Tree and Forestry Science and Biotechnology ©2007 Global Science Books Papaya (Carica papaya L.) Biology and Biotechnology Jaime A. Teixeira da Silva1* • Zinia Rashid1 • Duong Tan Nhut2 • Dharini Sivakumar3 • Abed Gera4 • Manoel Teixeira Souza Jr.5 • Paula F. Tennant6 1 Kagawa University, Faculty of Agriculture, Department of Horticulture, Ikenobe, 2393, Miki-cho, Kagawa, 761-0795, Japan 2 Plant Biotechnology Department, Dalat Institute of Biology, 116 Xo Viet Nghe Tinh, Dalat, Lamdong, Vietnam 3 University of Pretoria, Postharvest Technology Group, Department of Microbiology and Plant Pathology, Pretoria, 0002, South Africa 4 Department of Virology, Agricultural Research Organization, The Volcani Center, Bet Dagan 50250, Israel 5 Embrapa LABEX Europa, Plant Research International (PRI), Wageningen University & Research Centre (WUR), Wageningen, The Netherlands 6 Department of Life Sciences, University of the West Indies, Mona, Kingston 7, Jamaica Corresponding author: * [email protected] ABSTRACT Papaya (Carica papaya L.) is a popular and economically important fruit tree of tropical and subtropical countries. The fruit is consumed world-wide as fresh fruit and as a vegetable or used as processed products. This review focuses primarily on two aspects. Firstly, on advances in in vitro methods of propagation, including tissue culture and micropropagation, and secondly on how these advances have facilitated improvements in papaya genetic transformation. An account of the dietary and nutritional composition of papaya, how these vary with culture methods, and secondary metabolites, both beneficial and harmful, and those having medicinal applications, are dis- cussed. An overview of papaya post-harvest is provided, while ‘synseed’ technology and cryopreservation are also covered. This is the first comprehensive review on papaya that attempts to integrate so many aspects of this economically and culturally important fruit tree that should prove valuable for professionals involved in both research and commerce. _____________________________________________________________________________________________________________ Keywords: biolistic, papain, Papaya ringspot virus, postharvest management Abbreviations: ½MS, half-strength Murashige and Skoog (1962) medium; 2,4,5-T, 2,4,5-trichlorophenoxyacetic acid; 2,4-D, 2,4-dichlo- rophenoxyacetic acid; 2-iP, 6-(γ,γ-dimethylallylamino)-purine; AAC, 1-aminocyclopropane-1-carboxylic acid; ABA, abscisic acid; ACC, 1-aminocyclopropane-1-carboxylic acid; ACS 1 and ACS 2 1-aminocyclopropane-1-carboxylic acid synthase genes; AFLP, amplified fragment length polymorphism; AVG, aminoethoxyvinylglycine; AVG, aminoethoxyvinylglycine; BA, 6-benzyladenine; BAP, 6-benzyl- amino purine; BC, back-cross; CAPS, cleaved amplified polymorphic sequences; CaCl2, calcium chloride; CBF, C repeat binding factor; CoCl2, cobalt chloride; cp, coat protein gene; CPA, p-chlorophenoxyacetic acid; CP-ACO1 and CP-ACO2 1, aminocyclopropane-1- carboxylic acid oxidase genes; CPL, C. papaya lipase; CPNT, nontranslatable coat protein gene construct; CPT, translatable coat protein gene constructs; CSb, citrate synthase gene; CW, coconut water; DAF, DNA ampli-fication finger-printing; DmAMP1, Dahlia merckii defensin gene; EFE, ethylene forming enzyme; EST, expressed sequence tag; GA3, gibberellic acid; GFP, green fluorescent protein; GRAS, Generally Regarded As Safe; GUS, β-glucuronidase; IAA, indole-3-acetic acid; IBA, indole-3-butyric acid; KNO3, potassium nitrate; LED, light emitting diode; MA, modified atmosphere; Man, mannose; MSF, methanol sub-fraction; MSY, male-specific; Mt, million tones; NAA, α-naphthaleneacetic acid; NH, Nivun Haamir; nptII, neomycin phosphotransferase II; NSAIDs, non-steroidal anti- inflammatory drugs; PANV, Papaya apical necrosis virus; PBT, Papaya bunchy top; PCR, polymerase chain reation; PDB, Papaya dieback; PDNV, Papaya droopy necrosis virus; PM, Papaya mosaic; PMeV, Papaya meleira virus; PMI, phospho-mannose isomerase; PPT, phosphinothricin; PPT, phosphinothricin; PRSV HA 5-1, mild strain of Papaya ringspot virus; PRSV, Papaya ringspot potyvirus; PSDM, papaya sex determination marker; PYC, Papaya yellow crinkle; PLYV, Papaya lethal yellowing virus; RAF, randomly amplified DNA fingerprint; RAPD, random amplified polymorphic DNA; RP, viral replicase gene; SCAR, sequence characterized amplified region; STS, silver thiosulphate; TDZ, thidiazuron; TIBA, 2,3,5-triiodobenzoic acid); uidA, β-glucuronidase gene CONTENTS INTRODUCTION........................................................................................................................................................................................ 48 Geographic distribution and nomenclature.............................................................................................................................................. 48 BOTANY AND CULTIVATION.................................................................................................................................................................. 48 PESTS AND DISEASES ............................................................................................................................................................................. 49 GENETICS, CONVENTIONAL AND MOLECULAR BREEDING.......................................................................................................... 53 THE PLANT AND FRUIT: STRUCTURE, USES AND MEDICINAL PROPERTIES.............................................................................. 54 CHEMISTRY, PHYTOCHEMISTRY AND BIOCHEMISTRY .................................................................................................................. 55 POST-HARVEST MANAGEMENT OF PAPAYA ...................................................................................................................................... 56 Geographic distribution and nomenclature.............................................................................................................................................. 56 Harvesting, handling, heat treatment, storage and ripening..................................................................................................................... 57 Other post-harvest treatments.................................................................................................................................................................. 58 CONVENTIONAL PROPAGATION: SEEDS, SEEDLINGS AND SYNSEEDS ...................................................................................... 59 MICROPROPAGATION ............................................................................................................................................................................. 60 Shoot tip, axillary bud and single node culture........................................................................................................................................ 60 Organogenesis, anther and ovule culture, and regeneration from protoplasts.......................................................................................... 61 Callus induction and somatic embryogenesis .......................................................................................................................................... 61 Micropropagation and scaling-up ............................................................................................................................................................ 62 Rooting and acclimatization .................................................................................................................................................................... 62 GENETIC TRANSFORMATION................................................................................................................................................................ 63 Received: 1 November, 2006. Accepted: 1 November, 2007. Review Tree and ForestryPapaya Science biology and and Biotechnology biotechnology. 1(1) Teixeira, 47-73 da ©2007 Silva Globalet al. Science Books GENETICS AND GENOMICS ................................................................................................................................................................... 65 CONCLUDING REMARKS ....................................................................................................................................................................... 66 ACKNOWLEDGEMENTS ......................................................................................................................................................................... 66 REFERENCES............................................................................................................................................................................................. 66 _____________________________________________________________________________________________________________ INTRODUCTION comprise Cylicomorpha, Carica, Jacaratia, Jarilla, Horo- vitzia and Vasconcella), with Carica papaya the only spe- Geographic distribution and nomenclature cies within the genus Carica (Badillo 2002). The history of papaya appears to be first documented by Papaya, Carica papaya L., is one of the major fruit crops Oviedo, the Director of Mines in Hispaniola (Antilles) from cultivated in tropical and sub-tropical zones. Worldwide 1513 to 1525, where he describes how Alphonso de Val- over 6.8 million tonnes (Mt) of fruit were produced in 2004 verde took papaya