223:Tyrosine hydroxylase 2947:steroid hydroxylases 2185:10.1104/pp.109.2.525 1718:Mycological Research 935:"Polyphenol Oxidase" 531:compounds including 525:sodium metabisulfite 411:In plants, PPO is a 3173:iron–sulfur protein 2822:2016PNAS..113E1806M 2740:1996SciAm.275e..60B 2728:Scientific American 2638:1984PChem..23..501T 2585:2020NatSR..1010813P 2528:2020NatSR..10.1659P 1796:10.1104/pp.38.5.508 1703:10.1021/jf00006a007 1691:J. Agric. Food Chem 1625:2002Oecol.131..227T 1243:2017NatSR...7.8860K 1137:2014PChem..99...14M 1090:2006PChem..67.2318M 862:Aureusidin synthase 830:complement response 384:PPO is listed as a 3475:Enzyme superfamily 3408:Enzyme promiscuity 2972:Prolyl hydroxylase 2573:Scientific Reports 2516:Scientific Reports 1778:Palmer JK (1963). 1231:Scientific Reports 589:Enzymatic browning 406:multiple functions 265:contains a single 213:Polyphenol oxidase 47:Polyphenol oxidase 3631: 3630: 3340: 3339: 3006:Lysyl hydroxylase 2424:(37): 8203–8212. 2233:10.1071/pp9930099 2022:10.1042/bj2880063 1842:10.1021/jf0712301 1346:(18): 3860–3872. 1294:(22): 7475–7479. 1288:Angewandte Chemie 1188:(Pt 9): 2301–15. 879:isoliquiritigenin 877:activity against 760:prefer different 539:, in addition to 533:sodium dithionite 529:sulfur-containing 448:catechol oxidases 436:catechol oxidases 343:Agaricus bisporus 196: 195: 192: 191: 111:metabolic pathway 3678: 3648: 3647: 3639: 3511:Hanes–Woolf plot 3454:Enzyme activator 3449:Enzyme inhibitor 3423:Enzyme catalysis 3367: 3360: 3353: 3344: 3343: 2932: 2925: 2918: 2909: 2908: 2903: 2902: 2892: 2860: 2854: 2853: 2843: 2833: 2816:(13): E1806-15. 2801: 2795: 2794: 2766: 2760: 2759: 2725: 2716: 2710: 2709: 2664: 2658: 2657: 2621: 2615: 2614: 2604: 2564: 2558: 2557: 2547: 2507: 2501: 2500: 2490: 2458: 2452: 2451: 2441: 2409: 2403: 2402: 2400: 2398: 2387: 2381: 2380: 2352: 2346: 2345: 2325: 2316: 2315: 2305: 2277: 2271: 2270: 2268: 2266: 2260: 2251: 2245: 2244: 2216: 2207: 2206: 2196: 2173:Plant Physiology 2164: 2151: 2150: 2122: 2103: 2102: 2082: 2071: 2070: 2061:(32): 10435–41. 2050: 2044: 2043: 2033: 2001: 1995: 1994: 1966: 1960: 1959: 1931: 1925: 1924: 1895: 1889: 1888: 1860: 1854: 1853: 1824: 1818: 1817: 1807: 1784:Plant Physiology 1775: 1769: 1768: 1740: 1734: 1733: 1713: 1707: 1706: 1697:(6): 1043–1046. 1686: 1680: 1679: 1659: 1653: 1652: 1608: 1602: 1601: 1573: 1567: 1566: 1556: 1538: 1529:(7): 1161–1175. 1514: 1508: 1507: 1487: 1481: 1480: 1460: 1454: 1453: 1443: 1419: 1413: 1412: 1402: 1378: 1372: 1371: 1335: 1322: 1321: 1311: 1279: 1273: 1272: 1262: 1222: 1216: 1215: 1205: 1173: 1167: 1166: 1156: 1116: 1110: 1109: 1073: 1067: 1066: 1032: 1026: 1025: 997: 991: 990: 953: 947: 946: 944: 942: 931: 911:catechol oxidase 899:catechol oxidase 888: 826:Prophenoloxidase 793:catechol oxidase 659:, an example of 615:Catechol oxidase 530: 363:Persea americana 202:Catechol oxidase 149: 148: 22:Catechol oxidase 19: 18: 3686: 3685: 3681: 3680: 3679: 3677: 3676: 3675: 3656: 3655: 3654: 3642: 3634: 3632: 3627: 3539:Oxidoreductases 3525: 3501:Enzyme kinetics 3489: 3485:List of enzymes 3458: 3427: 3398:Catalytic triad 3376: 3371: 3341: 3336: 3290:- miscellaneous 3282: 3254: 3232: 3195: 3163: 3127: 3118:14α-demethylase 3027: 2954: 2939:Oxidoreductases 2936: 2906: 2861: 2857: 2802: 2798: 2767: 2763: 2723: 2717: 2713: 2665: 2661: 2622: 2618: 2565: 2561: 2508: 2504: 2459: 2455: 2410: 2406: 2396: 2394: 2389: 2388: 2384: 2353: 2349: 2326: 2319: 2278: 2274: 2264: 2262: 2258: 2252: 2248: 2217: 2210: 2165: 2154: 2123: 2106: 2083: 2074: 2055:ChemistrySelect 2051: 2047: 2002: 1998: 1967: 1963: 1932: 1928: 1896: 1892: 1861: 1857: 1836:(24): 9739–49. 1825: 1821: 1776: 1772: 1751:(5–6): 308–14. 1741: 1737: 1714: 1710: 1687: 1683: 1660: 1656: 1609: 1605: 1574: 1570: 1515: 1511: 1488: 1484: 1461: 1457: 1420: 1416: 1379: 1375: 1336: 1325: 1280: 1276: 1223: 1219: 1174: 1170: 1117: 1113: 1084:(21): 2318–31. 1074: 1070: 1033: 1029: 998: 994: 954: 950: 940: 938: 933: 932: 928: 924: 907: 886: 823: 821:Related enzymes 786: 784:In black poplar 740: 712: 677: 650: 623: 611: 591: 549: 528: 501: 471: 454:around the two 382: 353:Malus domestica 269:substituent to 238:PPO may accept 210: 204: 17: 12: 11: 5: 3684: 3674: 3673: 3671:Copper enzymes 3668: 3653: 3652: 3629: 3628: 3626: 3625: 3612: 3599: 3586: 3573: 3560: 3547: 3533: 3531: 3527: 3526: 3524: 3523: 3518: 3513: 3508: 3503: 3497: 3495: 3491: 3490: 3488: 3487: 3482: 3477: 3472: 3466: 3464: 3463:Classification 3460: 3459: 3457: 3456: 3451: 3446: 3441: 3435: 3433: 3429: 3428: 3426: 3425: 3420: 3415: 3410: 3405: 3400: 3395: 3390: 3384: 3382: 3378: 3377: 3370: 3369: 3362: 3355: 3347: 3338: 3337: 3335: 3334: 3329: 3324: 3319: 3314: 3309: 3302:Heme oxygenase 3299: 3297:Cyclooxygenase 3293: 3291: 3284: 3283: 3281: 3280: 3275: 3269: 3267: 3256: 3255: 3253: 3252: 3246: 3244: 3234: 3233: 3231: 3230: 3225: 3220: 3214: 3212: 3197: 3196: 3194: 3193: 3188: 3183: 3177: 3175: 3165: 3164: 3162: 3161: 3156: 3151: 3145: 3143: 3129: 3128: 3126: 3125: 3120: 3115: 3110: 3105: 3100: 3099: 3098: 3093: 3088: 3078: 3077: 3076: 3071: 3066: 3061: 3056: 3045: 3043: 3029: 3028: 3026: 3025: 3024: 3023: 3018: 3008: 3003: 3002: 3001: 2996: 2995: 2994: 2989: 2984: 2968: 2966: 2964:2-oxoglutarate 2956: 2955: 2935: 2934: 2927: 2920: 2912: 2905: 2904: 2855: 2796: 2761: 2711: 2659: 2632:(3): 501–505. 2626:Phytochemistry 2616: 2559: 2502: 2467:Food Chemistry 2453: 2404: 2382: 2347: 2330:Food Chemistry 2317: 2288:(4): 527–534. 2272: 2246: 2208: 2152: 2104: 2093:(3): 909–913. 2087:Food Chemistry 2072: 2045: 2016:(Pt 1): 63–7. 1996: 1961: 1926: 1890: 1855: 1819: 1770: 1735: 1724:(5): 552–556. 1708: 1681: 1670:(4): 381–385. 1654: 1619:(2): 227–235. 1603: 1568: 1509: 1498:(1): 106–112. 1482: 1455: 1414: 1373: 1323: 1274: 1217: 1168: 1125:Phytochemistry 1111: 1078:Phytochemistry 1068: 1049:(6): 677–683. 1027: 992: 965:(3): 274–308. 948: 925: 923: 920: 919: 918: 913: 906: 903: 822: 819: 785: 782: 739: 736: 726:products with 711: 708: 676: 673: 669:gene silencing 649: 646: 622: 619: 610: 607: 590: 587: 548: 545: 521:dithiothreitol 500: 497: 470: 467: 442:preference of 381: 378: 373:Lactuca sativa 313:fruit browning 301:polymerization 229:fruit browning 206:Main article: 200:Main article: 194: 193: 190: 189: 184: 178: 177: 172: 166: 165: 160: 154: 153: 145: 144: 133: 126: 125: 120: 114: 113: 108: 102: 101: 96: 90: 89: 84: 78: 77: 72: 66: 65: 60: 54: 53: 49: 48: 45: 41: 40: 35: 29: 28: 24: 23: 15: 9: 6: 4: 3: 2: 3683: 3672: 3669: 3667: 3664: 3663: 3661: 3651: 3646: 3641: 3640: 3637: 3623: 3619: 3618: 3613: 3610: 3606: 3605: 3600: 3597: 3593: 3592: 3587: 3584: 3580: 3579: 3574: 3571: 3567: 3566: 3561: 3558: 3554: 3553: 3548: 3545: 3541: 3540: 3535: 3534: 3532: 3528: 3522: 3519: 3517: 3514: 3512: 3509: 3507: 3504: 3502: 3499: 3498: 3496: 3492: 3486: 3483: 3481: 3480:Enzyme family 3478: 3476: 3473: 3471: 3468: 3467: 3465: 3461: 3455: 3452: 3450: 3447: 3445: 3444:Cooperativity 3442: 3440: 3437: 3436: 3434: 3430: 3424: 3421: 3419: 3416: 3414: 3411: 3409: 3406: 3404: 3403:Oxyanion hole 3401: 3399: 3396: 3394: 3391: 3389: 3386: 3385: 3383: 3379: 3375: 3368: 3363: 3361: 3356: 3354: 3349: 3348: 3345: 3333: 3330: 3328: 3325: 3323: 3320: 3318: 3315: 3313: 3310: 3307: 3303: 3300: 3298: 3295: 3294: 3292: 3289: 3285: 3279: 3276: 3274: 3271: 3270: 3268: 3265: 3261: 3257: 3251: 3248: 3247: 3245: 3243: 3239: 3235: 3229: 3226: 3224: 3221: 3219: 3216: 3215: 3213: 3210: 3209:BH4 dependent 3206: 3202: 3198: 3192: 3189: 3187: 3184: 3182: 3179: 3178: 3176: 3174: 3170: 3166: 3160: 3157: 3155: 3152: 3150: 3147: 3146: 3144: 3142: 3138: 3134: 3130: 3124: 3121: 3119: 3116: 3114: 3111: 3109: 3106: 3104: 3101: 3097: 3094: 3092: 3089: 3087: 3084: 3083: 3082: 3079: 3075: 3072: 3070: 3067: 3065: 3062: 3060: 3057: 3055: 3052: 3051: 3050: 3047: 3046: 3044: 3042: 3038: 3034: 3030: 3022: 3019: 3017: 3014: 3013: 3012: 3009: 3007: 3004: 3000: 2997: 2993: 2990: 2988: 2985: 2983: 2980: 2979: 2978: 2975: 2974: 2973: 2970: 2969: 2967: 2965: 2961: 2957: 2952: 2948: 2944: 2940: 2933: 2928: 2926: 2921: 2919: 2914: 2913: 2910: 2900: 2896: 2891: 2886: 2882: 2878: 2875:(3): 519–37. 2874: 2870: 2866: 2859: 2851: 2847: 2842: 2837: 2832: 2827: 2823: 2819: 2815: 2811: 2807: 2800: 2792: 2788: 2784: 2780: 2776: 2772: 2765: 2757: 2753: 2749: 2745: 2741: 2737: 2733: 2729: 2722: 2715: 2707: 2703: 2699: 2695: 2691: 2687: 2683: 2679: 2675: 2671: 2663: 2655: 2651: 2647: 2643: 2639: 2635: 2631: 2627: 2620: 2612: 2608: 2603: 2598: 2594: 2590: 2586: 2582: 2578: 2574: 2570: 2563: 2555: 2551: 2546: 2541: 2537: 2533: 2529: 2525: 2521: 2517: 2513: 2506: 2498: 2494: 2489: 2484: 2480: 2476: 2472: 2468: 2464: 2457: 2449: 2445: 2440: 2435: 2431: 2427: 2423: 2419: 2415: 2408: 2392: 2386: 2378: 2374: 2370: 2366: 2363:(2): 109–57. 2362: 2358: 2351: 2343: 2339: 2335: 2331: 2324: 2322: 2313: 2309: 2304: 2299: 2295: 2291: 2287: 2283: 2276: 2257: 2250: 2242: 2238: 2234: 2230: 2227:(1): 99–107. 2226: 2222: 2215: 2213: 2204: 2200: 2195: 2190: 2186: 2182: 2179:(2): 525–31. 2178: 2174: 2170: 2163: 2161: 2159: 2157: 2148: 2144: 2140: 2136: 2133:(1): 49–127. 2132: 2128: 2121: 2119: 2117: 2115: 2113: 2111: 2109: 2100: 2096: 2092: 2088: 2081: 2079: 2077: 2068: 2064: 2060: 2056: 2049: 2041: 2037: 2032: 2027: 2023: 2019: 2015: 2011: 2007: 2000: 1992: 1988: 1984: 1980: 1977:(2): 188–94. 1976: 1972: 1965: 1957: 1953: 1949: 1945: 1941: 1937: 1930: 1922: 1918: 1914: 1910: 1907:(1): 237–46. 1906: 1902: 1894: 1886: 1882: 1878: 1874: 1870: 1866: 1859: 1851: 1847: 1843: 1839: 1835: 1831: 1823: 1815: 1811: 1806: 1801: 1797: 1793: 1789: 1785: 1781: 1774: 1766: 1762: 1758: 1754: 1750: 1746: 1739: 1731: 1727: 1723: 1719: 1712: 1704: 1700: 1696: 1692: 1685: 1677: 1673: 1669: 1665: 1658: 1650: 1646: 1642: 1638: 1634: 1630: 1626: 1622: 1618: 1614: 1607: 1599: 1595: 1591: 1587: 1583: 1579: 1572: 1564: 1560: 1555: 1550: 1546: 1542: 1537: 1532: 1528: 1524: 1520: 1513: 1505: 1501: 1497: 1493: 1486: 1478: 1474: 1470: 1466: 1459: 1451: 1447: 1442: 1437: 1434:(6): 1593–9. 1433: 1429: 1425: 1418: 1410: 1406: 1401: 1396: 1393:: PC1489–91. 1392: 1388: 1384: 1377: 1369: 1365: 1361: 1357: 1353: 1349: 1345: 1341: 1334: 1332: 1330: 1328: 1319: 1315: 1310: 1305: 1301: 1297: 1293: 1289: 1285: 1278: 1270: 1266: 1261: 1256: 1252: 1248: 1244: 1240: 1236: 1232: 1228: 1221: 1213: 1209: 1204: 1199: 1195: 1191: 1187: 1183: 1179: 1172: 1164: 1160: 1155: 1150: 1146: 1142: 1138: 1134: 1130: 1126: 1122: 1115: 1107: 1103: 1099: 1095: 1091: 1087: 1083: 1079: 1072: 1064: 1060: 1056: 1052: 1048: 1044: 1043: 1038: 1031: 1023: 1019: 1015: 1011: 1007: 1003: 996: 988: 984: 980: 976: 972: 968: 964: 961: 960: 952: 936: 930: 926: 917: 914: 912: 909: 908: 902: 900: 896: 892: 884: 880: 876: 872: 871: 865: 863: 859: 857: 853: 849: 845: 843: 839: 835: 831: 827: 818: 816: 812: 808: 804: 800: 799: 798:Populus nigra 794: 791: 781: 779: 775: 771: 767: 763: 759: 755: 751: 747: 746: 745:Juglans regia 735: 733: 729: 725: 721: 720:cortex tissue 717: 707: 705: 701: 697: 693: 692:Ascorbic acid 689: 685: 681: 672: 670: 666: 662: 658: 657:Arctic apples 654: 645: 643: 639: 636: 632: 628: 618: 616: 606: 604: 600: 595: 586: 584: 580: 576: 572: 568: 566: 561: 559: 554: 544: 543:(vitamin C). 542: 541:ascorbic acid 538: 534: 526: 522: 518: 514: 510: 506: 496: 494: 489: 487: 486:caftaric acid 483: 479: 477: 466: 464: 460: 457: 453: 449: 445: 441: 437: 433: 429: 425: 420: 418: 414: 409: 407: 403: 399: 395: 391: 387: 377: 375: 374: 369: 365: 364: 359: 355: 354: 349: 345: 344: 339: 334: 332: 328: 324: 321: 316: 314: 311:) that cause 310: 306: 302: 298: 296: 291: 287: 283: 279: 275: 273: 268: 264: 260: 256: 255:hydroxylation 252: 248: 246: 241: 236: 234: 230: 226: 222: 218: 214: 209: 203: 198: 188: 185: 183: 179: 176: 173: 171: 167: 164: 161: 159: 155: 150: 146: 143: 140: 137: 134: 131: 127: 124: 121: 119: 115: 112: 109: 107: 103: 100: 97: 95: 91: 88: 87:NiceZyme view 85: 83: 79: 76: 73: 71: 67: 64: 61: 59: 55: 50: 46: 42: 39: 36: 34: 30: 25: 20: 3617:Translocases 3614: 3601: 3588: 3575: 3562: 3552:Transferases 3549: 3536: 3393:Binding site 3141:flavoprotein 2945:, including 2943:dioxygenases 2872: 2868: 2858: 2813: 2809: 2799: 2777:(8): 392–7. 2774: 2770: 2764: 2734:(5): 60–66. 2731: 2727: 2714: 2676:(1): 56–63. 2673: 2669: 2662: 2629: 2625: 2619: 2579:(1): 10813. 2576: 2572: 2562: 2519: 2515: 2505: 2470: 2466: 2456: 2421: 2417: 2407: 2395:. Retrieved 2385: 2360: 2356: 2350: 2336:(1): 23–30. 2333: 2329: 2303:10261/114868 2285: 2281: 2275: 2263:. Retrieved 2249: 2224: 2220: 2176: 2172: 2130: 2126: 2090: 2086: 2058: 2054: 2048: 2013: 2009: 1999: 1974: 1970: 1964: 1942:(2): 155–9. 1939: 1935: 1929: 1904: 1900: 1893: 1871:(1): 23–32. 1868: 1864: 1858: 1833: 1829: 1822: 1787: 1783: 1773: 1748: 1744: 1738: 1721: 1717: 1711: 1694: 1690: 1684: 1667: 1663: 1657: 1616: 1612: 1606: 1584:(1): 65–68. 1581: 1578:FEBS Letters 1577: 1571: 1526: 1522: 1512: 1495: 1491: 1485: 1471:(1): 29–44. 1468: 1464: 1458: 1431: 1427: 1417: 1390: 1386: 1376: 1343: 1339: 1291: 1287: 1277: 1234: 1230: 1220: 1185: 1181: 1171: 1128: 1124: 1114: 1081: 1077: 1071: 1046: 1040: 1030: 1005: 1001: 995: 962: 957: 951: 941:13 September 939:. Retrieved 929: 868: 866: 860: 846: 824: 803:caffeic acid 796: 787: 773: 765: 753: 749: 743: 741: 713: 678: 655: 651: 624: 612: 596: 592: 564: 557: 550: 502: 490: 480: 472: 431: 424:nomenclature 421: 410: 383: 371: 361: 351: 341: 335: 326: 317: 304: 294: 289: 271: 263:benzene ring 250: 244: 237: 227:involved in 220: 216: 212: 211: 197: 75:BRENDA entry 3388:Active site 2522:(1): 1659. 1523:ChemBioChem 1237:(1): 8860. 1008:(1): 1–11. 885:substrates 873:shows weak 813:at injured 770:monophenols 742:In walnut ( 665:trademarked 627:monophenols 463:active site 452:amino acids 444:tyrosinases 428:tyrosinases 404:, creating 331:tyrosinases 309:polyphenols 240:monophenols 63:IntEnz view 27:Identifiers 3666:EC 1.14.18 3660:Categories 3591:Isomerases 3565:Hydrolases 3432:Regulation 3273:Tyrosinase 3240:: reduced 3203:: reduced 3171:: reduced 3135:: reduced 2473:: 100053. 2397:5 November 922:References 916:Tyrosinase 883:tyrosinase 875:tyrosinase 852:tyrosinase 848:Hemocyanin 772:, whereas 762:substrates 758:isoenzymes 728:amino acid 684:maturation 675:In apricot 621:In avocado 603:harvesting 499:Inhibitors 392:, trimer, 320:amino acid 274:-diphenols 259:monophenol 247:-diphenols 208:Tyrosinase 132:structures 99:KEGG entry 44:Alt. names 3470:EC number 3242:ascorbate 3205:pteridine 2690:1432-1327 2654:0031-9422 2312:1097-0010 2241:1445-4416 1613:Oecologia 1545:1439-4227 1368:205692816 1131:: 14–25. 987:218831573 801:converts 790:monomeric 778:diphenols 752:PPO1 and 738:In walnut 710:In potato 635:polymeric 631:o-quinone 567:-quinones 560:-quinones 513:inhibitor 509:Tropolone 493:parasites 440:substrate 413:plastidic 402:dodecamer 386:morpheein 338:mushrooms 297:-quinones 286:oxidation 52:Databases 3494:Kinetics 3418:Cofactor 3381:Activity 2899:25697287 2850:26976571 2791:10916160 2706:29871864 2698:10499103 2611:32616720 2554:32015350 2497:31650127 2448:28812349 1885:12479831 1850:17958393 1814:16655824 1765:16814556 1649:25912204 1641:28547690 1598:84630585 1563:33108057 1477:18135760 1409:14284774 1360:27172067 1318:30825403 1269:28821733 1212:25195745 1163:24461779 1106:16973188 1063:10607672 979:32441137 905:See also 895:tyramine 891:tyrosine 811:melanine 732:petioles 696:protease 648:In apple 642:melanins 638:pigments 609:In mango 599:vacuoles 537:cysteine 505:Tentoxin 394:tetramer 358:avocados 323:tyrosine 282:hydroxyl 267:hydroxyl 233:tetramer 187:proteins 175:articles 163:articles 136:RCSB PDB 38:1.10.3.2 3650:Biology 3604:Ligases 3374:Enzymes 3288:1.14.99 3266:: other 3260:1.14.18 3238:1.14.17 3201:1.14.16 3169:1.14.15 3133:1.14.14 3033:1.14.13 2960:1.14.11 2890:4540782 2841:4822611 2818:Bibcode 2756:8875808 2736:Bibcode 2634:Bibcode 2602:7331820 2581:Bibcode 2545:6997208 2524:Bibcode 2488:6804514 2439:5609118 2377:8011143 2265:8 March 2203:7480344 2147:6794984 2040:1445282 2031:1132080 1991:5871805 1956:9178091 1921:8678307 1621:Bibcode 1554:8049008 1450:4975157 1309:6563526 1260:5562730 1239:Bibcode 1203:4157443 1154:3969299 1133:Bibcode 1086:Bibcode 1022:7873577 834:insects 807:quinone 724:quinone 700:apricot 680:Apricot 579:tritium 461:in the 438:). The 422:Enzyme 398:octamer 390:monomer 368:lettuce 366:), and 242:and/or 231:, is a 219:; also 123:profile 106:MetaCyc 3636:Portal 3578:Lyases 3137:flavin 3016:ALKBH1 2897:  2887:  2869:Planta 2848:  2838:  2789:  2754:  2704:  2696:  2688:  2652:  2609:  2599:  2552:  2542:  2495:  2485:  2446:  2436:  2375:  2310:  2239:  2201:  2194:157616 2191:  2145:  2038:  2028:  1989:  1954:  1919:  1883:  1848:  1812:  1805:549964 1802:  1763:  1647:  1639:  1596:  1561:  1551:  1543:  1475:  1448:  1407:  1366:  1358:  1316:  1306:  1267:  1257:  1210:  1200:  1161:  1151:  1104:  1061:  1020:  985:  977:  856:steric 688:latent 686:. 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Index

EC no.
1.10.3.2
IntEnz
IntEnz view
BRENDA
BRENDA entry
ExPASy
NiceZyme view
KEGG
KEGG entry
MetaCyc
metabolic pathway
PRIAM
profile
PDB
RCSB PDB
PDBe
PDBsum
PMC
articles
PubMed
articles
NCBI
proteins
Catechol oxidase
Tyrosinase
enzyme
fruit browning
tetramer
monophenols

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