Academic Press Rapid Manuscript Reproduction M e a s u r e m e nt T e c h n i q u es in P l a nt S c i e n ce Edited by YASUSHI HASHIMOTO PAUL J. KRAMER Department of Biomechanical Systems Department of Botany College of Agnculture, Ehime University Duke University Tarumi, Matsuyama, Japan Durham, North Carolina HIROSHI NONAMI BOYD R. STRAIN Department of Biomechanical Systems Department of Botany College of Agnculture, Ehime University Duke University Tarumi, Matsuyama, Japan Durham, North Carolina ACADEMIC PRESS, INC. Harcowt Brace Jovanovich, Publishers San Diego New York Boston London Sydney Tokyo Toronto This book is printed on acid-free paper. © Copyright © 1990 by Academic Press, 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 photo copy, recording or any information storage and retrieval system, without permission in writing from the publisher. Academic Press, Inc. San Diego, California 92101 United Kingdom Edition published by Academic Press Limited 24-28 Oval Road, London NW1 7DX Library of Congress Cataloging-in-Publication Data Measurement techniques in plant science / edited by Yasushi Hashimoto ... [et al.]. p. cm. Based on a symposium entitled "Instrumentation and Physi ological Ecology", held in Tokyo in 1985 ISBN 0-12-330585-3 1. Plant physiological ecology-Congresses. 2. Growth (Plants)- - Measurement-Congresses. 3. Crops-Growth--Measurement- -Congresses. I. Hashimoto, Yasushi, Date. QK905.M43 1990 581.3'l-dc20 90-49622 CIP Printed in the United States of America 90 91 92 93 9 8 7 6 5 4 3 2 1 C O N T R I B U T O RS Numbers in parentheses indicate the pages on which the authors' contributions begin. I. Aiga (343), College of Agriculture, University of Osaka, Prefecture, Sakai, Osaka 591, Japan /. S . Boyer (101), College of Marine Studies, University of Delaware, Lewes, Delaware 19958 W. Day (207), AFRC Institute of Engineering Research, Wrest Park, Silsoe, Bedford MK45 4HS, United Kingdom H. Eguchi (361), Biotron Institute, Kyushu University, Fukuoka 812, Japan E. Epstein (291), Department of Land, Air and Water Resources, Univer sity of California, Davis, California 95616 Y. Fares (265), Biosystems Technologies, Inc., Durham, North Carolina 27706-4851 C. B. Field (185), Department of Plant Biology, Carnegie Institution of Washington, 290 Panama Street, Stanford, California 94305-1297 JB. L. Fiscus (79), United States Department of Agriculture, Agriculture Research Service, 1701 Center Fort Collins, Colorado 80526 Avenue , J. D. Goeschl (265), Biosystems Technologies, Inc., Durham, North Caro lina 27706-4851 Y. Hashimoto (7,373), Department of Biomechanical Systems, College of Agriculture, Ehime University, Tarumi, Matsuyama 790, Japan S. E. Hetherington (229), Division of Horticulture, Sydney Laboratories, CSIRO, North Ryde, Sydney 2113, Australia C. R Jaeger (265), EPO Biology, University of Colorado, Boulder, Colorado 80309 T. Kaneko (277), Advanced Research Laboratory, Hitachi, Ltd., Kokubunji, Tokyo, Japan y. Kano (165), Department of Electrical Engineering, Faculty of Technol ogy, Tokyo University of Agriculture and Technology, Koganei, Tokyo, Japan M. JR . Kaufmann (69), United States Department of Agriculture Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado 80521 ix χ CONTRIBUTORS /· Kondo (343), Science Council of Japan, Roppongi, Tokyo 106, Japan P. J. Kramer (3,45,403), Department of Botany, Duke University, Durham, North Carolina 27706 S. Kuraishi (151), Department of Environmental Sciences, Faculty of In tegrated Arts and Sciences, Hiroshima University, Hiroshima 730, Japan /. S. MacFall (403), School of Forestry and Environmental Studies, Duke University, Durham, North Carolina 27706 C. £· Magnuson (265), Biosystems Technologies, Inc., Durham, North Carolina 27706-4851 H. Miyauchi (151), Department of Environmental Sciences, Faculty of Integrated Arts and Sciences, Hiroshima University, Hiroshima 730, Japan H. A. Mooney (185), Department of Biological Sciences, Stanford Univer sity, Stanford, California 94305 H. Nonami (7, 101), Department of Biomechanical Systems, College of Agriculture, Ehime University, Tarumi, Matsuyama 790, Japan K. Omasa (343,387), National Institute for Environmental Studies, Yatabe, Tsukuba, Ibaraki 305, Japan K. /. Parkinson (207), Analytical Development Company, Ltd., Pindar Road, Hoddesdon, Herts EN1 10AQ, United Kingdom N. Sakurai (151), Department of Environmental Sciences, Faculty of In tegrated Arts and Sciences, Hiroshima University, Hiroshima 730, Japan K. Shimazaki (387), College of General Education, Kyushu University, Ropponmatsu, Fukuoka 810, Japan /. ΛΓ. Siedow (403), Department of Botany, Duke University, Durham, North Carolina 27706 JR. M. Smillie (229), Division of Horticulture, Sydney Laboratories, CSIRO, North Ryde, Sydney 2113, Australia E. Steudle (113), Lehrstuhl fur Pflanzenokologie, Universitat Bayreuth, 8580 Bayreuth, Federal Republic of Germany B. R. Strain (265), Department of Botany, Duke University, Durham, North Carolina 27706 K. Supappibul (151), Mangrove Forest Management Unit, Lamngob, Namchew, Trat, Thailand M. Takatsuji (277), Advanced Research Laboratory, Hitachi, Ltd., Kokubunji, Tokyo, Japan R. M. Welch (319), United States Department of Agriculture, Agriculture Research Service, U.S. Plant, Soil & Nutrition Laboratory, Ithaca, New York 14853-0331 H. Yamasaki (25), Department of Mathematical Engineering and Infor mation Physics, Faculty of Engineering, University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113, Japan PREFACE Progress in plant science always has been and continues to be dependent on progress in the instrumentation required to measure plant processes and environmental factors. This book has its origins in the international symposium entitled "Instrumentation and Physiological Ecology" held in Tokyo in 1985. The symposium brought together scientists from labora tories in various countries to exchange information about existing in strumentation and provided opportunities to discuss new problems and new apparatus needed to solve them. The symposium covered the topics of water relations, photosynthesis, translocation, mineral nutrition, and image processing. Since the time of the symposium, there have been important developments and refinements in measurement techniques and technologies, such as intelligent sensing systems and nuclear magnetic resonance (NMR) computer tomography, which are included as topics in this book. In addition, articles have been revised and updated as deemed necessary by their authors. This book will be of interest not only to specialists in plant science, but also to students who may be interested in reading of the histories of the development of the instrumentations which are included in the reviews. It should also be of interest to engineers working toward practical appli cations of the techniques for crop management described in the book. Y. Hashimoto P. f. Kramer H. Nonami B. R. Strain xi INSTRUMENTATION AND PROGRESS IN SCIENCE Paul J. Kramer Department of Botany Duke University Advances in science depend on new ideas and on development of the instrumentation needed to investigate the ideas. Pioneers such as Newton, Darwin, Mendel, and Einstein created new concepts with revolutionary implications for various branches of science, but their concepts are still being explored and expanded as new instrumentation and new research methods permit the acquisition of new information. The importance of instrumentation is well recognized in the scientific world and several Nobel prizes have been awarded to developers of devices such as cyclotrons, transistors, lasers, partition chromatography, and the CAT X-ray scanning technique. Many good ideas have waited for decades because of the lack of instrumentation to investigate them, and on the other hand development of new instrumentation often results in a notable increase in research activity. The history of the study of photosynthesis provides an example of how progress depends on instrumentation. About 1771 Priestley observed that green plants changed the composition of the air in containers enclosing the plants. However, it was not until improved methods for analyzing the composition of the air and measuring changes in volume of its components were developed early in the 19th century that it was established by de Saussure that green plants in the light absorb CO2 and release O2· It was another 60 years before Sachs established that the chloroplasts were involved in this gas exchange and that carbohydrate was produced. It was not until after radioactive isotopes became available in the 1930s that the reductive carbon cycle was worked out, and only after the development of new instrumentation was it possible to study photosynthetic electron flow. Measurement Techniques in Plant Science Copyright © 1990 by Academic Press, Inc. All rights of reproduction in any form reserved. 3 4 Chapter 1: Introduction Few field measurements of photosynthesis were made during the 19th and early 20th century because there was no convenient method of measuring gas exchange. In early studies change in CO2 concentration of the air passed over leaves was measured by bubbling the air through tubes filled with dilute alkali and titrating the alkali or measuring change in conductivity to determine the amount that had been neutralized by the CO2. This method was slow, untidy, and not very accurate. During the 1940s infrared gas analyzers began to be used to measure change in concentration of gases and commercially built infrared gas analyzers sensitive to CO2 became available in the 1950s. However, the first instruments were large and heavy and required mobile laboratories for field measurements of photosynthesis. Improvements in electronics and microprocessors now make it possible to carry the equipment to the field in a suitcase. Infrared gas analyzers proved to be so useful, both in the laboratory and in the field, that there was a great increase in research on gas exchange of plants under various environmental conditions. Today infrared gas exchange measurements are being supplemented by measurements of oxygen production of leaf disks, chlorophyll fluorescence of attached leaves, and carbon isotope discrimination, to provide a broader understanding of photosynthesis. Some of these technologies are discussed in this volume. There was a similar lag between theory and practice in the field of plant water relations. The idea that the free energy status of water (now termed the water potential) was important was developed early in this century, but several decades passed before practical methods of measuring water potential were developed (see Chapter 2). Good physiological-ecological research often benefits from simultaneous measurement of CO2 exchange, stomatal conductance, and plant water status, all of which have become possible because reliable, portable gas analyzers, porometers, thermocouple psychrometers, and Scholander pressure chambers are available. As a result of these improvements in instrumentation research in this area has increased many fold in recent years. Improvements in the measurement of plant water status are discussed in Chapter 2. Instrumentation and Progress in Science 5 Improvements in equipment to control the plant environment also have contributed to progress in physiological and ecological research. Technological advances in air conditioning and control systems and improvements in artificial lighting have increased the usefulness of growth chambers and improved the reproducibility of controlled environmental regimes. New information often indicates new areas of ignorance and the need for more information, and this in turn creates the need for additional instrumentation. Looking to the future we are in a period of rapid development of instrumentation in which the combination of improved electronics and computer capacity combined with existing instrumentation permits measurements that were impossible a decade or two ago. An example is the development of nuclear magnetic resonance spectroscopy and imaging. The physical principles were understood before the technology made it useful in biology and medicine. Only after strong, stable, superconducting magnets and large computers to store and process data became available could the concept be fully exploited. Now improvements in gradient and radio frequency coils and pulse control are resulting in images in the microscopic range (see Chapter 6). Technology is advancing too rapidly for plant scientists to keep informed concerning new apparatus and new methods that might be useful in their research. An example is the development of a half dozen new scanning probes, following the appearance of the scanning tunneling microscope (STM) in 1981. The STM does not work well on biological material, but it seems possible that some of the new scanning probe microscopes such as the atomic force microscope or the scanning near- field optical microscope may become useful in biological research. Thus there is increasing need for interdisciplinary exchange and collaboration between plant scientists and designers of scientific equipment. Books such as this should increase the transfer of useful technology. OVERVIEW OF CURRENT MEASUREMENT TECHNIQUES FROM ASPECTS OF PLANT SCIENCE Yasushi Hashimoto Hiroshi Nonami Department of Biomechanical Systems College of Agriculture, Ehime University Tarumi, Matsuyama 790, Japan I . INTRODUCTION Advancements in sensor designs and developments in measurement techniques have been occurring rapidly in recent years owing to developments in electronics and computer science. Such progress has deeply influenced current measurement techniques in plant science. Agricultural industries, as well as scientists in the field of plant science, have an interest in the recent developments in sensors and measurement technologies, as attempts are being made to automate agricultural production. However, automation of agricultural production has not been widely attained a commercially feasible basis thus far. Developments in instrumentation have led to smaller and smaller instruments, enabling researchers to measure quantities more precisely. Also, the invention of instrumentation for use with intact living tissues has made it possible to measure physiological information without Measurement Techniques in Plant Science Copyright Ο 1990 by Academic Press, Inc. 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