Table Of ContentSTRESS PHYSIOLOGY AND FOREST PRODUCTIVITY
FORESTRY SCIENCES
Baas P, ed: New Perspectives in Wood Anatomy. 1982. ISBN 90-247-2526-7
Prins CFL, ed: Production, Marketing and Use of Finger-Jointed Sawnwood. 1982.
ISBN 90-247-2569-0
Oldeman RAA, et a!., eds:, Tropical Hardwood Utilization: Practice and Prospects. 1982.
ISBN 90-247-2581-X
Den Ouden P and Boom BK: Manual of Cultivated Conifers: Hardy in Cold and Warm-
Temperate Zone. 1982. ISBN 90-247-2148-2
Bonga JM and Durzan DJ, eds: Tissue Culture in Forestry. 1982. ISBN 90-247-2660-3
Satoo T and Magwick HAl: Forest Biomass. 1982. ISBN 90-247-2710-3
Van Nao T, ed: Forest Fire Prevention and Control. 1982. ISBN 90-247-3050-3
Douglas J: A Re-appraisal of Forestry Development in Developing Countries. 1983.
ISBN 90-247-2830-4
Gordon JC and Wheeler CT, eds: Biological Nitrogen Fixation in Forest Ecosystems:
Foundations and Applications. 1983. ISBN 90-247-2849-5
Nemeth MV: The Virus-Mycoplasma and Rikettsia Disease of Fruit Trees.
ISBN 90-247-2868-1
Duryea ML and Landis TD, eds: Forest Nursery Manual: Production of Bareroot Seed
lings. 1984. ISBN 90-247-2913-0
Hummel FC, ed: Forest Policy: A Contribution to Resource Development. 1984.
ISBN 90-247-2883-5
Manion PD, ed: Scleroderris Canker of Conifers. 1984. ISBN 90-247-2912-2
Duryea ML and Brown GN, eds: Seedling Physiology and Reforestation Success. 1984.
ISBN 90-247-2949-1
Staaf KAG and Wiksten NA: Tree Harvesting Techniques. 1984. ISBN 90-247-2994-7
Boyd JD: Biophysical Control of Microfibril Orientation in Plant Cell Walls. 1985.
ISBN 90-247-3101-1
Findlay WPK, ed: Preservation of Timber in the Tropics. 1985. ISBN 90-247-3112-7
Samset I: Winch and Cable Systems. 1985. ISBN 90-247-3205-0
Leary RA: Interaction Theory in Forest Ecology and Management. 1985.
ISBN 90-247-3220-4
Gessel SP: Forest Site and Productivity. 1986. ISBN 90-247-3284-0
Hennessey TC, Dougherty PM, Kossuth SV and Johnson JD, eds: Stress Physiology and
Forest Productivity. 1986. ISBN 90-247-3359-6
Stress physiology and
forest productivity
Proceedings of the Physiology Working Group Technical Session. Society of
American Foresters National Convention, Fort Collins, Colorado, USA,
July 28-31, 1985
edited by
THOMAS C. HENNESSEY
Oklahoma State University, Stillwater, Oklahoma, USA
PHILLIP M. DOUGHERTY
University oj Georgia, Athens, Georgia, USA
SUSAN V. KOSSUTH
USDA Forest Service, Gainesville, Florida, USA
JON D. JOHNSON
University oj Florida, Gainesville, Florida, USA
1986 MARTINUS NIJHOFF PUBLISHERS
a member of the KLUWER ACADEMIC PUBLISHERS GROUP
DORDRECHT / BOSTON / LANCASTER
Distributors
for the United States and Canada: Kluwer Academic Publishers, 190 Old Derby
Street, Hingham, MA 02043, USA
for the UK and Ireland: Kluwer Academic Publishers, MTP Press Limited,
Falcon House, Queen Square, Lancaster LAI lRN, UK
for all other countries: Kluwer Academic Publishers Group, Distribution Center,
P.O. Box 322, 3300 AH Dordrecht, The Netherlands
Library of Congress Cataloging in Publication Data
Stress physiology and forest productivity.
(Forestry sciences ; v.
Bibliography: p.
1. Trees--Physiology--Congresses. 2. Plants, Effect
of stress on--Congresses. 3. Forest ecology--Congresses.
4. Forest management--Congresses. 5. Forest productivity
--Congresses. I. Hennessey, Thomas C. II. Society of
American Foresters. Physiology Working Group.
III. Society of American Foresters. Convention (1985
Fort Collins, Colo.)
SD395.S77 1986 634.9'6 86-8644
ISBN-13: 978-94-010-8469-7 e-ISBN-13: 978-94-009-4424-4
DOl: 10.1007/978-94-009-4424-4
SAF 86-04
Copyright
© 1986 by Martinus Nijhoff Publishers, Dordrecht.
Softcover reprint of the hardcover 1s t edition 1986
All rights reserved. No part of this publication may be reproduced, stored in a
retrieval system, or transmitted in any form or by any means, mechanical,
photocopying, recording, or otherwise, without the prior written permission of
the publishers,
Martinus Nijhoff Publishers, P.O. Box 163, 3300 AD Dordrecht,
The Netherlands.
v
CONTENTS
PREFACE . . . . . .......................... VII
1. INTRODUCTION
S. G. Pallardy . 1
2. MOISTURE: EFFECTS OF WATER STRESS ON TREES
R. O. Teskey and T. M. Hinckley .... 9
3. MOISTURE STRESS MANAGEMENT: SILVICULTURE AND GENETICS
R. J. Newton, C. E. Meier, J. P. Van Buijtenen, and
C. R. Mc Kinley .•.••...•....•..•. . . . . . . 35
4. NUTRIENTS: USE OF FOREST FERTILIZATION AND NUTRIENT EFFICIENT GENOTYPES
TO MANAGE NUTRIENT STRESS IN CONIFER STANDS
J. L. Troth, R. G. Campbell, and H. L. Allen •••..•.. . . . . . . 61
5. FOREST PESTS: INFLUENCE OF FOREST MANAGEMENT PRACTICES ON PEST
POPULATION DYNAMICS AND FOREST PRODUCITVITY
T. Evan Nebeker, D. R. Hous ton, and J. D. Hodges • . . • • • · ...... 101
6. FOREST PESTS: THE ROLE OF PHLOEM OSMOTIC ADJUSTMENT IN THE DEFENSIVE
RESPONSE OF CONIFERS TO BARK BEETLE ATTACK
P. J. H. Sharpe, R. J. Newton, and R. D. Spence . . • . •• •. ....... 113
7. LOW TEMPERATURE: PHYSICAL ASPECTS OF FREEZING IN WOODY PLANT XYLEM
M. F. George and M. J. Burke. • • • . • . • • • • • • • • .. . ....... 133
8. MULTIPLE STRESS FACTORS: THE POTENTIAL ROLE OF SYSTEM MODELS IN
ASSESSING THE IMPACT OF MULTIPLE STRESSES ON FOREST PRODUCTIVITY
R. L. Graham, T. R. Fox, and P. M. Dougherty .... . . . . . . . . 151
9. FIRE: ITS EFFECTS ON GROWTH AND PHYSIOLOGICAL PROCESSES IN CONIFER
FORESTS
J. L. Chambers, P. M. Dougherty, and T. C. Hennessey · ...... 171
10. HERBICIDE STRESS: USE OF BIOTECHNOLOGY TO CONFER HERBICIDE RESISTANCE
TO SELECTED WOODY PLANTS
N. D. Nelson and B. E. Haissig .••...•••••••.• · . . . . . . 191
11. AIR POLLUTION: SYNTHESIS OF THE ROLE OF MAJOR AIR POLLUTANTS IN
DETERMINING FOREST HEALTH AND PRODUCTIVITY
1. J. Fernandez . • • • . . • . . . • • • . • . . . • • • . . . . . . . . . . 217
~I
PREFACE
Maintaining or increasing stand productivity is the
concern of forest land managers worldwide. Consequently,
there is increasing interest in understanding the impact of
environmental stress on productivity and the development of
management strategies that ameliorate or reduce the
deleterious effects.
Invited scientists gathered in Fort Collins, Colorado
on July 30, 1985, to present the current state of knowledge
regarding the impact of environmental stress on forest stand
productivity. Particular attention was given to elucidating
the mode of action by which individual stress elements
reduce productivity. Environmental factors and the levels
that constitute stressed (suboptimal) conditions in forest
stands were identified, and the effects of stress intensity
and duration on key stand parameters, including
photosynthesis, respiration, assimilate partitioning,
senescence and mortality, were emphasized.
The role of genetics and silvicultural treatments in
lessening the stress impact on stand productivity was
presented, particularly in regards to alternative methods
for environmental stress management. Modeling of stand
dynamics in response to environmental stress was explored as
an effective research and management tool.
VIII
Improved forest management practices will develop as we
improve our understanding of the nature of important
environmental stresses and as we comprehend their impact on
tree and stand performance, manifested through physiological
processes and genetic potential. This book is dedicated to
such an understanding and comprehension.
ThomM C. He.nnu-6e.y
Phittip M. Vough~y
SMa./'!. V. KOMuth
Jon V. John!.:,on
1. INTRODUCTORY OVERVIEW
S. G. PALLARDY
Associate Professor, School of Forestry, Fisheries, and
Wildlife, University of Missouri, Columbia, Missouri 65211
ABSTRACT
Environmental stresses, both biotic and abiotic,
commonly reduce plant growth below that which would be
indicated by genetic potential. Physiological processes of
plants integrate environment and heredity to control growth.
The responses of physiological processes to environmental
f~ctors are complex, but progress is being made in our
understanding of these responses and in the ability to pre
dict growth from knowledge of environmental modulation of
plant physiology. The most successful physiologically
oriented growth models have employed empirically determined
relationships between dominant environmental factors and
stand growth responses. The focus of this technical session
will center on research directed toward improving management
practices and growth modeling through a better understanding
of fundamental physiological relationships among environ
ment, hereditary potential of forest trees, and stand
growth.
1.1 INTRODUCTION
The environment presen~ed to forest stands, artifi
cially or naturally regenerated, managed or unmanaged,
commonly reduces growth and wood quality below the maximum
attainable level. This fact is not obvious because these
limitations are so pervasive that we have relatively few
Contribution of. the Missouri Agricultural Experiment
Station, Journal Series No. 9970
2
examples of unrestricted growth with which to compare the
norm. In this technical session, the nature of important
environmental limitations, how they influence stand perform
ance through effects on individual trees, and contemporary
research aimed at increasing forest stand growth and quality
by modifying both plant and environment will be discussed.
1.2 STRESS CONCEPTS
Although most people have an intuitive feeling for the
concept of stress in biology, exact definitions are elusive.
The majority of workers in stress research accept the
concepts, if not terminology, advanced by Levitt (6). In
Levitt's scheme, in which an analogy between biological and
physical stresses is drawn, stress is defined as "any envir
onmental factor capable of inducing a potentially injurious
strain in living organisms." The strain(s) induced are
physical and chemical responses within the organism that
eventually produce the suite of macroresponses we observe as
growth reduction, injury, or death. Continuing Levitt's
mechanical analogy, organisms generally exhibit a capacity
for elastic strain, where physiological responses are only
temporarily altered, but return to normal after the stress
is relieved, followed by plastic strain under greater levels
of stress, where permanent alteration (and often injury and
death) follow severe stress imposition and relief. The
point at which elastic strain turns to plastic strain is
called the "yield point" and it is important because it
marks the level of stress where severe and irreversible
effects begin. The yield point is not static but may change
with moderate stresses of longer duration, as is exemplified
by the process of osmotic adjustment (e.g., 8).
The use of the term "strain" for the physiological res
ponses of organisms to environmental factors has not been
widely adopted; if so, we would be talking about "strain"
management instead of stress management today, and as Paul
Kramer has pointed out (4), no one uses or is likely to use
the term~"plant water strain." In practice, stress is used
3
in a broad sense, describing both deleterious levels of
environmental factors and the condition of the plant.
Considerations of stress also include the influence of
biotic as well as abiotic factors in the plant environment.
Levitt's ideas do, however, provide a framework through
which one can understand the response of forests to
environmental influences. The forest scientist's and man
ager's jobs are to understand the nature of these elastic
and plastic responses and to work toward maintaining stands
under minimal "elastic strain."
1.3 ENVIRONMENTAL AND GENETIC INFLUENCES ON STAND GROWTH
The main objective of commodity forestry is to derive
raw materials by manipulating the natural growth process
through various management practices. In relating this
session to practical considerations, two questions are par
ticularly relevant: 1) What pattern of influence does the
environment (broadly defined) have on growth? and 2) How are
these environmental effects translated into observed changes
in growth? Justus von Liebig, arguably the original agric
ultural chemist, first addressed the former question when he
formulated his Law of the Minimum: "The growth of a plant is
dependent upon the amount of foodstuff [i.e., nutrients,
water, etc.] that is presented to it in minimum quantities."
A generalized response to an environmental factor is char
acterized by a region of direct response to the factor fol
lowed by a region of satiation or saturation (9). In the
early part of this century Blackman (1) developed this idea
further, demonstrating that the saturation level of a yield
factor can be increased by the addition of a second limiting
factor. These ideas provide a conceptual picture of the
influence of environmental factors (particularly nutrition
and light) on stand growth processes, but they by no means
cover all situations. For example, growth responses to
water availability would present an optimum response near
field capacity, with reduced growth at lower soil water
contents as water stress increases or at higher water
