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Genetic Variability in Carica Papaya and Related Species a Dissertation Submitted to the Graduate Division of the University Of

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Genetic Variability in Carica Papaya and Related Species a Dissertation Submitted to the Graduate Division of the University Of GENETIC VARIABILITY IN CARICA PAPAYA AND RELATED SPECIES A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN HORTICULTURE MAY 1996 By Maimunah Morshidi Dissertation Committee: Richard Manshardt, Chairperson James Brewbaker Francis Zee John Stiles Clifford Morden We certify that we have read this dissertation and that, in our opinion, it is satisfactory in scope and quality as a dissertation for the degree of Doctor of Philosophy in Horticulture. DISSERTATION COMMITTEE n ® Copyright 1996 by Maimunah Morshidi iii ACKNOWLEDGEMENTS I am grateful to Dr. Victor M. Badillo of Universidad Central, Maracay, Venezuela, and Dr. Victor M. Patino of Colombia for their generous advice on where to locate and collect Carica species in Central and South America. I gratefully acknowledge the assistance, and hospitality of friends and colleagues throughout Latin America, particularly Dr. Juan Jose Castillo Mont (Universidad de San Carlos; Guatemala); Dr. Raul Mosqueda-Vasquez (Veracruz, Mexico); Dr. Daniel Teliz-Ortiz and Dr. Maria Pilar Rodriguez de Gonzalez (Collegio de Postgraduados, Texcoco, Mexico); Dr. Raul Castillo, Dr. Jaime Estrella Engelmann, and Dr. Cesar Tapia (INIAP, Ecuador); Dr. Luiz Lopez (CIAT, Colombia); Dr. Katsuo A. Okada (IPGRI, Colombia); Mr. John Link (Central Farm, Belize); Dr. John Dickson (FHIA, Honduras); Dr. Panfilo Tabora (E.A.R.T.H., Costa Rica); Dr. Jorge Leon (CATIE, Costa Rica); Dr. Maria Luisa Garcia and Dr. Freddy Leal (Venezuela); and Dr. Miguel Jordan (Universidad Catolica, Chile). I also thank the following individuals and institutions for help in field accomodations and germplasm exchange: Dr. Michael Balick (NYBG), Dr. Calaway Dodson (Missouri Botanical Garden), and Dr. Richard Adams (University of Texas, San Antonio), Dr. Richard Hamilton (University of Hawaii, Honolulu); CIAT (Colombia), FHIA (Honduras), Rio Bravo Station (Belize), OTS-Las Cruces, (Costa Rica), and CENARGEN, Brazil. This study was supported in part by the Specific Cooperative Grant Agreement 58-5320-0-001 between the United States Department of Agriculture and the University of Hawaii. iv ABSTRACT As part of an ex situ conservation of Carica germplasm in Hawaii, expeditions in 1992 and 1993 resulted in the collection and analysis of seeds of 131 accessions of Carica papaya from 9 countries in Central and South America (Mexico, Belize, Guatemala, Honduras, Costa Rica, Panama, Venezuela, Colombia, and Ecuador). Populations of wild small-fruited C. papaya (previously designated as C peltata), and hybrids between the wild and domesticated were sampled from their native range on the Caribbean coast of Central America (Mexico, Belize, Guatemala, Honduras). Domesticated C. papaya specimens were also collected from Central and South America. The three types of C. papaya were examined for variability in nine enzyme systems (ACO, IDH, MDH, PGM, PGI, SKD, TPI, UGP, and 6-PGD) and 13 loci. Domesticated, hybrid, and wild C. papaya shared a Nei’s genetic identity. I, of greater than 0.9, which is consistent with conspecific populations. The intrapopulational genetic variability results indicate that the greatest diversity exists in the Yucatan-San Ignacio-Peten-Rio Motagua area in Central America. In general, the wild C. papaya populations have greater isozyme variability than the domesticated populations. The hybrid populations have intermediate variability between the domesticated and the wild populations. The least diverse populations are found in South America where no wild populations are found. \ Carica cauliflora, C. goudotiana, C. crassipetala, C. sphaerocarpa, C. pubescens, C. microcarpa ssp. baccata, C. microcarpa ssp. heterophylla, C. microcarpa ssp. microcarpa, C. stipulata, C. pulchra, and the natural hybrid C. X heilbornii n.m. pentagona (babaco) were collected in Venezuela, Colombia, and Ecuador. Carica glandulosa, C. monoica, C. parviflora, and C. quercifolia were obtained from germplasm banks in Latin America or were already present in the USDA- NCGR collection. Isozyme analysis indicated that C. papaya is distantly related to the other Carica species included in this study and that none of the other Carica species could have been a wild progenitor of C. papaya in the recent past. Recommendations on future collecting expeditions and conservation of Carica genetic resources are suggested. VI TABLE OF CONTENTS ACKNOWLEDGEMENTS................................................................................................. iv ABSTRACT........................................................................................................................... v LIST OF TABLES............................................................................................................... xii LIST OF FIGURES.............................................................................................................. xiii CHAPTER 1. INTRODUCTION.................................................................................. 1 CHAPTER 2. LITERATURE REVIEW....................................................................... 6 Plant genetic resources conservation.................................................... 6 Importance of plant genetic resources conservation.................. 6 Role of plant population genetics in conserving genetic resources............................................................................... 9 Allozymes: tools for assessing genetic variability............................. 10 Measures of genetic variation using isozymes............................ 14 Genetic similarity and distance.................................................... 17 Phenetic analyses........................................................................... 19 Use of isozymic evidence in the elucidation of maize’s wild progenitor..................................................................... 20 Ethnobotany of Carica papaya.............................................................. 21 Taxonomy of Caricaceae.............................................................. 21 Historical biogeography of Carica papaya................................. 22 Papaya’s introduction to Hawaii................................................... 28 vii Uses of Carica papaya................................................................... 29 Ethnobotany of other lesser known Carica species............................. 31 Carica pubescens Lenne et Koch (chamburo or ‘mountain papaya’).............................................................. 31 Carica stipulata Badillo (siglalon)..................................................32 Carica goudotiana Solms (payuelo or papayuelo).................... 33 Carica cauliflora Jdicq. (tapaculo)................................................ 34 Carica monoica Desf. (col de monte or “mountain cabbage”)........................................................... 34 Carica microcarpa Jacq. (col de monte).................................... 35 Carica pulchra Badillo (col de monte)...........................................35 Carica X heilbornii Badillo n.m. pentagona (Heilbom) (babaco)................................................................................. 36 Carica X heilbornii Badillo n.m. chrysopetala (Heilbom) (toronchi)............................................................................... 37 Other wild Carica species............................................................. 38 Disease resistance and conservation of Carica species....................... 39 CHAPTER 3. GERMPLASM COLLECTION OF CARICA PAPAYA AND RELATED SPECIES.................................................................... 41 METHODS............................................................................................... 41 RESULTS................................................................................................. 42 V lll Types of Carica papaya collected................................................ 42 Wild Carica papaya............................................................. 42 Domesticated Carica papaya.............................................. 50 Hybrid Carica papaya.......................................................... 50 Carica papaya populations........................................................... 53 Carica species collected................................................................... 82 CHAPTER 4. GENETIC VARIABILITY IN CARICA PAPAYA................................ 105 MATERIALS AND METHODS............................................................ 106 Carica papaya germplasm.............................................................. 106 Enzyme electrophoresis...................................................................107 Data analysis.....................................................................................I l l RESULTS................................................................................................. 113 Electrophoretic phenotypes and genetic control...........................113 Cluster analysis.................................................................................118 Principal component analysis....................................................... 127 Intrapopulational genetic variability.............................................. 129 Geographic patterns of genetic diversity in Carica papaya populations....................................................136 DISCUSSION...........................................................................................
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