Plant Physiological Ecology

Plant Physiological Ecology

Plant Physiological Ecology Field methods and instrumentation

Edited by

RW. PEARCY,J.R EHLERINGER, H.A. MOONEY and P.W. RUNDEL

KLUWER ACADEMIC PUBLISHERS DORDRECHT / BOSTON / LONDON

British Library Cataloging in Publication Data Plant physiological ecology: field methods and instrumentation. 1. Plants. Physiology. Ecological aspects I. Pearcy, R.w. (Robert W.), 1941581.1 ISBN-13:978-0-412-40730-7 e-ISBN-13:978-94-010-9013-1 DOl: 10.1007/978-94-010-9013-1 Library of Congress Cataloging-in-Publication Data Plant physiological ecology. Includes Index. 1. Plants physiological ecology - Field work. 2. Plants physiological ecology - Instruments. I. Pearcy, R.w. (Robert W.), 1941QK 905.P57 1989 581.5.222 88-30018 ISBN-13:978-0-412-40730-7 ISBN-13:978-0-412-40730-7 (PB)

Published by Kluwer Academic Publishers, P.O. Box 17, 3300 AA Dordrecht, The Netherlands. Sold and distributed in North, Central and South America by Kluwer Academic Publishers, 101 Philip Drive, Norwell, MA 02061, U.s.A. In all other countries, sold and distributed by Kluwer Academic Publishers, P.O. Box 322, 3300 AH Dordrecht, The Netherlands.

First edition 1989 Reprinted 1991

Printed on acid-free paper

All Rights Reserved © 2000 Kluwer Academic Publishers

No part of the material protected by this copyright notice may be reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording or by any information storage and retrieval system, without written permission from the copyright owner.

Contents

1

Contributors

xi

Introduction

xv

Principles of instrumentation for physiological ecology Arnold J. Bloom

1.1 1.2 1.3 1.4 1.5 2

Field data acquisition Robert W. Pearcy

2.1 2.2 2.3 2.4 2.5 3

Introduction Measurement and measurement errors Instrument organization Instrument initiation Postscript

Introduction Analog recorders Digital recorders Integrators Sampling considerations

Water in the environment Philip W. Rundel and Wesley M. Jarrell

3.1 Soil moisture 3.2 Atmospheric moisture 3.3 Moisture flux 4 Measurement of wind speed near vegetation

1

1 1 3 11

12

15

15 15 17 24 24 29

29 41 47 57

John Grace

4.1 4.2 4.3 4.4 4.5

Introduction Flow in wind tunnels, growth cabinets and ducts Weather stations and field survey Wind profiles above vegetation Boundary layer resistance

57 58 60 61 64

Contents

vi 4.6 4.7 4.8 4.9 4.10

Calibration Aerodynamic influence by masts Visualization Pressure measurements Some applications

5 Soil nutrient availability Dan Binkley and Peter Vitousek 5.1 5.2 5.3 5.4 5.5 5.6 5.7 5.8 5.9 5.10 5.11 5.12 5.13 5.14

Introduction Difficulties in measuring nutrient availability Nitrogen availability Phosphorus availability Sulfur availability Availability of essential cations Micronutrient availability Soil classification Bioassay of nutrient availability Soil acidity Soil salinity Soil redox potential Comments on sampling Index units

6 Radiation and light measurements Robert W. Pearcy

6.1 6.2 6.3 6.4 6.5 6.6 6.7 6.8 6.9

Introduction Definitions and units Energy versus photons as a measure of PAR Radiation sensors: general characteristics Determination of the diffuse and direct components of radiation Calibration of radiation sensors Sampling considerations Photographic estimations of light climate Spectral radiometry

7 Temperature and energy budgets James R. Ehleringer 7.1 7.2 7.3 7.4 7.5 7.6 7.7 7.8

Introduction Energy budget approach Variations in air and leaf temperatures with height Temperature and its measurement Orientation and its measurement Calculation of incident solar radiation on different surfaces Leaf absorptance and its measurement Boundary layer considerations

66 66 66 66 68 75 75 76 76 81 85 85 87 88 88 88 90 90 90 92 97

97 97 99 100 107 107 108 109 113 117 117 117 123 124 130 130 131 134

Contents 8 Measurement of transpiration and leaf conductance Robert W. Pearcy, E.-Detlef Schulze and Reiner Zimmermann

8.1 8.2 8.3 8.4 8.5 8.6 8.7 9

Introduction Leaf transpiration rate Leaf conductance to water vapor Instrumentation for transpiration measurements Calibration of water vapor sensors Systems for measuring transpiration and leaf conductance Whole-plant measurements of transpiration

Plant water status, hydraulic resistance and capacitance Roger T. Koide, Robert H. Robichaux, Suzanne R. Morse and Celia M. Smith

9.1 9.2 9.3 9.4 9.5

Introduction Water potential and its components Water content Hydraulic resistance and capacitance Conclusion

10 Approaches to studying nutrient uptake, use and loss in plants F. Stuart Chapin, III and Keith Van Cleve

10.1 10.2 10.3 10.4 10.5

Introduction Nutrient uptake Nutrient use and nutrient status Chemical analysis Nutrient loss

11 Photosynthesis: principles and field techniques Christopher B. Field, J. Timothy Ball and Joseph A. Berry

11.1 11.2 11.3 11.4 11.5 11.6 11.7 11.8 12

The system concept Principles of photosynthesis measurement Components of gas-exchange systems Real photosynthesis systems Matching instrument to objective Calibrating photosynthesis systems Calculating gas-exchange parameters List of symbols

Crassulacean acid metabolism

vii 137

137 137 139 143 147 148 153 161

161 161 174 174 178 185

185 185 195 200 202 209

209 210 215 231 238 240 244 248 255

C. Barry Osmond, William W. Adams III and Stanley D. Smith

12.1 12.2 12.3 12.4

Introduction Measurement of succulence Nocturnal acidification Nocturnal CO 2 fixation

255 256 257 258

Contents

viii

12.5 Analysis of day-night and seasonal patterns of CO 2 and H 2 0 vapor exchange 12.6 Measurement of photosynthesis and respiration by O 2 exchange 12.7 Water relations 12.8 Stress physiology

13 Stable isotopes James R. Ehleringer and C. Barry Osmond 13.1 13.2 13.3 13.4 13.5 13.6

Introduction Natural abundances of stable isotopes of ecological interest Stable isotope mass spectrometry Sample preparation Sample variability Application of stable isotopes in ecological studies

14 Canopy structure John M. Norman and Gaylon S. Campbell 14.1 14.2 14.3 14.4 14.5

Introduction Direct methods Semidirect methods Indirect methods Summary

15 Growth, carbon allocation and cost of plant tissues Nona R. Chiariello, Harold A. Mooney and Kimberlyn Williams 15.1 15.2 15.3 15.4

Introduction Growth analysis Fate of carbon Carbon and energy costs of growth and maintenance

16 Root systems Martyn M. Caldwell and Ross A. Virginia 16.1 Introduction 16.2 Assessing root system structure and biomass in the field - determining what is there 16.3 Determination of root length and surface area 16.4 Microscale distributions of roots 16.5 Root system turnover and production 16.6 Root phenology and growth 16.7 Root system function 16.8 Root associations 16.9 Conduding thoughts

260 264 267 269

281 281 281 282 285 289 290

301 301 303 308 311 323

327 327 328 345 350

367 367 367 371 372 373 376 379

390

392

Contents 17 Field methods used for air pollution research with plants William E. Winner and Carol S. Greitner 17.1 17.2 17.3 17.4 17.5 17.6

Introduction Studies of air pollution absorption Air pollution instrumentation Cuvettes Field fumigation systems and approaches Summary

ix 399

399 400 403 407 412 422

Appendix

427

Index

441

Contribu tors

William W. Adams III

Department of Environmental, Population and Organismic Biology, University of Colorado, Boulder, CO, USA

J. Timothy Ball

Biological Sciences Center, Desert Research Institute, University of Nevada, Reno, NV, USA

Joseph A. Berry

Department of Plant Biology, Carnegie Institution of Washington, Stanford, CA, USA

Dan Binkley

Department of Forest and Wood Science, Colorado State University, Fort Collins, CO, USA

Arnold J. Bloom

Department of Vegetable Crops, University of California, Davis, CA, USA

Martyn M. Caldwell

Department of Range Science and the Ecology Center, Utah State University, Logan, UT, USA

Gaylon S. Campbell

Department of Agronomy and Soils, Washington State University, Pullman, Washington, USA

F. Stuart Chapin III

Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska, USA

Nona R. Chiariello

Department of Biological Sciences, Stanford University, Stanford, CA, USA

Contributors

xii James R. Ehleringer

Department of Biology, University of Utah, Salt Lake City, UT, USA

Christopher B. Field

Department of Plant Biology, Carnegie Institution of Washington, Stanford, CA, USA

John Grace

Department of Forestry and Natural Resources, University of Edinburgh, Edinburgh, Scotland

Carol S. Greitner

Department of General Science, Oregon State University, Corvallis, OR, USA

Wesley M. Jarrell

Department of Environmental Sciences and Engineering, Oregon Graduate Center, Beverton, OR, USA

Roger T. Koide

Department of Biology, Pennsylvania State University, University Park, PA, USA

Harold A. Mooney


Suzanne R. Morse

Department of Botany, University of California, Berkeley, CA, USA

John M. Norman

Department of Soils Science, University of Wisconsin, Madison, WI, USA

C. Barry Osmond

Department of Botany, Duke University, Durham, NC, USA

Robert W. Pearcy

Department of Botany, University of California, Davis, CA, USA

Robert H. Robichaux

Department of Ecology and Evolutionary Biology, University of Arizona, Tuczon, AZ, USA

Philip W. Rundel

Laboratory of Biomedical and Environmental Sciences and Department of Biology, University of California, Los Angeles, CA, USA

Contributors E.-Detlef Schulze

Lehrstuhl fUr Pflanzen6kologie, Universitat Bayreuth, 0-8700 Bayreuth, Federal Republic of Germany

Celia M. Smith

Department of Botany, Smithsonian Institution of Washington, Washington DC, USA

Stanley D. Smith

Department of Biological Sciences, University of Nevada, Las Vegas, NV, USA

Keith Van Cleve

Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska, USA

Ross A. Virginia

Biology Department and Systems Ecology Research Group, San Diego State University, San Diego, CA, USA

Peter Vitousek


Kimberlyn Williams


William E. Winner

Department of General Science, Oregon State University, Corvallis, OR, USA

Reiner Zimmermann

Lehrstuhl fur Pflanzenokologie, Universitat Bayreuth, 0-8700 Bayreuth, Federal Republic of Germany

xiii

Introd uction

Physiological plant ecology is primarily concerned with the function and performance of plants in their environment. Within this broad focus, attempts are made on one hand to understand the underlying physiological, biochemical and molecular attributes of plants with respect to performance under the constraints imposed by the environment. On the other hand physiological ecology is also concerned with a more synthetic view which attempts to understand the distribution and success of plants measured in terms of the factors that promote long-term survival and reproduction in the environment. These concerns are not mutually exclusive but rather represent a continuum of research approaches. Osmond et al. (1980) have elegantly pointed this out in a space-time scale showing that the concerns of physiological ecology range from biochemical and organelle-scale events with time constants of a second or minutes to succession and evolutionary-scale events involving communities and ecosystems and thousands, if not millions, of years. The focus of physiological ecology is typically at the single leaf or root system level extending up to the whole plant. The time scale is on the order of minutes to a year. The activities of individual physiological ecologists extend in one direction or the other, but few if any are directly concerned with the whole space-time scale. In their work, however, they must be cognizant both of the underlying mechanisms as well as the consequences to ecological and evolutionary processes.

IMPORTANCE OF FIELD METHODS TO PHYSIOLOGICAL ECOLOGY Field research in physiological ecology is heavily dependent on appropriate methods and instrumentation. The writings of early plant geographers and ecologists, such as Schimper (1898), Warming (1896) and Clements (1907), were rich with hypotheses and speculations concerning the role of environment, physiology and morphology in the distributions of plants. Indeed as early as the 1880s attempts were made to measure physiological difference (transpiration) between populations at different elevations (Bonnier, 1890). The theory for water vapor and heat transfer from leaves, including the concept of stomatal resistance, was published in 1900 by Brown and Escombe and porometers

xvi

Introduction capable of providing at least a relative measure of stomatal aperture were first used shortly thereafter (Darwin and Pertz, 1911). The Carnegie Institution of Washington's Desert Research Laboratory in Tucson from 1905 to 1927 was the first effort by plant physiologists and ecologists to conduct team research on the water relations of desert plants. Measurements by Stocker in the North African deserts and Indonesia (Stocker, 1928, 1935) and by Lundegardh (1922) in forest understories were pioneering attempts to understand the environmental controls on photosynthesis in the field. While these early physiological ecologists were keen observers and often posed hypotheses still relevant today they were strongly limited by the methods and technologies available to them. Their measurements provided only rough approximations of the actual plant responses. The available laboratory equipment was either unsuited or much more difficult to operate under field than laboratory conditions. Laboratory physiologists distrusted the results and ecologists were largely not persuaded of its relevance. Consequently, it was not until the 1950s and 1960s that physiological ecology began its current resurgence. While the reasons for this are complicated, the development and application of more sophisticated instruments such as the infrared gas analyzer played a major role. In addition, the development of micrometeorology led to new methods of characterizing the plant environments. However, most of the instruments such as the infrared gas analyzer and recorders were primarily designed for industrial process control applications and required considerable ingenuity on the part of the physiological ecologist to adapt them for use in the field. There were no instruments manufactured specifically forecophysiological applications. Physiological ecologists had to be intimately involved with developing instrumentation systems, and be familiar with techniques from many fields, including meteorology, physiology and soil science. The last 15 years have brought about remarkable changes in the instrumentation available for physiological ecology. This has been in part driven by the electronics revolution that has allowed the development of highly sophisticated 'smart' instruments with microprocessors that control their performance and low-power circuits that allow operation on batteries in the field. In addition, the notebook and analog recorder have been largely replaced by data acquisition systems and solid-state memory, making at least the chore of collecting the numbers less demanding. However, at least as important is the increased demand for ecophysiological and environmental measurements. This has given rise to the development of commercially available instruments specifically designed for ecophysiological measurements and environmental monitoring in the field. For many research problems there is no longer a need for the physiological ecologists to design and assemble the instrumentation systems since commercially available systems function so well. Physiological ecologists have and will continue to play an important role in this development, and the strong cooperation between researchers and instrument companies that has developed in the past few years will lead to further progress. The increased availability of instruments, however, is a two-edged sword. Physiological ecologists who assembled their gas exchange system were familiar with its limitations. Those who purchase instrumentation systems may not be. It

Introduction is the responsibility of the investigator to become familiar with the system, its components and their limitations.

METHODOLOGICAL BOOKS FOR PHYSIOLOGICAL ECOLOGY Since 1960 many important books covering instrumentation of use in physiological ecology have appeared. Environmental measurements were covered by Slatyer and McIlroy (1961), Tanner (1963), Platt and Griffiths (1964) and Fritschen and Gay (1979). Tanner's volume was available only in mimeographed form but was widely utilized because of the practical information it contained on constructing instruments, including such details as the properties of solders and paints. Similarly, Slatyer and McIlroy had only a limited printing but also had enormous impact because of the practical information it provided. The first treatise covering methods of plant ecophysiology was Methodology of Plant Ecophysiology. Proceedings of the Montpellier Symposium edited by F.E. Eckhardt (1965). This volume contained probably the first detailed descriptions of the application of infrared gas analyzers in the field for measuring photosynthesis. Plant Photosynthetic Production, Manual of Methods by Sestak, Catsky and Jarvis followed in 1971 and served nearly as a bible for ecophysiologists measuring gas exchange of plants. Although it is clearly dated, the book still contains much valuable information. More recently, several books covering methods have appeared. Instrumentation for Environmental Physiology edited by Marshall and Woodward (1985) had the specific objective of covering developments in instrumentation since 1971. Techniques in Bioproductivity and Photosynthesis edited by Coombs, Hall, Long and Scurlock (1985) covers a wide array of techniques ranging from growth analysis to enzymology.

SCOPE OF THIS BOOK This book is intended as a guide to the methodology appropriate for field measurements in physiological ecology, including both measurements of the environment and of the physiological and morphological responses of the plants. In some cases, for example stable isotope measurements (Chapter 13), the measurement is done in the laboratory but on samples collected under field conditions and with the primary objective of understanding the ecological behavior of plants in the field. Other chapters, especially those on soils (Chapter 5) and growth and allocation (Chapter 15), similarly cover analytical techniques on field samples. There has been an attempt to avoid an exhaustive review of all techniques but rather to concentrate on those that have proved most useful to the authors. These techniques provide the basis of evaluating the acquisition of resources (e.g. carbon, light, water, nutrients) and the use of these'resources for plant growth. Methods for estimating some of the costs of allocation of resources to particular functions are also covered (Chapter 15). In addition, the methodological framework for extending from organ level (leaf, root) measurements to whole plants is provided in chapters on root systems (Chapter 16) and

xvii

Introduction

xviii

canopy structure (Chapter 14) and to ecosystem level processes (Chapter 13). There are many techniques not included in this book that might rightly be an important component of a research program in physiological ecology. The focus is on performance of plants in the field and therefore instrumentation systems and facilities such as growth chambers and phytotrons are not covered. We have also not covered the various biochemically based techniques such as analysis of leaf anatomy, carboxylation enzyme activities, chlorophyll content, electrontransport activities, hormones, or for that matter, root enzyme activities involved in such processes as flooding tolerance. These techniques can clearly be applied to leaf samples collected in the field but are more commonly conducted on greenhouse or growth chamber plants. Methods for biochemical analysis of the photosynthetic apparatus are given in Hipkins and Baker (1986) and Coombs et al. (1985) and there are many sources for general information such as the series Methods in Enzymology. Problems of experimental design, statistics, etc. are also not covered for reasons of space. We have also not covered the analytical framework necessary for much research in physiological ecology such as simulation models or cost-benefit analyses. Jones (1983) discusses the important role of models and their interrelationship with experiments in physiological ecology and Loomis et al. (1979) provides a discussion of modeling approaches. Biochemically based photosynthetic models such as the one of Farquhar and his colleagues (Farquhar et al., 1980; Farquhar and von Caemmerer, 1982; Farquhar and Sharkey, 1982) have indeed played an important role in linking biochemical events to an understanding of whole-leaf gas exchange. More empirical models have similarly been useful in understanding, for example, limitations on annual carbon gain in specific habitats (e.g. Schulze et al., 1976). Cost-benefit analyses also provide an important analytical framework for understanding plant function and adaptation. The reader is referred to Bloom et al. (1985), Givnish (1986) and Gutschick (1987) for thorough discussions of this approach.

REFERENCES Bloom, A.J., Chapin, F.S. and Mooney, H.A. (1985) Resource limitation in plants - an economic analogy. Ann. Rev. Eco/. System., 16, 363-92. Bonnier, C. (1890) Influence des hautes sur les fonctions des vegetaux. Compt. Rend., Ill, 377-80. Brown, H. and Escombe, F. (1900) Static diffusion of gases and liquids in relation the assimilation of carbon and translocation in plants. Philos. Trans. R. Soc. Ser. B, 193, 223-91. Clements, F.E. (1907) Plant Physiology and Ecology, Holt, New York. Coombs, J., Hall, D.O., Long, S.P. and Scurlock, J,M.O. (eds) (1985) Techniques in Bioproductivity and Photosynthesis, 2nd edn, Pergamon, Oxford. Darwin, F. and Pertz, D.F.M. (1911) On a new method of estimating the aperture of stomata. Proc. R. Soc., Ser. B, 84, 136-54. Eckhardt, F.E. (ed.) (1965) Methodology of Plant Ecophysiology. Proceedings of the Montpellier Symposium, UNESCO, Paris. Farquhar, C.D. and von Caemmerer, S. (1982) Modelling of photosynthetic response to environmental conditions. In Encyclopedia of Plant Physiology, new series, vol. 12b,

Introduction Physiological Plant Ecology (eds O.L. Lange, p.s. Nobel, CB. Osmond and H. Ziegler), Springer-Verlag, Heidelberg, Berlin and New York, pp. 549-88. Farquhar, G.D., von Caemmerer, S. and Berry, J.A. (1980) A biochemical model tor photosynthetic CO 2 assimilation in leaves of C3 species. Planta, 119, 78-90. Farquhar, G.D. and Sharkey, T.D. (1982) Stomatal conductance and photosynthesis. Annu. Rev. Plant Physiol., 33, 317-45. Fritschen, L.J. and Gay, L.W. (1979) Environmental Instrumentation, Springer-Verlag, Heidelberg. Givnish, T.J. (ed.) (1986) On the Economy of Plant Form and Function, Cambridge University Press, Cambridge. Gutschick, V.P. (1987) A Functional Biology of Crop Plants, Croom Helm, London. Hipkins, M.F. and Baker, N.R. (1986) Photosynthesis Energy Transduction a Practical Approach, IRL Press, Oxford. Jones, H.G. (1983) Plants and Microclimate, Cambridge University Press, Cambridge. Loomis, RS., Rabbinge, Rand Ng, E. (1979) Explanatory models in crop physiology. Annu. Rev. Plant. Physiol., 30, 339-67. Lundegardh, H. (1922) Zur physiologie und okologie der kohnlensaurassimilation. BioI. 2bl., 42, 337-58. Marshall, B. and Woodward, F.I. (eds) (1985) Instrumentation for Environmental Physiology, Cambridge University Press, Cambridge. Osmond, CB., Bjorkman, O. and Anderson, D.J. (1980) Physiological Processes in Plant Ecology: Towards a Synthesis with Atriplex, Springer-Verlag, Berlin. Platt, R. B. and Griffiths, J .F. (1964) Environmental Measurement and Interpretation, Reinhold, New York. Schimper, A.F.W. (1898) Pflanzen-Geographie auf physiologischer Grundlage, Fischer, Jena. Schulze, E.D., Lange, O.L., Evenari, M., Kappen, L. and Buschbom, U. (1976) An empirical model of net photosynthesis for the desert plant Hammada scoparia (Pomel) Iljin. I. Description and test of the model. Oecologia, 22, 355-72. Sestak, Z., Catsky, J. and Jarvis, P.G. (eds) (1971) Plant Photosynthetic Production, Manual of Methods, W. Junk, The Hague. Slatyer, RO. and McIlroy, I.C (1961) Practical Microclimatology, CS.I.RO. Plant Industry Division, Canberra. Stocker, O. (1928) Der wasserhaushalt agyptischer wusten-und salzpflanzen von standpunkt einer experimentellen un vergleichenden pflangeographie aus. Bot. Abh, 13, 1-200.

Stocker, O. (1935) Assimilation und atmung westjavanischer tropenbaume. Planta, 24, 402-45.

Tanner, CB. (1963) Basic Instrumentation and Measurements for Plant Environment and Micrometeorology, University of Wisconsin Soils Bulletin 6. Warming, E. (1896) Lehrbuch der okologischen Pflanzengeographie. Eine Einfuhrung in die Kenntnis der Pflanzenvereille, Borntraeger, Berlin.

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