Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald’s Road, London WC1X 8RR, UK Copyright # 2010 Markus Keller, Published by Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone: (þ44) 1865 843830, fax: (þ44) 1865 853333, E-mail: [email protected]. You may also complete your request online via the Elsevier homepage (http://elsevier.com), by selecting “Support & Contact” then “Copyright and Permission” and then “Obtaining Permissions.” Library of Congress Cataloging-in-Publication Data Keller, Markus. The science of grapevines : anatomy and physiology/Markus Keller. p. cm. Includes bibliographical references and index. ISBN 978-0-12-374881-2 (hard cover : alk. paper) 1. Grapes–Anatomy. 2. Grapes–Physiology. I. Title. QK495.V55K44 2010 634.8–dc22 2009033159 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library. ISBN: 978-0-12-374881-2 For information on all Academic Press publications visit our Web site at www.elsevierdirect.com Printed in China 091011987654321 Preface Grapes were among the first fruit species to stomatal pores may be an exception) are rooted be domesticated and today are the world’s in biochemistry. They are driven or at least most economically important fruit crop. facilitated by enzymes, which in turn are built According to 2009 statistics by the Food and based on blueprints provided by genes. I have Agriculture Organization of the United therefore taken it for granted that it is Nations, grapevines were planted on almost understood that a developmental process 7.3 million hectares producing more than 67 or change in chemical composition implies a million metric tons of fruit in 2007. This makes change in enzyme activity, which implies grapes the number 25 food crop in terms of a change in the activity of one or more genes. planted area and number 16 in terms of ton- This does not imply, as used to be thought, that nage. More than 70% of this crop was used to “one gene makes one enzyme,” nor that “one make wine, 27% consumed as fresh fruit (table enzyme makes one chemical,” but it merely grapes), 2% as dried fruit (raisins), and less means that all enzymatic processes have a than 1% was processed to grape juice or dis- genetic basis. tilled to brandy. Many biochemical and biophysical processes This book is an introduction to the physical apply to many or even all plants. Perhaps no structure of the grapevine, its various organs process is truly unique to grapevines. Chances and tissues, their functions, their interactions are that if grapes do it, some or many other with one another, and their responses to the species employ the same solution to a survival environment. It focuses essentially on the phys- issue because they share a common ancestor ical and biological functions of whole plants that invented the trick a long time ago. For rather than the metabolism and molecular example, microbes hit upon photosynthesis biology of individual cells. It is nonetheless and respiration long before these discoveries necessary to review some fundamental enabled some of them to combine forces and processes at the cell, tissue, and organ levels evolve into plants. Consequently, although this in order to build up an appreciation of whole- book is about grapevines, and primarily about plant function. The book covers those elements the wine grape species Vitis vinifera, I have bor- of physiology that will enhance our under- rowed heavily from research done with other standing of grapevine function and how they plant species, both wild and cultivated, peren- relate to grape production. Although of neces- nial and annual, woody and herbaceous, sity the text contains a plethora of technical including the “queen of weeds”—at least in terms and details, I have tried to resist the the fast-paced world of modern molecular biol- temptation to dwell in biochemical and molec- ogy—Arabidopsis thaliana, the otherwise incon- ular biological jargon. Most physiological spicuous thale cress. I have even taken the processes (water movement through the vine’s liberty to borrow insights gained using micro- hydraulic system and evaporation from the organisms, such as the yeast Saccharomyces ix x PREFACE cerevisiae that gives us wine and beer and The magnitude of the task of reviewing as bread, that enable us to think about these much of the pertinent literature as possible issues. often forced me to rely on review papers, This book aims to be global in scale. It covers where they were available. I apologize to those physiological aspects of tropical viticulture all friends and colleagues whose work I did not the way to those that pertain to the production cite or cited incompletely or incorrectly. of ice wine at the temperate northern margins Science—and scientists—can only ever hope to of grape growing. It moves from vineyards at approximate the truth. This and the simple fact sea level to vineyards at high altitude. It con- of “errare humanum est” will guarantee a number siders the humid conditions of cool, marine of errors throughout the text. These are entirely climates, the moist winters and dry summers my responsibility, and I would be grateful for of Mediterranean climates, as well as the arid any feedback that might help improve this book environment typical of continental climates in and further our understanding of the world’s the rain shadow of massive mountain ranges. most important and arguably most malleable Yet a book of this nature is necessarily incom- fruit crop. After all, the full quote from Seneca plete, and so is the selection of published infor- the Younger, who was a contemporary of mation included in the text. No one can read Columella, the Roman author of agriculture everything that has been and is being pub- and viticulture textbooks, reads “errare humanum lished, even in the admittedly relatively narrow est, sed in perseverare diabolicum” (“to err is field of grapevine anatomy and physiology. human, to persevere is devilish”). About the Author Markus Keller is the Chateau Ste. Michelle University in Wagga Wagga, Australia, before Distinguished Professor of Viticulture at coming to eastern Washington. Dr. Keller is Washington State University’s Irrigated Agri- the author of numerous scientific and technical culture Research and Extension Center in papers and industry articles in addition to Prosser, Washington. He received his master’s being a frequent speaker at scientific confer- degree in agronomy (plant science) in 1989 ences and industry meetings and workshops. and a doctorate in natural sciences in 1995 from He also has extensive practical experience in the Swiss Federal Institute of Technology in both the vineyard and the winery as a result Zu¨ rich. He has taught and conducted research of work in the family enterprise, and he was in viticulture and grapevine physiology on awarded the Swiss AgroPrize for innovative three continents, beginning at the Swiss Federal contributions to Switzerland’s agriculture Research Station for Fruit-Growing, Viticulture industry. His current research focuses on and Horticulture in Wa¨denswil (now Agro- environmental factors and management prac- scope Changins-Wa¨denswil), Switzerland, and tices as they influence crop physiology and then moving to Cornell University in Geneva, production of wine and juice grapes. New York, and from there to Charles Sturt vii Acknowledgments Completing this book would have been and encouragement from Nancy Maragioglio impossible without the help and support of and Carrie Bolger at Elsevier are greatly many individuals to whom I express my deep acknowledged. They provided numerous sug- gratitude. Many of the illustrations were gestions that improved this book and were skillfully drawn by Adrienne Mills, and Lynn always quick to answer my countless ques- Mills helped with data collection and assisted tions. Special thanks to my wife, Sandra Wran, with some of the most recalcitrant illustra- for scanning my entire slide collection and tions. I thank Gregory Gasic for reviewing supporting this project in many ways from the entire manuscript, identifying errors, and beginning to end. offering many insightful suggestions. The help xi Elsevier: The Science of Grapevines: Keller Pgina 1 de 2 The Science of Grapevines Keller Table of Contents Chapter 1 Botany and Anatomy 1.1 Botanical classification and geographical distribution 1.2 Cultivars, clones and rootstocks 1.3 Morphology and anatomy Chapter 2 henology and Growth Cycle 2.1 Seasons and daylength 2.2 Vegetative cycle 2.3 Reproductive cycle Chapter 3 Water Relations and Nutrient (ptake 3.1 Osmosis, water potential and cell expansion 3.2 Transpiration and stomatal action 3.3 Water and nutrient uptake and transport Chapter , hotosynthesis and Respiration ,.1 -ight absorption and energy capture ,.2 Carbon uptake and assimilation ,.3 hotorespiration ,., Respiration Chapter . artitioning of Assimilates ..1 hotosynthate translocation and distribution ..2 Canopy-environment interactions ..3 Nitrogen assimilation and interaction with carbon metabolism http:// .elsevierdirect.com/toc.jsp$isbn=9780123748812 5/4/2011 Elsevier: The Science of Grapevines: Keller Pgina 2 de 2 Chapter 0 1evelopmental hysiology 0.1 2ield formation 0.2 Grape composition and fruit 3uality Chapter 7 Environmental Constraints and Stress hysiology 7.1 Responses to stress 7.2 Water6 too much or too little 7.3 Nutrients6 deficiency and excess 7., Temperature6 too cold or too warm 7.. -iving with other organisms6 defense and damage Glossary References Internet Resources http:// .elsevierdirect.com/toc.jsp$isbn=9780123748812 5/4/2011 CHAPTER 1 Botany and Anatomy OUTLINE 1.1.