Cacti, Biology and Uses
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CACTI CACTI BIOLOGY AND USES Edited by Park S. Nobel UNIVERSITY OF CALIFORNIA PRESS Berkeley Los Angeles London University of California Press Berkeley and Los Angeles, California University of California Press, Ltd. London, England © 2002 by the Regents of the University of California Library of Congress Cataloging-in-Publication Data Cacti: biology and uses / Park S. Nobel, editor. p. cm. Includes bibliographical references (p. ). ISBN 0-520-23157-0 (cloth : alk. paper) 1. Cactus. 2. Cactus—Utilization. I. Nobel, Park S. qk495.c11 c185 2002 583'.56—dc21 2001005014 Manufactured in the United States of America 10 09 08 07 06 05 04 03 02 01 10 987654 321 The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48–1992 (R 1997) (Permanence of Paper). CONTENTS List of Contributors . vii Preface . ix 1. Evolution and Systematics Robert S. Wallace and Arthur C. Gibson . 1 2. Shoot Anatomy and Morphology Teresa Terrazas Salgado and James D. Mauseth . 23 3. Root Structure and Function Joseph G. Dubrovsky and Gretchen B. North . 41 4. Environmental Biology Park S. Nobel and Edward G. Bobich . 57 5. Reproductive Biology Eulogio Pimienta-Barrios and Rafael F. del Castillo . 75 6. Population and Community Ecology Alfonso Valiente-Banuet and Héctor Godínez-Alvarez . 91 7. Consumption of Platyopuntias by Wild Vertebrates Eric Mellink and Mónica E. Riojas-López . 109 8. Biodiversity and Conservation Thomas H. Boyle and Edward F. Anderson . 125 9. Mesoamerican Domestication and Diffusion Alejandro Casas and Giuseppe Barbera . 143 10. Cactus Pear Fruit Production Paolo Inglese, Filadelfio Basile, and Mario Schirra . 163 11. Fruits of Vine and Columnar Cacti Avinoam Nerd, Noemi Tel-Zur, and Yosef Mizrahi . 185 12. Forage, Fodder, and Animal Nutrition Ali Nefzaoui and Hichem Ben Salem . 199 13. Nopalitos, Mucilage, Fiber, and Cochineal Carmen Sáenz-Hernández, Joel Corrales-Garcia, and Gildardo Aquino-Pérez . 211 14. Insect Pests and Diseases Helmuth G. Zimmermann and Giovanni Granata . 235 15. Breeding and Biotechnology Brad Chapman, Candelario Mondragon Jacobo, Ronald A. Bunch, and Andrew H. Paterson . 255 Index . 273 CONTRIBUTORS Edward F. Anderson (Deceased), Desert Botanical Joel Corrales-Garca, Departamento de Ingeniería Garden, Phoenix, Arizona Agroindustrial, Universidad Autónoma de Chapingo, Mexico Gildardo Aquino-Prez, Insituto de Recursos Genéticos y Productividad, Montecillo, Mexico Rafael F. del Castillo, Centro Interdisciplinario de Investigacíon para el Desarrollo Integral Regional Giuseppe Barbera, Istituto di Coltivazioni Arboree, Unidad Oaxaca, Mexico Università degli Studi di Palermo, Italy Joseph G. Dubrovsky, Departamento de Biología Filadelfio Basile, Dipartimento Scienze Economico- Molecular de Plantas, Instituto de Biotecnología, Agrarie ed Estimativ, Universita degli Studi di Catania, Universidad Nacional Autónoma de México, Cuernavaca Italy Arthur C. Gibson, Department of Organismic Hichem Ben Salem, Institut National de la Biology, Ecology and Evolution, University of Recherche Agronomique de Tunisie, Laboratoire California, Los Angeles de Nutrition Animale, Ariana, Tunisia Hctor Godnez-Alvarez, Departamento de Edward G. Bobich, Department of Organismic Ecología Funcional y Aplicada, Instituto de Ecología, Biology, Ecology and Evolution, University of Universidad Nacional Autónoma de México, Mexico California, Los Angeles City Thomas H. Boyle, Department of Plant and Soil Giovanni Granata, Dipartimento di Scienze e Sciences, University of Massachusetts, Amherst Technologie Fitosanitartie, Università degli Studi di Catania, Italy Ronald A. Bunch, D’Arrigo Bros. Co., Salinas, California Paolo Inglese, Istituto di Coltivazioni Arboree, Palermo, Italy Alejandro Casas, Departamento de Ecología de los Recursos Naturales, Instituto de Ecología, Universidad James D. Mauseth, Department of Integrative Nacional Autónoma de México, Morelia Biology, University of Texas at Austin Brad Chapman, Plant Genome Mapping Laboratory, Eric Mellink, Centro de Investigación Cientifica University of Georgia, Athens y de Educación Superior de Ensenada, Mexico vii Yosef Mizrahi, Department of Life Sciences and Mnica E. Riojas-Lpez, Departamento de Institutes for Applied Research, Ben-Gurion University Ecología, Centro Universitario de Ciencias Biológicas of the Negev, Israel y Agropecuarias, Universidad de Guadalajara, Mexico Candelario Mondragon Jacobo, Programa de Carmen Senz-Hernndez, Departamento de Nopal y Frutales, Instituto Nacional de Investigaciones Agroindustria y Enología, Facultad de Ciencias Agrarias Forestales y Agropecuarias, Queretaro, Mexico y Forestales, Universidad de Chile, Santiago Ali Nefzaoui, Institut National de la Recherche Mario Schirra, Instituto per la Fisologia della Agronomique de Tunisie, Laboratoire de Nutrition Maturazione e della Conservazione del Frutto delle Animale, Ariana, Tunisia Specie Arboree Mediterranee, Oristano, Italy Avinoam Nerd, Institutes for Applied Research, Noemi Tel-Zur, Department of Life Sciences, Ben-Gurion University of the Negev, Israel Ben-Gurion University of the Negev, Israel Park S. Nobel, Department of Organismic Biology, Teresa Terrazas Salgado, Programa de Botánica, Ecology and Evolution, University of California, Los Colegio de Postgraduados, Montecillo, Mexico Angeles Alfonso Valiente-Banuet, Departamento de Gretchen B. North, Department of Biology, Ecología Funcional y Aplicada, Instituto de Ecología, Occidental College, Los Angeles, California Universidad Nacional Autónoma de México, Mexico City Andrew H. Paterson, Plant Genome Mapping Laboratory, University of Georgia, Athens Robert S. Wallace, Department of Botany, Iowa State University, Ames Eulogio Pimienta-Barrios, Departamento de Ecología, Centro Universitario de Ciencias Biológicas Helmuth G. Zimmermann, Plant Protection y Ambientales, Universidad de Guadalajara, Mexico Research Institute, Agricultural Research Council, Pretoria, South Africa viii Contributors PREFACE The Cactaceae, a family of approximately 1,600 species, is and their uses. Twelve authors are from Mexico, eleven from native to the New World but is cultivated worldwide. In re- the United States, five from Italy, three from Israel, two sponse to extreme habitats, cacti have evolved special phys- from Tunisia, and one each from Chile and South Africa. iological traits as well as distinctive appearances. The stem Most of the authors share my interests in basic research on morphology, spine properties, and often spectacular flowers the Cactaceae. Nearly half of the authors, especially those have caused hobbyists to collect and cultivate large numbers dealing with agronomic aspects, are involved with the of cacti. Both cactus form and function relate to nocturnal CactusNet sponsored by the Food and Agricultural Orga- stomatal opening and Crassulacean acid metabolism, which nization of the United Nations. Approximately 1,300 refer- lead to efficient use of limited soil water. Thus, cacti can ences are cited in the chapters, which not only indicate the thrive in arid and semiarid environments, where they are widespread interest in cacti but also should facilitate further often important resources for both wildlife and humans. investigations. The intended audience ranges from ecolo- Indeed, cacti have been consumed by humans for more gists and environmentalists to agriculturalists and con- than 9,000 years. Currently, Opuntia ficus-indica is culti- sumers to cactus hobbyists and enthusiasts. vated in over 20 countries for its fruit, and an even greater The point of departure is the evolution of the family in land area is devoted to its cultivation for forage and fodder. the broad sense, paying particular attention to new mo- The fruits of other cactus species, known as pitahayas and lecular and genetic approaches (Chapter 1). People recog- pitayas, and various other cactus products are appearing in nize cacti by their shoot morphology, which reflects vari- an increasing number of markets worldwide. ous cellular characteristics (Chapter 2). The uptake of Due to the high water-use efficiency and other adapta- water and nutrients from the soil by roots that sustains the tions of cacti, biological and agronomic interest in them has shoots has unique features as well (Chapter 3). Survival de- soared. From 1998 to 2000, more than 600 researchers pub- pends on adaptation to abiotic environmental conditions, lished over 1,100 articles on cacti, including papers in pro- which cacti have done in special ways (Chapter 4). In ad- ceedings of national and international meetings. Yet a cur- dition to enduring harsh conditions, cacti must reproduce, rent, synthetic, widely ranging reference is not available. for which many strategies have evolved (Chapter 5). Biotic This book, which consists of a series of authoritative, up-to- factors are also crucial for the success of cacti in natural en- date, review chapters written by established experts as well vironments (Chapter 6). Because of their ecological suc- as new contributors, emphasizes both the biology of cacti cess, cacti are important food resources for wild vertebrates ix (Chapter 7). The many unique characteristics of the De la Barrera assisted with the many Spanish citations. Cactaceae have attracted collectors and raised concerns Marian McKenna Olivas competently did line editing, and about issues of biodiversity and conservation (Chapter 8) Alicia Materi meticulously typed the developmental and as well as led to their ancient usage and subsequent wide line editing changes. Financial support for