Growth and Genetic Analysis of Pejibaye
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GROWTH AND GENETIC ANALYSIS OF PEJIBAYE {Bactris gasipaes KUNTH, PALMAE) IN HAWAII. 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 AUGUST 1995 By Charles R. Clement Dissertation Committee: Richard M. Manshardt, Chairperson Duane Bartholomew lames L. Brewbaker Philip Ito Francis Zee 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 VJJMJlJJiChairperson ( 9X^ IM © Copyright 1995 by Charles R. Clement 111 Acknowledgments I thank my wife, Rosa de Nazare Silva Clement, for her unfailing love, patience, understanding, support and assistance during the years of study and research in Hawai’i. I thank the Brazilian National Research Institute for Amazonia (INPA) and its directors, Drs. Herbert O. R. Schubart, Eneas Salati, Jose Seixas Louren^o, e Ozorio J. Menezes Fonseca, and the Brazilian National Council for Scientific and Technological Development (CNPq) for permission to study in Hawai’i and for the scholarship that made this physically possible. I thank the United States Department of Agriculture, Special Research Grants in Tropical Agriculture Program, for having funded the research project "Introduction and Evaluation of Pejibaye for Heart of Palm Production in Hawai’i," that includes the work done in this dissertation. Also, the Governor's Agricultural Coordinating Committee's Ad Hoc Committee on Agricultural Product Expansion (CAPE), for having funded research on weed control in pejibaye plantations, and its principal investigator. Dr. Joseph DeFrank, which made field management much easier. I thank all members of my dissertation committee for their unfailing patience in answering questions and offering suggestions to improve the quality of this research project and this dissertation. I thank Dr, Kaoru Yuyama, INPA, and M.S. Wanders B. Chavez Flores, ex-INPA, for the seed of the Benjamin Constant population of pejibaye. I thank Mr. John Mood, owner of the farm in Ninole, Hawai’i, where the principal progeny x density trial was planted, for use of his land and his frequent assistance with IV field management. Also, Mr. Dennis Ida, Waiakea Experiment Station manager, and Mr. Richard Nakano, Poamoho Experiment Station manager, Mr. Craig Okazaki, Magoon Nursery Facility manager, and their staffs for valuable nursery and field management assistance without which all phases of the project would have been much more difficult. I thank my colleagues in the Fruit Genetics Laboratory, especially Dr. Mallikaijuna Aradhya, for his patience in teaching me the art and science of starch gel electrophoretic analysis and discussing its implications, and Ms. Maimunah Morshidi, for her assistance with the electrophoretic analysis. Also, Dr. Vincent Lebot, Dr. Suresh Sondur, M.S. Kevin Crosby, M.S. Lius Suwenza, Mr. Leren Zhang, Mr. Xiaohu Wang, and Ms. Wendy Jones for their unfailing humor and comradeship in the ups and downs of university life, bureaucracy, class, laboratory and field work. Finally, but by no means least important, I thank the native peoples of tropical America who domesticated the pejibaye in Amazonia and elsewhere, without whom it would never have occurred to me to study this species nor introduce it to Hawai’i as a potential new crop. Abstract Pejibaye or peach palm {Bactris gasipaes, Kunth) was introduced into Hawai’i to supply the gourmet market with fresh heart of palm. New crop introduction requires evaluation of crop adaptation to its new environment and planning for future development, including genetic improvement. Leaf number of open-pollinated Benjamin Constant (Putumayo landrace) progenies was lower at harvest (6-8) than elsewhere (8- 10), and offshoot number dropped dramatically from first harvest (6.5) to second harvest (2). Allometric equations for estimating whole plant leaf area and biomass were developed, using height and leaf number predictors. No significant plant population (density) effects on individual plant dimensions or growth were found over the range of 3333 to 6666 plants/ha. Relative growth rate (RGR) and unit leaf rate (E ^ between nursery and first harvest were highly correlated (r = -0.99 and -0.95, respectively) with earliness (days to harvest). The early progenies partitioned photoassimilates differently; two had high E^, while one had moderate E^ and partitioned preferentially to leaf area, resulting in a higher leaf area ratio. Heart of palm yields were close to 900 kg/ha after 12 months of harvest and 1400 kg/ha after 18 months, both comparable to tropical American yields. When edible stem and leaf were added to the yields, these increased to 2.8 and 4.5 t/ha of marketable product, respectively. Quantitative genetic analysis of growth parameters suggested high levels of inbreeding in the germplasm studied, since the narrow-sense heritabilities were double those observed in other perennials. Additive genetic variances for RGR and earliness suggested the potential for significant response to selection, but phenotypic variation varied depending on the interval over which RGR VI was estimated. The lowest estimate of RGR (over an entire development phase) provided the smallest response to selection but is similar to the response observed in other crops. Allozyme heterozygosity was remarkably low, ranging from 0.038 to 0.099, with a mean of 0.074, on par with inbred crops, rather than outbreeders. There was a lack of correlation between allozyme heterozygosity and growth parameters and morphological traits. Vll Table of Contents Acknowledgments............................................................................................ iv Abstract.............................................................................................................. vi List of Tables..................................................................................................... xi List of Figures.................................................................................................... xiii List of Abbreviations...................................................................................... xiv Preface............................................................................................................... xv Chapter 1: Introduction.................................................................................... 1 Chapter 2: Literature Review......................................................................... 7 2.1. Cytotaxonomic Background............................................................ 7 2.2. Morphology of the Mature Pejibaye............................................... 11 2.3. Phases of Growth in Palms................................................................ 13 2.4. Germination of Pejibaye.................................................................... 15 2.5. Allometric Relationships in Plants................................................... 17 2.5.1. Allometric Relationships in African Oil Palm..................... 19 2.5.2. Allometric Relationships in Pejibaye.................................... 21 2.6. Growth Analysis..................................................................................26 2.6.1. Growth Analysis in African Oil Palm................................... 30 2.6.2. Growth Analysis in Pejibaye.................................................. 31 2.7. Genetic Analysis..................................................................................33 2.7.1. Genetic Analysis in Economically Important Palm s 36 2.8. Plant Isoenzymes.................................................................................40 2.8.1. Isoenzymes in Palms............................................................... 40 2.8.2. Correlations of Isoenzymes with Other Traits....................... 43 Chapter 3; Material and Methods.................................................................. 44 3.1. Germplasm......................................................................................... 44 3.2. Germination...................................................................................... 46 3.3. Nursery Conditions.......................................................................... 47 3.4. Localities..............................................................................................48 3.4.1. Ninole.........................................................................................48 3.4.2. Waiakea......................................................................................49 3.4.3. Poamoho.................................................................................... 50 3.5. Plantation Densities.......................................................................... 51 3.6. Experimental Designs........................................................................ 52 3.7. Field Preparation and Planting.......................................................... 55 3.8. Field Management............................................................................. 56 3.9. Fertilization........................................................................................ 57 3.10. Morphological Measurements........................................................ 58 3.11. Plant Morphology - Analysis.......................................................... 63 3.12. Leaf Area and Biomass Allometry................................................