Synthesis and Integration
of the
Forest Response Program:
The Path from Projects
to
Major Program Outputs
Synthesis and Integration Report Number 14


tea
ivimnmenta! Protection
Agency







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Synthesis and Integration
of the
Forest Response Program:
The Path from Projects
to
Major Program Outputs
Synthesis and Integration Report Number 14
A Report of the Synthesis and Integration Project
Ross Kiester, Project Leader
Authors: Jeff Brandt, Ross Kiester, Lew Ladd, Chas Peterson,
Greg Reams, Paul Schroeder, Doug Shadwick, Karl Stoszek, Bill
Warren
EPA Environmental Research Laboratory - Corvallis
200 SW 35th St.
Corvallis, OR 97333
Notice
This document is an internal report. It has neither been peer
reviewed nor approved by the U.S. Environmental Protection
Agency. It is being circulated for comment on technical merit
and policy implications. Do not release. Do not guote or cite.
February, 1988

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Table of Contents
1	Introduction: The Three Components of the Path 		1
2	The Projects and Project Deliverables 		2
3	Outlines for Each Major Program Output 		3
3.1	Major Program Output #1		4
3.2	Major Program Output #2 		6
3.3	Major Program Output #3 		8
3.4	Major Program Output #4		11
3.5	Major Program Output #5		14
3.6	Project Result Due Dates		14
4	Synthesis and Integration: How do FRP Projects Address the Policy
Questions? 		15
4.1	Scientific Questions 1.1 and 1.2: Changes and Patterns in
Forest Condition		15
4.1.1	Overall Structure 		15
4.1.2	General Strategy 		16
4.1.3	Specific Strategies 		16
4.1.3.1	Western Conifers 		16
4.1.3.2	Southern Pines		17
4.1.3.3	Northeastern Spruce-Fir Complex		17
4.1.3.4	Eastern Hardwoods Complex		18
4.2	Scientific Question 2.1: Soil Mediated Effects 		19
4.2.1	Introduction 		19
4.2.2	Research 		19
4.3	Scientific Question 2.2: Foliar Leaching 		21
4.3.1	Introduction 		21
4.3.2	Research 		21
4.4	Scientific Question 2.3: Carbon Allocation 		23
4.4.1	Introduction 		23
4.4.2	Research 		24
4.4.2.1	Relative sensitivity rankings 		24
4.4.2.2	A carbon allocation model 		25
4.4.2.3	Mechanisms of Damage 		25
4.5	Scientific Question 2.4: Winter Injury 		27
4.5.1	Introduction 		27
4.5.2	Research 		27
4.5.2.1	Current Winter Injury 		27
4.5.2.2	Influence of Air Pollutants on Winter Injury 		28
4.5.3	Mechanisms of Winter Injury 		28
4.6	Scientific Question 2.5: Reproduction/Regeneration 		28
4.6.1	Introduction 		28
4.6.2	Research 		28
4.7	Scientific Question 2.6: Pests/Pathogens 		29
4.7.1	Introduction 		29
4.7.2	Research 				29
4.8	Atmospheric Exposure Support 		30
4.8.1	Information Needs 		30
4.8.2	Available Data - The Data Book 		30
4.8.2.1	Geographic areas 		30
4.8.2.2	Time intervals 		31
4.8.2.3	Quality assurance 		31
4.8.2.4	Statistics 		31
4.8.3	Format 		32
5	The Overall Path to Major Program Output #4 		33
6	Specification for New Milestones for Individual Projects 		34
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7	Appendix I Categories of Projects in the Forest Response Program ... 35
8	Appendix II Forest Response Program Project Summaries 	 43
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1 Introduction: The Three Components of the Path
The purpose of this Report is to show in some detail how Synthesis and
Integration of the Forest Response Program will be achieved. For this purpose
we recognize three components of the Path (see Figure 1.1):
1.	The Projects and project deliverables
2.	The Major Program Outputs
3.	Synthesis & Integration Activities
The overall plan of this Report is to begin with the projects, describe
the Major Program Outputs (MPOs) as currently envisioned, and then discuss how
the projects relate to one another and the MPOs (S&I activities). The MPOs
are a critically important part of the FRP because they are the tangible means
by which the program will address the policy questions behind it. Addressing
those questions is the objective of the FRP.
The heart of the Report is the discussion of Synthesis and Integration
Activities which include both those of the Synthesis and Integration Project
itself as well as the work of the Research Cooperatives. This discussion is
an analysis of how the projects are coordinated to address the program's
Policy Questions and provide information for development of the MPOs.
Addressing the Policy Questions is the program's ultimate objective, and the
MPOs are the tangible vehicles for accomplishing that.
This Report builds on virtually all of the planning work in the Program
that has preceded it. Knowledge of the National Plan of the Forest Response
Program and of the Synthesis and Integration Plan is a prerequisite for
understanding this Report. Knowledge of the Cooperative plans is important as
well.
By the same token we would like to acknowledge all of those who have
participated in planning in the FRP. You will recognize that we have borrowed
heavily from your work, and we thank you for your previous efforts.
This Report should be treated as a dynamic object. Our intent has been
to create a tool, much like the Project Tracking System, which can be updated
and corrected. Please help us with this effort by sending us your comments and
corrections.
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Project
[MAJOR PROGRAM
1 OUTPUT #3
MAJOR PROGRAM
OUTPUT #4
MAJOR PROGRAM
OUTPUT #5
MAJOR PROGRAM
OUTPUT #2	
MAJOR PROGRAM
/ OUTPUT #1
Figure 1.1 The three components of the path

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2 The Projects and Project Deliverables
At the present time there are approximately 95 Projects within the FRP.
As a first step in Synthesis and Integration, we assign each Project to one or
more of twelve categories. Categorization is an organizational aid. Of
course, many Projects fall into more than one category. These categories,
together with the name of the Synthesis and Integration staff member who is
the primary contact for that category, are listed in Table 1. A listing of
the projects in each category appears in Appendix I. Projects may appear in
more than one category. A brief summary of Projects and their deliverables is
contained in Appendix II. Throughout the remainder of this report, as well as
in the Appendices, projects are referenced by principal investigator and
project number.
TABLE 1. CATEGORIZATION OF PROJECTS. Each project is assigned to one
or more of the following categories. Assignment of S&I staff
responsibility for each category is also given.
Policy Question 1
Is there a problem?
1.	Spatial Studies - Greg Reams
2.	Dendrochronology Data - Bill Warren
3.	FIA Data - Bill Warren
4.	Fixed Plots Data - Ross Kiester
Policy Question 2
What are the mechanisms?
5.	Soil Mediated Effects - Paul Schroeder
6.	Foliar Leaching - Jeff Brandt
7.	Carbon Allocation - Chas. Peterson
8.	Winter Injury - Jeff Brandt
9.	Reproduction and Regeneration - Chas. Peterson
10.	Pests and Pathogens - Greg Reams
Policy Question 3
What are the dose/response relationships?
11.	Seedling/Branch/Tree Integration - Ross Kiester
12.	Atmospheric Exposure - Jeff Brandt
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3 Outlines for Each Major Program Output
This section presents outlines of the reports that will be produced for
each of the five Major Program Outputs. Table 2 lists the MPOs. Please see
pages 5-8 of the Synthesis and Integration Plan for a description of these
outputs and their motivation. There is necessarily more detail for the earlier
MPOs than the later. In particular, MPO #5 will be described in more detail as
the modeling effort continues.
TABLE 2. The Major Program Outputs.
1.	Evaluation of the extent and magnitude of recent changes in forest
condition.
2.	Evaluation of the role of non-air pollution factors in growth
reduction and visible decline.
3.	Quantitative estimates of seedling responses to S, N, and
associated pollutants at ambient conditions.
4.	Evaluation of the roles of S, N, and associated pollutants in
forest decline.
5.	Projection of responses under alternative deposition scenarios.
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3.1 Major Program Output #1.
Extent and Magnitude of Recent Changes ("Trends")
S&I Contact: W. Warren.
1	Introduction
1.1 Goal for MPO #1
Major Program Output #1 is the natural lead into MPO #2. In MPO #1 we
investigate whether any of the case studies of the Forest Response Program
indicate trends in forest condition that may result from forest dynamics or
environmental factors (Sections 2.1 & 2.2 of MPO #2). MPO #1 thus
identifies case studies that indicate changes in forest condition and MPO
#2 then attempts to interpret the results of the various case studies as to
whether the trends in condition are the result of expected trends of forest
dynamics or from changing environmental factors and their interaction with
forest dynamics.
2	Trends in Condition: The Cases and Regions
The various projects can be classified in several ways. The most
meaningful appears to be by species of tree which, by and large, coincide
with research cooperative responsibility. Although there are commonalities
of approach over the four cooperatives, there are some notable
distinctions. Data sources can be classified as dendrochronological
studies, FIA plots, fixed plots and gradient studies. These categories are
by no means exclusive, for example some gradient studies have a
dendrochronological component, some dendrochronological studies are
associated with FIA plots as are some gradient studies, etc. The Western
Conifers Coop is dominated by dendrochronological data bases, whereas FIA
and fixed plot data are the major component of the Southern Commercial
Coop. The Spruce-Fir Coop has a number of broad surveys complemented by
intensive studies at certain sites. The Eastern Hardwood Coop is dominated
by gradient studies with a complement of intensive site studies with some
gradient studies.
2.1 Southern Pines
2.1.1	FIA data
(1)	McClure (VS07)
(2)	deSteiger/Ruark (VS04)
(3)	Saucier (VS08)
(4)	Dell (VS12)
2.1.2	Fixed Plot data
(1)	Dell (VS12)
(2)	Christiansen (SC10)
2.1.3	Dendrochronological data
(1) Saucier (VS08)
2.1.4	Regionalization
(1) Marx (VS01), with input from above studies
2.2 Spruce-Fir
2.2.1	Fixed Plot data
(1)	Johnson (SF08)
(2)	Zedacker (SF25)
(3)	Solomon (SF28)
2.2.2	Dendrochronological data
(1)	Hornig (SF05)
(2)	Hornbeck (VS06)
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2.2.3	Remote Sensing data
(1)	Rock (VS15)
(2)	Dull (SF26)
(3)	Ward (VS02)
2.2.4	Historical and Bibliographic data
(1) Wentworth (SF18)
2.2.5	Regionalization
2.3	Eastern Hardwoods
2.3.1	Fixed Plot/FIA data
(1)	Brooks (VS11)
(2)	Ohman (VS10)
2.3.2	Dendrochronological data
(1)	Brooks (VS11)
(2)	Ohman (VS10)
(3)	Davis (EH04)
(4)	Loucks (EH05)
2.3.3	Intensive Site data
(1)	Davis (EH04)
(2)	Witter (EH03)
(3)	Loucks (EH05)
2.3.4	Historical and Bibliographic data
(1) Millers (EH02)
2.3.5	Regionalization
(1)	Ohman (VS10)
(2)	Brooks (VS11)
2.4	Western Conifers
2.4.1	Fixed Plot data
(1) Graybill/Swetnam (WC24)
2.4.2	Dendrochronological data
(1)	Brubaker (WC25)
(2)	Graybill (WC24)
(3)	Peterson (WC26)
2.4.3	Regionalization
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3.2 Major Program Output #2
The role of non-air pollution factors in forest condition changes S&I
Contact: Karl Stoszek
1	Introduction
1.1	Goals
The goal of Major Program Output #2 (MPO #2) is to assess whether
changes in forest conditions, identified through MPO #1 efforts, are
explainable in terms of natural and management-induced causes.
1.2	Approach
Two sources of information are available. The first is information on
non-pollution factors of the forest environment collected as part of a
given forest trend study. This information will be utilized in evaluation
of forest condition trend-cases.
The second source of information is the general knowledge from
ecological sciences, soil sciences, forest sciences, and allied efforts in
acidic deposition research (e.g., EPRI, NCASI, and CSRS).
These general sources will serve a two-fold purpose:
First, for development of conceptual models of natural factor
interactions with tree and rhizosphere processes. These will be concept
(box and arrows) models patterned somewhat after both Fritts (1962) and
Manion (1981); the models will include mechanisms associated with
hypothesized causes of change.
Second, for development of a general scenario of forest development
dynamics in which effects of multiple factor interactions on tree
regeneration, growth, and mortality will be portrayed (see Figure 3.1).
Envisioned is a scenario of the successional development dynamics of a
subalpine forest, and a scenario of developmental dynamics of an
intensively managed plantation. In the scenarios, we will generalize the
possible effects of major natural and management-induced factors on the
systems' phytomass productive capacity, tree species habitat suitability,
and resilience to disturbances.
Our general strategy is to use the case information in a sort of
"results" section and the general information in both "background" and
"discussion" sections.
2	Forest and Natural Factor Interactions
This section develops a common framework for the specific and general
analysis of natural factor effects. Environmental factors are legion, only
the most important will be analyzed in terms of effects on tree and
rhizosphere processes, and forest developmental dynamics.
2.1	Physical and Biotic Factors in Tree and Rhizosphere Processes
2.1.1	Climatic
2.1.2	Fire
2.1.3	Nutrients and Related Soil Chemistry and Biota
2.1.4	Pest and Pathogens
2.1.5	Management Practices
2.2	Multiple Stress Factors in Forest Development Dynamics
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Population
Parameters
• Reproduction
& Regeneration
Growth
• Mortality
Major
Program
Output §2
Use
Associational
Methods and
General Forestry
Background
Major
Program
Output #4
& #5
Builds on #2 with
Whole tree
Process Models
coupled with
individual tree
stand dynamics
models
Environmental
Parameters
("site")
•	Climate
•	Weather
•	Fire
•	Water & Hydrology
•	Nutrients & Soil
•	Biotic Factors
•	Management
Practices
Figure 3.1. Influence of natural factors
on stand dynamics.

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Effects of multiple stress factors on composition, structure, and
environmental attributes and "health" of forests will be portrayed through
generalized scenarios of successional or management-induced developmental
dynamics.
2.2.1	Natural Stand Dynamics and Secondary Forest Succession
2.2.2	Managed Forests and Plantations
3	Natural Factors and Forest Condition: The Cases and Regions
The cases are grouped by Research Cooperatives. Further subdivision will
be undertaken as necessary (for example, dividing Spruce-Fir into northern
and southern). The analysis in this section stay close to the data
collected for each case and their statistical combination into regions.
Emphasis will be on statistical association between measures of forest
condition and natural factors such as climatic influences, competition,
aging, and pest organisms. This section corresponds to a "Results" section.
3.1	Southern Pines
3.2	Spruce-Fir
3.3	Eastern Hardwoods Complex
3.4	Western Conifers
4	Natural Factors and Forest Condition: Interpretation of the Cases
For most of the cases, there will be data on only a few of the many
possible natural factors. To properly interpret them, we must make use of
the general knowledge of that forest type which exists in the literature
and of generalized information in 2.1 and 2.2 above. This section
corresponds to a "Discussion" section.
4.1	Southern Pines
4.2	Spruce-Fir
4.3	Eastern Hardwoods Complex
4.4	Western Conifers
5	Conclusion and Recommendation
This will be a synopsis highlighting key observations and recommendations.
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3.3 Major Program Output #3
Quantitative estimates of seedling responses to S, N, and associated
pollutants at ambient conditions. (Seedling Report)
S&I contact: Charley Peterson
1	Introduction
This report will be the first of several annual reports, documenting
results from Coop-funded experiments which study the impact of pollutant
exposures on seedling material. Seedlings are emphasized over mature trees
primarily because they are easier to manipulate and present fewer sampling
problems in experimental designs which require chamber fumigation systems.
They also represent the size of plant material whose mechanisms can be
measured and modeled over a range of experimental treatments, something
which is currently infeasible with mature trees. With this in mind, the
seedling reports will address the following:
1.1	WHY we are doing these studies?
If the purpose of these studies is to determine sensitivity, then
sensitivity to what, and how is it measured? What do we mean by
dose/response, screening/sensitivity, and mechanisms? What are the
issues?
1.2	WHAT do we expect from these studies (i.e. visible or latent
response)?
What issues, if any, will be resolved?
1.3	HOW will these results affect future research?
That is, what does seedling response mean to both the current seedling
population and future seedling development taken to a mature tree
(i.e. under different sensitivity-exposure combinations)? This is not
the same as extrapolation to existing mature trees.
Section III below reflects broad classes of research objectives which
seedling studies might address. Since most of the FRP studies emphasize
screening and dose/response research, the successive seedling reports
should reflect the amount of research completed or ongoing in each of those
areas.
2	Methods/Materials
This section will define the similarities and differences among
experiments in plant material, pollutant, exposure regime, facility, plant
parameters, and meteorological parameters. It will also describe the
analytical methods which were used.
2.1 Variables for consideration
species (age, source, etc.)
plant material
pollutant (S, N, Ozone, etc.)
exposure regimen (level, duration, season, etc.)
facility (open-top, CSTR, greenhouse)
study type (controlled field, controlled lab)
length of study
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foliar injury: -- due to its subjective nature (i.e. how is
"standard" determined?), each PI must offer a precise description
as to the degree of subjectivity (judgement based on experience)
involved in combination with whatever qualitative classification
or quantitative measure is employed.
Plant parameters and interval of measure/sample:
plant diameter
plant height
plant dry weight
foliage dry weight
root dry weight
stem dry wt
root length
needles
primaries (number and length)
secondaries (number and length)
leaf area
Physiological parameters:
Nutrient concentrations (leaves, stems, roots)
C02 exchange (photosynthesis)
stomatal conductance
C-allocation
transpiration
water potential
Meteorological parameters
air temperature
solar radiation
precipitation
2.2 Statistical issues
Explanation of the statistical considerations involved in combining
experiments -- if necessary, S&I will append a white paper on "combining
results from several studies." We should be able to document our
assumptions, methods, and resulting decisions, to show that our analyses
are both justifiable and credible.
3	Results
3.1	Mechanisms
3.2	Design/model
3.3	Preliminary effects
3.4	1986 loblolly pine exposures
3.5	Visible injury
3.6	Plant material
4	Conclusions/Recommendations
Particularly recommendations for future research needs.
5	Appendix A: Role of mechanisms as scientific questions in the FRP
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6	Appendix B
7	Appendix C
8	Appendix D
Summary of projects
Detailed lists/arrays of methods and materials
Format for results requested from Pis by S&I
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3.4 Major Program Output #4.
Evaluation of the roles of S, N, and associated pollutants in forest
decline. (Forest Response)
S&I Contact: Ross Kiester
1	Introduction
1.1	Goal for MPO #4
MPO #4 is a comprehensive report for the Forest Response Program. It
is intended to summarize all of the work of the Program (with the
exception of projection of response under alternative deposition
scenarios) and to lead to an assessment of the effects of current
levels of sulfur, nitrogen, and associated pollutants.
1.2	Strategy for MPO #4
Major Program Output #4 will be produced as a series in 1989, 1990 and
1991. We see a phased implementation of this output over the three
versions. Earlier versions of MPO #4 will bear some resemblance to
MPO #3, but later will achieve their own identity.
2	Are forests being adversely affected?
Here we review MPO #1. Trends and variability in forest condition
regardless of causes. Case studies.
3	Can Sulfur, Nitrogen, and associated pollutants affect forests? How?
This is a review of plant physiology and the knowledge gained by the
Program on the applicability of specific mechanisms of pollutant effects.
These are mostly experimental results.
3.1	Basic Physiology
Here we review plant physiology (briefly) to set the stage for the
possible mechanisms of air pollutants. We also introduce the M1/M2
concept (levels of mechanisms) here.
3.2	Hypothesized Mechanisms
These are the six scientific questions of PQ #2 with some sort of
yes/no/maybe evaluation of potential effect.
3.2.1
Soil Mediated Effects
3.2.2
Foliar Leaching
3.2.3
Carbon Allocation
3.2.4
Winter Injury
3.2.5
Reproduction/Regeneration
3.2.6
Pests/Pathogens
extent
are air pollutants responsible for the effects seen?
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Here we have to tie laboratory and field studies together. We set this up
by asking how do we know that a given natural factor (say temperature)
affects forests? Then we use a parallel line of logic to address air
pollutants. This is an expository and educational device.
4.1 "Everywhere and Nowhere": Generalization
Here we have a discussion of our knowledge of the general effects of
air pollutants. We set it up so that we have competing explanatory
factors for decline: natural or anthropogenic or both.
4.1.1	Summary of MPO #2: Natural Factors
4.1.2	Summary of mechanisms from section 3 above
4.1.3	General summary of deposition
4.1.4	Models to resolve the competition
Here we partial out the importance of the different factors
4.2 "Somewhere": Specific cases
Now using the general information at our disposal from 4.1 (considered
as a sort of toolkit) we proceed with a series of case studies. These
are on specific pieces of real estate and specific forest types. Some
are specific to an area covered by a research project and some are
regionalizations.
4.2.1 Specific Site Case Studies
4.2.1.1	Camels Hump (for example)
4.2.1.2	...
4.2.2 Regional Case Studies
4.2.2.1	NE Lowland Red Spruce
4.2.2.2	...
5 Conclusions
Here we summarize the program by going through two hypotheses and a
parameter estimation for a series for cases
5.1	HO: The "simple" null hypothesis
All forest decline is due to natural factors
5.2	HI: x% of decline is due to S and N
Cannot answer this in general but must estimate x for each case.
5.3	Estimate x for a series of cases both specific and regional
For those really well understood cases from IV.2 above we try to press
on to this.
5.3.1	Specific Site Case Studies
5.3.2	Regional Case Studies
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5.4 Implications for assessment: Our knowledge and its limitations
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3.5 Major Program Output #5.
Projection of Forest Response under Alternative Deposition Scenarios
("Proj ection")
S&I Contact: Ross Kiester
This output is the modeling effort described in the S&I Plan. It will
summarize the outcome of various modeling activities aimed at predicting
forest responses to pollutants. An outline has not yet been developed.
3.6 Project Result Due Dates.
The development of each MPO will involve a synthesis of results from
individual research projects, a rigorous peer review, and a final revision.
This lengthy process means that for project results to contribute to the
outputs they must be received by S&I several months (sometimes as many as
six) before the outputs are actually due. Table 3, below, lists the final
dates that results must be received.
TABLE 3. Project result due dates
for each Major Program Output.
Results
Due
MPO 1
8/01/88
MPO 2
8/01/88
MPO 3
12/01/87

12/01/88
MPO 4
4/01/89

4/01/90

4/01/91
MPO 5
7/01/91
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4 Synthesis and Integration: How do FRP Projects Address the Policy
Questions?
This section consists of pathway diagrams and discussion for research
related to each of the Scientific Questions under Policy Questions 1 and 2.
For Policy Question 1 the two Scientific Questions (1.1 and 1.2) are treated
together because of the way the individual projects overlap and because
grouping them by data type and region has proved more useful.
Discussions under Policy Question 2 are of the six Scientific Questions
pertaining to possible mechanisms for effects of air pollutants on trees.
However, we need to describe what we mean by "mechanism." Everyone recognizes
that what is one person's mechanism is another's effect. Under each
Scientific Question are a variety of studies that are looking at mechanisms at
different levels of biological organization. Understanding how to fit these
studies together requires some classification of the kinds of mechanisms with
regard to different levels of organization. While most researchers perceive
many levels of mechanism, we have found it convenient to group the mechanisms
being studied in the FRP into two broad classes: Mechanisml and Mechanism2 (Ml
and M2).
Mechanisml contains the mechanism level which is believed to result in
tree or forest damage. Mechanism2, then are the proximate mechanisms by which
Ml work. For example, under Scientific Question 2.4, Winter Injury, the Ml
would be winter injury itself, while the M2 would be those particular
mechanisms causing winter injury such as cuticular wax degradation. In the
discussions that follow we separate these two kinds of mechanisms whenever
possible. The strategy of the research then is to work on both classses of
mechanism in parallel, but to come to a decision on the Ml as soon as
possible. If the Ml does not produce an effect then research on that Ml and
its associated M2 would be curtailed. For some Scientific Questions most of
the work is at the Ml level, but for those about which there is more
information work is proceeding at both levels.
4.1 Scientific Questions 1.1 and 1.2: Changes and Patterns in Forest
Condition.
Scientific Questions 1.1 and 1.2, namely "Are changes in forest
condition greater than can be attributed to typical trends and levels of
natural variability" and "What spatial patterns exist in forest condition
and how do these patterns relate to spatial patterns of pollutant
exposure", are intimately connected; indeed, both are addressed
simultaneously in many of the current projects. For these reasons a clear
separation of the questions is not attempted in the construction that
follows. Such separation seems unnecessary and would not be easily achieved
without appreciable redundancy.
4.1.1 Overall Structure
The various projects can be classified in several ways, the most
meaningful, perhaps, being by species which, by and large, coincide with
geographic regions and/or Cooperative responsibilities. We thus
establish four divisions, namely (1) western conifers, (2) southern
pines, (3) the northeast spruce- fir complex and (4) the eastern
hardwood complex.
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While there are certainly commonalities of approach over these
divisions there are some notable distinctions. Data sources can be
classified as dendrochronological studies, FIA plots, fixed plots and
spatial studies. These categories are by no means exclusive. For
example, some spatial studies have a dendrochronological component, some
dendrochronological studies are associated with FIA plots as are some
spatial studies, etc. Currently the western conifers projects are
dominated by dendrochronological data bases whereas FIA data form the
major component of the southern pine projects. Spatial studies prevail
in the eastern hardwood complex. The northeastern spruce-fir complex is
somewhat of a mixed bag of broad surveys complemented by intensive
studies at certain sites.
Accordingly, while the flow charts for each division have been
constructed from the same principles, on the surface they appear to be
rather different.
4.1.2 General Strategy
The first step is to assess from the appropriate data bases whether
there has been a change in forest condition that cannot be accounted for
by natural factors such as stand dynamics, pests and pathogens etc. and
most notably, climate. This is far from a simple task; several such
factors may be acting simultaneously and, apart from some experimental
areas, the relevant records for forest stands are rarely complete.
In addition, there are acknowledged limitations with the
conventional methodology used for analyzing these data, especially in
the interpretation of dendrochronological records. Also, FIA plots, a
major source of available data, were designed for other purposes and are
lacking in the type of information now required.
The Dell/Van Deusen (VS09) and Warren (SI02) projects, in
particular, are intended to reduce, if not eliminate, these
methodological limitations. Recognition of the inadequacies of the FIA
data for southern pines in this respect prompted Dell's (VS12)
compilation and analysis of a variety of permanent plot data.
If it can be concluded that forest condition has declined beyond
what would be consistent with variation in natural factors, an attempt
will be made to relate, as quantitatively as possible, the deviations to
patterns of atmospheric deposition.
4.1.3 Specific Strategies
4.1.3.1 Western Conifers
The specific strategy for western conifers is summarized in
Figure 4.1. Coverage of western conifers includes
dendrochronological studies of Douglas-fir in the Puget Sound area
(Brubaker WC25), Douglas-fir and ponderosa pine in Central Arizona
(Graybill WC24) and ponderosa and jeffery pine in the Sierra Nevada
(Peterson WC26). These have been planned as thorough
dendrochronological studies and, while not formally spatial studies,
cover known ranges of deposition so that there is a good possibility
that any changes that exist in excess of what can be accounted for by
climate etc. can be related to deposition magnitudes. The Edmonds
16

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Fixed plots/FIA data
Graybiil/Swetnam
Dendrochronology
methodology
(1)	Dell
(2)	Warren
MPO #1
Adjust for Stand
Characteristics
Cloud Chemistry
(1)	Miller, Co.
(2)	Larson, Wa.
Relate to pollution
gradient/site differences
Puget Sound Gradient
Edmonds
Adjust for climate
from pollutant disturbances
Remote Sensing
Ustin
Wet deposition mapping
Networks
(2) Svoboda
Dendrochronological Data
(1)	Brubaker. D. fir, Puget sound
(2)	Graybill. (a) D. fir, Central Az
(b) Ponderosa pine, Central Az.
(3)	Peterson (a) Ponderosa Pine, Sierra Nevada
(b) Ponderosa pine + Jeffery pine, Sierra Nevada
Figure 4.1. The path to MPO #1 for Western Conifers.

-------
(WC18) project was planned as a spatial study in the Puget Sound area
and should complement the Brubaker project. Ustin (WC16, WC22) is
working on the development of remote sensing techniques that could
ultimately be applied to broad surveys forest of condition in the
west.
Several projects are involved in monitoring air quality in the
west. These include Larson (WC03) at Mt. Rainier, WA, Miller (WC04)
at Mt. Werner, CO, and Svoboda (WC05) in the Rocky Mountians. Cronn
(WC06) is developing a passive ozone monitor that could find
widespread use. These projects, in conjunction with monitoring data
from existing networks, and the other projects mentioned above, will
provide data to compare forest condition and air quality for some
regions of the west.
4.1.3.2 Southern Pines.
The approach to southern pines is shown in Figure 4.2. The
emphasis in the south has been on utilization of FIA data (McClure
VS07) and the application of multiple regression procedures to
account for stand density, age, etc. Although extensive, such data
are not well suited to the present objectives, but should supplement
the more sophisticated analysis of fixed plot data being undertaken
by Dell (VS12). Another fixed plot study by Christensen (SC10) will
provide a case study of patterns of tree growth and mortality at Duke
Forest. Dendrochronological data, however, would seem to provide the
best means for accounting for climatic effects, especially drought,
but the only study with a dendrochronological component is that of
Saucier (VS08), and this aspect seems not to have been emphasized.
A regional evaluation of Scientific Questions 1.1 and 1.2 for
the south depends on integrating Dell's extensive fixed plot data
with information compiled by the Marx (VS01) Atlas project. The
Atlas contains information on climate, soils, atmospheric deposition,
and distributions of major tree species. The Cowling project (SC01)
has also assembled regional information including atmospheric
exposure data, de Steiguer and Ruark (VS04) are engaged in an
assessment of tree growth models that could lead to the
identification of dependent and independent variables for a broad
survey of forest condition.
4.1.3.3 Northeastern Spruce-Fir Complex.
The project structure in the northeast is multi-tiered (Figure
4.3); one could also make a division between New England and the
Southern Appalachians. There are broad surveys of forest condition
(Ward VS02, Wentworth SF18, Dull SF26, Millers VS14, DeHayes SF24)
with technological input on remote sensing by Rock (VS15). There are
the more intensive studies of Hombeck (VS06) in New England and
Zedaker (SF25) and Wells and Robarge (SF21) in the southern
Appalachians. There are also localized but very intensive studies
of Hornig (SF05) at Mt. Moosilauke, NH and Johnson (SF08) at
Whiteface Mt., NY. The projects of both Hornig and Hornbeck contain a
strong dendrochronological component. Dell (VS06) includes
development of statistical methods that will allow a more thorough
analysis and interpretation of tree ring data. The Mountain Cloud
Chemistry Project (Mohnen AE01), which includes Mueller (SF03)
provides atmospheric monitoring for the intensive site studies.
17

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Dendrochronological
Data
Saucier, GA., SC, NC
FIA Data

(1) McClure
(2)deSteiger/Ruark
(3) Saucier
(4) Dell



Fixed Plo
(1)	Dell
(2)	Chrisl
t data
tensen


Develop
appropriate
analytical tools
(Dell)
Account for stand characteristics
(density, age, composition, etc.)
by conventional methods or models
i
Dendrochronological
Methodology
(1)	Dell
(2)	Warren
Incorporate the effects
of climate, especially droughl
1)	Dell
2)	Zahner
MPO #1
Exposure Data
1)	Marx
2)	Cowling
Figure 4.2. The path to MPO #1 for Southern Pines

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MPO #1
Stand Dynamics
Solomon
Remote Sensing
Methodology
Rock
Patterns of Deposition
Mohnen
Mueller
Dendrochronology
Methodology
Dell
(Van Deusen)
Warren
Patterns and Trends in
Forest Condition
Spatial Studies
(Surveys)
Ward
Wentworth
Dull
Millers
DeHayes
Dendrochronological
Data
Hornbeck
Hornig
Fixed Plot/Intensive Study
Hornig
Johnson
Zedaker
Wells/Roparge
Bruck
Figure 4.3. The Path to MPO #1 for
Northeastern spruce—fir.

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The situation is thus somewhat analogous to a multiphase
sampling system and, thus, a reasonable overall picture of the state
of the spruce-fir forest should be obtainable, albeit with relatively
wide error bounds.
There are a variety of projects that provide information
specific to certain aspects, for example Bruck (SF02) on pests and
pathogens, and Solomon (SF28) on stand dynamics, which should aid in
the interpretation of the broader studies.
Some information will be obtained for species other than red
spruce and balsam and Fraser fir, for example red and white pine, the
last mentioned being individually surveyed by Bennett (VS16).
4.1.3.4 Eastern Hardwoods Complex.
The research strategy for eastern hardwoods is shown in Figure
4.4.
The projects in the "midwest" (Minnesota, Wisconsin, Michigan,
Illinois, Ohio, Pennsylvania) largely address northern and eastern
hardwoods forests, with the exception of Ohmann's (VS10) study which
addresses both hardwoods and softwoods.
There are two studies (Brooks VS11 and Ohmann) that can be
categorized as spatial/gradient studies. These studies cover large
geographic areas where a pollution deposition gradient is known to
exist. Both projects involve the use of FIA fixed plot data and tree
core data. Ohmann's study is using an individual- tree growth model
(STEMS) to account for stand dynamics, with the intent that any
remaining variation in the tree growth data will be correlated to
climatic and pollution variables. Although the Brooks' study does not
account for stand dynamics at this time, it is likely a similar
approach can be taken since the necessary data have been recorded.
There are four studies (Brooks, Davis EH04, Loucks EH05, and
Ohmann) that will be collecting tree core data. It appears likely
that at least two of these studies will be using advanced
dendrochronological methodology in addressing growth response of
individual trees to stand dynamics, climate and pollution.
The studies of Davis, Witter (EH03) and Loucks are intensive
site studies that are attempting to relate changes in forest
community to a pollution gradient. Because Davis's study covers a
subregion of Brooks' larger spatial/gradient study there is a natural
connection between an extensive and intensive study. The Atmospheric
Exposure Cooperative is working closely with these projects to
install deposition monitoring systems at the study sites.
The Groton project (EH08) is determining the long-term growth
patterns of trees in the Tennessee Valley region. It is also
attempting to statistically partition out factors that influence
growth, such as stand density, topographic location, and climate.
In a cooperative project with the Canadian Forestry Service,
Millers (EH07) will study the rate of change in condition of sugar
maple over the next two years.
18

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MPO #1
Forest
Community
Analysis
Adjust for stand
characteristics
Historical forest
decline review
for Eastern U.S.
Miller
Adjust for Climate
A non pollution disturbances
Dendrochronology
Methodology
1) Dell
(Van Oeusen)
2.) Warren
Relate to pollution gradient/
site differences (pollution)
Development of
Spatial Statistics
for Brooks ft Ohman
studies
Warren (Reams)
Intensive Site/Spatial
(1)	Davis.
Eastern Hardwood, Pa.
(2)	Witter.
Northern Hardwoods,
Michigan
(3)	Loucks.
Oak/Hickory,
S. Ill, S. Ohio
(4)	Groton.
Tennessee Valley
(3) Millers.
(2) Ohman.
Eastern Hardwood, Pa.
Northern Hardwoods ft
Softwoods, Minn, Wis, Mich.
Dendrochronological Data
(1)	Brooks.
Eastern Hardwoods, Pa.
(2)	Davis
Eastern Hardwoods, Pa.
(3)	Loucks.
Oak/Hickory, S. Ill, S. Oh.
0+) Ohmann.
Northern Hardwood A
Softwoods
Figure 4.4 The path to MPO #1 for
eastern hardwoods.

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4.2 Scientific Question 2.1: Soil Mediated Effects
4.2.1 Introduction
There are three facets to this scientific question that cover:
1.	Direct toxicity to roots, mycorrhizae, or soil microbial
populations by mobilized metal ions in acidified soil water;
2.	Increased leaching of soil nutrients resulting in reduced
nutrient availability; and,
3.	Nitrogen toxicity to mycorrhizae.
Although three main problem areas have been distinguished, they have
been included in a single question to demonstrate possible
interrelations and to encourage integrative research.
The overall approach to addressing effects of acidic deposition and
other air pollutants via soil mediated mechanisms is illustrated (Figure
4.5). Specific projects involved in each step are indicated. Different
techniques for implementing this approach are appropriate for each of
the two major tree species being studied: red spruce and southeastern
pine species, primarily loblolly. There was a strong scientific
consensus for red spruce ecosystems that it was at least plausible that
acid deposition could contribute to forest decline through soil mediated
processes. No such consensus existed for southeastern pine ecosystems.
Therefore, a series of literature reviews was undertaken to summarize
existing knowledge.
4.2.2 Research
Johnson et al. (SF08) have established a network of permanent study
plots in the high elevation spruce-fir forest at Whiteface Mountain, NY.
Plots have been located at random within strata based on elevation,
exposure, and disturbance history. Research addresses a number of
questions including determination of the nutrient, available aluminum,
and heavy metal content of the soil. Based on previous work, standard
error values of 15 to 20% of the mean values for soil pools of N and C
and exchangeable H+, Al, Ca, Mg, Na, and K are expected. Similar
results are expected for Pb, Cu, Zn, Ni, and Cd content of the forest
floor. Historical data are available that will allow limited
comparisons of nutrient and heavy metal concentration changes over time.
Combined with available deposition data, these studies will contribute
to the evaluation of metal mobilization and accumulation as well as to
determining the effects of different levels of acid input on
biogeochemical cycles and nutrient availability. A companion study also
by Johnson (SF30) uses controlled irrigation treatments to determine the
effects of different levels of acid inputs on soil solution chemistry.
The Integrated Forest Study at the Howland, ME site includes work
by Fernandez (SF04) on effects of acid deposition on the rate of cation
leaching from soil, the balance of critical soil elements influencing
forest productivity, exchangeable aluminum and accumulation of trace
elements that may affect tree growth and decomposition of organic
matter, as well as natural stress factors related to soil properties in
commercial, low elevation red spruce and balsam fir forests of northern
19

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Characterize and evaluate
Changes in tree and
Forest Condition
Wells/Robarge
Determine Nitrogen
Effect on Mycorrhizae
Cline
Cowling
Hodges
Accumulation
Johnson
Fernandez
Binkley
Cowling
Hodges
Evaluate Metal
Determine Toxic Effects
on Roots, Mycorrhizae
and Soil Microbes
Cowling
Wong
Determine Effects on
Biogeochemical Cycles
Wells/Binkley
Johnson
Fernandez
Binkley
Cowling
Gholz
Figure 4.5. Soil mediated effects.

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New England. Intensive soil sampling and analyses over time, including
work on soil solution chemistry, deposition monitoring, and biomass
accumulation will allow evaluation of the contribution of acid
deposition to these processes. Estimates of mineral weathering rates
will help determine changes in process rates as well as their relation
to fluxes of hydrogen, sulfate, and nitrate in the system. This project
is also coordinated with two high elevation EPRI IFS sites.
The Gholz (SC17) project in Florida is also associated with the
EPRI IFS program. Conducted in a slash pine plantation, the project
concentrates on characterizing nutrient cycling from measurements of
deposition, throughfall, stemflow, and soil and soil water chemistry.
Wells and Robarge (SF21) are collecting baseline information in the
southern Appalachians and evaluating relationships between soil, root,
and foliage characteristics and symptoms, location, elevation, slope
position, and exposure. This project is coordinated with Bruck (SF02)
and Zedaker (SF25).
Wells and Binkley (SF17) have completed a project to study nitrogen
input and transformations in soils of spruce-fir ecosystems at sites on
Mt. Mitchell, NC, Mt. Rogers, VA, and Clingman's Dome in the Great Smoky
Mountains National Park. Work will concentrated on ammonium and nitrate
levels and dynamics as related to elevation, exposure, and stand
characteristics and on denitrification, which may be a major process in
the balance of nitrogen input and output in spruce-fir ecosystems.
Among other things, they found that nitrogen mineralization was high
relative to other forests and nitrate concentrations were large enought
o induce high cation leaching.
As mentioned above, there is little agreement among scientists as
to the plausibility of effects of acid deposition on southeastern pine
ecosystems via soil mediated processes. The Hodges (SC03) review
contains reference material on potential effects on soil processes. In
his review, Cowling (SC01) concluded from available evidence with regard
to soil mediated mechanisms that these hypothetical mechanisms can be
listed in the following order of decreasing likelihood:
i)	Leaching of essential nutrients from soils by acid deposition
ii)	Mobilization of toxic metals in soils by acid deposition
iii)	Inhibition of mycorrhizae by nitrogen deposition
In order to evaluate this situation better, Cline (SC09) produced a
comprehensive critical review of the literature on the influence of
atmospheric deposition of nitrogen compounds on mycorrhizal development
in southern forests. He found that no studies in the literature showed
nitrogen to be directly toxic to mycorrhizae at current levels of
atmospheric deposition in the southeastern U.S.
A similar critical review of effects of sulfate and nitrogen
deposition on soil nutrient status in southern commercial forests is
being carried out by Binkley et al. (SC16). Their approach is to
synthesize available information on the biogeochemical impacts of
sulfate and nitrogen and to examine available empirical evidence
relating to the effects of sulfate and nitrogen in southern forest
20

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ecosystems. In addition they will use computer simulation models to
explore soil sensitivity to sulfate deposition and relate these criteria
to major types of forest soils in the South.
Presently, Wong et al. (SC08) are carrying out the only laboratory
and greenhouse studies on the effects of acid deposition on southern
pine species via soil mediated mechanisms. Their work addresses the
problem of enhanced solubility and mobility of metal ions in acidified
soil solution. They are concentrating on whether mycorrhizal fungi
increase or provide no barrier to the uptake of trace metals into the
root and how acidity and metal concentration affect the uptake of metals
by the mycorrhizal fungus.
4.3 Scientific Question 2.2: Foliar Leaching
4.3.1 Introduction
Leaching of nutrients from foliage is a common and natural
phenomenon in humid ecosystems. It is a normal part of nutrient cycling
and does not adversely affect the plant as long as the critical internal
nutrient balance is maintained by uptake of nutrients from the substrate
or atmosphere. Exposure to atmospheric pollutants or acidic deposition
may accelerate normal leaching rates and increase net leaching,
primarily of base cations. Extreme climate events, such as drought or
cold temperatures, also may accelerate leaching.
The primary mechanism (Ml; Figure 4.6) in question is whether loss
of cations from foliage in amounts greater than under natural conditions
results in nutrient deficiency and deleterious effects on growth and
yield of trees. In Central Europe, for example, increased leaching of
Mg from needles is thought to be a major factor contributing to
chlorosis of Norway spruce. Extensive chlorosis of older needle sets is
a main symptom of "decline" of Norway spruce stands.
4.3.2 Research
There are three main methods used for evaluating leaching of
elements from foliage:
1.	Collection and analysis of throughfall water either in natural
stands or in controlled experiments is useful for biogeochemical
studies and does not require harvest of plant material. A
short-coming of this method is that it is difficult to quantify
amounts of throughfall water as well as its element content because
some rain does not contact any leaves while other drops may contact
more than one leaf and remain on the surface for differing periods
of time. In addition, elements previously deposited to foliage can
be washed off during precipitation events. This causes
difficulties in evaluating relative amounts of these elements that
are leached from foliage or deposited from the atmosphere.
2.	Analysis of nutrient levels in foliar tissues after periods of
exposure to pollutants is useful to study nutrient cycles. However,
substantial losses of elements from foliage may occur without
measurable decreases in tissue levels due to redistribution of
element pools within the plant or compensatory uptake from the
substrate.
21

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Estimate growth, mortality,
	symptomatology
Characterize changes in
tree water/nutritional
status
Determine normal
Foliar leaching
(1)	Jenson et al.
(2)	Jacobsen &
Lassoie
(3)	Tingey &
Turner
Determine leaching
mechanisms
(1)	Jenson et al.
(2)	Jacobsen &
Lassoie
(3)	Tingey &
Turner
Evaluate increased
foliar leaching due
to pollutant impact
(1)	Jenson et al.
(2)	Jacobsen &
Lassoie
(3)	Tingey &
Turner
(4)	Cowling
Figure 4.6. Foliar leaching.

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3. Immersion of shoots or foliage in aqueous solution for fixed
periods of time and analysis of elements leached provides an
opportunity to examine possible causes of leaching under controlled
conditions and a more exact measurement of the amount of foliar
tissue subjected to leaching.
Cowling et al. (SC01) have prepared a critical review of the
present state of scientific knowledge on a number of scientific
questions including leaching of nutrients from foliage as a potential
contributor to changes in forest condition in southeastern United
States. They concluded that present evidence is not sufficient to
accept or reject any of the hypothetical mechanisms of the action of
acid deposition by foliage mediated mediated avenues. They could not
rule out the possibility that acid deposition could affect soil
chemistry. They also concluded, however, that the most likely of the
mechanisms is increased leaching of nutrients from foliage by acid
deposition following injury to leaf cell membranes by ambient ozone or
hydrogen peroxide.
Jensen et al. (SF07) have designed a study to address the secondary
mechanism (M2; Figure 4.7) of ozone and acid deposition causing an
increase in foliar nutrient leaching with a subsequent growth loss due
to nutrient deficiency. Experiments are carried out in Continuously
Stirred Tank Reactor (CSTR) fumigation chambers. Seedlings are treated
weekly with ozone and acid rain. After several weeks of treatment,
runoff water of individual seedlings are collected and analyzed for
nutrients and other elements. In a second approach, single shoots of
the seedlings from the runoff collection study are immersed for 24 hours
in a solution of the same pH as the acidic rain in the runoff study.
The shoot is dried and analysis of the element content of both the
leaching solution and the dry tissue carried out. These studies will
yield valuable information on whether ozone has damaged foliar
membranes, leading to increased leaching of nutrients from tissues by
acidic precipitation.
The effects of acidity and sulfate and nitrate content of simulated
acidic mist are being studied by Jacobson and Lassoie (SF06) to
determine if these factors alter growth, development, water relations,
or cold tolerance of red spruce seedlings in ways that may contribute to
decline. The simulated mist contains the major inorganic components of
precipitation. Seedling provenance from low and high elevation sources
in eastern United States and the concentration of hydrogen, sulfate, and
nitrate ions in mist are varied in different tests. Experimental
conditions approximate conditions occurring at high elevation sites by
adjusting the number of hours of exposure per week and average rates of
deposition of water, sulfuric, and nitric acids to actual field data.
During the summer, experiments are carried out in outdoor mist chambers
which are covered to exclude ambient rain. Exposures take place in
greenhouse facilities during winter. These studies allow the evaluation
of whether acidity and chemical composition of mist, comparable to that
at high elevation sites, change amounts or structure of cuticular waxes
or chlorophyll in needles leading to an increase in leaching of
nutrients from foliage.
Tingey and Turner (WC07) are testing the hypothesis that tree
seedlings can tolerate leaching of cations from foliage due to impact of
acidic precipitation only if they can compensate for these losses by
increased uptake of cations by the roots. Acidified mist is applied in
the greenhouse to hydroponically grown seedlings of Douglas-fir and
22

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Impact of
or air poll
climate
utants


Disruption of cuticular wax
and membrane integrity
(1)	Jensen et al.
(2)	Jacobson & Lassoie
Precipitation
Increased leaching of
nutrients from foliaae
Changes in nutrient/
water uptake from soil
Tingey 6c Turner
Net change in foliage
nutrient content
(1)	Jensen et al.
(2)	Jacobson & Lassoie
(3)	Tingey & Turner
Expression of
decline symptoms
(e.g. chlorosis)
(1)	Jensen et al.
(2)	Jacobson 6c Lassoie
(3)	Tingey 6c Turner
Net change in plant
water relations	
I
Change in
foliaae biomass
Figure 4.7. Mechanisms of foliar leaching.

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Engelmann spruce. The mist and throughfall water are analyzed for
electrical conductivity, pH, and concentrations of calcium, magnesium,
and potassium. Cation uptake (measured by cation losses from the
nutrient solutions) and changes in biomass, root/shoot ratio, and foliar
concentrations of Ca, Mg, and K are also monitored. This will allow the
evaluation of the interaction of foliar cation leaching and root uptake
of cations in the responses of two coniferous tree species of the
western United States to simulated acidic precipitation.
In addition to the literature study, which produced background
information and up-to-date status of the state of the science on several
different scientific questions, the empirical studies described here
address all major facets of the hypothesis of increased foliar leaching
(influence of ozone, importance of acidity and chemical composition of
mist, foliar leaching vs. root uptake). The relative importance of
these variables is difficult to assess in the field. The understanding
of their importance that will come from these studies is critical to
assessing effects of acidic precipitation on vegetation and to
developing management strategies to minimize potential losses in forest
productivity.
4.4 Scientific Question 2.3: Carbon Allocation
4.4.1 Introduction
This is the broadest Scientific Question within the Forest Response
Program; it involves the effects of air pollutants on basic
physiological processes. Therefore, it requires a broad strategy and a
wide variety of kinds of research. Carbon allocation is the movement of
photosynthate between the various compartments that make up plants. In
its simplest form, the study of carbon allocation would involve
measuring the production of roots and shoots and comparing the two
compartments. Obviously, carbon allocation studies can become very
complex as the number of compartments increases. The FRP has tried to
find a middle ground that allows a realistic depiction of plant
processes, but is still relatively simple.
Our approach is based on the conceptual model depicted in Figure
4.8. The model has been adopted because it clearly demonstrates the
interactions of the various processes and the possible outcomes of
modifications in one or more. For example, from Figure 4.8 it is
apparent th&t if repairing pollutant damaged tissue increases
respiration, less energy will be available for root and foliage
maintenance and growth (even if photosynthetic rate remains constant).
Reduced root growth can affect photosynthesis via disruptions^of water
and nutrient fluxes. Reduced foliage area will result in a diminished
capacity to fix carbon and a decrease in energy available for all plant
processes. The conceptual model also shows that plants must maintain a
balance between the various compartments.
A central challenge for the Forest Response Program is the
integration of the results of fundamentally different kinds of research.
How can information be pulled together in a cohesive way to address the
program's policy questions? To answer this challenge, the FRP adopted
an approach based on biological levels of organization: seedling,
branch, tree, stand, region. The thinking behind such an approach is
two-fold. First, each level can be studied and analyzed in and of
23

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Foliage
area
Growth,
storage,
respiration,
reproduction
Nutrient
flux
Carbohydrate
reserve
growth
Root
Water flux
Photosynthetic
rate
Figure 4.8. Conceptual model of carbon allocation

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itself. Second, information at one level can be synthesized and used as
input to the next higher level (Figure 4.9). One advantage of this
modular approach is that it provides a framework that can accommodate
and assimilate results from both survey and experimental types of
research. The levels of organization approach is useful for all aspects
of the FRP, but it is especially valuable for questions related to
carbon allocation.
As Figure 4.9 illustrates, one of the ways that information is
integrated within the FRP is through the use of computer simulation
models. These may either apply to a single level of organization or may
serve to link two levels. For example, much of the experimental work in
the program is conducted on seedlings for obvious practical reasons.
But it is important that the connection between seedling physiology and
mature tree physiology be made. Although this is a very difficult
problem, models may help. In this particular case, some of the tools
the program will use to make the seedling-mature tree connection are
physiological process models. Focusing on basic processes, rather than
simply growth response to exposure, should allow some of the seedling
results to also pertain to mature trees. The model for carbon
allocation is discussed below in Section 4.4.2.2.
Figure 4.9 also illustrates the parallel use of branch studies in
developing an understanding of mature tree response. Branch studies
also contribute to process level models and can help to improve the
interpretation of seedling results.
At the stand level, a modular system of simulation models will be
employed. The approach is to construct a series of individual tree
based stand models. This means a model in which individual trees are
well described, perhaps by the physiological process models, and then
groups of these trees are made to interact to produce the patterns of
stand dynamics. This approach allows the incorporation of physiological
mechanistic information in the specification of the individual trees.
It also recognizes that a stand is not simply the sum of its individual
parts, but the result of many complex interactions between individual
trees. Natural factors like drought or pest/pathogen interactions can
also be included. Once these stand models are in place, they can be
used in conjunction with spatial statistical techniques to undertake the
appropriate regional aggregations
4.4.2 Research
4.4.2.1 Relative sensitivity rankings
In the cases of two broad categories of tree species, eastern
hardwoods and western conifers, the first step has been to begin to
identify the species that are most sensitive to acidic deposition and
associated air pollutants. Davis (EH01) and Jensen (EH06) are
determining the relative sensitivities of a variety of hardwood
species: red oak, white oak, yellow poplar, white ash, sweetgum, red
maple, sugar maple, American beech, black cherry, yellow birch, and
shagbark hickory. Miller (WC09) is studying the sensitivities of
five western coniferous species: ponderosa pine, Douglas-fir, white
fir, Englemann spruce, and sub-alpine fir. A companion project
conducted by Hogsett (WC08) is doing the same for ponderosa pine,
Douglas-fir, western red cedar, western hemlock, and lodgepole pine.
Future work on eastern hardwood and western coniferous species will
be based on the results of these projects.
24

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Deposition
Data
Natural
Factors
Spatial
Statistics
Deposition
Data
Natural
Factors
Individual Tree Growth
and Yield Models
Experiments
Inference
Physiological
Process -
Models
Regional Growth
and Yield
Stand Growth
and Yield
Seedling Physiology
Whole Tree Physiology:
Growth, Mortality
Branch Physiology
Figure 4.9. Levels of organization.

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4.4.2.2 A carbon allocation model
A schematic of the carbon allocation model being developed by
Ford (SI01) is shown in Figure 4.10. It describes the dynamics of
tree growth through the same basic physiological processes as
McMurtrie and Wolf (1983); however, the structure of the model is
more complex and incorporates features necessary to model some
theories of pollution influence. Foliage is distributed by age, with
each age having a different photosynthetic rate and mortality
function. The priority in allocation is given first to satisfy
respiration, CI, which is modeled separately from mortality. Each
tissue category can have a different respiration rate. The second
priority is to foliage. Three limits are placed upon its growth: i)
An annual increment may only exceed the previous year's by a certain
factor, C2. This is an attempt to simulate the branch support
requirement. C2 declines as canopy size increases. ii) To simulate
light or water limitation, a maximum foliage amount is assumed, C3.
Annual foliage mortality increases proportionally as the maximum
value is approached. iii) Foliage increment can never exceed a
certain proportion of the photosynthate that remains after the
respiration requirement is satisfied. This simulates the balance of
photosynthate distribution attained through seasonality.
Allocation to fine roots maintains a specific fine root:total
foliage ratio, C4. This assumes a necessary balance is maintained
between root and shoot. Allocation to fine root will increase if
root mortality increases, C5, but canopy size stays large. The
photosynthate that remains after all of these processes are satisfied
is allocated to woody tissue.
The most difficult aspect of the forest growth mechanism to
simulate is foliage development. This model attempts to describe
mechanisms at the stand level that are really properties of the
individual tree and crown. The behavior of the model can be
critically influenced by values used for C2, the limits set by
branching structure, and C3, maximum size. These constraints on
growth of foliage have a rational basis, but further modeling of this
aspect of forest growth is essential at the branch and crown
development level and is in progress.
Table 4 lists the model compartments and the FRP projects that
should contribute to each. It is important that this table be
accurate. Program scientists should review it carefully. If they
note an error regarding their project's place in the table, or if
changes occur in the future, they should contact S&I.
4.4.2.3 Mechanisms of Damage
Only one mechanism2 level project is currently in progress.
Richardson (SF11) is studying the generation of oxygen-based free
radicals by photochemical oxidants in the tissues of affected plants.
The results of the mechanisml level studies summarized above will
help to identify appropriate mechanisms of damage for further study.
25

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Photosynthetic rate, by age.
Foliage\
mortality!
•v rate /
Foliage by age category
^ Total \
Canopy
v Size/
C 7
Dead i
Foliage [
Dead
Fine
Root
Fine
Root
C5
Photosynthate
Branch, bole
and thick
root
C4
Respiration
Figure 4.10 Schematic of the carbon allocation
model.

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Table 4. Projects contributing to the carbon allocation model.
Photosynthesis
Greenwood (SF32)
Jacobson (SF06)
Jensen (SF07)
Kohut (SF31)
McLaughlin (SF10)
Seiler (SF13)
Thornton (SF27)
Unsworth (SF14)
Weinstein (SF16)
Jacobson (SF06)
Jensen (SF07)
Kohut (SF31)
McLaughlin (SF10)
Seiler (SF13)
Thornton (SF27)
Unsworth (SF14)
Weinstein (SF16)
Houpis (WC20)
Fong (SC02)
Kossuth (SC11)
McLaughlin (SC04)
Reinert (SC05)
Richardson (SC07)
Teskey (SC18)
Houpis (WC20)
Foliage Amount
Flagler (SC14)
Fong (SC02)
Johnson (SC13)
Kossuth (SC11)
Kress (SC06)
Lockaby (SC15)
McGregor (SC12)
McLaughlin (SC04)
Reinert (SC05)
Richardson (SC07)
Foliage Mortality
No FRP projects are studying foliage mortality at this time.
Respiration
Greenwood (SF32)
Jacobson (SF06)
Jensen (SF07)
McLaughlin (SF10)
Seiler (SF13)
Thornton (SF27)
Unsworth (SF14)
Weinstein (SF16)
Fong (SC02)
Kossuth (SC11)
McLaughlin (SC04)
Reinert (SC05)
Richardson (SC07)
Teskey (SC18)
Houpis (WC20)
Branch. Bole. Thick Root
No FRP projects are studying thick root,
measure shoot weight of seedlings.
The projects listed here
Jensen (SF07)
Seiler (SF13)
Thornton (SF27)
Unsworth (SF14)
Houpis (WC20)
Flagler (SC14)
Fong (SC02)
Johnson (SC13)
Kress (SC06)
Lockaby (SCI5)
McGregor (SC12)
McLaughlin (SC04)
Reinert (SC05)
26

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Table 4 continued.
Fine Roots
Jensen (SF07)
Seller (SF13)
Thornton (SF27)
Unsworth (SF14)
Flagler (SC14)
Fong (SC02)
McLaughlin (SC04)
Reinert (SC05)
4.5 Scientific Question 2.4: Winter Injury
4.5.1 Introduction
The approach to addressing Scientific Question 2.4 follows two
tracks (Figure 4.11). The first, which is depicted on the left-hand
side of Figure 4.11, seeks to determine if there is reason to believe
that winter injury, in and of itself, contributes to forest decline.
The second track, on the right in Figure 4.11, is to determine if
current levels of winter injury are caused by air pollutants or if they
are about what we would expect to occur naturally. In the interest of
timeliness, both tracks are being pursued simultaneously.
4.5.2 Research
4.5.2.1 Current Winter Injury
Winter injury results from the exposure of susceptible tissues
to damaging temperatures. Because both susceptibility, or its
inverse hardening, and temperature follow seasonal patterns, it is
the dynamic interaction of these two patterns that we must
understand.
The seasonal pattern of temperature for the two major parts of
the red spruce range, north and south, are being developed by
Unsworth (SF14) from weather records. The result will be a
probability distribution of temperature throughout the winter that
will be useful in the interpretation of all of the other winter
injury projects.
Two projects, Unsworth and Williams (SF20), are characterizing
present patterns of winter hardening. These are the patterns
actually occurring in the field regardless of ambient air quality or
deposition. The results of these projects will also be probability
distributions. Here they will be distributions of depth of hardening
throughout the winter.
The seasonal development of frost hardiness will then be
compared with the temperature pattern to quantify the probability of
winter injury in the field. This comparison will not tell us
anything about the possible involvement of air pollutants, but it
will provide an indication of whether winter injury is likely to be a
contributing factor in forest decline.
27

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Define Pollutant
Modified
Injury Pattern
Define Normal
Injury Pattern
Evaluate Importance of
Winter Injury Under
Present Condition*
Determine Pollutant Effect
on Susceptibility
Uneworth
Williams
Jacobeon
Weinetein
Determine Normal* Winter
Injury Susceptibility
Uneworth
William*
Jacobeon
Weinetein
Characterize Seasonal
Weather Pattern*
Uneworth
Characterize Preeent
Hardening Pattern*
Uneworth
Williams
Compare to Determine
Pollutant Effect
•Normal susceptibility refers to injury that would be
expected in the absence of all pollutants.
Figure 4.11. Winter injury.

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Effects on
needle
cuticle,
waxes, etc.
Freezing or
Desiccation?
Normal Winter
Condition
Abnormal Winter
Condition:
Morphological
Wilkinson
Normal Summer
Condition
Abnormal Winter
Condition:
Physiological/Biochemical
Unsworth
Figure 4.12. Mechanisms of winter injury.

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4.5.2.2 Influence of Air Pollutants on Winter Injury
Several projects are conducting work to determine the effect of
air pollutants on the occurrence of winter injury: Unsworth,
Williams, Jacobson (SF06), and Weinstein (SF16). The strategy is to
use exposure studies to characterize the hardening pattern of
seedlings in response to various levels of air pollutants. The
hardening pattern that could be expected in the absence of pollutants
(the "normal" pattern in Figure 4.11) will be inferred from the
behavior of the experimental control treatments.
The seasonal temperature patterns developed by Unsworth will
then be used to define normal and pollutant modified patterns of
winter injury that could be expected to occur in the field. The
comparison of these two patterns will provide the basis for an
evaluation of the pollutant impact on winter injury.
4.5.3 Mechanisms of Winter Injury
Our concept of the potential mechanisms of winter injury is
illustrated in Figure 4.12. Instead of the development of normal winter
hardened condition, trees can attain an abnormally susceptible condition
via morphological or physiological/biochemical effects or both.
Morphological effects refer to an actual physical alteration in the
plant or leaf. This could be such effects as damage to the foliar
cuticle or erosion of cuticular waxes. Trees suffering this kind of
damage might undergo the normal physiological processes of winter
hardening, but be otherwise physically unable to resist freezing or
desiccation. One project, Wilkinson (SF19), is studying this type of
mechanism.
Physiological/biochemical effects refer to disruptions of the
normal hardening process itself or other key processes. For example,
the timing of winter hardening could be affected so that it occurs too
late in the season or ends to early in the spring. Another possibility
is that the depth of winter hardening attained is inadequate to protect
the plant from injury. Unsworth is testing the hypothesis that
absorption of air pollutants during winter results in phytotoxic
accumulations of sulfite and nitrite.
4.6 Scientific Question 2.5: Reproduction/Regeneration
4.6.1 Introduction
Fundamental processes such as reproduction and regeneration of
forest species are essential to sustaining a healthy forest succession.
Yet relatively little research has addressed the effect of air
pollutants on these processes and the potential impact on the forest
ecosystem. In fact, it is the exception when these processes are
incorporated into stand growth projection systems.
4.6.2 Research
As shown in Figure 4.13, once a regeneration pattern is
hypothesized as being modified from air pollutant effects, it must be
compared with the expectation of a "normal" regeneration pattern (i.e
occurring in absence of pollution), in order to determine a pollutant
28

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Determine expected
regeneration pattern
Compare to
determine pollutant
effect
Determine pollutant
effect on
seed germination
Determine pollutant
effect on
regeneration pattern
Determine pollutant
effect on
quality/quantity
of seed production
Determine expected
seed germination
(1)	Feret
(2)	Bonner
Determine pollutant
effect on
seedling survival
and development
Define impact of
seed quantity/quality
and survival on
regeneration
pattern
Determine expected
quality/quantity
of seed production
(1)	Feret
(2)	Bonner
(3)	Zedaker
Define pollutant
impact on
seed quantity/quality
and survival on
regeneration
pattern	
Determine expected
seedling survival
and development
(1)	Bonner
(2)	Bruck
(3)	Zedaker
(4)	Christensen
Figure 4.13. Reproduction and regeneration.

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effect. That is to say, we wish to establish whether or not a
disturbance of reproduction or regeneration is outside the bounds of
what we expect to occur naturally, both temporally and spatially.
In the Spruce-Fir Cooperative, Bonner (SF01) is studying both seed
quality at the time of dispersal, and retention of seed quality in
declining spruce-fir stands. Feret (SF22) is studying Fraser fir pollen
germination and flower production in open-top chambers and at Mt. Rogers
and Mt. Mitchell. The Zedaker (SF25) project is evaluating regeneration
success in relation to site characteristics in the southern
Appalachians. A specific objective of Zedaker's project is to first
characterize existing stand conditions and subsequently determine the
regeneration success of spruce-fir.
In the Southern Commercial Cooperative, Christensen (SC10) is
taking advantage of a decades-old system of permanent plots at Duke
Forest to characterize patterns of seedling establishment and mortality.
It may, however, not be appropriate to initiate a more extensive
investigation of reproduction/regeneration in the south. The Cowling
critical review determined that disruption of reproduction and
regeneration was the least likely mechanism of forest decline in the
south.
4.7 Scientific Question 2.6: Pests/Pathogens
4.7.1 Introduction
Interactions of pest/pathogens with anthropogenic air pollutants
must take into account other biotic and abiotic stress relationships
(e.g. drought, frosts, successional patterns, stand characteristics).
At the same time, attempts must be made to discuss and measure
biological interactions under more controlled conditions of
experimentation.
4.7.2 Research
As indicated in Figure 4.14, the approach to detecting increased
susceptibility of forest stands to insects and/or disease, as a result
of exposure to air pollutants, is to first characterize both what might
be considered "normal" pest/pathogen outbreak patterns and the kinds of
environmental stresses that can lead to outbreaks. Three projects, now
complete, provided information on patterns of pest/pathogen occurrence.
The Hodges (SC03) literature review discusses major insects and diseases
in the south. Millers (EH02), in a review of recorded declines in
eastern hardwood species, covers pests and pathogens as sometime causes
of forest decline. He concluded that there was no evidence that any of
these outbreaks had been induced by air pollutants. Dull (VS13)
produced a key and aerial photo interpretation guide for for visual
assessment of forest condition in the southern Appalachians that
includes pest and pathogen impacts. Hodges and Millers also include
information, as does Cowling (SC01) about the types of stresses that can
predispose a forest to outbreaks of pests and pathogens.
Most of the projects working on Scientific Question 2.6 are
involved in studying the ways in which pests and pathogens may alter
forest condition or be associated with declining forests. Hornig
(SF05), Johnson (SF08), and Bruck (SF02) are all intensive field studies
29

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Compare to
determine
pollutant effect
Characterize
major environmental
stress events
(1)	Hodges
(2)	Cowling
(3)	Millers
Identify change
in forest condition
along deposition
gradients
(1)	Hornig
(2)	Johnson
Characterize normal
pest/pathogen
patterns
(1)	Hodges
(2)	Millers
(3)	Dull
Identify pests and
pathogens and
stress events
along gradient
(1)	Hornig
(2)	Johnson
Determine potential
for altering
forest condition
(1)	Hornig
(2)	Johnson
(3)	Wargo
(4)	Millers
(5)	Bruck
(6)	Jensen/Schier
(7)	Skelly
Figure 4.14. Pests and pathogens.

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that include surveys of pest/pathogen occurrence for correlations with
forest condition. Wargo (SF15), also a field study, compares crown and
root condition of healthy and declining trees and attempts to isolate
root pathogens. Skelly (VS05) has developed a manual for diagnosing
injury arising from a variety of causes including pests, pathogens, and
air pollutants. Jensen & Schier (SF07) have the only experimental study
that directly investigates pollutant exposure on pest/pathogen
susceptibility. They use red spruce seedlings and spruce budworm
larvae.
The Hornig and Johnson projects also have spatial components that
may make it possible to analyze interactions between pest/pathogen
occurrence and forest conditions at varying levels of atmospheric
deposition.
4.8 Atmospheric Exposure Support
A.8.1 Information Needs
The Atmospheric Exposure Cooperative has identified two information
needs of the Forest Response Program to be met in acquiring air quality
data derived from monitoring activities:
1)	Information on the deposition of nitrogen and sulfur compounds
in forested areas. The monitoring data that is available to
estimate the total deposition (wet deposition plus dry deposition)
of nitrogen and sulfur compounds is not summarized in a way that is
suitable for use in the FRP.
2)	Information on the ambient concentrations of gases containing
nitrogen and sulfur. In order that the contribution of sulfur and
nitrogen gases to forest damage may be adequately assessed and that
effects due to other gaseous pollutants are not confounded with the
sulfur and nitrogen gas effects, it is also necessary to compile
summary statistics for other gaseous pollutants such as ozone.
4.8.2 Available Data - The Data Book
The combination of accumulated data and the assessment of the
usefulness of the data for estimates of total deposition and
concentrations of ambient gasses will be refered to as a "Data Book".
The Data Book is to contain only summaries of existing monitoring data.
That is, the data summaries will be on a site basis over whatever years
of recorded data are available. A more detailed account of the
accumulated data is given below. That is, the geographic areas under
consideration, the time intervals for the accumulation of statistics,
quality assurance of the data, and the statistics to be accumulated will
be discussed briefly.
4.8.2.1 Geographic areas
The geographic areas to be considered are those near or
containing forested areas. Particular attention is to be given to
areas that are used directly in the FRP as study sites. The gradient
areas (Minnesota-Michigan, Pennsylvania, and the Ohio River corridor)
of the Eastern Hardwoods Cooperative will recieve intensive
examination. To the extent possible, the areas covered by the Western
Conifers Cooperative, Spruce-Fir Cooperative, and the Southern
Commercial Cooperative will be included in the DataBook. The Mountain
Cloud Chemistry study sites will also be included.
30

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4.8.2.2	Time intervals
In order that the Data Book does not become too large, data
summaries will be done on a monthly basis whenever possible. The
monthly summaries provide a flexible summary interval for the data.
That is, longer term summaries (e. g. of the growing season) may be
constructed from the basic summaries to fit the needs of
investigators. A yearly summary will also be included for a quick
overview of the data summaries. Given the paucity of monitoring data
before about 1978, the years of data covered will be 1978 through
1986.
The data bases that will be accessed to provide data include:
EPA SAROAD/AIRS, EPRI-SURE/ERAQS, National Park Service, and the
Tennessee Valley Authority for ambient gasses; and NADP, UAPSP-EPRI,
and MAP3S for precipitation data. Each agency involved in the
compilation of the data bases has a quality assurance plan that will
be referenced in the Data Book to aid in providing relevant quality
assurance information to the quality assurance program within the
FRP. The accumulation of such quality assurance information will
proceed with the accumulationof the Data Book.
4.8.2.3	Quality assurance
As a part of the the quality assurance aspect of the DataBook
and as an aid in statistical interpretation, estimates of the
standard errors of the summary statistics will be provided. The
extent to which reliable standard error estimates can be provided is
an effort that will continue throughout the time the Data Book is
being compiled. One section of the written summary text for the Data
Book will be devoted to how the standard error estimates were arrived
at and how they might be used in data interpretation. There will also
be an effort made to explain how to combine summaries in the data
book to arrive at other than monthly summaries (e. g. growing season
means). For the longer term summaries, the synthesis of appropriate
standard errors will be discussed.
4.8.2.4	Statistics
At present, there is a lack of the detailed information
necessary for the calculation of dry deposition of sulfur and
nitrogen. However, information about some of the component
pollutants necessary for the dry deposition calculation is available:
1)	Monthly and yearly summary statistics will be given for the
ambient concentrations of the gases S02 and NOX (in most cases
N02). These concentrations will provide a rough guide to the
availability of monitoring information that might be used at a
later date for the dry deposition estimate. In addition, the
concentration summaries will give an indication of the
availability of monitoring data as a guide for 'real world'
patterns of ambient gas concentrations that can be used in
fumigation studies. Percentile and/or fractile summaries of
hourly concentrations and the number of hours represented in the
summaries will constitute the Data Book entries.
2)	Summaries of particulate sulfate and nitrate will add to the
information about the components needed in the drydeposition
calculation.
31

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The wet deposition data will be supplied as: a volume weighted mean
and/or median; the total volume used to compute the mean and/or
median; and an a real estimate of wet deposition. The summaries will
be done for selected ions (at least: lab hydrogen, pH, sulphate,
nitrate, ammonium; and possibly: chloride, sodium, calcium,
magnesium, potassium).
4.8.3 Format
The final form of the Data Book will consist of two sections:
1)	The summary statistics will appear in a form that can be
accessed by computer. That is the summaries will be provided on a
diskette or magnetic tap'e, for example.
2)	The assessment of the summary statistics will be provided
inhard copy form as a report.
The details of the final presentation form of the Data Book will be
decided as the Data Book is accumulated. During the course of the
assembly of the Data Book, investigators with a use for the completed
sections will be provided with summary statistics. The completion date
for the Data Book is December 1988.
32

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5 The Overall Path to Major Program Output #4
The key output of the FRP is MPO #4. Here the Program will summarize the
effects of air pollutants on trees and forests. A first version of this
Output will be completed in 1989 using whatever information is available. This
will be updated in 1990 and a final version produced in 1991 which will use
all of the information produced by the FRP.
The overall strategy for producing MPO #4 (and indeed for the FRP) is
given in Figure 5.1. The idea of parallel work on Policy Questions 1 and 2 is
detailed in the National Plan. Figure 5.2 gives a more detailed account of
how the components of the FRP will articulate to form MPO #4. The dates in the
boxes are either the dates that the MPOs will be completed or the dates by
which the program will need the appropriate inputs. It is important to realize
that work will be proceeding in parallel on many activities. For example, the
whole tree modeling is already underway and is not waiting for the results of
the mechanism decisions to begin.
Figure 5.2 shows that Policy Question 1 studies will result in answers to
the extent and magnitude questions and will provide stand and region level
inputs to MPO #4. Meanwhile Policy Question 2 studies on mechanisms are
determining if a given mechanism is a mechanism of decline (Ml) and, if so,
what the corresponding underlying mechanisms (M2) are. These studies are
primarily, but by no means exclusively, conducted on seedlings so a parallel
activity is understanding the nature of inferences that an be made from
seedlings to mature trees. The Atmospheric Exposure Cooperative provides
support for linking pollution exposure directly to field sites and also
provides regional estimates of exposure based on data from monitoring
networks. The three lines of activity (PQ #1, PQ #2, and atmospheric exposure)
come together in a toolkit of models designed to assess the impact of
pollution on trees and forests. Figure 5.2 also shows that MPO #5, projection
of response under alternative deposition scenarios, follows from MPO #4 and
from the use of projections of deposition derived from deposition models.
The first version of MPO #4 will be ready in time to be of use for the
1990 NAPAP assessment. Other, earlier, assessments will derive from the
ongoing parallel work that is complete at the time the assessment is required.
33

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Atmospheric
Exposure
S and N
forests?
affect
Can
S and N
forests?
affect
Does
Policy Question #1
Studies
Policy Question §2
Studies
Figure 5.1. Overall strategy for MPO #4

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PQ 1
Studies
Data Types

Dendro,

FIA.

Fixed Plot,

Gradient

PQ 2
Studies
Atmospheric
Studies


New data
Collected
on Site
f
Mechanism j
Studies
Mechanism 2
Studies
Natural
Factors
Data 4c
Analysis
Existing
Monitoring
Data
Model
Generated
Data
MPO #3
Decision:
Is this a
mechanism of
damage?
Caee Studiee
Regionalized trends
analysis
MPO #2
Intermediate
outputs:
Mechanism
White Papers
Seedling to
Tree
MPO #1
Individual
Tree stand
dynamics
Extrapolation
Methods
Whole Tree
Physiology
Process Model
MPO #4
MPO #5
Figure 5.2. Paths to all Major Program Outputs.

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6 Specification for New Milestones for Individual Projects
This report is the first of what will be a series throughout the life of
the Forest Reponse Program. It represents our understanding at this point in
time. Further development will follow from a two-way interaction between this
document and the individual Projects and Cooperatives. On the one hand,
Synthesis and Integration needs to have continuing and detailed input from the
Projects as to their activities, results, and interpretations. On the other,
the Projects need to be aware of the overall components and timelines of the
Program (see Figure 5.2) and to structure their outputs accordingly.
We hope that this version of the report will stimulate the Projects and
Cooperatives to suggest possible new milestones and dates for their delivery
in light of the overall schedule. We recognize that science does not proceeed
on a schedule easily, but that this must be balanced against the need to
produce the Major Program Outputs on time. In most cases Projects have
intermediate outputs identified; we will welcome others.
34

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Appendix I
Categories of Projects in the
Forest Response Program
35

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POLICY QUESTION 1
1. SPATIAL STUDIES	S&I Contact: Greg Reams
Brooks	Relationship between forest conditions and atmospheric
VS11	deposition across a deposition gradient in Pennsylvania
Davis	Measurement of forest condition and response along an
EH04	atmospheric deposition gradient
DeHayes	Genetic uniformity: A fundamental mechanism for red
SF24	spruce decline
Dull	Detection and evaluation of spruce-fir decline in the
SF26	southern Appalachians using remote sensing
Edmonds	Pollution gradient studies in the Puget Sound region and
WC18	evaluation of existing conifer plot data for correlative
studies
Groton	Assessment of factors contributing to forest condition in
EH08	mixed species hardwood stands in the Tennessee Valley
region
Hornbeck	Analysis and interpretation of tree cores in New England
VS06
Hornig	Studies of spruce-fir decline on Mt. Moosilauke in Hew
SF05	Hampshire and Berry Pond in Maine
Johnson	Quantifying spruce decline and associated forest
SF08	characteristics at Whiteface Mountain, NY
Loucks	Investigation of pollutant effects patterns in
EH05	oak/hickory forests from southern Ohio to southern
Illinois
Marx	Design and pilot test of a long-term monitoring study of
VS01	eastern U.S. forest for response to atmospheric
deposition
Millers	Cooperative survey of red spruce and balsam fir decline
VS14	and mortality in the northeast: symptoms and trends
Millers	North American sugar maple decline project
EH07
Ohmann	The relation between forest condition and atmospheric
VS10	deposition across the Minnesota to Michigan deposition
gradient
Rock	Vegetation survey pilot study: detection and
VS15	quantification of forest decline damage using remote
sensing techniques
Ustin	Remote sensing of forest condition in the western United
WC16	States
Ustin	Spectral characteristics of conifer species exposed to
WC22	simulated pollutant regimes.
Ward	Pilot test of the application of geographic information
VS02	system technology for monitoring boreal montane forest
decline
Warren	Development of innovative statistical procedures for the
SIQ2	FRP's Synthesis and Integration Project.
38

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Wentworth
SF18
Witter
EH03
Compilation and interpretation of the vegetation data
base and disturbance history of southern Appalachian
spruce-fir
Effects of an air pollution gradient on northern
hardwoods forests in the southern Great Lakes
2. DENDROCHRONOLOGY
S&I Contact: Bill Warren
Brubaker
WC25
Davis
EH 04
Dell
VS09
Graybill
WC24
Hornbeck
VS06
Hornbeck (Jagels)
VS06
Hornig
SF05
Marx
VS01
Peterson
WC26
Warren
SI02
Growth variations in old-growth Douglas-fir forests of
the Puget Sound area
Measurement of forest condition and response along an
atmospheric deposition gradient
A review of statistical methods for evaluation of
atmospheric deposition influences on forests
Analysis of growth trends and variation in conifers from
central Arizona
Analysis and interpretation of tree cores in New England
Finger printing radial increment data for red spruce
using morphometric analysis
Studies of spruce-fir decline on Mt. Moosilauke in New
Hampshire and Berry Pond in Maine
Design and pilot test of a long-term monitoring study of
eastern U.S. forests for respone to atmospheric
deposition
Growth trends ain the mixed conifer forest of the Sierra
Nevada
Development of innovative statistical procedures for the
FRP's Synthesis and Integration Project
3. FIA	S&I Contact: Bill Warren
Dell	Southern forest growth trends
VS12
McClure	Formulation and testing of non-deposition growth loss
VS07	hypotheses with existing southeast forest inventory data
Saucier	Near-term survey of eastern forest conditions/atmsopheric
VS08	deposition
de Steiguer	Synthesis and integration of tree growth models to
and Ruark	identify variables for surveying and monitoring eastern
VS04	forest ecosystems for atmospheric deposition effects
de Steiguer	Methods for economic assessment of atmospheric pollution
VS03	impacts on forests of the eastern U.S.
Warren	Development of innovative statistical procedures for the
S102	FRP's Synthesis and Integration project.
37

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k. FIXED PLOT
S&I Contact: Ross Kiester
Bennett
VS16
Christensen
SC10
Dell
VS12
Saucier
VS08
Solomon
SF28
Warren
SI02
The status of eastern white pine affected by air
pollutants in the Eastern U.S.
Analysis of Duke Forest permanent plots: Patterns
production and mortality of seedlings and trees
Southern forest growth trends
of
Near-term survey of eastern forest condition/atmospheric
deposition
Modeling stand dynamics of spruce-fir forests in the
northeast
Development of innovative statistical procedures for the
FRP's Synthesis and Integration Project.
POLICY QUESTION g
5. SOILS: SQ 2.1
S&I Contact: Paul Schroeder
Binkley
SC16
Cline
SC09
Cowling
SC01
Fernandez
SF04
Gholz
SC17
Hodges
SC03
Johnson
SF08
Johnson
SF30
Wells &
SF17
Wells &
SF21
Wong
SC08
Binkley
Robarge
Impacts of sulfate and nitrogen deposition on southern
forest soils
Influence of atmospheric deposition of nitrogen compounds
on mycorrhizal development in the southern forest: A
comprehensive critical review
Critical assessment of the probable role of acid
deposition and other airborne sulfur and nitrogen-derived
pollutants in the forests of eastern North America
An integrated study of atmospheric deposition and
nutrient cycling In a commercial northeast spruce-fir
ecosystem
The development of a location in north Florida as an
Intensive Forest Study (IFS) site
Critical review of the effects of natural and airborne
chemical stresses on the growth and development of
individual trees and forest
Quantifying spruce decline and associated forest
characteristics at Whiteface Mountain, NY
Response of spruce-fir forest soils at Whiteface Mt. to
acidic deposition
Nitrogen transformation in soils of the spruce-fir
ecosystem
Soil and tissue chemical properties associated with stand
characteristics of spruce-fir in the southern
Appalachians
Heavy metal content of tree roots and stems: Role of
mycorrhizae in the accumulation of trace metals under
different pH conditions
38

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6. FOLIAR LEACHING:
SQ 2.2	S&I Contact: Jeff Brandt
Cowling	Critical assessment of the probable role of acid
SC01	deposition and other airborne sulfur and nitrogen-derived
pollutants in the forests of eastern North America
Jacobson	Test of the nitrogen fertilization hypothesis of red
SF06	spruce decline
Jensen & Schier	Impact of ozone and acid deposition on foliar leaching
SF07	and growth of red spruce seedlings
Tingey	Foliar leaching and root uptake of Ca, Mg, and K in
WC07	relation to acid mist effects on conifers
7. CARBON ALLOC:
SQ 2.3	S&I Contact: Charley Peterson
Cowling	Critical assessment of the probable role of acid
SC01	deposition and other airborne sulfur and nitrogen-derived
pollutants in the forests of eastern North America
Davis	Testing the sensitivity of ten eastern hardwood species
EH01	to sulfur dioxide and ozone alone and in combination with
pre- and post-exposures to acidic precipitation
Eckert	Genetic variation in red spruce pollution response
SF23
Flagler	Response of shortleaf pine families to acidic
SC14	precipitation and ozone in East Texas
Fong	Growth responses of loblolly pine seedlings to ozone and
SC02	low-water stress
Greenwood	Evaluation of the impact of atmospheric deposition on red
SF32	spruce seedlings to open top chambers
Hodges	Critical review of the effects of natural and airborne
SC03	chemical stresses on the growth and development of
individual trees and forests
Hogsett	Sensitivity of important western conifer species to S02
WC08	and seasonal interaction of acid fog and ozone
Jacobson	Test of th« nitrogen fertilization hypothesis of red
SF06	spruce decline
Jensen & Dochinger Testing the sensitivity of eleven eastern hardwood
EH06	species to sulfur dioxide and ozone alone and in
combination with pre- and post-exposures to acidic
precipitation
Jensen & Schier	Impact of ozone and acid deposition on foliar leaching
SF07	and growth of red spruce seedlings
Johnson, J.	Response of slash pine families to acidic precipitation
SC13	and ozone in North Florida
Kohut	Comparison of the responses of seedling and sapling red
SF31	spruce exposed to ozone and acidic precipitation under
field conditions
39

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Kossuth
SC11
Kress
SC06
Lockaby
SC15
McGregor
SC12
McLaughlin
SC04
McLaughlin
SF10
Miller, P.
WC09
Reinert & Wells
SC05
Use of clonal trees for acid deposition studies
Response of loblolly pine to acidic precipitation and
ozone stress
Response of loblolly pine families to acidic
precipitation and ozone in Alabama
Response of shortleaf pine families to acidic
precipitation and ozone in South Carolina
Comparative sensitivity mechanisms, and whole plant
physiology implications of responses of loblolly pine
genotypes to ozone and acid rain.
Interactive effects of natural and anthropogenic factors
on the growth and physiology of red spruce
Testing the sensitivity of five western conifer species
to S02 alone, and ozone followed by acidic fog
Comparative responses of loblolly pine families to ozone
adn simulated acid rain
Richardson
SC07
Richardson
SF11
Seiler
SF13
Thornton
SF27
Weinstein
SF16
Unsworth
SF14
Effects of gaseous pollutatns and acid deposition on
open-top chambered loblolly pine seedlings
Effects of atmospheric deposition on red spruce: A free
radical based approach
The impact of ozone and simulated acid rain on the
growth, physiology, and water relations of Fraser fir
A field chamber study of the response of red spruce to
cloud interception and ozone
Effect of ozone and soil nutrient status on the
physiology of photosynthesis, carbohydrate allocation,
nutrition and winter hardiness in red spruce
Frost hardiness of red spruce in relation to forest
decline and effects on red spruce of winter exposure to
S02 and N02
8. WINTER INJURY
SQ 2.4
S&I Contact: Jeff Brandt
DeHayes
SF20
Jacobson
SF06
Unsworth
SF14
Weinstein
SF16
Wilkinson
SF19
Mechanisms of winter injury to red spruce foliage and
possible complcations from supplemental inputs of
nitrogen
Test of the nitrogen fertilization hypothesis of red
spruce decline
Frost hardiness of red spruce in relation to forest
decline and effects on red spruce of winter exposure to
S02 and N02
Effect of ozone and soil nutrient status on the
physiology of photosynthesis, carbohydrate allocation,
nutrition, and winter hardiness of red spruce
Geographic and genetic variation in foliar cuticle
development and biochemistry of epicuticular waxes of red
spruce in relation to winter damage and decline
40

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9. REPRO/REGEN:
SQ 2.5	S&I Contact: Charley Peterson
Bonner	Quality of seed produced in declining spruce-fir forests
SF01	of the eastern U.S.
Christensen	Analysis of Duke Forest permanent plots: Patterns of
SC10	production and mortality of seedlings and trees
Cowling	Critical assessment of the probable role of acid
SC01	deposition and other airborne sulfur and nitrogen-derived
pollutants in the forests of eastern North America
Feret	The impact of environment and genotype on the
SF22	reproductive fitness of Fraser fir from Mt. Rogers and
Mt. Mitchell
Zedaker	Site and stand characteristics associated with potential
SF25	decline and regeneration success of spruce-fir stands in
the Southern Appalachians
10. PEST/PATH:
SQ 2.6	S&I Contact: Greg Reams
Bruck	Interactions of spruce-fir pathogens, insects, and
SF02	ectomycorrhizae on the etiology and epidemiology of
boreal and montane decline in the southern Appalachian
Mountains
Cowling	Critical assessment of the probable role of acid
SC01	deposition and other airborne sulfur and nitrogen-derived
pollutants in the forests of eastern North America
Dull	Developing an aerial photographic technique for
VS13	assessment of visual spruce-fir decline resulting from
air pollution or other environmental stress in the
southern Appalachians
Hodges	Critical review of the effects of natural and airborne
SC03	chemical stresses on the growth and development of
individual trees and forests
Hornig	Studies of spruce-fir decline on Mt. Moosilauke in New
SF05	Hampshire and Berry Pond in Maine
Jensen & Schier	Impact of ozone and acid deposition on foliar leaching
SF07	and growth of red spruce seedlings
Johnson	Quantifying spruce decline and associated forest
SF08	characteristics at Whiteface Mountain, NY
Millers	Case studies of declines of major eastern hardwood forest
EH02	species
Skelly	A manual for diagnosing injury to eastern forest trees:
VS05	Air pollutants, pathogens, insects, and abiotic stresses
Wargo	Relationships of root vitality to decline in spruce-fir
SF15	forests in the Northeastern United States


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POLICY QUESTION 3
11. SEEDLING/
BRANCH/TREE
INTEGRATION
Ford
SI01
Houpis
WC20
Kohut
SF31
Maguire
SF09
Taylor
SC19
Teskey
SC18
Winner
WC32
12. ATMOSPHERIC
EXPOSURE
Cowling
SC01
Cronn
WC06
Larson
WC03
Marx
VS01
Miller, D.
WC04
Mohnen
AE01
Mueller
SF03
Svoboda
WC05
S & i Contact: Ross Kiester
Analysis and modeling of the effect of pollutants on
forests
Comparison of the response of seedlings and mature
branches of Ponderosa pine to air pollution
Comparison of the resonses of seedling and sapling red
spruce exposed to ozone and acidic precipitation under
field conditions
Experimental design and forest modeling for spruce-fir
decline in the southern Appalachians
Air pollutant exposure of mature forest trees:
methodology development and performance evaluation
Field investigations of the impacts of ambient air,
ozone, and tree water relations on net carbon exchange
and growth of loblolly pine trees
Workshop - The response of trees to air pollution: the
role of branch studies
S&I Contact: Jeff Brandt
Critical assessment of the probable role of acid
deposition and other airborne sufur and nitrogen-derived
pollutants in the forests of eastern North America
Development and testing of an improved ozone monitor for
outdoor monitoring
Characterization of fall cloud chemistry at a high
elevation site, Mt. Rainier, WA
Design and pilot test of a long-term monitoring study of
Eastern U.S. forest for respone to atmospheric deposition
Characterization of cloud chemistry and frequency of
canopy exposure to clouds at Mt. Werner, CO
Mountain cloud chemistry project
Acid inputs to high elevation forests in the southeastern
U.S. - cloud contact, wet and dry deposition
Rocky Mountain deposition monitoring project
42

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Appendix II
Forest Response Program
Project Summaries
A3

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Project Number: AE01
Principal Investigator: V. Mohnen
Category: Atmospheric Exposure
Cooperative: Atmospheric Exposure
Scientific Question(s): 1.1, 1.2
Title: Mountain Cloud Chemistry Project (MCCP)
Tree Species: High-elevation spruce-fir forests
Objectives: Characterize the exposure of montane forested ecosystems to
chemical, physical, and climatic atmospheric inputs: a) Determine the
elevational gradients in wet and dry deposition, b) Determine the relative
significance of various deposition mechanisms to the fluxes of chemical
materials into and through forest canopies, c) Determine the frequency
distributions of chemical, physical, and climatic exposure.
Deliverables: Elevational gradients in wet and dry deposition, 9/87.
Relative significance of deposition mechanisms in mountain forests: A
comparison of deposition rates of pollutants in rainfall, cloud impaction and
dry deposition, 9/87. Exposure of forests to gaseous air pollutants and
clouds, 5/87. Primary assessment reports, 12/89.
Summary: Preliminary results indicate for high-elevation forests that:
-	dry deposition of SO- and nitric acid probably does not contribute an
appreciable fraction or total deposition of sulfate and nitrate to mountain
forests subjected to a high frequency of cap clouds;
-	wet deposition of sulfate and nitrate is important at all elevations, and in
some cases contributes the dominant fraction to total deposition, especially
at lower elevations;
-	for mountains with high frequency of cap clouds, cloudwater deposition of
sulfate and nitrate becomes more important as elevation increases and in some
cases is equal to or exceeds the wet deposition contribution to the total;
-	chemical ionic concentrations in cloudwater are usually 3-5 times greater
than those in corresponding rain;
-	hydrogen peroxide concentrations are generally greater than sulfur dioxide
at summit sites during warm seasons, making it possible to oxidize rapidly all
of the sulfur dioxide in clouds to acid, with excess hydrogen peroxide
absorbed in the droplets;
-	during the summer, ozone at the summit of Whiteface Mountain is about 10-20
ppbv greater than at nearby valley sites.
S&I Contact: Jeff Brandt
44

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Project Number: EH01
Principal Investigator: D.D. Davis and J. Skelly
Category: Carbon allocation
Cooperative: Eastern Hardwoods
Scientific Question(s): 2.3
Title: Testing the Sensitivity of Ten Eastern Hardwood Species to Sulfur
Dioxide and Ozone Alone and in Combination with Pre- and Post-exposures to
Acidic Precipitation.
Tree Species: 10 hardwood species
Objectives: The specific objective of this study is to screen seedlings of
eastern hardwood species for relative sensitivity to 03, S02, and acid
precipitation.
Deliverables: Documentation of relative sensitivities of species to S02, 03
and acid precipitation, 12/87, 12/88, spring/89.
Summary: Conducted in close coordination with the Jensen and Dochinger
project. Controlled environment chambers are used to expose seedlings to
various levels of ozone, sulfur dioxide and acidic precipitation. Response
variables include leaf area, stem height,, and stem, root, and leaf dry
weight.
S&I Contact: Charley Peterson
45

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Project Number: EH02
Principal Investigator: I. Millers
Category: Pest/Pathogen
Cooperative: Eastern Hardwoods
Scientific Question(s): 2.6
Tree Species: Eastern hardwood species
Title: Case Studies of Declines of Major Eastern Hardwood Forest Species.
Objective: Examine the historic occurrence of tree mortality events in the
eastern hardwood forests to determine their relationship to known
concentrations of air pollutants.
Deliverables: A report documenting decline case histories. Maps and reports
analyzing possible correlations between declines and deposition patterns.
Summary: The approach was a comprehensive literature review which has been
completed, although the final report must still be peer reviewed. The review
found that many hardwood species have experienced declines and mortality
events during the last century with apparent increase during the last few
decades. The apparent increase may be due to intensification of reporting and
to the maturation of the forest itself. Most of the mortality events are
attiributed to various abiotic and biotic stress factors such as wealther,
silviculture, and damage by insects and diseases. Firm evidence of
atmospheric pollutant damage is present from point source pollution such as
smelters and from ozone damage on white pine. No evidence was found for
hardwood mortality from regional or long range atmospheric pollution.
S&I Contact: Greg Reams
46

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Project Number: EH03
Principle Investigator: J.A. Witter
Category: Spatial Study
Cooperative: Eastern Hardwoods
Scientific Questions: 1.1, 1,2
Title: Effects of an Air Pollution Gradient on Northern Hardwood Forests in
the Southern Great Lakes.
Tree Species: sugar maple, and associated northern hardwood species
Objectives: 1) Establish and characterize analogous research plots in northern
hardwood forests along the pollution gradient in the southern Great Lakes
region. 2) Determine whether atmospheric deposition could be causing
fundamental changes in forest productivity and health. This objective is to
address the following: (a) determine patterns of nutrient cycling that are
related to the regional pollution gradient; (b) determine if growth efficiency
(stemwood production/unit leaf area) and carbon allocation to fine roots and
leaves are related to the pollution gradient; (c) determine if seedling
recruitment and species diversity are related to the pollution gradient; (d)
determine if seedlings from the more polluted sites are more tolerant of
laboratory exposures to S02 and 03 than those from less polluted sites.
Deliverable: Annual report 9/87, 9/88, 9/89, 9/90, Other manuscripts include:
Emissions and Deposition, 3/88; Stand and Plot Descriptions, 4/88; Genetic
Changes in Aspen, 4/89; Root Elongation, 4/89; Nutrient Cycling, 4/90.
Summary: This study is evaluating the role of air-pollutant deposition in
producing significant ecological changes in forest ecosystems.
S&I Contact: Greg Reams
47

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Project Number: EH04
Principal Investigator: D.D. Davis and J. Skelly
Category: Spatial Study
Cooperative: Eastern Hardwood
Scientific Question(s): 1.1, 1.2
Title: Measurement of Forest Condition and Response Along an Atmospheric
Deposition Gradient.
Tree Species: oak-hickory forest types (white and red oaks, hickories, and
associated species).
Objectives: Determine if forest productivity measures (species diversity and
growth) are negatively associated to the level of atmospheric deposition.
Deliverable: Evaluation of deposition gradient on forest growth, competition,
xylem structure and chemistry, and insect and disease incidence and severity.
Annual reports, 9/87, 9/88, 9/89, final report 9/90.
Summary: This study is evaluating forest condition and response along a 100
mile atmospheric gradient in Pennsylvania. They have established 11
atmospheric monitoring stations at 10 mile intervals. The following types of
quantitative analyses have been proposed:
1.	Community analysis (species diversity),
2.	Dendroecology,
3.	Height growth rates.
S&I Contact: Greg Reams
46

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Project Number: EH05
Principal Investigator: 0. Loucks
Category: Spatial Study
Cooperative: Eastern Hardwood
Scientific Question(s): 1.1, 1.2
Title: Investigation of Pollution Effect Patterns in Oak-Hickory Forests,
Arkansas to Southern Ohio.
Tree Species: oak-hickory (not itemized).
Objectives: 1) To establish a set of forest sites in a transect from
southwestern Ohio to Arkansas, along a gradient of wet S04 deposition, but
within a region of similar 03 dose. 2) To select for each study plot a
parallel plot similar in soil type and forest structure located in proximity
to a local fossil-fuel generating source, so the paired plots are exposed to a
significantly different dry deposition and gaseous sulfur loading. 3) To
conduct standardized, intensive measurements at each study area and plot, each
year, in order to detect measurable change in the forest condition along the
study transect, between study areas and plots and annually, so as to quantify
natural variability. (Forest condition is indicated by foliar injury, branch
dieback, intensity of biotic and abiotic stresses, basal area, species
composition, evidence of past disturbances and other characteristics). 4) To
create a data base of these conditions that can be analyzed for possible time
trends over the duration of the study. 5) To characterize the pollutant
environment of each study area for each year of the study through site
monitoring of wet and dry deposited pollutants and synthesis of available
pollutant data. 6) To analyze the collected data for evidence of altered
forest condition along all of part of the study transect attributable to air
pollution effects alone or in combination with other stresses. Determine if
study sites at the high end of the wet sulfate gradient or sites located near
local pollution sources exhibit altered forest conditions, altered stand
community structure or increased incidence of stress responses to pollution in
comparison to study areas at the low end of the gradient.
Deliverable: Annual report 9/87. A fully documented database on pollutant
concentrations and deposition in the region. Report on temporal and geographic
variability and loadings, 12/89.
Summary: The research questions addressed are presented as a comparative
analysis In the form of multiple ANOVA's and multiple regression hypothesis
testing, and the use of various dendrochronological techniques.
S&I Contact: Greg Reams
49

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Project Number: EH06
Principal Investigator: K.F. Jensen and L.S. Dochinger
Category: Carbon allocation
Cooperative: Eastern Hardwoods
Scientific Question(s): 2.3
Title: Testing the Sensitivity of Eleven Hardwood Species to Sulfur Dioxide
and Ozone Alone and in Combination with Post-exposures to Acidic
Precipitation.
Tree Species; 11 hardwood species
Objectives: Screen seedlings of selected tree species for relative
sensitivity to ozone, sulfur dioxide, and acidic precipitation.
Deliverables: Documentation of relative sensitivities of species to S02, 03,
and acid precipitation, 12/87, 12/88, spring/89.
Summary: Conducted in close coordination with the Davis and Skelly project.
CSTR's are used to expose seedlings to various levels of ozone, sulfur dioxide
and acidic precipitation. Response variables include leaf area, stem height,,
and stem, root, and leaf dry weight.
S&I Contact: Charley Peterson
so

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Project Number: EH07
Principal Investigator: I. Millers and D. LaChance
Category: Spatial Studies
Cooperative: Eastern Hardwoods
Scientific Question(s): 1.1, 1.2
Title: North American Sugar Maple Decline Project
Tree Species: Sugar Maple
Objectives: 1. To determine the rate of change in sugar maple tree condition
ratings during the 1988 to 1990 period. 2. To determine if the rate of change
in sugar maple tree condition ratings are different between a) high and low
pollutant (sulfate and nitrate) deposition zones; b) sugar bush and trees in
the forest; and c) in initially rated heavy, moderate and light decline class
stands. 3. To determine the possible causes of sugar maple decline and the
geographical relationship between causes and extent of decline.
Deliverables: Establishment report showing location of plots, and for those
sampled in 1988, the assessment of stand and tree conditions, 1/89. Annual
progress reports. Final report, 6/91.
Summary: This is a cooperative project in conjunction with the Canadian
Forestry Service, Laurentian Forestry Centre, Quebec. The draft work plan and
a Field Manual is in final review and will be completed 1/15/88. Study plots
will be located in the four cooperating Provinces/States. In addition, plots
may also be established in New Brunswick, Maine, Michigan, New Hampshire, and
Wisconsin. The general study areas are being selected and plot establishment
in the field will begin in the summer of 1988. First field tree vigor ratings
will be made in July/August, 1988.
S&I Contact: Greg Reams
51

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Project Number: EH08
Principal Investigator: E.S. Groton and J.M. Kelly
Category: Spatial Studies
Cooperative: Eastern Hardwoods
Scientific Question(s): 1.1, 1.2
Title: Assessment of Factors Contributing to Forest Condition in Mixed
Species Hardwood Stands in the Tennessee Valley Region
Tree Species: Mixed hardwoods
Objectives: 1. To determine the long-term growth patterns of trees on a
variety of sites, including sites with different disturbance histories, sites
with varying degrees of site quality and sites classified as sensitive and
non-sensitive to acidic deposition. 2. To statistically partition out the
factors that influence tree growth, such as increasing stand densities,
topographic location, and climate. 3. To produce maps of gradients in forest
productivity using TVA's Geographic Information System (GIS). 4. To produce
an extensive database on forest productivity and environmental factors for use
in subsequent studies.
Deliverables: 1. Identification of patterns of decline existing in the
region. 2. Characterization of relationship of changing patterns of growth to
naturally occurring factors. 3. Assessment of relationship of forest
productivity to soil parameters, particularly in soils hypothesized to be
sensitive to acidic deposition, 12/88.
Summary: A total of 306 forest plots at 3 study sites have been identified
for remeasurement using the past measurements to identify similarities in
species composition and size of individual trees. Forest inventory fieldwork
on the plots is approximately 80% complete. Disturbance histories have been
completed for 2/3 of the plots. Soil series have been identified for all of
the plots for which surveys have been done. Soil chemical and physical
characterizations for these series have been obtained from ORNL.
S&I Contact: Greg Reams
52

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Project Number: SC01
Principal Investigator: E.B. Cowling
Category: Soils, foliar leaching, carbon allocation, reproduction/regeneration
Cooperative: Southern Commercial
Scientific Question(s): 2.1, 2.2, 2.3, 2.5
Title: Critical Assessment of the Probable Role of Acid Deposition and Other
Airborne Sulfur and Nitrogen-Derived Pollutants in the Forests of Eastern
North America
Tree Species: Southern pines, spruce-fir
Objective: Critically review the literature on the effects of atmospheric
pollutants on eastern forests, with a determination of the likelihood of
effects on southern forests in particular. The SQ's will be covered, but the
soils and carbon allocation mechanisms will be concentrated on. Part of this
effort is a review of emissions data and air quality data.
Deliverables: Critical review of the present state of understanding of sulfur
and nitrogen derived pollutants and prediction of expected increase/decrease
on forests, 9/87.
Summary: The approach is a comprehensive literature review. The final report
has been completed and is being peer reviewed. Among the report's conclusions
were:
1.	Sulfur dioxide in eastern North America currently occurs in concentrations
sufficient to cause visible injury to vegetation only in certain
urban-industrial areas and in rural areas near major pollution sources.
2.	Nitrogen oxides alone have not been detected in concentrations sufficient
to cause visible injury to vegetation more than 50 km. distant from major
pollution sources.
3.	Ozone occurs in concentrations sufficient to cause injury to vegetation in
near-urban, rural, and high elevation forests in large parts of eastern
North America.
4.	Sulfur dioxide and ozone have both been shown to disturb photosynthesis and
to alter carbon allocation between roots and shoots.
5.	Ambient deposition of sulfur and nitrogen compounds provides amounts of
nutrient sulfur and nitrogen that may actually be beneficial for growth of
forest trees in eastern North America.
6.	Ambient deposition of total nutrient nitrogen is more likely to cause
detrimental effects on most eastern North American forests than ambient
deposition of total nutrient sulfur.
7.	Ambient acid deposition has not been proven to cause detrimental effects on
forests by either foliage-mediated or soil-mediated mechanisms.
S&I Contact: Paul Schroeder
53

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Project Number: SC02
Principal Investigator: F. Fong
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Growth Response of Loblolly Pine Seedlings to Ozone and Low-water
Stress.
Tree Species: Loblolly pine
Objectives: 1) Assess responsiveness of loblolly pine as a species to ozone;
quantify the genetic variation in response to ozone, and characterize;
symptomology and mechanisms (physiological responses) of ozone phytotoxicity;
2) characterize physiological responses of seedlings to ozone x low- water
stress interactions.
Deliverables: Reports on responses of loblolly pine to ozone and water
stress, 7/87, 12/88.
Summary: Uses fumigation chambers to quantify the genetic variation in the
response of loblolly pine to ozone and to characterize the symptomology and
mechanisms of ozone phytotoxicity. 30 half-sib families are exposed to
different levels of ozone and moisture stress. Response variables include
plant height, root collar diameter, total fresh and dry weight of needles,
stem, and root, photosynthesis, respiration, and total non-structural
carbohydrate.
S&I Contact: Charley Peterson
34

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Project Number: SC03
Principal Investigator: J.D. Hodges
Category: Soils, carbon allocation, pests and pathogens
Cooperative: Southern Commercial
Scientific Question(s): 2.1, 2.3, 2.6
Title: Critical Review of The Effects of Natural and Airborne Chemical
Stresses on Growth and Development of Individual Trees and Forests.
Tree Species: Loblolly pine
Objectives: Review the literature characterizing the effects of natural
stresses and pollutant stresses on loblolly pine. The natural stresses
include water stress, nutrient stress, 4 major insects/pathogens, and
competition. The major air pollutants and the information known about their
mechanisms will be reviewed. There will also be some effort to describe
interactions between natural stresses and pollutants.
Deliverables: Review of the present understanding of effects of natural and
man-made stresses on forest growth and development, 8/87.
Summary: The approach is a comprehensive literature review which has been
completed, but not yet peer reviewed. Conclusions of the draft report
included:
1.	Indirect effects of acid precipitation on forest productivity may occur
through alterations in forest soils as: (1) a fertilizer effect caused by
the deposition of nitrogen and, under specific conditions, sulfur, (2) an
acidification effect caused by accelerated leaching of base cations, and
(3) an aluminum toxicity effect in cases where soil acidity is considerably
increased.
2.	Although acid precipitation has the capability to accelerate the leaching
of base cations and enhance the solubilization of aluminum in soils, field
studies have failed to demonstrate that these potential effects have
occurred to any great magnitude or over a widespread area in North America.
3.	Heavy metals deposited from the atmosphere or mobilized in soils are of at
least local importance because they may accumulate in soils or in plant
tissues, they have long residence times, and they aretoxic to some plant
species at concentrations that may occassionally occur in the environment.
4.	Direct effects of acid precipitation on physiological processes have been
demonstrated in a variety of species, but only by treatment with simulated
acid precipitation of pH 3.0 or less, which is significantly more acid than
mean ambient precipitation pH.
5.	There is no conclusive evidence that acid precipitation has caused, or is
currently causing detrimental effects on forest productivity, under field
conditions, in North America. However, there is also no evidence that acid
precipitaiton is not a significant stress factor in forests.
6.	Short-term exposures to high concentrations of gaseous pollutants are more
detrimental to photosynthesis and growth than long-term exposures to low
concentrations, at equal doses.
35

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7.	Within broad species groups, reductions in net photosynthesis and growth
due to ozone are linear with respect to either ozone dose or ozone uptake.
8.	Ambient ozone concentrations above 5 pphra are probably sufficient to cause
reductions in net photosynthesis and growth of most types of vegetation,
even in the absence of visible injury symptoms.
S&I Contact: Charley Peterson
58

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Project Number: SC04
Principal Investigator: S.B. McLaughlin
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Comparative Sensitivity, Mechanisms, and Whole Plant Physiological
Implications of Responses of Loblolly Pine Genotypes to Ozone and Acid
Deposition
Tree Species: Loblolly pine
Objectives: 1) Quantify growth and physiological responses of 53 loblolly
pine genotypes to ozone and acid rain in the field and laboratory; compare lab
vs. field results. 2) Develop protocols to quantify physiological and growth
responses of large trees in the field.
Deliverables: Interim report during summer of 1987. Final report 12/88.
Summary: The objectives are met by implementing closely related field and
laboratory studies designed to incorporate many common cultural and
experimental protocols both within the studies at ORNL and across
collaborating sites within the Southern Commercial Coop. Laboratory studies
have focused on testing the physiological responses of 8 common families to
ozone while using the approximate ambient rainfall pH level as a common
background irrigant. Field studies have utilized open top chambers to examine
individual and interactive effects of ozone and simulated acid rain. Response
variables include growth, photosynthesis, carbon metabolism, and mycorrhizal
development.
S&I Contact: Charley Peterson
57

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Project Number: SC05
Principal Investigator: R. Reinert & C. Wells
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Comparative Responses of Loblolly Pine Families to Ozone and Simulated
Acid Rain
Tree Species: Loblolly pine
Objectives: Evaluate loblolly pine "sensitivity" to ozone and acid rain.
Deliverables: Dose-response model, 3/88; Quantification of growth and
physiological responses of various half-sib families to ozone and acid rain,
12/88
Summary: This is a genotype screening study using short-term seedling
exposures in the greenhouse. It seeks to determine relative responses of
different half-sib families of loblolly pine. The primary response variables
are needle injury, stem diameter and height, and biomass.
S&I Contact: Charley Peterson
58

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Project Number: SC06
Principal Investigator: L. Kress
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Response of Loblolly Pine to Acidic Precipitation and Ozone Stress
Tree Species: Loblolly pine
Objectives: 1) Determine responses of a number of loblolly pine families to
ozone exposure. 2) Estimate effects of ambient ozone concentrations on young
loblolly pine. 3) Study ozone x acid rain interactions.
Deliverables: Quantification of loblolly response to ozone and acid rain.
Development of 15' open top chambers. Assessment of exposure effects on
internal nutrient content and mycorrhizal development. Interim report 12/87.
Final report 12/90.
Summary: The approach is to study plant responses over a range of ozone
concentrations. A range of acidic precipitation treatments is utilized in a
factorial design with the ozone concentrations. Phase I uses 10' chambers to
test experimental protocols, assess physiological responses, and provide
initial dose-response estimates. Phase II will use 15' diameter chambers to
study larger trees for longer experimental periods.
S&I Contact: Charley Peterson
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Project Number: SC07
Principal Investigator: C.J. Richardson
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Effects of Gaseous Pollutants and Acid Deposition on Open-top
Chambered Loblolly Pine Seedlings
Tree Species: Loblolly pine
Objectives: 1) Characterize physiological effects of loblolly genotypes
exposed to ozone and acid rain. 2) Establish dose-response relationships:
ozone vs. physiological and biochemical responses. 3) Develop mathematical
relationships between physiology, biochemical responses, and growth across 5
levels of ozone and 2 levels of acid rain. 4) Study relationship between
ozone exposure and status of the antioxidant defense system. 5) Develop
diagnostic measurements for ozone exposure (e.g. light response curves).
Deliverables: Annual Reports 1/88, 12/88; Quantification of physiological and
biochemical responses and relationship to growth.
Summary: Uses open-top chambers, a range of ozone treatments, and two levels
of simulated acid rain. Physiological measurements include survey
measurements of photosynthesis, transpiration, and stomatal conductance.
Monthly measurements of photosynthetic responses at both high and low
irradiation levels for each treatment group will be coupled with measurements
of transpiration, stomatal conductance, chlorophyll and carbohydrates. Dose
response relationships will also be developed.
S&I Contact: Charley Peterson
60

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Project Number: SC08
Principal Investigator: B. Wong
Category: Soils
Cooperative: Southern Commercial
Scientific Question(s): 2.1
Title: Heavy Metal Content of Tree Roots and Stems: Role of Mycorrhizae in
the Accumulation of Trace Metals under Different pH Conditions.
Tree Species: Loblolly pine
Objectives: Characterize and quantify trace metal uptake and accumulation by
mycorrhizae of loblolly pine seedlings, as well as the effects on seedling
growth.
Deliverables: Evaluation of trace metal accumulation and the role of acid
deposition, 3/88.
Summary: Mycorrhizal and non-mycorrhizal seedlings were treated with three
levels of Zn, Pb, or Al at pH 3.0, 4.0, or 5.0. Observed response variables
were per cent mycorrhizal infection, root and shoot lengths, root and shoot
fresh and dry weights, root morphology, and metal concentration.
Pb uptake was the same for mycorrhizal and non-mycorrhizal seedlings.
Seedling Pb concentration increased as treatment Pb concentration increased.
Root Pb content was lower at pH 3.0 than at 4.0 or 5.0.
S&I Contact: Paul Schroeder
61

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Project Number: SC09
Principal Investigator: M. Cline
Category: Soils
Cooperative: Southern Commercial
Scientific Question(s): 2.1
Title: Influence of Atmospheric Deposition of Nitrogen Compounds on
Mycorrhizal Development in the Southern Forest: a Comprehensive Critical
Review
Tree Species: Loblolly pine
Objectives: To develop a comprehensive critical review of all available
knowledge which addresses the problem of toxic effects of atmospheric
deposition of nitrogen compounds on mycorrhizal development in southern
forests.
Deliverables: Critical review of understanding of the effect of nitrogen
deposition on mycorrhizal development, 9/87.
Summary: The approach is a comprehensive literature review which has been
completed, but not yet peer reviewed. To date, no studies have adequately
addressed the direct impact of atmospherically deposited N on mycorrhizae.
Therefore, for this analysis the influence of N on mycorrhizae was assessed by
reviewing the many studies examining the effect of N fertilization on
mycorrhizal development. Field, nursery, greenhouse, and pure culture
experiments fail to show direct toxicity of high levels of N to mycorrhizal
fungi. Reported decreases in the percentage of short roots converted to
mycorrhizae are probably atrlbutable to changes in host root growth rate
and/or methods used to quantify mycorrhizal infection.
The complexity of the mycorrhizal association prevents the use of generalized
statements regarding the effects of any factor on mycorrhizal development.
Variables influencing mycorrhizal response to N deposition include: fungal
species, host species, chronological age of the host, edaphic and climatic
conditions, and host and fungal succession. However, regardless of the number
of combinations of factors considered in the literature, no studies revealed N
to be directly toxic to mycorrhizae at current levels of atmospheric
deposition in the southeastern U.S.
S&I Contact: Paul Schroeder
62

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Project Number: SC10
Principal Investigator: N. L. Christensen
Category: Fixed Plot
Cooperative: Southern Commercial
Scientific Question(s): 1.1, 1.2
Title: Analysis of Duke Forest Permanent Plots: Patterns of Production and
Mortality of Seedlings and Trees
Tree Species: Various hardwood and softwood species
Objectives: 1. Resurvey seedling mapped plots located in 20 forest stands
stratified by site conditions and successional age. 2. Analysis of seedling
data including comparisons of growth and mortality among sites, using profile
analysis to examine year-to-year patterns. 3. Resurvey Duke Forest (and
nearby) Permanent Sample Plots (PSP). 4. Analyze PSP data for changes in
aboveground production during various intervals over the past 50 years,
focusing on changes in the last decade relative to previous periods. 5.
Collaboration in development of forest growth models and projections.
Deliverables: Final report on seedling plots, 6/87. Final report on PSP
plots, 12/87.
Summary: The project documents changes in production in Duke Forest stands
over the past 50 years, taking into account changes related to stand age,
initial stocking, site conditions, and management interventions. Patterns of
growth and mortality of tree seedlings during the past 8 years in relation to
stand successional status, site conditions, and year-to-year variations in
climate will also be documented. An important product will be a model of
production which can be applied to other data sets to determine whether
changes in production observed during any time interval are related to the
factors described or may be due to changes in atmospheric deposition. Data
will also provide a baseline for future studies of patterns of forest
regeneration.
S&I Contact: Ross Kiester
63

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Project Number: SC11
Principal Investigator: S. Kossuth
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Use of Clonal Trees for Acid Deposition Studies
Tree Species: Slash pine
Objectives: Evaluate the potential for use of clonal and grafted southern
pine material in exposure research.
Deliverables: Evaluation of the use of clonal material for air pollution
studies 1/88.
Summary: Ramets from pine clones are grafted onto a half-sib rootstock. The
grafts are then exposed to various levels of ozone in open top chambers.
Response variables include height, photosynthesis, chlorophyll, and carbon
partitioning.
S&I Contact: Charley Peterson
6 *

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Project Number: SC12
Principal Investigator: D. McGregor
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Response of Shortleaf Pine Families to Acidic Precipitation and Ozone
in South Carolina.
Tree Species: Shortleaf pine
Objectives: Determine the influences of acidic precipitation and ozone on the
growth, nutrition, and physiology of shortleaf pine under field conditions.
Deliverables: Quantification of Shortleaf Pine Response to Acid Precipitation
and Ozone, 6/91.
Summary: One year old loblolly pine seedlings are grown in 15' open top
chambers. They are exposed to 3 levels of pH and 4 ozone concentrations
curing the growing season. Treatment effects are quantified on plant dry
weight, stem diameter, cumulative height growth, fascicle length, and leaf
area.
S&I Contact: Charley Peterson
63

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Project Number: SC13
Principal Investigator: J. Johnson
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Response of Slash Pine Families to Acidic Precipitation and Ozone in
North Florida.
Tree Species: Slash pine
Objectives: Determine the influences of acidic precipitation and ozone on the
growth, nutrition, and physiology of slash pine under field conditions.
Deliverables: Quantification of slash pine response to acid precipitation and
ozone, 6/91.
Summary: One year old seedlings are grown in open top chambers. They are
exposed to 3 levels of pH and 4 ozone concentrations. Treatment effects are
quantified on plant dry weight, stem diameter, total height, average fascicle
length of the current flush, total leaf area, and visible damage.
S&I Contact: Charley Peterson
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Project Number: SC14
Principal Investigator: R.B. Flagler
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Response of Shortleaf Pine Families to Acidic Precipitation and Ozone
in East Texas
Tree Species: Shortleaf pine
Objectives: Determine the influence of acidic precipitation and ozone on the
growth, nutrition, and physiology of shortleaf pine under field conditions.
Deliverables: Quantification of shortleaf pine response to acid precipitation
and ozone, 6/91.
Summary: One year old seedlings are grown in open top chambers. They are
exposed to 3 levels of pH and 4 ozone concentrations. Treatment effects are
quantified on plant dry weight, stem diameter, total height, fascicle length,
and visible injury,
S&I Contact: Charley Peterson
87

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Project Number: SC15
Principal Investigator: B.G. Lockaby and A.H. Chappelka
Category: Carbon allocation
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Response of Loblolly Pine Families to Acidic Precipitation and Ozone
in Alabama.
Tree Species: Loblolly
Objectives: Determine the influences of acidic precipitation and ozone on the
growth, nutrition, and physiology of loblolly pine under field conditions.
Deliverables: Quantification of loblolly pine response to acidic
precipitation and ozone, 6/91.
Summary: One year old loblolly pine seedlings are grown in 15' open top
chambers. They are exposed to 3 levels of pH and 4 ozone concentrations
during the growing season. Treatment effects are quantified on above ground
plant biomass, by component part, stem diameter, cumulative height growth, and
visible injury.
S&I Contact: Charley Peterson
68

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Project Number: SC16
Principal Investigator: D. Binkley
Category: Soils
Cooperative: Southern Commercial
Scientific Question(s): 2.1
Title: Impacts of Sulfate and Nitrogen Deposition on Southern Forest Soils
Tree Species: Loblolly pine
Objectives: 1) Synthesize biogeochemical impacts of sulfate and nitrogen; 2)
examine available empirical evidence relating to 'the effects of sulfate and
nitrogen on southern forest ecosystems; and 3) use computer simulation models
to explore soil sensitivity to sulfate deposition.
Deliverables: Critical review of effects of acid deposition on soil nutrient
status in southern commercial forests, 12/87.
Summary: This project is assessing the potential effects of sulfate and
nitrogen deposition by:
1. Synthesizing the biogeochemical impacts of sulfate and nitrogen.
.2. Examining available empirical evidence relating to the effects of sulfate
and nitrogen in southern forest ecosystems.
3.	Using computer simulation models to explore soil sensitivity to sulfate
deposition.
4.	Relating the the results of these three analyses to major types of southern
forest soils.
S&I Contact: Paul Schroeder
69

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Project Number: SC17
Principal Investigator: H. Gholz
Category: Soils
Cooperative: Southern Commercial
Scientific Question(s): 2.1
Tree Species: Slash pine
Title: The Development of a Location in North Florida as an Intensive Forest
Study (IFS) Site
Objective: Evaluate the effects of acid deposition on forest nutrient
cycling.
Deliverables: Evaluation of the effects of acid deposition on forest nutrient
cycling, 6/90.
Summary: This is a field study conducted in a 20 year old stand of slash
pine. Nutrient cycling is determined from measurements of deposition,
throughfall and stemflow, and soil and soil water chemistry. Soil budgets
will account for spatial and temporal variation of the soil properties which
influence water and solute movement. Laboratory studies are conducted to
characterize ion movement and exchange when needed. Co-located with this
site, is an EPA project which provides air quality data. Also closely
coordinated with this study is an EPRI project.
S&I Contact: Paul Schroeder
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Project Number: SC18
Principal Investigator: R. Teskey and P. Dougherty
Category: Carbon allocation, seedling/branch/tree integration
Cooperative: Southern Commercial
Scientific Question(s): 2.3
Title: Field Investigation of the Impacts of Ambient Air, Ozone, and Tree
Water Relations on Net Carbon Exchange and Growth of Loblolly Pine Trees
Tree Species: Loblolly pine
Objectives: 1) Compare impacts of pollutants and environmental stresses on
physiology and growth of loblolly pine trees in a plantation; 2) Examine
effects of ozone and water availability on leaf area and physiology to
determine if there is an interaction between ozone and water stress; 3)
Determine to what extent ozone and other environmental factors are limiting
tree growth under ambient conditions; and 4) Develop a model relating carbon
gain to pollutant impacts and associated environmental factors.
Deliverables: Evaluation of the effect of ambient air, ozone, and water
stress on net carbon exchange and growth of loblolly pine, 3/90.
Summary: This study is conducted on paired irrigated and non-irrigated plots
on an 11 year old loblolly pine plantation. Small open top chambers (0.6m x
1.5m) are mounted on the terminal end of selected branches. Branches inside
the chambers are exposed to various levels of ozone throughout the entire
year. Response variables include net carbon exchange, dark respiration, leaf
conductance, xylem pressure potential, needle growth, needle senescence,
branch growth, and tree height and diameter growth.
S&I Contact: Charley Peterson
71

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Project Number: SC19
Principal Investigator: G. Taylor
Category: Seedling/branch/tree integration
Cooperative: Southern Commercial
Scientific Question(s): 3.1
Title: Air Pollutant Exposure of Mature Forest Trees: Methodology
Development and Performance Evaluation.
Objective: Develop a range of methodologies for the field exposure of mature
forest trees to wet and dry deposition of airborne chemicals.
Deliverables: Evaluation of methodologies for air pollutant exposure of
mature trees. 12/89
Summary: Uses wind tunnel facilities to evaluate the feasibility of various
design options including open top chamber and forced air systems.
S&I Contact: Ross Kiester
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Project Number: SF01
Principal Investigator: F.T. Bonner
Category: Reproduction/Regeneration
Cooperative: Spruce-fir
Scientific Question(s): 1.2, 2.5
Title: Quality of Seed Produced in Declining Spruce-Fir Forests of the
Eastern United States.
Tree Species: red spruce, balsam fir, and fraser fir.
Objective: Evaluate and compare seed "quality" in declining and healthy
stands of red spruce, balsam fir, and fraser fir.
Deliverables: Comparison of seed quality in healthy and declining forests,
6/88
Summary: Evaluation of seed quality proceeds along two lines: 1) seed
condition at dispersal, and 2) potential for retention of quality. Initial
seed condition is evaluated by size and weight of the embryos and germination
response to standard tests. Description and measurement of embryo development
is augmented with radiographs of seeds. Seed potential for retention of
quality is evaluated by the use of seed vigor tests: electrolyte leakage,
initial seedling growth, and accelerated aging.
S&I Contact: Charley Peterson
73

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Project Number: SF02
Principal Investigator: R.I. Bruck
Category: Pests/pathogens
Cooperative: Spruce-fir
Scientific Question(s): 2.6
Title: Interactions of Spruce-Fir Pathogens, Insects, and Ectomycorrhizae on
the Etiology and Epidemiology of Boreal and Montane Decline in the Southern
Appalachian Mountains.
Tree Species: red spruce, fraser fir
Objective: Develop techniques for continuous monitoring of pathogens,
insects, and ectomycorrhizae. Develop a statistical assessment of the impact
of biotic pests ana pathogens on spruce-fir ecosystems.
Deliverables: Key of pest/pathogen/ectomycorrhizae in southern Appalachians;
assess pest/pathogen damage to spruce-fir; sampling technique for
ectomycorrhizae , 6/88
Summary: Dominant and co-dominant spruce and fir are selected and visually
assessed for gross pathology and insect damage. Decline classes and foliar
damage (chlorosis/necrosis) are assessed. Roots and foliage are returned to
the lab for microbiological and ectomycorrhizal analysis.
S&I Contact: Greg Reams
7A

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Project Number: SF03
Principal Investigator: S.F. Mueller
Category: Atmospheric Exposure
Cooperative: Spruce-Fir
Scientific Question(s): 1.1, 1.2
Title: Acidic Inputs to High Elevation Forests in the Southeastern United
States - Cloud Contact, Wet and Dry Deposition.
Tree Species: High-elevation spruce-fir forests
Objectives: 1) Characterize form, frequency, and magnitude of pollutant
deposition to typical high elevation forest systems in southeastern United
States, in particular to determine the level of exposure to clouds and cloud
water-borne pollutants, precipitation-borne pollutants, and dry deposited
gaseous and particulate pollutants. 2) Characterize chemical and physical
processes occurring in precipitating and non-precipitating cloud systems.
Deliverables: Deposition database for high-elevation spruce-fir forests,
1/87, 1/88, 1/89. Characterization of deposition climatology, 3/89.
Comparison of SE and NE deposition climatology, 9/89.
Summary: Preliminary estimates of the seasonal dry, wet, and cloud-water
deposition of S02, SO*, 0,, N03, and other species have been completed.
Results suggest substantial deposition of ozone (approaching 100 kg/ha/yr),
peaking in the spring of 1986. Sulfate deposition appears to be between 275
and 550 kg/ha/yr. Roughly 90% of the total sulfate and nitrate deposition
were by cloud impaction. Likewise, one-half to two-thirds of the summit
canopy gross water input comes from cloud impaction.
S&I Contact: Jeff Brandt
75

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Project Number: SF04
Principal Investigator: I. Fernandez
Category: Soils
Cooperative: Spruce-Fir
Scientific Question(s): 2.1
Title: An Integrated Study of Atmospheric Deposition and Nutrient Cycling in
Commercial Spruce-fir Ecosystems.
Tree Species: Red spruce, balsam fir
Objectives: At the Howland, Maine intensive site, determine: 1) the effect of
acid deposition on cation leaching, mineral weathering, and the balance of key
elements in the soil; 2) the relationship between soil properties and natural
stress factors; 3) the relationship between soil chemical properties and the
quality of throughfall and 4) the extent and accumulation of trace metals in
forest floor and mineral soil.
Deliverables: Evaluation of effects of atmospheric deposition on nutrient
cycling In commercial spruce-fir forests, 12/89, 12/90.
Summary: This project uses an intensive process-level approach to
quantitatively evaluate the influence of atmospheric deposition on nutrient
cycling. This approach will provide information necessary to assess
transformations and fluxes of key elements such as sulfur and nitrogen, as
well as to quantify element pools. Two stands were selected to represent
relatively mature, low elevation, commercial spruce-fir forests that can be
compared to the high elevation stands being studied by EPRI. Components being
monitored are throughfall, stemflow, soil solution (with lysimeters), plant
tissue, soils, literfall, roots, and nitrogen mineralization (with buried bag
incubations).
S&I Contact: Paul Schroeder
76

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Project Number: SF05
Principal Investigator: J.F. Hornig
Category: Dendrochronology Data, spatial studies, pest/pathogen
Cooperative: Spruce-Fir
Scientific Questions(s): 1.1, 1.2, 2.6
Title: Studies of Spruce-fir Decline on Mt. Moosilauke in NewHampshire and
Berry Pond in Maine.
Species: Red spruce, balsam fir
Objective: 1. To ground truth interpretations of aerial photographs, locate
major spruce stands, and provide information needed to establish the main
study area for intensive permanent plots;
2. To establish permanent intensively measured study plots to (i) document the
present structure, composition and condition of the forest vegetation and (ii)
monitor changes in growth rate, structure, composition, condition through
time.
Deliverables: Evaluation of the relationship of spruce-fir growth and vigor
to climate, pest/pathogen occurrence, natural cycles of disturbance, and site
specific forest stand dynamics, 2/90.
Summary: This is an extensive project that embodies mensuration, pathology,
entomology and soils. A major component is a dendroecological study. Reference
is made to both the standard dendrochronological literature and some of the
more recent developments. This includes techniques to study intra-ring
variations in wood density.
S&I Contact: Bill Warren
77

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Project Number: SF06
Principal Investigator: J.S. Jacobson and J. Lassoie
Category: Foliar leaching, carbon allocation, winter injury
Cooperative: Spruce-Fir
Scientific Question(s): 2.2, 2.3, 2.4
Title: Test of the Nitrogen Fertilization Hypothesis of Red Spruce Decline.
Tree Species: Red spruce
Objectives: Determine if combinations of sulfate and nitrate acidic mist
alter the growth, development, cold tolerance, or water relations of red
spruce seedlings in ways that might contribute to premature decline.
Deliverables: Dose-response relationship of acid mist vs. seedling growth,
development, phenology, water relations, nutrient, balance, and biochemistry,
3/89.
Summary: Red spruce seedlings are exposed repeatedly, for extended durations,
to simulated acidic mist at levels of acidity, sulfate, and nitrate
concentrations that range from above to below those found in ambient wet
deposition at high elevations in spruce-fir forests. Cold tolerance tests are
performed in the fall as seedlings enter dormancy and in the early spring as
they break dormancy. Needles and roots are analyzed for total nitrogen and
sulfur concentration. Seasonal measurements are taken of needle water
potential. Other measurements include needle diffusive conductance and
transpiration, total chlorophyll, and cuticular wax content.
S&I Contact: Charley Peterson
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Project Number: SF07
Principal Investigator: K.F. Jensen and G.A. Schier
Category: Foliar leaching, carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.2, 2.3
Title: Impact of Ozone and Acid Deposition on Foliar Leaching and Growth of
Red Spruce Seedlings
Tree Species: Red spruce
Objectives: 1) Determine effects of acid rain and ozone on photosynthesis and
water relations of spruce seedlings. 2) Determine if acid rain and ozone
reduce root tip growth of red spruce.
Deliverables: Dose-response of acid rain and ozone on red spruce needle
leachate, 3/88. Dose-response of ozone and acid rain on red spruce needle
development and physiology, 9/88. Effect of ozone and acid rain on spruce
budworm development, 9/88.
Summary: Red spruce seedlings are exposed to ozone and acid rain treatments
in CSTR's. Photosynthesis is measured periodically and the composition of
foliar leachate is analyzed. At harvest, needle, stem, and root dry weights,
total starch, and sugar contents are measured. Supplemental tests will
involve infestation of treated and untreated seedlings with spruce budworm
larvae.
S&I Contact: Charley Peterson
79

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Project Number: SF08
Principal Investigator: A. Johnson
Category: Spatial Study, soils
Cooperative: Spruce-Fir
Scientific Question: 1.1, 1.2, 2.6
Title: Quantifying Spruce Decline and Associated Forest Characteristics at
Whiteface Mt., New York.
Tree Species: Red spruce, balsam fir
Objectives: 1) Determine the nature and extent of dieback and decline in
stands of varying age, exposure and elevation (40 permanent plots installed
and measured). 2) Determine of nutrient content, available Al content or
degree of decline (30 quantitative soil pits excavated). 3) Determine if
deficiencies in foliar nutrients are related to the nature and degree of red
spruce decline (foliar tissue sampled and preserved). 4) Determine (a) which
pathogens are present in twigs, bark, and foliage and if those pathogens can
cause apical dieback; (b) how these organisms are associated with the nature
and degree of the decline (summer and fall sampling completed).
Deliverable: Determination of extent of dieback and decline. Evaluation of
effects of soil and foliar chemistry and pathogens of spruce decline. 12/90
Summary: A series of 305 extensive plots will be established to stratify the
area for placement of 30 permanent plots. The 305 plots can be used to reflect
conditions over the study area. The research includes soil surveys, insect and
disease surveys, assessment of forest history, dendrochronology, and
development of growth models for red spruce and balsam fir.
S&I Contact: Greg Reams
80

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Project Number: SF09
Principal Investigator: L. Maguire
Category: Seedling/branch/tree integration
Cooperative: Spruce-Fir
Scientific Questions: 3.1
Title: Experimental Design and Forest Modeling for Spruce-Fir Decline in the
Southern Appalachians.
Tree Species: Red spruce, Fraser fir
Objectives: Implement a FORET-type model of spruce-fir forest growth and
composition as a tool for synthesizing existing information about southern
Appalachian spruce-fir,
Deliverables: Document analyzed version of FORET to represent effects of air
pollution on spruce-fir forests. 5/88.
Summary: Simulates hypothetical changes in species growth, reproduction, and
mortality which might be caused by air pollution and its effects on forest
growth and composition.
S&I Contact: Ross Kiester
81

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Project Number: SF10
Principal Investigator: S.B. McLaughlin
Category: Carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: Interactive Effects of Natural and Anthropogenic Factors on Growth and
Physiology of Red Spruce
Tree Species: Red spruce
Objectives: 1) Determine dose/response of HN03 H202 Al, and Mn on spruce
seedlings, 2) Characterize differences in gas exchange, carbon allocation, and
growth patterns across a "gradient" in decline and presumed deposition.
Deliverables: 1) Characterization of physiological changes associated with
declining red spruce. 2) Determination of effects of nitric acid vapor,
hydrogen peroxide, Al, and Mn, 10/88.
Summary: This project includes both field and laboratory components. The
field study is a comparison of one high and one low elevation site in the
Great Smoky Mountain National Park. Growth is estimated and monitored on
canopy and sapling trees at each site. Photosynthetic capacity, respiration,
and water relations are estimated on saplings. Carbon metabolism studies are
also conducted using C14 techniques.
In the laboratory, red spruce seedlings are exposed to various levels of H202
and NO,. Height, diameter, photosynthesis, and nitrogen reductase activity
are all measured. Screening techniques are being developed for examining the
toxicity of red spruce to individual and combined trace metals.
S&I Contact: Charley Peterson
82

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Project Number: SF11
Principal Investigator: C.J. Richardson
Category: Carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: Effects of Atmospheric Deposition on Red Spruce: a Free Radical Based
Approach
Tree Species: Red spruce
Objectives: To determine if the key mechanism for atmospheric imposed stress
in forest vegetation is the generation of oxygen-based free radicals by
photochemical oxidants in the tissues of affected plants.
Deliverables: Evaluation of radical formation in leaf tissue of red spruce,
11/87.
Summary: Seedlings from the Weinstein project that have previously been
exposed to different ozone treatments are used. Analyses include activities
of superoxide dismutase and peroxidase, and concentrations of glutathione and
malondialdehyde.
S&I Contact: Charley Peterson
83

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Project Number: SF13
Principal Investigator: J.R. Seller and B.I. Chevone
Category: Carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: The Impact of Ozone and Simulated Rain on the Growth, Physiology, and
Water Relations of Fraser fir
Tree Species: Fraser fir
Objectives: 1) Determine growth response of Fraser fir to ozone, acid rain,
and water stress. 2) Evaluate the effect of moisture stress conditioning on
the growth response of Fraser fir to ozone and water stress. 3) Examine the
impact of moisture stress, ozone, and acid rain on photosynthesis,
transpiration, leaf conductance, and needle water potential parameters.
Deliverables: Evaluation of changes in physiology, growth, and carbon
allocation due to moisture stress, ozone, and acid rain. Evaluation of leaf
water potential components and changes due to ozone and moisture stress, 9/88.
Summary: Uses CSTR's to study the effects of ozone, acid rain, and water
stress on four year old Fraser fir seedlings. Growth is measured on root
collar diameter and height. Photosynthesis, transpiration, and needle
conductance are measured at the beginning, middle, and end of experiments.
Root, shoot, and needle dry weights, root length, and water potential are also
measured.
S&I Contact: Charley Peterson
84

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Project Number: SF14
Principal Investigator: M.H. Unsworth
Category: Winter injury
Cooperative: Spruce-Fir
Scientific Question(s): 2.4
Tree Species: Red spruce
Title: Frost Hardiness of Red Spruce in Relation to Forest Decline and
Effects of Winter Exposure to S02 and N02.
Objectives: Determine if S02, N02, S04 acidic mist and 03: i) alter frost
hardening and ii) result in winter accumulation of phytotoxic substances that
alter metabolism and growth of red spruce.
Deliverables: Evaluation of risk of frost injury based on weather data and
field observations. Experimental evaluation of relationship of S, N, and
acidity to physiology and biochemistry of frost hardiness. Model relating
deposition to risk of frost injury, 1/89.
Summary: Shoots excised from red spruce growing at Whiteface Mt., NY, were
tested in Scotland for frost hardiness. Results indicate that susceptible
trees were likely to be damaged in 6 of the last 7 years at Newfound Gap and
in 4 of the last 7 years at Whiteface Mt. An exposure facility for simulated
acid mist has been established. Filtered and unfiltered chambers will be used
to test for interactions with air quality. Preliminary results indicate that
shoots exposed to S02 + N02 were significantly more damaged than controls
after freezing to -4 and -7°C. Ion chromatographic analyses of xylem sap from
red spruce shows increased nitrate, nitrite, sulfite, and sulfate in sap from
trees exposed to S02 + NO,. The study has also developed NMR techniques to
measure intracellular pH in spruce needles.
S&I Contact: Jeff Brandt
85

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Project Number: SF15
Principal Investigator: P. Wargo
Category: Pest/Pathogen
Cooperative: Spruce-fir
Scientific Questlon(s): 2.6
Title: Relationship of Root Vitality to Decline in Spruce-Fir Forests in the
Northeastern United States.
Tree Species: Spruce-fir
Objective: Characterize root systems of declining trees, and determine if
crowns are deteriorating before, with, or after root mortality. Determine if
pathogens are associated with woody roots or fine roots, including
predisposition to root pathogens.
Deliverables: Evaluation of relationships between crown condition and root
condition, carbohydrate reserves, elevation, mycorrhizal infection, and
elevation, 12/88.
Summary: This study is an intensive description and comparison of crown and
root characteristics of trees exhibiting varying degrees of decline. The
crown description includes number of branches snowing dieback, chlorosis,
necrosis, or brooming. Portions of root systems are excavated and their
morphology characterized. Pathogen isolations are also carried out and
carbohydrate reserves are estimated.
S&I Contact: Greg Reams
86

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Project Number: SF16
Principal Investigator: L. Weinstein
Category: Carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: Effect of Ozone and Soil Nutrient Status on the Physiology of
Photosynthesis, Carbohydrate Allocation, Nutrition, and Winter Hardiness in
Red Spruce
Tree Species: Red spruce
Objectives: Evaluate effects of ozone on the physiology, growth, and
development of spruce seedlings using open-top chambers. Study effects on
chloroplast function, respiration, carbon assimilation and allocation.
Deliverables: Dose-response relationships between ozone exposure and various
physiological response parameters, 1/88.
Summary: Red spruce seedlings are exposed to various levels of ozone in open
top chambers. Variables measured include those related to photosynthesis,
carbohydrate production and translocation, and shoot and root growth.
S&I Contact: Charley Peterson
87

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Project Number: SF17
Principal Investigator: C. Wells & D. Blnkley
Category: Soils
Cooperative: Spruce-Fir
Scientific Question(s): 2.1
Title: Nitrogen Transformation in Soils of the Spruce-fir Ecosystem.
Tree Species: Red spruce, balsam fir, Fraser fir.
Objectives: Determine the relationship of elevation and stand characteristics
to mineral nitrogen pools, nitrogen mineralization, and denitrification in the
southern Appalachians.
Deliverables: Seasonal characterization of nitrogen cycle in relation to
elevation, exposure, and stand characteristics. Evaluation of potential for
soil nitrogen to affect tree nutrition, seasonal nitrogen availability, cold
hardening, and excess Nitrogen leaching losses, 9/87.
Summary: Nitrogen transformation data were collected from several plots at
high and low elevations on a number of southern Appalachian mountains. Higher
atmospheric inputs were expected at higher elevations. Nitrogen
mineralization was determined for four sequential intervals from 11/85 to
10/86.
N mineralization was high relative to other forests and nitrate concentrations
were large enough to induce high cation leaching. Nitrate and ammonium
concentrations were approximately constant from June to October. There was
also an indication that the nitrate/ammonium ratio on Mt. Mitchell was higher
on the exposed high elevation west slope than on the low elevation east slope.
Denitrification was undetectable or very low at all locations.
This project has been completed
S&I Contact: Paul Schroeder
88

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Project Number: SF18
Principle Investigator: T.R. Wentworth
Category: Spatial Study
Cooperative: Spruce-Fir
Scientific Question(s): 1.1, 1.2
Title: Compilation and Interpretation of the Vegetation Data Base and
Disturbance History of Southern Appalachian Spruce-Fir.
Tree Species: spruce-fir
Objectives: 1) Provide a thorough bibliography of research on southern
Appalachian spruce-fir vegetation, 2) assemble the historic data base for the
southern Appalachian spruce-fir ecosystem, 2b) assemble field evidence of
disturbance history and integrate with historic records, 3) assemble historic
disturbance records for the 3 intensive study sites - Great Smoky Mountains
National Park, Mt. Mitchell areas, and Mt. Rogers area, 4) Synthesize
information from the above objectives, with emphasis on: (a) natural and human
impacts on stand structure and stand dynamics in southern Appalachian
spruce-fir forests, (b) the implications of such impacts for testing of
hypotheses regarding recent effects of environmental perturbations, 5)
establish photo-monitoring plots for documentation of future change in
spruce-fir forests and comparison with historic photos where possible.
Deliverable: Bibliography of research on southern spruce-fir historical data
base. Photo-monitoring network to document future changes. 6/88.
Summary: The major component of this study is the compilation of an annotated
bibliography of research conducted on southern Appalachian spruce-fir forests.
Written records, historical maps, aerial photographs, and oral history
accounts were used to compile disturbance histories of the SARRMC intensive
study sites.
S&I Contact: Greg Reams
89

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Project Number: SF19
Principal Investigator: R. Wilkinson
Category: Winter injury
Cooperative: Spruce-Fir
Scientific Question(s): 2.4
Title: Genetic Variation in Foliar Cuticle Development and Biochemistry of
Epicuticular Waxes of Red Spruce in Relation to Winter Damage and Decline.
Tree Species: Red spruce
Objectives: Define the range of genetic expression in anatomical and
biochemical needle characteristics of red spruce related to winter desiccation
injury along elevational and ecological zones of winter injury susceptibility
to provide 1) information on possible natural biological basis for spruce
decline 2) the means for evaluating the potential for adaptive evolutionary
change and future resurgence of red spruce.
Deliverables: Influence of needle anatomy and epicuticular waxes on red
spruce transpiration rates and winter injury, 10/88.
Summary: Geographic seed sources of red spruce of known susceptibility to
winter desiccation injury growing in a range-wide provenance test plantation,
and field-planted half-sib progeny and their female parents from stands
growing at a series of different elevations in northern Vermont will be
examined for:
1.	Anatomical development of the needle cuticle and cuticular layers in the
epidermis.
2.	Gravimetric rate of water loss from needles.
3.	Quantity of epicuticular wax on needle surfaces.
4.	Qualitative and quantitative concentrations of the homologous series of
n-alkanes in epicuticular wax.
S&I Contact: Jeff Brandt
90

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Project Number: SF20
Principal Investigator: D. DeHayes
Category: Winter Injury
Cooperative: Spruce-Fir
Scientific Question(s): 2.4
Title: Mechanisms of Winter Injury to Red Spruce Foliage and Possible
Complications from Supplemental Inputs of Nitrogen
Species: Red spruce
Objectives: 1) Examine seasonal/developmental patterns of cold tolerance,
photosynthesis, and foliar nitrogen content. 2) Determine the extent to which
elevated and/or untimely inputs of nitrogen perturb seasonal patterns of
photosynthesis and cold tolerance.
Deliverables: Description of the mechanism of winter injury in red spruce and
possible alterations in spruce cold hardiness due to the excess or untimely
input of nitrogen, 6/88.
Summary: This project is assessing potential effects of sulfur and nitrogen
through studies of the developmental physiology of cold tolerance and
evaluations of seasonal patterns of photosynthesis and foliar nitrogen content
in red spruce and balsam fir. Preliminary results suggest large tissue
differences in susceptibility to winter injury, and refute the hypothesis that
damage results from direct injury to twig Dark (and subsequent injury to
foliage), since even severely injured shoots are capable of supporting healthy
growth the following season.
S&I Contact: Jeff Brandt
91

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Project Number: SF21
Principal Investigator: C. Wells and W. Robarge
Category: Soils
Cooperative: Spruce-Fir
Scientific Question(s): 2.1
Title: Soil and Tissue Chemical Properties Associated with Stand
Characteristics of Spruce-Fir in the Southern Appalachians.
Tree Species: Red spruce
Objective: The general objective is to evaluate soil, foliar, and root
chemistry and provide baseline data for future comparisons. Secondary
objectives are to relate data on foliar and root variables to soil data and,
in coordination with scientists working on stand characterization,
mycorrhizae, pests, and pathogens, to relate tree and stand variables to soil,
foliar, and root variables.
Deliverables: Evaluation of the relationship between soil, root, and foliage
characteristics and symptoms, location, elevation, slope position, and
exposure. 9/88
Summary: This project is coordinated with the Zedaker and Bruck projects.
Zedaker provides data on site and stand characteristics for 130 permanent
plots. Bruck has collected soil and foliage samples from 115 plots and root
samples from 30 plots. This study conducts additional intensive soil and
foliage sampling
S&I Contact: Paul Schroeder
92

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Project Number: SF22
Principal Investigator: P. Feret
Category: Reproduction/Regeneration
Cooperative: Spruce-fir
Scientific Question(s): 1.2, 2.5
Title: The Impact of Environment and Genotype on the Reproductive Fitness of
Fraser Fir From Mr. Rogers and Mt. Mitchell
Tree Species: Fraser fir
Objective: Measure the environmental and genetic sources of variation in the
reproductive fitness of Fraser fir, and measure the impact of pollutants on
Fraser fir reproductive attributes.
Deliverables: Quantification of environmental and genetic sources of
variation, and the effect of ozone and ambient acid rain on the reproductive
fitness of Fraser fir, 6/88
Summary: Replicated reciprocal plantings of known genotypes of Fraser fir
were established in the subalpine environments of Mt. Rogers and Mt. Mitchell.
To test the response of the same genotypes to controlled pollutant exposures,
they were also planted in open top chambers near Mt. Lake, VA.
Reproductive fitness will be analyzed by measuring the fertile plant parts
including: analysis of pollen viability, male strobilus production, female
strobilus production during the pollination season, and cone attributes.
Extracted seed will also be analyzed with x-ray tests.
S&I Contact: Charley Peterson
93

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Project Number: SF23
Principal Investigator: R.T. Eckert, M.E. Demeritt and D.M. O'Malley
Category: Carbon Allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: Genetic Variation in Red Spruce Pollution Response
Tree Species: Red Spruce
Objectives: 1. Determine the level of relationship between seedling genotype
and carbohydrate allocation response to ozone stress in red spruce seedlings.
2. Determine the strength of relationships between decline symptomatology,
hybridization level with black spruce, and within stand genetic structure.
Deliverables: Correlation distribution of genotypes with site and vegetation
measurements, 11/88.
Summary: Methods for establishing true identity of red and black spruce
samples were established. Identifying either species with certainty is
especially difficult in disturbed habitats. Species are discriminated based
on branch morphology and on allozyme differences. Analysis of foliar isozymes
from 17 samples of 30 trees each nas been completed. Study of 18 loci from
approximately 500 red spruce trees revealed that trees in these plots have
reduced genetic variability, in comparison with other conifer species.
S&I Contact: Charley Peterson
84

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Project Number: SF24
Principal Investigator: D.H. DeHayes
Category: Spatial Study
Cooperative: Spruce-Fir
Scientific Question(s): 1.2
Title: Genetic Uniformity: A Fundamental Mechanism for Red Spruce Decline.
Tree Species: Red spruce, balsam fir, fraser fir.
Objectives: 1) Compare the extent and distribution of genetic diversity of
red spruce and balsam fir/fraser fir growing sympatrically throughout eastern
North America, 2) determine the relationship between relative tree vigor and
the degree of genetic variability within and among natural populations of red
spruce and balsam fir/fraser fir, 3) insure the preservation of red spruce
germ plasm as a provision for long-term genetics and physiological research,
and the maintenance of genetic diversity.
Deliverables: Assessment/comparison of spruce-fir vigor throughout the
northeast. Evaluation of relationship of vigor to genetic diversity,
environmental characteristics, and pollution. 5/89.
Summary: This research will attempt to determine if the 'widespread decline'
of red spruce throughout eastern North America is a function of a biological
phenomenon unique to the species which places it in jeopardy when confronted
with relatively rapid changes (climatic- or pollution-induced) in
environmental conditions.
S&I Contact: Greg Reams
95

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Project Number: SF25
Principal Investigator: S. Zedaker
Category: Reproduction/Regeneration
Cooperative: Spruce-fir
Scientific Question(s): 1.1, 1.2, 2.5
Title: Site and Stand Characteristics Associated with Potential Decline and
Regeneration Success of Spruce-Fir Stands in the Ssouthern Appalachians.
Tree Species: Spruce-fir
Objective: Characterize existing stand conditions, and determine regeneration
success of spruce-fir. Monitor long-term changes in stand composition and
growth as affected by atmospheric deposition.
Deliverables: Description of current stand conditions and relationship to
site characteristics, 6/87. Evaluation of regeneration success, biomass
relationships, and long-term changes in composition and growth of southern
spruce-fir stands, 2/90.
Summary: 133 permanent plots have been established in three geographically
distinct areas in the southern Appalachians. At each plot, the vegetation
structure and composition is intensively characterized. Vegetation data is
recorded for overstory, understory, and herb strata. Every year, overstory
stem crown condition, seed fall, seed viability, and seedling recruitment are
assessed.
Near selected permanent plots, smaller destructive sampling plots are also
established. Stem analysis is conducted on overstory trees. Dimension
analysis is performed to determine biomass of tree boles, branches, and
foliage. Foliage subsamples are taken for specific leaf area analysis. Leaf
area-sapwood area ratios, and biomass equations using foliage, branch, and
bole components will be developed using regression analysis of biomass
components.
S&I Contact: Charley Peterson
96

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Project Number: SF26
Principal Investigator: C.W. Dull, J.D. Ward, H.D. Brown and W. Shain
Category: Spatial Studies
Cooperative: Spruce-Fir
Scientific Question(s): 1.1, 1.2
Title: Detection and Evaluation of Spruce-Fir Decline in the Southern
Appalachians using Remote Sensing
Tree Species: Red spruce, Fraser fir
Objectives: 1. Delineate the boundaries of the spruce-fir forests of the
southern Appalachians. 2. Provide baseline data for the assessment of the
geographic extent of the mortality of the forest.
Deliverables: 1. Color infrared aerial photography (1:12000) during the
summer growing season and true color transparencies (1:12000) during the
winter season. Aerial photographic coverage of 20 research plots (1:4000).
9/86. 2. Aerial photo indices showing the locations of the center of each
exposure. 9/86. 3. A series of USGS quad sheet delineating the boundaries of
the spruce-fir type in the Southern Appalachians. 9/86. 4. A stratification
within the spruce-fir type boundaries showing mortality. Analysis of these
maps including statistical data. 9/86.
Summary: Photo interpretation delineated the boundaries of the spruce-fir as
well as the amount of mortality stratified within this type. Three mortality
classes were determined for study sites on Mt. Mitchell, Roan Mt., Mt. Rogers,
Grandfather Mt., Great Smoky Mtns., and the Blue Ridge Parkway. Based on tree
counts made on the aerial photography and verified on the ground, light (<30%)
mortality ranged at these sites from 17% (Roan) to 100% (Rogers); heavy
(30-70%) mortality from 4% (Mitchell) to 38% (Roan); severe (>70%) mortality
from 16% (Grandfather) to 53% (Blue Ridge Pkwy).
S&I Contact: Greg Reams
97

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Project Number: SF27
Principal Investigator: F. Thornton
Category: Carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: A Field Chamber Study of the Response of Red Spruce to Cloud
Interception and Ozone
Tree Species: Red spruce
Objectives: Determine the effects of acidic cloud water and 03, alone and in
combination, on red spruce seedling root and shoot growth, photosynthetic
rates and tissue nutrient concentrations.
Deliverables: Evaluation of use of chambers for cloud exclusion studies,
11/87. Results of exclusion experiments, 1/88, 1/89.
Summary: Uses open top chambers on Whitetop Mt., VA. The chambers are used
to exclude ambient ozone, clouds, or both. Throughout the growing season
periodic evaluation of physiological response is determined by measuring
photosynthetic rate, stomatal conductance, and needle water potential.
Periodic event sampling of foliar interception of cloud water is also
conducted. Root and shoot biomass and tissue nutrient concentrations are
determined from a subset of destructively sampled seedlings.
9&I Contact: Charley Peterson
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Project Number: SF28
Principal Investigator: D.S. Solomon
Category: Fixed Plot
Cooperative: Spruce-Fir
Scientific Question(s): 1.1, 1.2
Title: Modeling Stand Dynamics of Spruce-Fir Forests in the Northeast
Tree Species: Red spruce
Objectives: 1. Analyze the temporal and spatial variability of growth and
mortality within the spruce-fir forest through application and modification of
the FIBER model. 2. Evaluate the influence of stress on the temporal and
spatial variability of tree growth and mortality within the model FIBER using
changes in foliar conditions.
Deliverables: Updated version of the growth and yield model FIBER. 9/89.
Summary: Different versions of the model will account for growth differences
by species for different elevations on an annual basis. The model will be
tested for differences between regression coefficients developed prior to 1965
and between 1965 and the impact of the spruce budworm in 1975. These
differences should provide some indications of changes in growth on trees on
the same plots across time.
S&I Contact: Ross Kiester
98

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Project Number: SF30
Principal Investigator: A. H. Johnson
Category: Soils
Cooperative: Spruce-Fir
Scientific Questions: 2.1
Title: Response of Spruce-Fir Forest Soils at Whiteface Mountain, NY to
Acidic Deposition.
Tree species: Red spruce, balsam fir
Objectives: 1) Determine the effects of different levels of acid input and
water flux rates on soil solution chemistry and base cation leaching in a high
elevation spruce-fir ecosystem. 2) Utilize results of experimental trials to
test or calibrate available bulk soil, soil solution chemistry models for
application to the spruce-fir soils of the Adirondacks. 3) Use models,
experimental results, and information from related research projects to
evaluate the long term effects of changing atmospheric inputs.
Deliverables: 1) Determination of the degree to which aluminum, base cations,
and selected heavy metals are mobilized. 2) Evaluation of the effect of water
flux rates on soil solution chemistry. 3) A detailed chemical desctiption of
soil solution held at ambient tensions.
Summary: Uses controlled irrigations of 3m x 3m plots in the field on
Whiteface Mountain. Treatments consist of discrete rainfall events that vary
in intensity, depth, and duration representative of the 15 year historical
record on the mountain. Samples are obtained of saturated flow from the
downslope face of the soil blocks. Variable tension lysimeters are used to
extract samples of near ambient tension soil water during the course of
profile wetting and drying.
S&I Contact: Paul Schroeder
100

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Project Number: SF31
Principal Investigator: R. Kohut
Category: Carbon allocation, seedling/branch/tree integration
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: Comparison of the Responses of Seedling and Sapling Red Spruce Exposed
to Ozone and Acidic Precipitation Under Field Conditions.
Tree Species: Red spruce
Objectives: 1) Assess effects of acid precipitation and ozone on
photosynthesis and growth of red spruce seedlings. 2) Produce dose/response
functions using measures of photosynthesis and growth as response variables
for seedlings and saplings. 3) Develop quantitative assessments of the
effects of previous exposure and tree age on the dose/response function.
Deliverables: Comparison of dose/response relationships of 1 to 3 year old
spruce seedlings and saplings for acid and ozone treatments, 9/88.
Summary: Red spruce seedlings and saplings are exposed to ozone and acidic
deposition in open top chambers over a three year period. Response variables
include foliar and root pathology, net photosynthesis, stomatal conductance,
growth, carbon allocation, projected leaf area, leaf dry weight per age class,
and chlorophyll content.
S&I Contact: Charley Peterson
101

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Project Number: SF32
Principal Investigator: M.S. Greenwood and K.F. Jensen
Category: Carbon allocation
Cooperative: Spruce-Fir
Scientific Question(s): 2.3
Title: Evaluation of the Impact of Atmospheric Deposition on Red Spruce
Seedlings Using Open Top Chambers
Tree Species: Red spruce, balsam fir
Objectives: 1) Determine, under controlled field conditions, the effects of
ozone on the growth and morphology of red spruce and balsam fir seedlings, via
the mechanism of altered carbon allocation and physiological processes; 2)
Determine the influence of tissue maturation on red spruce response to ozone
alone or in combination with a natural environmental stress (i.e. water
stress).
Deliverables: Dose-response of ozone on spruce seedling development and
physiology, 12/88, 12/89. Dose-response of ozone and water stress on red
spruce as a function of tissue maturation, 12/90. 12/91.
Summary: Open top chambers are used to expose seedlings to various levels of
ozone. A later phase of the project will use mature and juvenile grafted
scions of red spruce. Height, diameter, foliage color, photosynthetic rate,
and stomatal conductance will be measured. Seedlings will then be
destructively sampled and analyzed for root and shoot biomass, chlorophyll,
and leaf area.
S&I Contact: Charley Peterson
102

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Project Number: SI01
Principal Investigator: D. Ford
Category: Seedling/branch/tree integration
Cooperative: Synthesis and Integration
Scientific Question(s): 3.1
Title; Analysis and Modeling the Effect of Pollutants on Forests
Tree Species: All
Objectives: Provide scientific and technical support in the creation of
models which relate how pollutants may influence tree and forest growth
through the effects they have on eco-physiological processes.
Deliverables: A library of C functions to create models of forest
eco-physiological processes. A coordinated program of functions to simulate
photosynthesis and carbon allocation. Program of functions to simulate stand
water relations and budgets. 9/88.
Summary: Work has started on developing models for predicting water stress in
whole trees as they age. A model has been developed to relating pollution
influence specified in general terms as decline in rates of physiological
processes. In addition, a model for the branching structure of trees was
developed and can be used with experimental data being collected in the FRP.
The essential feature of this model is that it predicts export of
photosynthate to the trunk and calculates this as a function of photosynthetic
rate, foliage amount, branch thickening and branch morphology in its influence
on foliage amount.
S&I Contact: Ross Kiester
103

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Project Number: SI02
Principal Investigator: W.G. Warren
Category: Dendrochronology, FIA, Fixed plot, spatial studies
Cooperative: Synthesis and Integration
Scientific Questions: 1.1, 1.2
Title: Development of Innovative Statistical Procedures for the FRP's
Synthesis and Integration Project
Objectives: To provide statistical support in all aspects of the FRP, in
particular the critical review of proposals (especially experimental designs
and analysis), the critical review of project reports (primarily with respect
to statistical analyses), and the application and/or development of innovative
statistical methodology as the situation demands, in particular in the areas
of the analysis of dendrochronological data and spatial analysis (Reams).
Deliverables: Reports will appear periodically. The following have been
completed.
Consequences of various growth models. S&I Report No. 5 (An abbreviated
version published in the Proceedings of the IUFRO Conference on Growth
Modeling Prediction).
Some novel statistical analyses relevant to the reported growth decline of
pine species in the southeast. S&I Report No. 9.
Review and evaluation of dendrochronological methods. S&I Report No. 10.
Evaluation of tree-ring studies on northeastern red spruce. S&I Report No.
11.
On the combining of independent tests of the same hypothesis. S&I Report No.
12 .
Initial results of the FIBER matrix model. S&I Report No. 13.
Reports on (i) significance in stepwise regression, (ii) model based
standardization of tree-ring studies, (iii) the interpretation of complex
multifactor analyses of variance, and (iv) the spatial analysis of gradient
studies are in various stages of preparation.
S&I Contact: Bill Warren
104

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Project Number: VS01
Principal Investigator: D.H. Marx
Category: Spatial Study
Cooperative: National Vegetation Survey
Scientific Question(s): 1.2
Title: Design and Pilot Test of a Long-term Monitoring Study of Eastern United
States Forests for Response to Atmospheric Deposition.
Tree Species: All major softwood and hardwood species of the Southeast.
Objectives: The project encompasses the development of four atlases for the
southeast, (1) the WEATHER ATLAS, (2) the SOILS ATLAS, (3) the TREE
DISTRIBUTION ATLAS, and (4) the ATMOSPHERIC DEPOSITION ATLAS. The tree
distribution atlas is derived from FIA data.
Deliverable: Electronic atlas for the south with information on past climate,
drought and acid deposition prone soils, current distribution of major tree
species, and chemical climatology of the region, 9/87.
Summary: The completed atlas provides a foundation for testing atmospheric
deposition effects on forests throughout the south.
S&I Contact: Greg Reams
105

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Project Number: VS02
Principal Investigator: C.W. Dull, J.D. Ward, W. Clerke and H.D. Brown
Category: Spatial Studies
Cooperative: Vegetation Survey
Scientific Question(s): 1.1, 1.2
Title: Evaluation of Spruce-Fir Mortality in the Southeast Utilizing Remote
Sensing and Geographic Information System Technologies
Tree Species: Red spruce, Fraser fir
Objectives: 1. Develop and implement a geographic database using existing
photography on extent and intensity of mortality in the spruce-fir type in
North Carolina, Tennessee, and Virginia. 2. Provide for comparison between
current and future extent and intensity of mortality and decline through the
computer analysis, storage, and display of spatial (geographic) information.
Deliverables: 1. Complete aerial photographic stereo coverage at a scale of
1:12000 in color infrared and true color transparencies of the spruce-fir
range in the southeast. 2. Spruce-fir type maps (1:24000). 3. Maps
displaying the extent of mortality within the spruce-fir range classified as
light, heavy, and severe. 4. Final report (in progress).
Summary: Analysis revealed that 65,672 acres of spruce-fir forest exist on
higher elevation peaks throughout the Southeast which form a series of
island-like areas in VA, NC, and TN. Within this area, 24% of the total area
was classified as severe mortality (>70% of the dominant and co-dominant
standing trees dead; 6% of the area showed heavy mortality (30-70% standing
dead); 70% of the area showed light mortality (<30% standing dead). Analysis
of the percent of standing dead timber showed that severe mortality occupied a
greater proportion of the area at higher elevations where the larger
percentage of the stand component was Fraser fir. Conversely, red spruce
occupied a greater percentage of the stand component at lower elevations. A
very small percentage of Fraser fir was found at the lower elevations.
S&I Contact: Greg Reams
106

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Project Number: VS03
Principal Investigator: J.E. de Steiguer
Category: FIA Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Methods for Economic Assessment of Atmospheric Pollution Impacts on
Forests of the Eastern United States.
Species: No species specified.
Objective: 1. To determine candidate forest-related market and nonmarket goods
and services which may be affected by atmospheric pollution, and to suggest
economic models which might be used to assess impacts upon them.
2. To investigate and develop the capability to project future atmospheric
pollution impacts using TRIM and to provide linkages to the economic
evaluation model TAMM.
Deliverables: Evaluation of Existing Models and Recommendations for Future Use
of Both Models and Existing Data, 12/87.
S&I Contact: Bill Warren
107

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Project Number: VS04
Principal Investigator: J.E. de Steiguer & G. Ruark
Category: FIA Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Synthesis and Modification of Tree Growth Models to Identify Variables
for Surveying and Monitoring Eastern Forest Ecosystems for Atmospheric
Deposition Effects
Species: Loblolly pine (initially)
Objective: 1. To identify models which express forest stand conditions as a
function of pollution-related and non-pollution- related variables.
2. To provide a list of the dependent and independent variables (from #1
above) to assist in designing the National Vegetation Survey.
Deliverables: Evaluation of models and their appropriateness for the FRP,
12/89
S&I Contact: Bill Warren
108

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Project Number: VS05
Principal Investigator: J. Skelly
Category: Pest/Pathogen
Cooperative: Spruce-fir
Scientific Question(s): 2.6
Title: Manual for Diagnosing Injury to Eastern Forest Trees
Tree Species: Eastern forests
Objective: Produce a manual for diagnosing injury to eastern forest trees.
Deliverables: Manual for diagnosing injury to eastern forest trees.
Summary: This project has been completed. It accomplished its objective of
producing a manual for diagnosing forest injury. The manual includes both
pollutant induced injury and injury resulting from other major diseases,
insects, and abiotic stresses.
S&I Contact: Greg Reams
109

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Project Number: VS06
Principal Investigator: J.W. Hornbeck
Category: Dendrochronology Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Analysis and Interpretation of Tree Cores in New England.
Species: red spruce, white pine, balsam fir (sugar and red maple, red and
white oak, yellow birch).
Objective: 1) to determine whether regional growth decline can be detected in
other species besides red spruce;
2)	to analyze the dependence of growth decline on site factors and on tree or
stand age and history;
3)	to determine the effects of climate on growth decline of red spruce and
other species;
4)	to quantify the relation between growth decline in red spruce and spruce
budworm damage.
Deliverables: Analysis of increment core data bases with associated location
and stand data, 9/87
Summary: The project contains a major dendroecological component. The cores
are obtained from the Forest Survey program of the U.S. Forest Service (FIA).
The investigator appears to be attempting a more sophisticated approach than
commonly taken.
S&I Contact: Bill Warren
no

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Project Number: VS06
Principal Investigator: Jagels (Hornbeck)
Category: Dendrochronology Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Analysis and Interpretation of Tree Cores in New England.
(Fingerprinting radial increment data for red spruce using morphometric
analysis).
Species: Red spruce
Objective: Test the hypothesis that during periods of radial increment change
(stimulation or decline) morphometric fingerprints of individual growth rings
will mirror the unique conditions associated with the decline or stimulation,
and that these fingerprints will, therefore, be different depending upon the
conditions which initiated the radial increment change.
Deliverables: Analysis of increment core data bases with associated location
and stand data, 9/87
S&I Contact: Bill Warren
in

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Project Number: VS07
Principal Investigator: J.P. McClure
Category: FIA Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Formulation and Testing of Nondeposition Growth Loss Hypotheses with
Existing Southeast Forest Inventory Data
Species: All southern pines
Objective: The listed objectives are: (1) Computerize the FIA data collected
during the 1957 to 1967 period: determine growth rate for this period for
comparison to two subsequent remeasurements.
(2)	Determine if changes in stand characteristics (such as density or age),
singly or in combination, can explain any of the observed reduction in tree
diameter growth.
(3)	Determine if average diameter growth is significantly different in pine
stands with varying degrees of competing non- pine vegetation.
(4)	Identify possible differences in the development of old field pine stands
contrasted with the development of pine stands originating on cutover forest
land. Assess possible differences in individual tree growth trends for these
two sets of plots.
(5)	Evaluate the amount of observed loss that can be attributable to drought
and support the development of a forest drought index across the Southeast.
(6)	Evaluate patterns and trends in tree mortality in the Southeast that are
not attributed to known causes and carry out field explorations of individual
plots and trees in search of clues as to causes.
Deliverables: Analysis of FIA data for the southeast for the last three
inventory periods with respect to a possible decline in growth and evaluation
of potential causes, 12/87
Summary: The FIA data are, a priori, inadequate to resolve all of the
questions asked because FIA was not designed to address them. This project
seeks to get as much out of the FIA data as possible.
S&I Contact: Bill Warren
112

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Project Number: VS08
Principal Investigator: J.R. Saucier
Category: FIA Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Near-term Survey of Eastern Forest Conditions/Atmospheric Deposition.
Species: Loblolly pine
Objective: To determine if there is any current or past relationship between
selected parameters of forest condition and atmospheric deposition, while
considering other (natural and man- caused) factors that affect forest
conditions.
Deliverables: Assessment of pilot study results and recommendations for future
inventory design; field trials and analyses of 200 plots related to the
southern pine growth reduction, 9/87
Summary: Although the objective is sweeping, in execution the project seems
not far removed from that of McClure. To date it focuses on a selection of 200
FIA plots from Georgia, North and South Carolina.
S&I Contact: Bill Warren
113

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Project Number: VS09
Principal Investigator: T. Dell
Category: Dendrochronological Data
Cooperative: National Vegetation Survey
Scientific Questions: 1.1, 1.2
Title: A Review of Statistical Methods for Evaluation of Atmospheric
Deposition Influences on Forests.
Tree species: Southern species
Objectives: Evaluate existing statistical methods for studying the effects of
air pollutants on forests, and develop mew methods where appropriate. Of
special interest are methods for analyzing dendrochronological and FIA data.
Deliverables: Evaluation of statistical methods for their appropriateness to
the FRP. Includes increment core data sets and independent analyses using a
variety of procedures, 12/89.
S&I Contact: Bill Warren
114

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Project Number: VSlO
Principal Investigator: L.F. Ohmann
Category: Spatial Study
Cooperative: National Vegetation Survey
Scientific Question(s): 1.1, 1.2
Title: Study of the Relationships Between Various Aspects of Forest Condition
and Atmospheric Deposition Across the Northwestern Minnesota to Southeastern
Michigan Deposition Gradient.
Tree Species: Red pine, white pine, jack pine, balsam fir, aspen, paper birch,
basswood, white oak, sugar maple, red maple.
Objectives: 1) Analyze the relationship between tree residual growth increment
and sulfur content of tree woody tissue and soils across an established acid
precipitation gradient, 2) identify the macroclimate and acid deposition
variables which have a significant relationship with the rate of diameter
growth in the Lake States Region; the study will (a) use the regional growth
model STEMS to eliminate the effect of stand, tree and site factors on
diameter growth; (b) analyze the resulting diameter prediction residuals by
species across the major climate and acid deposition variables; (c) stratify
data to account for other local site factors, such as topography and drainage;
(d) evaluate major trends using average prediction errors by prevalent species
and weather zone (county).
Deliverable: Analysis of stand growth along a known deposition gradient.
Includes a data set of remeasured plot variables. 12/87
Summary: The general hypothesis Is that a wet sulfate deposition gradient is
reflected in the amount of accumulated sulfur in the forest floor-soil system
and tree tissue. Tree radial increment is also associated with the deposition
gradient.
S&I Contact: Greg Reams
115

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Project Number: VS11
Principal Investigator: R. Brooks
Category: Spatial Studies
Cooperative: National Vegetation Survey
Scientific Question(s): 1.1, 1.2
Title: Analysis of Forest Survey and Atmospheric Deposition Data in
Pennsylvania.
Tree Species: Red maple, sugar maple, chestnut oak, northern red oak, black
oak, white oak, black cherry, black birch, american beech, eastern hemlock.
Objectives: 1) Analyze tree measurement and tree core data to identify
possible spatial and temporal patterns of tree and forest stand growth while
controlling certain site and stand influences on growth; 2) Analyze any
identified spatial and temporal tree and forest growth pattern for
relationship to established or estimated atmospheric deposition patterns.
Deliverable: Analysis of remeasured plot variables with associated increment
core data and soil chemical data, 12/87.
Summary: This project is investigating whether there is a temporal or spatial
pattern to tree or stand growth, and if so is this pattern related to
atmospheric deposition.
S&I Contact: Greg Reams
116

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Project Number: VS12
Principal Investigator: T.R. Dell
Category: FIA Data
Cooperative: NVS
Scientific Question(s): 1.1, 1.2
Title: Southern Forest Growth Trends
Species: Loblolly pine, slash pine, shortleaf pine, longleaf pine, southern
hardwoods.
Objective: Develop data and prediction processes to independently determine if
there is evidence of growth decline in the Southern U.S. region.
Deliverables: Analysis of southern pine plantation reraeasurement data with
emphasis on possible growth impacts related to atmospheric pollution, 12/87.
Summary: Unlike the studies of McClure, Saucier and de Steiguer this study
focuses on research growth and yield (RGY) plots while taking what it can from
FIA data. It also plans to incorporate the work of Zahner on the development
of processes and software to take into account standard weather records. The
study can also be viewed as a complement to the Marx et al. "Atlas" study.
This appears to be a comprehensive and well considered study that employs
statistical methods with care and discrimination. Zahner's approach is
apparently a step forward from multiple regression on monthly temperature and
precipitation records.
S&I Contact: Bill Warren
117

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Project Number: VS13
Principal Investigator: C.W. Dull, J.D. Ward and H.D. Brown
Category: Pest/Pathogen
Cooperative: Vegetation Survey
Scientific Question(s): 2.6
Title: Developing an Aerial Photographic Technique for Assessment of Visual
Spruce-Fir Decline Resulting From Air Pollution or Other Environmental Stress
in the Southern Appalachians.
Tree Species: Red spruce, Fraser fir
Objectives: To identify key characteristics for aerial detecting visual
symptoms of declining red spruce and Fraser fir on conventional 9x9
photography. To identify key characteristics for separating red spruce and
Fraser fir in the southern Appalachians.
Deliverables: Key and photo interpretation guide for spruce and fir in the
southern Appalachians.
Summary: Color infrared photography has been acquired for sites on Roan Mt.
and Mt. Mitchell. Plots have been selected on the ground with a) pure spruce,
b) pure fir, c) mixed spruce-fir, d) various age classes of these species.
Characteristics of the various species compositions as well as different types
of tree damage in the study area are being identified. A dichotomous key
using the various attributes and ecological factors found on the aerial photos
will be developed. A pictorial guide showing both aerial and ground views of
these species and damage symptoms will be prepared for training
photointerpreters in the study.
S&I Contact: Greg Reams
118

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Project Number: VS14
Principal Investigator: I. Millers
Category: Spatial Study
Cooperative: National Vegetation Survey
Scientific Question(s): 1.2
Title: Cooperative Survey of Red Spruce and Balsam Fir Decline and Mortality
in the Northeast: Symptoms and Trends.
Tree Species: Red spruce and balsam fir.
Objectives: 1) To determine the frequency and geographic variability of visual
crown symptoms on deteriorating red spruce and balsam fir in the northeastern
United States, 2) to determine the progression of symptoms on the trees and in
the forest as the severity of damage increases, 3) to identify and describe
symptoms caused by historically important damage agents and to differentiate
them from those for which causal agents cannot be determined.
Deliverable: Inventory of visual symptoms and indication of the frequency of
occurrence on trees. 12/89.
Summary: A large variety of symptoms is expected. Similar symptoms may be
produced by different agents, or a variety of symptoms may be produced by a
single causal agent, or symptoms may be produced by a combination of agents.
S&I Contact: Greg Reams
119

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Project Number: VS15
Principal Investigator: B.N. Rock
Category: Spatial Study
Cooperative: National Vegetation Survey
Scientific Question(s): 1.2
Title: Vegetation Survey Pilot Study: Detection and Quantification of Forest
Decline Damage Using Remote Sensing Techniques.
Tree Species: Red spruce and balsam fir.
Objectives: 1) Use remote sensing techniques to accurately detect, quantify,
and map the spatial extent and relative levels of damage in high-elevation
spruce/fir forests in the eastern United States, 2) provide accurate relative
damage rating values which allow comparison of forest condition (damage)
within and among the study sites, 3) assess whether spatial patterns of forest
damage exist which coincide with spatial patterns of pollutant exposure.
Deliverable: Technical report that includes damage assessment maps, damage
rating values, and discussion of techniques used. 12/87.
Summary: This study was developed to demonstrate the feasibility of using
remote sensing techniques to detect and map levels of forest damage at
selected sites in the eastern United States.
S&I Contact: Greg Reams
120

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Project Number: VS16
Principal Investigator: J.P. Bennett, R.L. Anderson and M. Mielke
Category: Fixed Plot
Cooperative: Vegetation Survey
Scientific Question(s): 1.1, 1.2
Title: The Status of Eastern White Pine affected by Air Pollutants in the
Eastern United States
Tree Species: Eastern White Pine
Objectives:
Deliverables: List of white pine data sets with atmospheric pollution data,
5/87. Assessment of quality and utility of data sets for evaluation of
pollution impacts on white pine, 5/87. Recommendations for an eastwide
inventory of white pine to detect damage, 5/87.
Summary: Summary of data sets has been compiled. Assessment of quality and
development of inventory recommendations are in progress.
S&I Contact: Ross Kiester
121

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Project Number: WC03
Principal Investigator: T. Larson
Category: Atmospheric Exposure
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Fall and Winter Cloudwater Chemistry Measurements at Mount Rainier
Tree Species: Subalpine forest: Abies lasiocarpa. Tsupa mertensiana
Objectives: Field study to evaluate the chemical composition of cloudwater at
a montane site in the Cascades to establish realistic fog and cloudwater
exposure protocols for western conifer seedlings.
Deliverables: Comparison of cloudwater composition at low and high elevation
sites in Western Washington, 6/87. Final report, 6/87.
Summary: Cloudwater composition at Mt. Rainier during this experiment
provides the first measurements of this parameter in the Cascades. These data
will allow testing of the hypothesis that during fall and winter events the
composition of cloudwater as measured by the equivalent fraction of acidic
species does not vary significantly from event to event at this site, nor
significantly from the values measured during the winter at the polluted urban
Seattle site.
S&I Contact: Jeff Brandt
122

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Project Number: WC04
Principal Investigator: D.F. Miller and R.D. Borys
Category: Atmospheric Exposure
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Characterization of Cloud Chemistry and Frequency of Canopy Exposure
to Clouds at Mt. Werner, Colorado
Tree Species: High-elevation, subalpine forest in U.S. Rocky Mountains
Objectives: Determine the chemical composition of clouds and the frequency of
forest canopy exposure to clouds at a high elevation site in the Rocky
Mountains during the summer growing season.
Deliverables: Characterization of cloud chemistry and frequency of canopy
exposure to clouds in the Rocky Mountains, Data tape, 4/88
Summary: Information provided by this study will include:
-	chemistry and liquid water content of all cloud events for the summer of
1987 at Mt. Werner, Colorado;
-	measurement of precipitation amount and frequency at the mountain field
sites;
-	measurement of meteorological conditions during the study period;
-	determination of the chemical composition of rainwater during periods when
precipitating clouds impact the mountains;
-	determination of the average concentration of acidic gases and airborne
particulates during periods when clear air conditions prevail at the study
sites.
S&I Contact: Jeff Brandt
123

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Project Number: WC05
Principal Investigator: L. Svoboda
Category: Atmospheric Exposure
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Rocky Mountain Deposition Monitoring Project
Tree Species: n.a.
Objectives: 1) Characterize current levels of deposition at ten
high-elevation sites in the Rocky Mountains of Colorado and Wyoming. 2)
Evaluate the effects of elevation on measured wet deposition levels. 3)
Determine the spatial variability of precipitation chemistry at high
elevations within the Colorado Rocky Mountain region.
Deliverables: Report on the influence of elevation on rates of wet deposition
to high-elevation systems of the Rocky Mountains, 5/88
Summary: This project established and operates a network of paired wet
deposition monitoring stations at five different high-elevation sites in the
Rocky Mountain region. The data obtained from this network will be used to
evaluate current levels of wet acidic deposition at high elevations and to
determine if wet acidic deposition varies at each of the four study areas as a
function of elevation.
S&I Contact: Jeff Brandt
124

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Project Number: WC06
Principal Investigator: D. Cronn and M. Campbell
Category: Atmospheric Exposure
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Development and Testing of a Passive Ozone Monitor using Dry
Chemiluminescence for Outdoor Monitoring
Tree Species: n.a.
Objectives: Develop a small, inexpensive, and reliable ozone exposure monitor
using a proven active method of dry chemiluminescence in a new passive design
incorporating photographic film.
Deliverables: Fully operational prototype, 12/87. Results of field and lab
evaluation, 6/88. Final data processing and report, 8/88.
Summary: The project focuses on development of a passive monitor of simple
design, moderate accuracy and low operating cost, together with exhaustive
testing of prototypes for accuracy and interferences under realistic
laboratory and field conditions.
S&I Contact: Jeff Brandt
125

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Project Number: WC07
Principal Investigator: D.T. Tingey and D. Turner
Category: Foliar leaching
Cooperative: Western Conifers
Scientific Question(s): 2.1, 2.2
Title: Foliar leaching and root uptake of Ca, Mg, and K in relation to acid
mist effects on conifers.
Tree Species: Douglas fir, Engelmann spruce
Objectives: To evaluate the role of foliar cation leaching and cation
availability to roots of Douglas-fir and Engelmann spruce in determining
seedling response to acidified mist.
Deliverables: Report on first year effects of simulated acid mist on
throughfall chemistry, and seedling biomass and nutrient content, 10/87.
Final report, 10/88.
Summary: Results of a pilot study on Douglas fir include:
-	Biomass per plant increased at the end of the 12 week experiment in response
to higher levels of nutrient availability.
-	Content of Ca, Mg, and K in second year foliage increased in response to
higher levels of nutrient availability, but did not show a significant
response to fog pH.
-	Leaching of Ca, Mg, and K from foliage was considerably higher with fog of
pH 3.1 than with fog at pH 5.6, with K being most susceptible to leaching.
However, the amounts of nutrients removed at pH 3.1 are small relative to the
observed uptake rates of these trees.
-	Epicuticular waxes were not affected by either treatment.
Further studies will focus on foliar leaching in Engelmann spruce and on the
cation exchange capacity and buffering capacity of conifer needles.
S&I Contact: Jeff Brandt
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Project Number: WC08
Principal Investigator: W. Hogsett and D.T. Tingey
Category: Carbon allocation
Cooperative: Western Conifers
Scientific Question(s): 2.3
Title: Sensitivity of Important Western Conifer Species to S02 and Seasonal
Interaction of Acid Fog and Ozone
Tree Species: Douglas fir, ponderosa pine, lodgepole pine, western hemlock,
western red cedar
Objectives: Assess relative sensitivity of western species to: 1) a seasonal
pattern of acid fog and ozone, and 2) S02 exposure during fall and winter.
Deliverables: Interim sensitivity ranking, 10/87. Final sensitivity
rankings, 10/88.
Summary: The screening of various species for sensitivity in growth and
visible needle injury is accomplished with two deposition exposure scenarios:
l)Acid fog (winter)/ozone (summer) as a seasonal combination of pollutants,
and 2) gaseous S02 deposition (winter). Seedling sensitivities are being
assessed as a growth response over two growth periods with year-round
exposures. The fumigation regimes reflect the seasonality of deposition
duration, frequency of events, fog chemistry, and the seasonality of the
frequency and distribution parameters of ozone and S02 characteristic of
selected regions of the west. A range of treatment concentrations is employed
for each pollutant which are representative of the possible air quality
conditions of the west. The fumigation periods are those months when these
deposition patterns occur. Seasonal interaction of the pollutants, rather
than concurrent pollutant combinations, represent a realistic exposure
scenario for much of the climatic conditions of the coastal western U.S. and
Cascade and Sierra foothills.
S&I Contact: Charley Peterson
127

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Project Number: WC09
Principal Investigator: P. Miller
Category: Carbon allocation
Cooperative: Western Conifers
Scientific Question(s): 2.3
Title: Testing the Sensitivity of Five Western Conifer Species to S0Z Alone,
and Ozone Followed by Acidic Fog
Tree Species: Douglas fir, ponderosa pine, white fir, subalpine fir, Engelmann
spruce
Objectives: Using open-top chambers, determine the sensitivity of the 5
conifer species to S02 alone, and acid fog and 03 in combination..
Deliverables: Report of preliminary rankings, 10/87. Report of final
rankings, 10/88.
Summary: This project is conducted in coordination with the Hogsett and
Tingey project. Exposures are conducted year-round in modified open top
chambers under natural environmental conditions. Simulated ambient exposure
profiles for acidic fog, S02, and ozone are employed. Exposure regimes for
the gaseous pollutants were developed by averaging air quality characteristics
from a number of sites across a region. These average values were then used
to construct a 30 or 60 day hourly concentration regime that reflects the
ambient air quality characteristics of the region of interest. A range of
treatments for each pollutant is created by building additional hourly
concentration profiles within 1-2 standard deviations of the base profile.
Development of fog regimes follows a similar methodology reflecting the
frequency, chemistry, and deposition volume of selected regions.
S&I Contact: Charley Peterson
128

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Project Number: WC16
Principal Investigator: S. Ustin
Category: Spatial Study
Cooperative: Western Conifers
Scientific Question(s): 1.2
Title: Remote Sensing of Forest Condition in the Western United States.
Tree Species: Ponderosa pine and associated western conifers.
Objectives: 1) Develop spectral response curves for several conifer species
(Douglas fir, lodgepole pine, western hemlock, western red cedar) and various
atmospheric pollutants (ozone, S02, acid foe) under controlled exposure and
environmental conditions, 2) determine if the pattern of spectral signature
variation is separable from that produced by other sources of variation, e.g.
soil-vegetation mixtures, community variation, 3) determine if pollutant
specific or species specific spectral responses exists under controlled
conditions, 4) assess the relationship between spectral features and potential
physiological mechanisms, 5) determine whether spectral evidence exists for in
situ vegetation stress caused by atmospheric pollutants, 6) determine the
relationship between field and lab spectra and suspected pollutants, 7)
establish a preliminary association between spectral responses and
physiological mechanisms.
Deliverable: Final report, 12/88.
Summary: The proposed research will collect and analyze: (1) spectral
signature measurements on seedlings under controlled atmospheric pollution
regimes, (2) ground-based spectral measurements on seedlings and trees growing
along known atmospheric pollution gradients selected to coincide with other
WCRC research, (3) Advanced Visible Infrared Imaging Spectrometer (AVIRIS)
data across these gradients, (4) Thematic Mapper satellite data. Field
collected ground-based, aircraft, and satellite spectral data will coincide to
provide the best basis for extrapolating between experimental levels.
S&I Contact: Greg Reams
129

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Project Number: WC18
Principle Investigator: R.L. Edmonds
Category: Spatial Studies
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Pollution Gradient Studies in the Puget Sound Region and Evaluation of
Existing Conifer Plot Data for Correlative Studies.
Tree Species: Douglas-fir
Objectives: 1) Identify regions of relatively high (potentially deleterious to
tree growth) and low (background) pollutant exposures through the use of
existing monitoring data and modeling, 2) confirm these subregions by
developing an improved characterization of ambient pollutant levels at
forested sites including monitoring of acidic sulfates and nitrates in fog,
monitoring of ozone, and at a subsample of sites monitoring of dry air species
including S02, particulate sulfate, and 3) assess growth, yield and other
existing data from plots located in western Washington and southwestern
British Columbia for suitability for correlating forest condition and
pollutant levels.
Deliverable: Final report, 4/89.
Summary: Describe the temporal and spatial patterns of atmospheric
concentrations of sulfur and nitrogen species, and ozone in forested areas of
the Puget Sound region, and to assess suitability of existing forest plots for
correlating pollutant levels with forest stand conditions.
S&I Contact: Greg Reams
130

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Project Number: WC20
Principal Investigator: J. Houpis
Category: Carbon allocation, seedling/branch/tree integration
Cooperative: Western Conifers
Scientific Question(s): 2.3
Tree Species: Ponderosa pine
Title: Comparison of the Response of Seedlings and Mature Branches of
Ponderosa Pine to Air Pollution.
Objective: Develop and test a branch exposure chamber. Document if a branch
exposure chamber is a valid research method by testing for branch autonomy in
mature Ponderosa pine. Test the difference in responses of mature branches
and seedlings.
Deliverables: Report on autonomy characterization and effects of ozone using
branch chambers, 10/88. Preliminary results of exposure experiments and
autonomy study, 10/89. Final report, 1/91.
Summary: This project undertakes to both design and utilize experimentally,
branch exposure chambers. The chambers will be designed and tested after an
extensive literature search and meetings with other scientists involved in
branch chamber design. After the chambers have been designed and tested, they
will be used in conjunction with open top chambers to compare the responses of
mature branches and seedlings to varying levels of ozone exposure. Response
variables include gas exchange, water potential, pigmentation, and growth.
1aC02 will also be used to study branch autonomy and pollutant effects on
autonomy.
S&I Contact: Ross Kiester
131

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Project Number: WC22
Principal Investigator: S. Ustin
Category: Caibon allocation, spatial studies
Cooperative: Western Conifers
Scientific. Question(s): 1.2, 2.3
Title: Spectral characteristics of conifer species exposed to simulated
pollutant regimes.
Tree Species: Douglas fir, ponderosa pine, Engelmann spruce, western hemlock,
western red cedar
Objectives: 1) Evaluate spectral response of western conifers to controlled
S02,_ acid fog/03 exposure; and 2) evaluate the use of aircraft and satellite
derived remote sensing data to characterize regional forest condition as
influenced by air pollution.
Deliverables: Evaluation of 0,, acid mist, and S02 effects on spectral
changes 6/87. Re-evaluation of progress and approaches, Summer/87.
Summary: Looked at spectral changes of seedlings from the Hogsett project
that had been exposed to S02, 03, and acid mist.
S&I Contact: Charley Peterson
132

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Project Number: WC24
Principal Investigator: D.A. Graybill
Category: Dendrochronology Data
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Analysis of Growth Trends and Variation in Conifers from Central
Arizona.
Species: Ponderosa pine, pinyon pine, Douglas fir
Objective: To address the question "are changes in forest condition greater
than can be attributed to typical trends and levels of natural variability?"
by applying a series of analytical strategies to newly developed tree ring
chronologies cross-dated from the extensive data base held by the Laboratory
of Tree-Ring Research, and to analyze archived forest plot measurement data.
Deliverables: Final report, 9/88.
Summary: This project employs, primarily, methodology that has become
conventional in dendroclimatology, although an auto- regressive - moving
average component is included and exploration of Kalman filtering, in
cooperation with P. van Deusen, is contemplated.
The study plan gives the impression that reliance will be placed on large
scale analyses of variance and covariance coupled with ultra conservative
multiple comparisons procedures. The investigators are also performing a
series of simpler analyses, e.g. examining the trend over diameter for each
inventory separately and thence whether such trends are stable over time of
change in a systematic manner.
S&I Contact: Bill Warren
133

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Project Number: WC25
Principal Investigator: L.B. Brubaker
Category: Dendrochronology Data
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Growth Variations in Old-growth Douglas-fir Forests of the Puget Sound
Area.
Species: Douglas fir.
Objective: 1. To quantify the temporal variability of annual growth of
Douglas-fir stands (ca. 1800 to 1980) from existing dendrochronological
records.
2. To examine growth records from ten new sites for evidence of growth
variations since the time of the original collections in the Puget Sound area.
Deliverables: Final report, 9/88.
Summary: Collecting data from new sites involves:
(1)	Selecting and visiting new sites for collections. These collections
will come from contrasting areas where pollution levels are thought to be
relatively high and low in the Puget Sound region.
(2)	Using the information gathered in (1) to describe the statistical
properties of ring-width sequences using time series analysis for periods
prior to and following any hypothesized point calculated.
(3)	Calculating multivariate models in the time domain, with climatic
data as inputs and standardized ring-width sequences as outputs. These
models will be fitted to periods prior to and following the hypothesized
change point.
S&I Contact: Bill Warren
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Project Number: WC26
Principal Investigator: D.L. Peterson
Category: Dendrochronology Data
Cooperative: Western Conifers
Scientific Question(s): 1.1, 1.2
Title: Growth Trends in the Mixed Conifer Forest of the Sierra Nevada
Species: Ponderosa pine
Objective: 1. Quantify growth trends and natural variability in growth for
ponderosa pine in the mixed conifer forest of the Sierra Nevada: (a) at the
stand level, (b) at the regional level.
2.	Evaluate recent growth trends with respect to expected patterns: (a) by
comparing recent to previous growth with the use of multivariate generalized
least squares analysis using Kalman filter techniques (b) by comparing growth
of trees with symptomatic injury to trees without injury.
3.	Account for sources of variance in growth that can be attributed to injury
level, climate, and other environmental factors.
Deliverables: Final report, 9/88.
Summary: This project explores several new approaches to dendrological
studies. The investigator, by sampling trees within the typical dense stands
of the region, in contrast to the dominant open-grown trees traditionally
selected for dendro- chronological studies, is facing a challenging problem,
but one that sooner or later has to be addressed.
S&I contact: Bill Warren
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Project Number: WC32
Principal Investigator: W. Winner
Category: Seedling/branch/tree integration
Cooperative: Western Conifers
Scientific Question(s): 3.1
Title: Workshop - The response of trees to air pollution: the role of branch
studies.
Tree species: All
Objectives: Review current research on development and use of branch
chambers. Identify criteria for optimal branch chamber design. Discuss the
relationship between branch responses and whole tree responses. Discuss
branch autonomy theory. Identify research needs not currently addressed.
Deliverables: Workshop proceedings. 3/88.
Summary: The workshop was conducted in November, 1987. The conclusions were:
1)	Ideal chambers should allow manipulating air pollution exposure and
measuring gas exchange in order to calculate photosynthesis and conductance.
They should be used in the field on trees of all sizes.
2)	Branches can be studied in a way relevant to understanding the entire
canopy. Both allometric and modeling approaches are useful for studying
branch-tree relationships.
3)	Branches may be independent units after they mature, at least with respect
to carbon. However, the carbon allocation processes between branches of
conifers are still not understood. Compensation between branches is apparent
for other resources such as nitrogen and water.
4)	Future research: a) Clarigy the issue of branch-branch allocation
processes with the canopy, b) Pursue chamber design opportunities which
minimize leaf heating in short experiments, c) Develop techniques to scale
from branch studies to whole trees.
S&I Contact: Ross Kiester
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