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.), 1941-
581.1
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.), 1941-
QK 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
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retrieval system, without written permission from the copyright owner.
 Contents

Contributors xi

Introduction xv

1 Principles of instrumentation for physiological ecology 1

Arnold J. Bloom
1.1 Introduction 1
1.2 Measurement and measurement errors 1
1.3 Instrument organization 3
1.4 Instrument initiation 11
1.5 Postscript 12

2 Field data acquisition 15

Robert W. Pearcy
2.1 Introduction 15
2.2 Analog recorders 15
2.3 Digital recorders 17
2.4 Integrators 24
2.5 Sampling considerations 24

3 Water in the environment 29

Philip W. Rundel and Wesley M. Jarrell
3.1 Soil moisture 29
3.2 Atmospheric moisture 41
3.3 Moisture flux 47

4 Measurement of wind speed near vegetation 57

John Grace
4.1 Introduction 57
4.2 Flow in wind tunnels, growth cabinets and ducts 58
4.3 Weather stations and field survey 60
4.4 Wind profiles above vegetation 61
4.5 Boundary layer resistance 64
vi Contents
4.6 Calibration 66
4.7 Aerodynamic influence by masts 66
4.8 Visualization 66
4.9 Pressure measurements 66
4.10 Some applications 68

5 Soil nutrient availability 75

Dan Binkley and Peter Vitousek
5.1 Introduction 75
5.2 Difficulties in measuring nutrient availability 76
5.3 Nitrogen availability 76
5.4 Phosphorus availability 81
5.5 Sulfur availability 85
5.6 Availability of essential cations 85
5.7 Micronutrient availability 87
5.8 Soil classification 88
5.9 Bioassay of nutrient availability 88
5.10 Soil acidity 88
5.11 Soil salinity 90
5.12 Soil redox potential 90
5.13 Comments on sampling 90
5.14 Index units 92

6 Radiation and light measurements 97

Robert W. Pearcy
6.1 Introduction 97
6.2 Definitions and units 97
6.3 Energy versus photons as a measure of PAR 99
6.4 Radiation sensors: general characteristics 100
6.5 Determination of the diffuse and direct components
of radiation 107
6.6 Calibration of radiation sensors 107
6.7 Sampling considerations 108
6.8 Photographic estimations of light climate 109
6.9 Spectral radiometry 113

7 Temperature and energy budgets 117

James R. Ehleringer
7.1 Introduction 117
7.2 Energy budget approach 117
7.3 Variations in air and leaf temperatures with height 123
7.4 Temperature and its measurement 124
7.5 Orientation and its measurement 130
7.6 Calculation of incident solar radiation on different surfaces 130
7.7 Leaf absorptance and its measurement 131
7.8 Boundary layer considerations 134
 Contents vii

8 Measurement of transpiration and leaf conductance 137

Robert W. Pearcy, E.-Detlef Schulze and Reiner Zimmermann
8.1 Introduction 137
8.2 Leaf transpiration rate 137
8.3 Leaf conductance to water vapor 139
8.4 Instrumentation for transpiration measurements 143
8.5 Calibration of water vapor sensors 147
8.6 Systems for measuring transpiration and leaf conductance 148
8.7 Whole-plant measurements of transpiration 153

9 Plant water status, hydraulic resistance and capacitance 161

Roger T. Koide, Robert H. Robichaux, Suzanne R. Morse
and Celia M. Smith
9.1 Introduction 161
9.2 Water potential and its components 161
9.3 Water content 174
9.4 Hydraulic resistance and capacitance 174
9.5 Conclusion 178

10 Approaches to studying nutrient uptake, use and loss in plants 185

F. Stuart Chapin, III and Keith Van Cleve
10.1 Introduction 185
10.2 Nutrient uptake 185
10.3 Nutrient use and nutrient status 195
10.4 Chemical analysis 200
10.5 Nutrient loss 202

11 Photosynthesis: principles and field techniques 209

Christopher B. Field, J. Timothy Ball and Joseph A. Berry
11.1 The system concept 209
11.2 Principles of photosynthesis measurement 210
11.3 Components of gas-exchange systems 215
11.4 Real photosynthesis systems 231
11.5 Matching instrument to objective 238
11.6 Calibrating photosynthesis systems 240
11.7 Calculating gas-exchange parameters 244
11.8 List of symbols 248

12 Crassulacean acid metabolism 255

C. Barry Osmond, William W. Adams III and Stanley D. Smith
12.1 Introduction 255
12.2 Measurement of succulence 256
12.3 Nocturnal acidification 257
12.4 Nocturnal CO 2 fixation 258
viii Contents
12.5 Analysis of day-night and seasonal patterns of CO 2
and H 2 0 vapor exchange 260
12.6 Measurement of photosynthesis and respiration
by O 2 exchange 264
12.7 Water relations 267
12.8 Stress physiology 269

13 Stable isotopes 281

James R. Ehleringer and C. Barry Osmond
13.1 Introduction 281
13.2 Natural abundances of stable isotopes of ecological interest 281
13.3 Stable isotope mass spectrometry 282
13.4 Sample preparation 285
13.5 Sample variability 289
13.6 Application of stable isotopes in ecological studies 290

14 Canopy structure 301

John M. Norman and Gaylon S. Campbell
14.1 Introduction 301
14.2 Direct methods 303
14.3 Semidirect methods 308
14.4 Indirect methods 311
14.5 Summary 323

15 Growth, carbon allocation and cost of plant tissues 327

Nona R. Chiariello, Harold A. Mooney and Kimberlyn Williams
15.1 Introduction 327
15.2 Growth analysis 328
15.3 Fate of carbon 345
15.4 Carbon and energy costs of growth and maintenance 350

16 Root systems 367

Martyn M. Caldwell and Ross A. Virginia
16.1 Introduction 367
16.2 Assessing root system structure and biomass in the field
- determining what is there 367
16.3 Determination of root length and surface area 371
16.4 Microscale distributions of roots 372
16.5 Root system turnover and production 373
16.6 Root phenology and growth 376
16.7 Root system function 379
16.8 Root associations 390
16.9 Conduding thoughts 392
 Contents ix

17 Field methods used for air pollution research with plants 399
William E. Winner and Carol S. Greitner
17.1 Introduction 399
17.2 Studies of air pollution absorption 400
17.3 Air pollution instrumentation 403
17.4 Cuvettes 407
17.5 Field fumigation systems and approaches 412
17.6 Summary 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
xii Contributors
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 Department of Biological Sciences,
Stanford University,
Stanford, CA, USA
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 xiii

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 Department of Biological Sciences,
Stanford University,
Stanford, CA, USA
Kimberlyn Williams Department of Biological Sciences,
Stanford University,
Stanford, CA, USA
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
 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 under-
stand 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 equip-
ment 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 instrument-
ation 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 xvii

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 physio-

logical 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 ecophysio-
logists 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 develop-
ments in instrumentation since 1971. Techniques in Bioproductivity and Photosyn-
thesis 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) measure-
ments to whole plants is provided in chapters on root systems (Chapter 16) and
xviii Introduction
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, electron-
transport 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.

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 Introduction xix
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