C R C R E V I V A L S C R C R E V I V A L S E P d r it e e d d i c b y ti n J g o h P n h D o Predicting Photosynthesis for . t H o s e Ecosystem Models s y k n e t h t h e Volume I , s J i a s m f e o s r W E c . J o o s n y Edited by e s s t e John D. Hesketh, James W. Jones m M o d e l s ISBN 978-1-315-89686-1 ,!7IB3B5-ijgigb! www.crcpress.com Predicting Photosynthesis for Ecosystem Models Volume I Editors J. D. Hesketh, Ph.D. Crop Physiologist Agricultural Research, Scientific and Education Administration U.S. Department of Agriculture University of Illinois Urbana, Illinois James W. Jones, Ph.D. Associate Professor Agricultural Engineering University of Florida Gainesville, Florida Boca Raton London New York CRC Press, Inc. CRC Press is an imprint of the Taylor &B Foracnaci sR Graotuop, nan, Finlfoorrmiad abu siness First published 1980 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1980 by CRC Press, Inc. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. 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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging in Publication Data Main entry under title: Predicting photosynthesis for ecosystem models. Bibliography: p. Includes indexes. 1. Photosynthesis — Simulation methods. I. Hesketh, John D. II. Jones, James Wigington, 1944- QK882.P73 581.1’3342’0184 79-4047 ISBN 0-8493-5335-1 (v. 1) ISBN 0-8493-5336-X (v. 2) A Library of Congress record exists under LC control number: 79004047 Publisher’s Note The publisher has gone to great lengths to ensure the quality of this reprint but points out that some imperfections in the original copies may be apparent. Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-1-315-89686-1 (hbk) ISBN 13: 978-1-351-07596-1 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com PREFACE Research and development agencies in the United States have emphasized the need for methods of predicting ecosystem behavior, so as to minimize pollution (insecticide, herbicide, and fertilizer usage) and conserve energy (petroleum products) while maxi- mizing productivity and profits. Some of this pressure may have evolved from the successful use of system analysis techniques by the National Aeronautics and Space Administration in their missions in the 1960s. The editors of these volumes collabo- rated in the late 1960s and early 1970s in a regional cotton project using such system analysis techniques while located at Mississippi and North Carolina State Universities with the U.S. Department of Agriculture. Dr. W. G. Duncan (Universities of Kentucky and Florida) and Dr. D. N. Baker (USDA, Mississippi State) played important roles in these collaborations. It was an exciting period involving several disciplines in an uniquely exploratory effort. For example, at one point we were interested in the role of the crop in regulating the density and physiological state of the boll weevil, an insect pest. Therefore, we studied the nitrogen and carbon compositions of various plant parts with respect to the nutritional requirements of the boll weevil larvae and adult~. Diets high in protein induced reproduction while diets high in carbohydrates induced storage of fats and preparation for overwintering (diapause). Photosynthate, as recently formed reductant in photosynthetic cells or as reduced nitrogen or carbon, plays a key role in supplying the energy needs for an ecosystem. In the context of predicting ecosystem behavior, photosynthesis must be predicted as it interacts with other plant physiological processes as well as with animal behavior, such as the effect of insects feeding on leaves. Likewise a study of other aspects of ecosystem behavior must take into account their interactions with photosynthesis activ- ities. Therefore our coverage is broad, including descriptions of how one might inte- grate a photosynthetic model into an ecosystem predictor. The understanding and de- scription of interactions among processes is vital to a successful ecosystem model. We invited several full-time experimentalists unassociated with any prediction effort (one of us, JDH, is a full-time experimentalist) to review problems one frequently encounters when attempting to predict field photosynthesis. Some of these problems may only be resolved using a whole plant approach and studying photosynthesis in relation to other plant physiological processes. In analyzing the recent literature, it would seem that experimentalists in the various specialties of plant physiology have developed, independently, the same techniques as those using the multidisciplinary ap- proach to study photosynthesis for ecosystem predictions. Photosynthesis is being studied in many laboratories as it relates to whole plant behavior. In our opinion, the experimentalists authoring chapters in these volumes had no problem identifying rele- vant subject matter for discussion. Indeed, a major limiting factor in ecosystem pre- diction efforts is the participation of experimentalists in reviewing the status of various biological problems involved. We recommend more of this in the near future. It is likely that participation will only come with the invitation to review a carefully defined problem. We go into greater detail justifying this effort in the first chapter. There are now four methods for predicting photosynthesis, each with limitations and strengths. Their present usage depends upon the purpose of the prediction to be made, as well as im- provements in associated techniques that seem to revolutionize the methodology. At this point in time, comparisons among methods are valuable, justifying a state-of-the- art review for all photosynthetic scientists. We were faced with the need to develop general notation for all the chapters but the subject matter was so diverse, with so many traditions in notation, that we gave up. Perhaps our failure in this respect will focus on a solution to the problem. Rarely has so much material been brought together in one place. We can't help but feel that progress in this general area of research will be rapid in the near future. We hope these volumes speed up the process. J. D. Hesketh J. W. Jones July 1979 CONTRIBUTORS Randall S. Alberte, Ph. D. John D. Hesketh, Ph.D. Assistant Professor Crop Physiologist Department of Biology AR-SEA-USDA University of Chicago University of Illinois Chicago, Illinois Urbana, Illinois Larry G. Brown, Ph.D. Associate Professor, Industrial Engineering James W. Jones, Ph.D. Mississippi State University Associate Professor Mississippi State, Mississippi Department of Agricultural Engineering University of Florida J. Robert Cooke, Ph.D. Gainesville, Florida Director of Instruction College of Agriculture and Life Sciences Kenneth R. Knoerr, Ph.D. Professor of Agricultural Engineering Professor of Forest Meteorology Cornell University Department of Botany Ithaca, New York Duke University Durham, North Carolina Gary L. Cunningham, Ph.D. Professor of Biology New Mexico State University Harry R. Leffler, Ph.D. Las Cruces, New Mexico Plant Physiologist AR-SEA-USDA Cotton Physiology and C. D. Elmore, Ph.D. Genetics Laboratory Plant Physiologist Stoneville, Mississippi AR-SEA-USDA Southern Weed Science Laboratory Stoneville, Mississippi Jackson R. Mauney, Ph.D. Plant Physiologist David M. Gates, Ph.D. AR-SEA-USDA Western Cotton Professor of Botany Research Laboratory Director, Biological Station Phoenix, Arizona University of Michigan Ann Arbor, Michigan Gene Guinn, Ph.D. Bobby L. McMichael, Ph.D. Plant Physiologist (Research Leader) Plant Physiologist AR-SEA-USDA Western Cotton AR-SEA-USDA Plant and Soil Science Research Laboratory Department Phoenix, Arizona Texas Technical University Lubbock, Texas Peter C. Harley, M.S. Research Assistant John M. Norman, Ph.D. Biological Station Professor of Agronomy University of Michigan The University of Nebraska Ann Arbor, Michigan Lincoln, Nebraska David T. Patterson, Ph.D. James F. Reynolds, Ph.D. Plant Physiologist Assistant Professor AR-SEA-USDA Southern Weed Department of Botany Science Laboratory North Carolina State University Stoneville, Mississippi Raleigh, North Carolina John W. Radin, Ph.D. Boyd R. Strain, Ph.D. Plant Physiologist AR-SEA-USDA Western Cotton Professor of Botany Research Laboratory Duke University Phoenix, Arizona Durham, North Carolina Richard H. Rand, Sc.D. Associate Professor John D. Tenhunen, Ph.D. Department of Theoretical and Applied Research Associate Mechanics Biological Institute Cornell University University of Wurzburg Ithaca, New York Wurzburg, West Germany THE EDITORS J. D. Hesketh was born in Maine and attended the University of Maine and Cornell University. He has worked in Connecticut, Arizona, Canberra (Australia), Mississippi, North Carolina, and Illinois with various governmental agencies. His early research was concerned with gas exchange aspects of the so-called C_,-C phenomenon, as well 4 as the effect of environment on crop photosynthesis. His interest in the latter has con- tinued to date, as well as collaborations in various efforts to forecast crop yields. J. W. Jones was born in Mississippi and received degrees from Texas Tech, Mississippi State, and North Carolina State Universities. He worked in Mississippi and in North Carolina with the U.S. Department of Agriculture and in Florida with the University of Florida. His early research focused on analysis of cotton production systems with emphasis on modeling cotton crop growth and yield as well as crop-pest interactions. Currently, his interests are in modeling crop growth processes as affected by water and other stresses. He has taught several courses related to systems analysis of agricultural and biological systems. TABLE OF CONTENTS VOLUME 1 Chapter 1. What is Modeling and Why Model Photosynthesis? ............................. 1 J.D. Hesketh and J. W. Jones Chapter 2. Micrometeorological Methods for Predicting Environmental Effects on Photosyn- thesis .................................................................... 9 J. M. Norman and J. D. Hesketh Chapter 3. Predicting Canopy Photosynthesis from Gas Exchange Studies in Controlled Environ· ments ................................................................... 37 J. D. Hesketh Chapter 4. Integrating Traditional Growth Analysis Techniques with Recent Modeling of Carbon and Nitrogen Metabolism .................................................. Sl J.D. Hesketh and J. W. Jones ChapterS. Diffusion Resistance Models ................................................ 93 J. R. Cooke and R. H. Rand Chapter 6. Leaf Photosynthesis Models ............................................... 123 J.D. Tenhunen, J.D. Hesketh, and D. M. Gates Chapter 7. Water Stress Adaptation .................................................. 183 B. L. McMichael Chapter 8. Light and Temperature Adaptation ......................................... 205 D. T. Patterson Index .................................................................. 237 VOLUME II Chapter 1 Analysis of CO, Exchange Assumptions: Feedback Control ...................... 1 G. Guinn and J. Mauney Chapter 2. Modeling C Respiration in the Light ......................................... 17 3 J.D. Tenhunen J.D. Hesketh, and P. C. Harley Chapter 3. Interfacing Leaf and Canopy Light Interception Models ........................4 9 J. Norman Chapter4. Predicting Dark Respiration in the Soil-Plant System .........................6 9 J.D. Hesketh, R. S. Alberte, and J. W. Jones Chapter 5. Predicting Leaf Expansion ................................................. 85 J. W. Jones and J.D. Hesketh Chapter 6. Predicting Flowering and Subsequent Leaf Expansion ......................... 123 J.D. Hesketh, C. D. Elmore, and J. W. Jones Chapter 7. Leaf Growth and Senescence .............................................. 133 H. Leffler Chapter 8. Concepts of Translocation with Special Reference to the Assimilation of Nitrogen and its Movement into Fruits .................................................. 143 J. Radin and C. D. Elmore Chapter 9. The Paradox of No Correlation Between Leaf Photosynthetic Rates and Crop Yields .................................................................. 155 C. D. Elmore Chapter 10. Predicting Primary Productivity for Forest and Desert Ecosystem Models ........ 169 J. F. Reynolds, B. R. Strain, G. L. Cunningham, and K. R. Knoerr Chapter 11. COTCROP: A Computer Model for Cotton Growth and Yield ................. 209 J. W. Jones, L. G. Brown, and J.D. Hesketh Index .................................................................. 243