Investigations into the Morphological, Agronomic, and Nutritional Diversity within Breadfruit (Artocarpus, Moraceae) as a Resource for Food Security by Andrew Maxwell Phineas Jones A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSIPHY In THE COLLEGE OF GRADUATE STUDIES (Biology) THE UNIVERSITY OF BRITISH COLUMBIA (Okanagan) December, 2010 ©Andrew Maxwell Phineas Jones, 2010 Abstract Global food security is one of the most pressing issues facing humanity in the 21st century, with the number of undernourished people reaching an unprecedented high of over 1 billion. The problem is most acute in rural areas in tropical climates. Breadfruit (Artocarpus, Moraceae), a high-yielding tropical staple food crop, has been identified under the International Treaty on Plant Genetic Resources for Food and Agriculture for its potential to impact food security. The Breadfruit Institute at the National Tropical Botanical Garden houses the largest breadfruit collection in the world which represents a vast diversity of botanical and nutritional characters developed through millennia of traditional breeding. Breadfruit exhibits a great degree of morphological variability with cultivars that produce small 500g seeded fruit to those that produce large 3.5 kg seedless fruit. Variation is also expressed in the 57 characteristics evaluated in this study, with deep implications regarding the history of breadfruit domestication and the utilization of this crop to bolster food security. Evaluation of agronomic diversity has classified breadfruit into 10 seasonality groups, including non-seasonal, early, and late season cultivars. Informed cultivar selection based on these data will allow the fruiting season to be extended, and year round production will be possible. Further, 94 cultivars of breadfruit along with two related species, Artocarpus camansi and Artocarpus mariannensis, were evaluated for nutritional quality in fresh fruit and flour. Breadfruit is a good source of calcium, copper, iron, potassium, magnesium, has similar levels of protein as many other tropical staple crops, and many cultivars produce pro-vitamin A carotenoids. Individual cultivars have been identified that would provide 20%-25% of the recommended daily adult requirement for protein, approximately 23.5% calcium, 97.4% copper, 19.2% iron, 48.1% potassium, 115.8% magnesium, 33.6% manganese, 0.6% sodium, 53.5% phosphorous, and 21.0% zinc of the recommended daily intake ii of a female between 19-30 years old, and enough pro-vitamin A carotenoids to fulfill over 60% of the minimum daily vitamin A requirement of adults. Together these data show the immense diversity present within breadfruit germplasm and provide a foundation to utilize this variability to provide food security in the tropics. iii Preface The work contained in this thesis was conducted as part of a collaborative project between the Breadfruit Institute at the National Tropical Botanical Garden and the University of British Columbia Okanagan. Versions of Chapter 2-7 are in preparation, or have been submitted and are in various stages of the publication process. The submission information and my contribution to each Chapter are presented below. Chapter 2 has been accepted for publication: Jones, A.M.P., Ragone, D., Tavana, N.G., Bernotas, D., Murch, S.J. Beyond the Bounty: Breadfruit (Artocarpus altilis) for Food Security and Novel Foods in the 21st Century. Ethnobotany Research and Applications. In press. This Chapter is a comprehensive review of breadfruit with an emphasis on its nutritional content and potential for food security. I was responsible for writing the majority of the manuscript. The co-authors provided specific expertise with respect to ethnobotany, economics, breadfruit germplasm and ecology, and general editorial review of the manuscript. Chapter 3 has been submitted and is currently under review: Jones, A.M.P., Murch, S.J., Wiseman, J., and Ragone, D. Morphological diversity in breadfruit (Artocarpus, Moraceae): Insights into domestication, conservation, and cultivar identification. American Journal of Botany. The data for this Chapter were collected by Dr. Ragone at the Breadfruit Institute of the National Tropical Botanical Garden. I was responsible for developing the approaches to data analysis, the statistical analysis of the data, interpreting the results, composing the manuscript, and developing the multi-access cultivar identification key. iv . Chapter 4 has been accepted for publication: Jones, A.M.P, Murch, S.J., Ragone, D. Diversity of Breadfruit (Artocarpus altilis, Moraceae) Seasonality: A Resource for Year- Round Nutrition. Economic Botany: In Press The data for this Chapter were collected by Dr. Ragone at the National Tropical Botanical Garden over the period 1996-2005. I was responsible for designing the approaches to data analysis, conducting the statistical analysis, interpreting the results, and composing the manuscript. Chapter 5 has been submitted and is currently under review: Jones, A.M.P., Ragone, D., Lane, A., Murch, S.J. High Yield and Protein Cultivars of Breadfruit (Artocarpus altilis): Developing an Underutilized Crop for Food Security and Novel Foods. Journal of Food Composition and Analysis. I was responsible for the experimental design, sample collection, quantification of protein, statistical analysis and interpretation, and composing the final manuscript for publication. My co-authors provided valuable assistance in designing the experiments, sample collection, and revising the final manuscript. Chapter 6 has been submitted and is currently under review: Jones, A.M.P., Ragone, D., Aiona, K., Murch, S.J. Breadfruit (Artocarpus altilis): An underutilized crop with potential to address the 'hidden hunger' of micronutrient deficiency. Journal of Food Composition and Analysis. Soil analysis data were coordinated by Mr. Aiona at the National Tropical Botanical Garden. I was responsible for the experimental design, sample collection, analysis of fruit mineral content, statistical analysis and interpretation, and composing the final manuscript for v publication. My co-authors provided valuable assistance in designing the experiments, sample collection, and revising the final manuscript. Chapter 7 is currently in preparation and will be submitted in the near future. vi Table of Contents Abstract ........................................................................................................................................... ii Preface............................................................................................................................................ iv Table of Contents .......................................................................................................................... vii List of Tables ................................................................................................................................. ix List of Figures ................................................................................................................................. x Acknowledgements ....................................................................................................................... xii Chapter 1: Introduction ................................................................................................................... 1 Chapter 2: Breadfruit (Artocarpus altilis) for Food Security and Novel Foods in the . 21st Century ................................................................................................................... 6 Breadfruit: An Historical Perspective ........................................................................... 6 Ecological Requirements .............................................................................................. 9 Taxonomy and Botanical Description ........................................................................ 10 Genetic Diversity and Germplasm Conservation ....................................................... 14 Breadfruit Phytochemistry .......................................................................................... 16 Nutritional Composition ............................................................................................. 16 Agronomic Considerations ......................................................................................... 20 The Potential of Breadfruit .......................................................................................... 24 Chapter 3: Morphological Diversity in Breadfruit (Artocarpus, Moraceae): . Insights into Domestication, Conservation, and Cultivar Identification....................................... 35 Introduction ................................................................................................................. 35 Materials and Methods ................................................................................................ 37 Results ......................................................................................................................... 40 Discussion ................................................................................................................... 48 Chapter 4: Diversity of Breadfruit (Artocarpus, Moraceae) Seasonality: . A Resource for Year-Round Nutrition .......................................................................................... 68 Introduction ................................................................................................................