EPA-R3-73-030
    EFFECTS OF SULFUR OXIDES
                    IN THE
               ATMOSPHERE
            ON VEGETATION;
Revised Chapter 5 for Air Quality Criteria for Sulfur Oxides
                        Task 16
                     ROAP No. 26AAA
                   Program Element 1A1001
          NATIONAL ENVIRONMENTAL RESEARCH CENTER
           OFFICE OF RESEARCH AND DEVELOPMENT
           U.S. ENVIRONMENTAL PROTECTION AGENCY
        RESEARCH TRIANGLE PARK, NORTH CAROLINA 27711
                    September 1973

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               RESEARCH REPORTING SERIES

   Research reports of the Office of Research and Development,
Environmental Protection Agency,  have been grouped into five
series.  These five broad categories were established to facili-
tate further development and application of environmental tech-
nology .  Elimination of traditional grouping was consciously
planned to foster  technology transfer and a maximum interface
in related fields.  The five series are:

   1. Environmental Health Effects Research
   2. Environmental Protection Technology
   3. Ecological Research
   4. Environmental Monitoring
   5. Socioeconomic Environmental Studies

   This report has been assigned to the ECOLOGICAL RESEARCH
series.  This series describes research on the effects of pollution
on humans, plant and animal species, and materials.  Problems
are assessed for their long- and short-term influences. Investi-
gations include formation, transport,  and pathway studies to
determine the fate of pollutants ai~A their effects. This work
provides the technical basis for setting  standards to minimize
undesirable changes in living organisms Li the aquatic, terres-
trial, and atmospheric environments.
                            11

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                            PREFACE

   Air quality criteria state what science has thus far been able to measure of
the obvious as well as the insidious effects of air pollution on man and his
environment.  Criteria provide the most useful basis presently  available for
determining the levels of air pollutants that will protect the public health
and  welfare.  The  Clean Air Act  states:  "Air quality  criteria for an air
pollutant shall accurately  reflect the latest scientific knowledge  useful in
indicating the kind and extent of all identifiable  effects on public health or
welfare which may be expected from the presence of such pollutant in the
ambient air, in varying quantities."
   Air Quality Criteria  for  Sulfur  Oxides*  was issued under  the  1967
amendments to  the Clean  Air Act.  National ambient air quality standards
were  proposed,  based on  these  criteria, and  were promulgated under the
1970 amendments to the Clean Air Act.
   The Clean  Air  Act states that the  Administrator  of the Environmental
Protection  Agency (EPA)  "shall  from  time  to  time review,  and, as
appropriate, modify, and reissue any criteria .  . ." Limitations in the criteria
for secondary standards in Air Quality Criteria for  Sulfur Oxides,* which
became ^apparent  since the adoption  of Air Quality Standards,  prompted
review  and revision of Chapter  5,  "Effects  of  Sulfur  Oxides in  the
Atmosphere on Vegetation." This document presents the revision to Chapter
5, and  also  includes  revised  portions of  Chapter  10, "Summary  and
Conclusions," that relate to effects on vegetation.
   This  revision includes  a  number  of studies completed  since initial
publication in 1969. All data expressed in micrograms per cubic meter are
referenced to  25°  C and a pressure of 1013.2 mb  (760 mm Hg) unless stated
otherwise.
   Following the initial revision by EPA personnel, there was a  sequence of
review and  revision by (1) the National Air Quality  Criteria  Advisory
Committee,  which has  a membership broadly representative  of  industry,
universities,  conservation interests, and all levels of government, and by (2)
individuals specially selected  for their  competence,  expertise, or special
interest in the effects of air pollutants on vegetation. The efforts of these
reviewers, without which this  document could  not  have been completed
successfully, are acknowledged individually on the following pages.
   As  required by  the Clean  Air  Act, appropriate Federal departments  and
agencies  were consulted prior to issuing this criteria document. A Federal
consultation committee, comprised of members designated by the heads of
departments and agencies,  reviewed the document and met with EPA staff
members  to  discuss -their  comments.  These  representatives  are also listed
following this discussion.
*Air Quality Criteria for Sulfur Oxides. U.S. Department of Health Education and
 Welfare, National Air Pollution Control Administration. Washington, D.C. Publication
 No. AP-50. January 1969. 178 p.
                                   ill

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  The EPA is pleased to  acknowledge the efforts of each of the persons
specifically named, as well  as the many  not named, who contributed to
producing this  document. Their participation does not necessarily imply
complete endorsement of all the conclusions presented herein; in the last
analysis,  the  Environmental Protection  Agency  alone retains full respon-
sibility for its contents.
                                      Russell E. Train
                                      Administrator
                                      Environmental Protection Agency
                                  IV

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       NATIONAL AIR  QUALITY  CRITERIA ADVISORY
                                  COMMITTEE

                                      CHAIRMAN
                               Dr. John F. Finklea, Director
                          National Environmental Research Center
                          Research Triangle Park, North Carolina
 Dr. Mary O. Amdur
 Associate Professor of Toxicology
 School of Public Health
 Harvard University
 Boston, Massachusetts

 Dr. David M. Anderson
 Manager of Environmental Quality Control
 Bethlehem Steel Corporation
 Bethlehem, Pennsylvania

 Dr. Anna M. Baetjer
 Professor Emeritus of Environmental
   Medicine
 Department of Environmental Medicine
 School of Hygiene and Public Health
 The Johns Hopkins University
 Baltimore, Maryland

 Dr. Samuel S. Epstein
 Swetland Professor of Environmental
   Health and Human Ecology
 School of Medicine
 Case Western Reserve University
 Cleveland, Ohio

 Dr. Arie J. Haagen-Smit
 Professor and Director
 Plant Environment Laboratory
 California Institute of Technology
 Pasadena, California

 Dr. John V. Krutilla
 Director
 Natural Environment Program
 Resources for the Future, Inc.
 Washington,  D.C.

 Dr. Frank J. Massey, Jr.
Professor, School of Public Health
 University of California
 Los Angeles, California
Dr. James McCarroll
Professor and Chairman
School of Public Health
   and Community Medicine
Department of Environmental Medicine
Department of Environmental Health
   University of Washington
Seattle, Washington

Dr. Eugene P. Odum
Director, Institute of Ecology
University of Georgia
Athens, Georgia

Mr. Morton Sterling
Director, Wayne County Air Pollution
   Control Division
Wayne  County Department of Health
Detroit, Michigan

Mr. Arthur C. Stern
Professor of Air Hygiene
University of North Carolina
Chapel Hill, North Carolina

Dr. Raymond R. Suskind
Director, Kettering Laboratory
College of Medicine
University of Cincinnati
Cincinnati, Ohio

Mr. Elmer P. Wheeler
Manager, Environmental Health
Medical Department
Monsanto Company
St. Louis, Missouri

Dr. John T. Wilson, Jr.
Professor and Chairman
Department of Community Health
   Practice
College of Medicine
Howard University
Washington, D.C.

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                CONTRIBUTORS AND  REVIEWERS
Dr. Thomas W. Barrett
Professor of Agronomy
Arizona State University
Division of Agriculture
Tempe, Arizona

Dr. C. Stafford Brandt
Bureau of Air Quality Control
Department of Health and Mental
  Hygiene
Environmental Health Administration
Baltimore, Maryland

Dr. Harris M. Benedict
Staff Scientist
Stanford Research Institute
Menlo Park, California

Dr. Leon S. Dochinger
Principal Plant Pathologist
United States Department of
  Agriculture
Forest Service
Delaware, Ohio

Dr. William A, Feder
Professor
University of Massachusetts
Suburban Experiment Station
Waltham, Massachusetts

Dr. Jay Jacobson
Boyce Thompson Institute for
  Plant Research, Inc.
Yonkers, New York
Dr. Wilhelm Knabe
Landesanstalt fuer Immissions und
  Bodennutzungsschutz
Des Landes Nordrhein-Westfalen
Essen, West Germany

Dr. Emanuel Landau
Chief, Epidemiologic Studies Branch
Division of Biological Effects
Bureau of Radiological Health
Department of Health, Education, and
  Welfare
Public Health Service
Food and Drug Administration
Rockville, Maryland

Dr. S. N. Linzon
Chief, Phytotoxicology Section
Ministry of the Environment
Air Management Branch
Toronto, Ontario, Canada

Dr. O. Clifton Taylor
Associate Director
University of California, Riverside
Statewide Air Pollution Research Center
Riverside, California


Dr. Leonard H. Weinstein
Program Director
Environmental Biology
Boyce Thompson Institute for Plant
  Research, Inc.
Yonkers, New York
                                           VI

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       FEDERAL  AGENCY LIAISON  REPRESENTATIVES
Department of Agriculture
Dr. Theodore C. Byerly
Assistant Director of Science
   and Education
Office of the Secretary

Department of Commerce
Dr. James R. McNesby
Manager, Measures for Air Quality
Institute for Materials Research
National Bureau of Standards

Department of Defense
Commander Harvey A. Falk, Jr.
Director for Environmental Media
Office of the Assistant Secretary of
   Defense

Department of Health, Education, and
Welfare
Dr. Charles H. Powell
Director, Office of Research and
   Standards Development
National Institut   of Occupational
   Safety and Health

Department of Housing and Urban
Development
Mr. Richard H. Broun, Deputy Director
Office of Community and Environmental
   Standards
Community Planning and Management

Department of the Interior
Mr. Harry Moffet
Deputy  Assistant Director - Minerals
   and Energy Policy
U.S. Bureau of Mines

Department of Justice
Mr. Walter Kiechel, Jr.
Deputy Assistant Attorney General
Land and Natural Resources Division

Department of Labor
Mr. F.A. Van Atta
Office of Compliance
Occupational Safety and Health
   Administration
Department of Transportation
Dr. Richard L. Strombotne
Office of the Assistant Secretary
   for Systems Development and
   Technology

Department of the Treasury
Mr. Gerald M. Brannon
Director, Office of Tax Analysis

U.S. Atomic Energy Commission
Dr. Martin B.  Biles
Director, Division of
   Operational Safety

Federal Power Commission
Mr. T.A. Phillips
Chief, Bureau of Power

General Services Administration
   Mr. Harold J. Pavel
Director, Repair and Improve-
   ment Division

National Aeronautics and Space
Administration
Mr. Ralph E. Cushman
Special Assistant
Office of Administration (Code B)

National Science Foundation
Dr. O.W.  Adams
Program Director for
   Structural Chemistry

Tennessee Valley Authority
Dr. F.E. Gartrell
Director of Environmental
   Research and Development

U.S. Postal Service
Mr. Robert Powell
Assistant Program Manager

Veterans Administration
Mr. Gerald M. Hollander
Director of Architecture and
   Engineering
Office of Construction
                                          VII

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                       INTRODUCTION

  Pursuant  to  authority  delegated to  the  Administrator of the Environ-
mental  Protection Agency, Revised Chapter 5 of Air Quality Criteria for
Sulfur Oxides,  Effects of Sulfur Oxides in the Atmosphere on Vegetation,
including  revisions to related parts of Chapter  10,  "Summary  and Con-
clusions," is issued in accordance with Section 108 of the Clean Air Act (42
U.S.C. \857etseqJ.
  Air quality criteria are an expression of the scientific knowledge of the
relationship between various concentrations of pollutants in the air and their
adverse  effects  on  man  and  his environment.  Air quality criteria  are
descriptive;  that is,  they describe  the  effects that have  been observed to
occur when the ambient air  level of a pollutant has reached or exceeded a
specific figure for a  specific  time period. In developing and using criteria,
many factors have to be  considered. The  chemical and  physical charac-
teristics of the pollutants and the techniques available for measuring these
characteristics  must  be considered,  along  with exposure  time, relative
humidity, and  other  conditions  of  the environment.  The  criteria must
consider the contribution of all such variables to the effects of air pollution.
Further, the individual characteristics of the receptor must  be  taken into
account.
  The criteria in this document serve as the bases for National Secondary
Ambient Air Quality Standards.  National Secondary Ambient Air Quality
Standards specify a level of air quality, the  attainment and maintenance of
which in the judgment of the Administrator, based on criteria, are requisite
to protect the public welfare from any known or anticipated adverse effects
associated with the presence of such air pollutant in the ambient air.
  Upon promulgation  of the  standards,  each State  must prepare  im-
plementation plans that describe how these standards will be met.  The Clean
Air Act has provisions ensuring that  a plan is prepared and carried out by
each State or by EPA where States default or otherwise are judged incapable
of meeting the standards.
                                  vin

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                       CONTENTS

                                                         Page

A.  GENERAL	   1

B.  SYMPTOMS OF  THE EFFECTS  OF SULFUR DIOXIDE ON
    VEGETATION  	   1

    1.   Visible Effects	   1
        a.   Acute Injury	   2
        b.   Chronic Injury	   2
    2.   Subtle Effects  	   2
    3.   Physiological Effects  	   3
        a.   Photosynthesis   	   3
        b.   Stomatal Relationships	   3
        c.   Changes in Chemical Composition   	   3
        d.   Growth and Yield Reductions  	   4

C.  MECHANISM OF ACTION   	   4

D.  FACTORS  AFFECTING  RESPONSE OF  VEGETATION TO
    SULFUR DIOXIDE   	   4

    1.   Environmental Factors  	   4
        a.   Temperature 	   4
        b.   Humidity	   5
        c.   Light  	   5
        d.   Edaphic Factors	   5
        e.   Diurnal Changes	   6
        f.   Interaction with Other Pollutants	   6
    2.   Genetic Factors  	   6
    3.   Stage of Development   	   6

E.  PROBLEMS IN  DIAGNOSIS  AND ASSESSMENT OF THE
    ECONOMIC IMPACT OF  SULFUR DIOXIDE  	   7

F.  EFFECTS ON LOWER ORGANISMS  	   8

G.  ACID PRECIPITATION  	   9

H.  MISCELLANEOUS ASPECTS OF SULFUR DIOXIDE EFFECTS
    ON VEGETATION  	  10

    1.   Vegetation as a Sulfur Dioxide Sink   	  10
    2.   Effects of Sulfuric Acid Mist on Vegetation  	  10

I.   EFFECTS ON BIOMASS AND YIELD   	  10

J.   DOSE-INJURY RELATIONSHIPS  OF SULFUR DIOXIDE TO
    VEGETATION RESPONSE  	  12
                             ix

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                                                        Page

    1.   Mathematical Equations  	14
    2.   Dose-Injury Data   	 16

K.  SUMMARY   	32

L.  CONCLUSION  	 34

M.  REFERENCES  	 35

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                     LIST OF  TABLES

Table                                                            Page

5-1   Sulfur  Dioxide Concentrations and  Associated  Vegetational
     Effects in Research Areas at Biersdorf, Germany   	    13

5-2   Changes in Net Tree Volume of Eastern White Pine Associated
     with Sulfur Dioxide  Concentrations near a Smelter Complex in
     Sudbury, Canada	    14

5-3   Growth Reduction in Vegetation Exposed to Sulfur Dioxide for
     Long and Short Time Periods	    15

5-4   Seasonal  Average Sulfur  Dioxide  Concentrations  Associated
     with Foliar Injury to Spruce  in Two Locations in  Czechoslo-
     vakia  	    17

5-5   Concentrations of Sulfur Dioxide Causing Injury to Sensitive
     Vegetation   	    18

5-6   Sulfur Dioxide Concentrations Causing Injury to Agricultural
     and Forest Species   	    19

5-7   Vegetational Response to  Sulfur Dioxide in Combinations with
     either Ozone or Nitrogen Dioxide  	    21

5-8   Projected  Sulfur Dioxide Concentrations  that Will  Produce
     Threshold Injury to Vegetation for Short-Term Exposures   ...    24

5-9   Lists of Plants in Three Susceptibility Groups by Sensitivity to
     Sulfur Dioxide   	    25
                                  XI

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   EFFECTS OF SULFUR  OXIDES  IN THE  ATMOSPHERE
                               ON  VEGETATION;
  Revised  Chapter 5 for Air Quality  Criteria for Sulfur Oxides
              A.  GENERAL

  The sulfur oxides represent one category of
pollutants that affect  plant life. Within  this
category of pollutants, sulfur dioxide  (SO2)
appears to be the major causal agent of plant
injury, although plants may respond to other
compounds  of sulfur  such  as sulfuric acid
aerosols.  In  one  of  the  earliest reports
concerning  SO2  injury to   plants,  Stoeck-
hardt,1  in 1871,  discussed smoke damage to
forest  trees in  Germany. Since  this early
report,  extensive  experiments and  observa-
tions of the  effects of SO2 on vegetation have
been made by investigators in  all parts  of the
world. Thomas,2 >3 Brandt and  Heck,4  Katz
and  McCallum,5  and  Daines,6 have written
reviews of these studies. Studies of the effects
of sulfur oxides upon  vegetation need to be
continued if  the manner in  which  sulfur
oxides cause plant injury is to be understood.
  Plants vary greatly in their  response to
SO2.  This variation in response by plants is
due  to their  genetic  composition,  to  their
response to  environmental factors, individual-
ly and in  combination,  and  to the  time-
concentration  relationship of SO2  by itself
and  in  combination with other atmospheric
pollutants.   Variation  in  any  one  of  the
complex of  interacting factors will result in a
variation in plant  response.
  This  document is not intended as a com-
plete, detailed literature review, and it does
not  cite every published article relating to
effects of sulfur oxides in the ambient atmos-
phere upon  vegetation. However,  the  litera-
ture, comprising more  than  700 articles, has
been  reviewed  thoroughly  for  information
related  to the development of  criteria. The
document,  based  on  both professional  and
scientific judgment, not only summarizes the
current  scientific  knowledge of air pollution
effects by sulfur oxides upon vegetation, but
also points up the major deficiencies in that
knowledge.
   B. SYMPTOMS OF THE EFFECTS OF
   SULFUR DIOXIDE ON VEGETATION

   The  effects  of SO2 upon plants can be
classified into two general categories:  visible
effects  and subtle effects. Visible effects are
identifiable   pigmented  or  necrotic  foliar
patterns that result from major physiological
disturbances to plant cells. Subtle effects are
those that are  not  visibly identifiable  but
result in measurable growth or physiological
changes in the plant.  Subtle effects are not
visibly identifiable and can be identified only
when  measurable growth  or physiological
changes occur in the plant. Both  visible and
subtle  effects are physiological effects  and
result from the  disturbance of biochemical
processes at the molecular level.  Whether or
not the biochemical disturbances give  rise to
visible symptoms determines the  category to
which they are assigned.

             1.  Visible Effects

   Visible  effects  to plants  can  be  further
classified into acute and chronic injury. Acute
injury is severe injury that occurs within a few
hours after exposure and  is characterized by
the  collap»?  of  cells  with  the  subsequent
development  of  necrotic  patterns.  It is as-
sociated with high, short-term SO2  concentra-
tions, although severe injury, similar to acute
injury, may develop  from chronic exposures.
Chronic injury  is  light to  severe  injury that
developes  from  exposure  over an  extended
time period.  It is associated with long-term
exposures where the pollutant concentration
is  sufficiently high  to produce   some  cell
destruction or disruption. It is identifiable by
chlorotic or other pigmented patterns and in
some  instances  is associated  with  necrotic
markings. Acute  injury symptoms  are general-
ly  more characteristic  of a specific pollutant
than those of chronic  injury,  which are  not
necessarily  specific  for a  particular toxic

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agent. Insects, nutrition, microbiotic diseases,
and  other factors  can  produce leaf injury
patterns similar to those induced by SO2.
  Foliar  symptoms  in plants  have  often
provided the first indication  of a pollution
problem; however, since SO2 type symptoms
may result from other abiotic and/or biotic
influences, related  evidence  must  be  con-
sidered before attributing injury to SO2. The
related evidence should include a knowledge
of  SO2  sources and  observations  showing
decreased injury levels with increasing distance
from  the  source.  Monitoring  of  pollution
concentrations,  consideration   of  meteoro-
logical conditions, and observation of several
plant  species, especially  when symptoms are
not characteristic, can also aid in the diagnosis
of injury.  The  use  of field chambers from
which pollution has been removed may also
contribute  to identifying the cause of injury.
  Descriptions  of  SO2  injury  are found in
numerous  publications.2'3'6"9  These reports
also  consider  symptoms  caused  by  other
pollutants  and  various  biotic  and environ-
mental stresses that may produce symptoms
resembling those caused by SO2. Summaries
of effects  on a large number of plant species
susceptible to SO2  are given by Wood10  and
by  Middleton and  Taylor.11  Three pictorial
atlases document visual SO2 symptoms12"14
and  include  detailed  descriptions  of injury
with a listing of SO2 susceptible species.
 a. Acute Injury
               12.13
   Acute symptoms of SO2 injury result from
 the  rapid absorption of toxic concentrations
 of the gas.  In broad-leaved plants, tissues in
 sharply defined marginal and  interveinal areas
 take on a  dull water-soaked  appearance im-
 mediately after exposure.  These areas  subse-
 quently  dry  and  may bleach  to  ivory or
 become brown to reddish-brown in color. The
 separation of injured areas from surrounding,
 apparently  healthy, tissue is  usually distinct.
 Injury seldom extends across  leaf veins unless
 the injury is severe.
   The basic bleached and collapsed  blotches
 described on broad-leaved plants are,  how-
 ever, also  typical  of grass foliage. The final
 bleached pattern between the parallel veins of
 grass leaves  gives a streaked effect.
  Acute injury of conifers  usually occurs in
bands on needle tips, with injured areas taking
on  a  red-brown  color.  Injured- areas change
from  the  usual dark green  color to a lighter
green,  and lesions develop yellow-brown and
finally red-brown coloration. In severe  cases,
discoloration may involve the  whole needle.
The affected trees usually cast their needles
prematurely.

b.  Chronic Injury12 >13

  Low  concentrations   of sulfur  dioxide
require several days  or  weeks to  cause  the
development of  the  yellowing or chlorotic
symptoms of chronic injury.  The chlorotic
effect,  with  varied  color  patterns,  often
resembles  premature  senescence.  Necrosis
may develop in some plants, resulting in white
bleached  areas or red to brown  coloration,
which  may resemble acute injury. Chronic
injury  may  be followed by  leaf abscission. A
large amount of sulfate is found in leaves with
chronic symptoms,  whereas leaves that  are
acutely injured show  only a small increase in
sulfate content.  However, large quantities of
sulfate may accumulate in leaf tissue without
visible leaf  symptoms.15'16 Both  acute and
chronic injury symptoms may  develop upon
the same  plant. The period of development
and the sensitivity of the plant to particular
sulfur dioxide concentrations are important in
differentiating the type of injury.
  Chronic injury, when exhibited on plants
exposed to  SO2, is due  to  either short-term
peaks  or  long-term  average concentrations.
The general consensus of most investigators is
that short-term  peaks  are  more  important
than long-term averages.

             2. Subtle Effects

  Subtle  effects, as a concept, implies that
SO2  can interfere  with physiological and
biochemical processes and with plant growth
and yield without attendant development of
visible  symptoms.  The  processes are  mi-
croscopic  or molecular in nature. Therefore,
in  order to determine their  existence, studies
have to be conducted that can detect whether
measureable  changes  in  the rate  of photo-
synthesis, in stomatal  behavior, and in growth
or yield have occurred.

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          3. Physiological Effects

   Physiological  effects include both  visible
and subtle effects. Both types of effects result
from   the  disturbance  of  physiological
processes at the molecular level.  Whether or
not  the  physiological  changes give  rise to
visible symptoms  determines to which cate-
gory they are assigned.

a.  Photosynthesis

   Wislicenus17   indirectly  related  SO2  to
photosynthesis  in  demonstrating  that  the
sensitivity of spruce to SO2 was proportional
to light  intensity.  Thomas and  Hill18  and
Katz9  reported  that  exposures of  alfalfa to
high concentrations of SO2  for short time
periods resulted in a  transitory reduction in
carbon dioxide (CO2) assimilation; recovery
began within an hour after treatment.  In the
latter  experiments,9  the  CO2 suppression
response was recorded for about 2 days. Katz
and  Lathe8 and Katz9  reported  that SO2
concentrations of  262 to 524 jug/m3  (0.1 to
0.2  ppm) did  not  affect  photosynthesis,
respiration, stomatal behavior, or growth but
that  concentrations above  1048 jug/m3 (0.4
ppm)  did  affect  sensitive  plants,  if  the
stomata were open.
   These  results  and   the  research   of
others1 9 >2 ° indicate that the rate of photo-
synthesis is reduced soon after sensitive plants
are exposed to SO2. If visible injury does not
occur,  the  photosynthetic rate  returns  to
normal   after  exposure  terminates,  but if
injury  results,   complete  recovery  is  not
attained.  The   magnitude of  the  photo-
synthetic response varies  with  respect  to
pollutant  concentrations,   environmental
influences, and plant sensitivity.21"24
   Recent information indicated that SO2  was
reduced to hydrogen sulfide (H2 S) by several
plant  species  during   and   after  fumiga-
tion.25"27  This reaction may  be  associated
with photosynthesis,  since the response  was
obtained only in the presence  of light.2 7

b.  Stomatal Relationships

   Stomata are  the principal avenue of SO2
entrance  into  plant leaves. Conditions that
 favor open stomata result  in increased SO2
 assimilation  and   increased   plant   sensi-
 tivity.3'9'28   Majernik and Mansfield29'30
 demonstrated a stimulation  of stomatal open-
 ing in bean plants when they were exposed to
 SO2 at relative humidities above  40 percent
 but  a suppression  of  stomatal  opening at a
 relative humidity of 32 percent.
   Katz9  found  a slight  reduction in  the
 number of open stomata after exposing alfalfa
 to  2358  /ug/m3  (0.9  ppm) of  SO2  and a
 significant reduction after exposure to 2620
 jug/m3  (1  ppm). Continuous fumigation at an
 average concentration of  1050  /xg/m3  (0.4
 ppm) did not influence stomatal opening until
 acute  injury  symptoms  developed.  Vogl19
 reported that  stomata of pine remained open
 after plants  were  injured.  Neither  of these
 reports  included humidity conditions.
   Spedding31  presented  information  that
 suggested   that humidity  influenced  the as-
 similation  of  SO2 when stomata were closed
 and  that  SO2  also  entered  plant  tissues
 through the cuticle.

 c.  Changes in  Chemical Composition

   Sulfur  dioxide  exposures  may result  in
 changes  in  the  chemical   composition  of
 plants.  Materna3 2  found  increases in sulfur
 and  potassium  levels  when spruce  needles
 were exposed in  spring,  but  calcium and
 magnesium  levels  were   not affected.  For
 citrus leaves,  calcium  and  potassium  levels
 decreased  during  winter exposures; however,
 in  the  summer,  calcium  levels were not
 affected and  potassium content  increased.33
 Materna34  reported increases in  the silicic
 acid content  of  spruce  needles injured  by
 SO2.
   Arndt3 5 demonstrated  increases in  amino
 acid concentrations of herbaceous plants after
 exposure to 660 jug/m3 (0.25 ppm) SO2. The
 amount  of increase  depended   on specific
 amino acids and  plant species. In exposures
 producing chronic injury, Boertitz2! reported
no significant  change in pH, carbohydrate, or
 amino acid content of extracts  from spruce
 needles; however, in more recent field studies
of the Ore Mountain area, he found increases
in the carbohydrate levels and pH values of
needle extracts.36

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   Injury resulting from chronic SO2 exposure
can usually be  confirmed by the presence of
high sulfur content in leaves, although varia-
tion  in  normal  sulfur  content  must   be
considered.7'37 In Japan,38  sulfur content of
citrus leaves correlated with atmospheric SO2
concentrations.   Katz9  demonstrated that
sulfur   content  increased  with  time   of
exposure. Under natural conditions,  analysis
can reflect the  degree of pollution to which
vegetation  has  been  exposed.   In  some
industrial  areas,  McCool   and Johnson39
found a  decrease in sulfur content  of vegeta-
tion with an increase in distance  from  the
SO2 source.

d.  Growth and  Yield Reductions

   Growth and yield reductions  may result in
the absence of  visible injury. Tingey  et a/.40
demonstrated reduced root weights of radish
when  exposed to SO2  concentrations of 131
to  160   jug/rrf  (0.05 to  0.06  ppm)  for
40 hr/week for  5   weeks   in greenhouse
exposure chambers; however, the second run
of the experiment did not produce the same
plant  response. Other studies by Reinert et
al.41  showed  reductions  in several  growth
parameters for Bel W3  tobacco when exposed
to 262  jug/m3  (0.1 ppm) SO2,8  hr/day,  5
days/week,  for  4  weeks   in  greenhouse
exposure chambers. Bel W3  is a variety  of
tobacco extremely sensitive to injury  by SO2
and has been used as a plant monitor.  Both of
the above studies were  conducted under
conditions  which would seldom, if ever,  be
found in the ambient air.  More studies  are
needed to verify the results.
   In   ambient  air   studies,  Bleasdale42
reported  that growth of rye grass was reduced
when  the maximum average SO2  concentra-
tions for 24 hours were between 262 and 524
jug/m3  (0.1 and 0.2 ppm) for 2 and 3 days
during experiments of 63 and 73 days. In this
study, interactions with pollutants other than
SO2  were  not considered  and  may have
contributed to the observed growth response.

      C. MECHANISM OF  ACTION

   The mechanism by which plants are injured
by SO2  is not understood. Transient  physio-
logical effects, subtle growth reductions, and
acute injury symptoms may result from  the
formation of sulfite ions and their effect on
membrane  integrity. Acute injury does not
occur if the rate of SO2 absorption does not
exceed the capacity of the plant to oxidize
the sulfite  to  sulfate  ions. Under long-term
SO2   stress, sulfates  thus  formed  may  ac-
cumulate with the subsequent development of
chronic injury symptoms.33

D. FACTORS AFFECTING RESPONSE OF
   VEGETATION TO SULFUR DIOXIDE
   The response  of a given variety or species
of plants to a specific air pollutant cannot be
predetermined on the basis  of the known
response of related plants to  the same pol-
lutant.  Neither  can  the response   be  pre-
determined by a given response of a  plant to
similar  doses  of  different  pollutants.  The
interplay  of genetic  susceptibility,  growth
stage, and environmental influences  must  be
considered  for each plant and pollutant.  No
one  factor  may  be considered independently
of the other factors.

         1. Environmental Factors

a.  Temperature

   Plants are  more  resistant  1o   SO2   at
temperatures below 40° F.43'47  Setterstrom
and  Zimmerman43  reported that buckwheat
was  equally susceptible to injury at  65° and
105°F.   Several  investigators9'48  have
reported greater resistance in conifers during
the  winter and  have related  this to lower
physiological activity  of plants. Resistance
may  increase  during  winter dormancy with
low gas exchange rates; however, even at low
levels of physiological activity,  conifers may
be injured, especially in areas with higher SO2
concentration  during  winter months. In  ad-
dition, temperatures are often near 40° F
during winter seasons in many  areas. Van
Haut and Stratmann13 indicated that conifers
remain sensitive during the winter when water
is  available  to  them. In  the  spring, with
increases in physiological activity, sensitivity
to SO2 also increases. On the basis of studies
with  Douglas fir  and yellow  pine, Katz9

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reported that in  the spring, with increases in
physiological activity,  sensitivity to SO2 also
increases. Exposures to SO2 at concentrations
of  1965  Mg/m3  (0.75 ppm) for  147  hours
near the end  of  the winter dormancy period
resulted  in foliar injury  of 55  percent; how-
ever,  in  early autumn  this concentration was
applied for 334  hours without the develop-
ment of injury. For spruce, experiments have
demonstrated increased  sensitivity  in the
spring and  autumn   when  compared  with
summer and winter seasons.49

b. Humidity

   Sensitivity to  SO2  tends to increase with
increasing humidity.43^*5 Wells45 noted that
in the Salt Lake  Valley  70 percent appeared
to be  the critical humidity level.  Above 70
percent,  plants were much more susceptible
to  injury  by  sulfur  dioxide  than  below.
Swain44  concurred but  stated  that increases
in relative humidity (RH)  from 70  to 100
percent  did  not  result in much increase  in
sensitivity. In 1-hour exposures at an  average
concentration of  3537  jug/m3   (1.35  ppm)
SO2, Zimmerman and  Crocker5 ° found that
variations between 50 and 75 percent RH had
little effect on plant sensitivity.  Setterstrom
and  Zimmerman43 concluded  that for RH
values  above  40  percent,  differences of 20
percent RH are required to produce detect-
able differences in  sensitivity; however, they
mention  that O'Gara in an address before the
American  Institute  of  Chemical  Engineers
stated  that plants were  three times as sensitive
at an  RH of 100 percent as at 30 percent.
Thomas   and  Hendricks28  reported  a  90
percent  loss  in  sensitivity  when  RH  was
reduced  from 100 to  0 percent. Generally,
resistance to injury by  sulfur dioxide seems to
be   associated   with  decreasing  relative
humidity; however, variations associated with
a particular  plant  species or with environ-
mental conditions do exist.
  Under  conditions  of  high   humidity,  a
sulfuric acid mist  may  form. In fog,5 * >5 2 this
acid  mist may cause leaf spotting on several
plant species.

c.  Light

  Setterstrom and Zimmerman43  reported
that buckwheat  was  more susceptible  to
injury from  SO2  when  grown  under condi-
tions of reduced light intensity.  A 65 percent
reduction in  light intensity (to approximately
3000 foot-candles) resulted in greater suscep-
tibility  than that for plants grown in full
sunlight  (approximately  10,000  to 12,000
foot-candles) or under conditions of 25 and
35 percent reduction in light intensity.  Light
received prior to treatment affected sensitivity,
since plants  kept in the  dark for 2  hours
preceding SO2  exposure were more resistant
than  comparable plants  kept  in the light.50
Plants are 5 to  6 times as resistant to SO2  in
the dark as in  the light.13'45 Since stomata
of most plants are closed  during darkness,
plants are  more  tolerant  of  SO2  in the
absence of light. In studies conducted at night
with exposure to 2227 jug/m3  (0.85  ppm) for
4  hours,  foliar injury  of alfalfa,  tomato,
buckwheat, sweet clover, oats, rye, and barley
was  not observed; however, injury resulted
during daylight exposures of  2096 Mg/m3
(0.8  ppm) of  SO2  for  periods of 1  or 2
hours.50 In  experiments with  bush beans,
SO2  exposures that produced moderate injury
during the  day had no effect at night.13-53
However,  when plants were exposed during
the day following a night exposure, injury was
greater  than the  single  day  exposure.  The
reverse order also resulted in increased injury,
as did exposure under continuous light for 24
hours.

d. Edaphic Factors

   Plants  are more  sensitive  to  SO2  when
adequate soil moisture is available for normal
plant  growth. Minor  variations  in  soil
moisture have  no  detectable effect  on  sensi-
tivity; however, when moisture content ap-
proaches the wilting point or if wilting occurs,
plant resistance increases.9 >4 3 >5 ° >5 4 Brandt
and  Heck4 recommended withholding  water
from greenhouse and irrigated  crops during
periods of   high  pollution  potential  :>s  a
preventative measure for reducing damage.
   Investigations of the effect of soil nutrient
levels  in  relation to plant  sensitivity have
involved comparisons of plants growing under
various nutritional or fertilization levels. Such
studies  indicate an increased  resistance with
increased  fertilization in  rape,  spinach, and
radish.54  Varied  results  have been  recorded

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for  pine.55"57  In  the Ore Mountains  of
Czechoslovakia,34   nutrient   applications
resulted in an increase in resistance to SO2 for
several tree species. In field  and laboratory
experiments, Cotrufo  and Berry58  found  a
reduction in SO2 induced  needle necrosis of
several white  pine  clones  after fertilizer ap-
plications.   In  contrast,   deficiencies   of
nitrogen  and  sulfur  were  correlated with
increased   resistance  for  tobacco  and
tomato.59  With alfalfa,43  nutrient  deficient
conditions increased sensitivity, but with oats,
increases in  nitrogen and other nutrients v/ere
associated with increased sensitivity.54
   Few studies have considered the influence
of soil  structure, soil temperature,  aeration,
and  the  biotic complex on plant sensitivity.
Brandt  and Heck4  state  that sensitivity is
reduced when plants are grown in heavy soils.
This  may be  the  result  of lower oxygen
tensions.  In  studies  involving  three  plant
species and  four soil types, Guderian60 found
that plant injury varied in respect to soil type,
•iitrogen application, and species of plant.

e.  Diurnal Changes

   The sensitivity of plants to SO2 may vary
during  the  day.  Factors  that  favor open
stomata and photosynthesis also favor SO2
assimilation. Under these  conditions,  plants
are more sensitive  during the morning than
during   the   afternoon.  Thomas  and
Hendricks28 concluded that on  a  cloudless
day exposure of alfalfa early in  the morning
resulted  in  only slight injury, while  exposure
later  in the  morning  resulted  in increased
injury; plants exposed between late morning
and mid-afternoon had decreased injury, with
the  most  rapid  decrease  later  in  the day.
Although climatic  conditions and  stomatal
movements are important factors in diurnal
injury patterns, decreases in sensitivity during
afternoon  periods  may be  related to  the
accumulation  of carbohydrates in leaves13 '2 8
or  to an increase  in buffering  capacity  of
plant tissues.54

/  Interactions with OtherPollutants

   Ambient  air is composed of many different
pollutants. A few studies have considered this
fact, but  most studies deal -with single pol-
lutant  effects. The interaction of ozone (03)
with  SO2  was  first  reported on  Bel  W3
tobacco  by  Menser  and  Heggestad.6*  The
interaction  of these  air pollutants  affecting
injury  and growth in  several other species has
been   reported.62"64  Injury   to six  plant
species from  the interaction of  nitrogen
dioxide (NO2) and  SO2  has recently been
reported.6 s   Growth  and  injury results  are
further discussed  in  Section  J.2. The inter-
actions shown between  SO2  and other pol-
lutants offers  a  partial  explanation for oc-
casional  inconsistencies   between   results
obtained in  laboratory studies in which only
single  pollutants   were used and the  results
obtained  in  the  natural  environment. For
example,  growth reductions   of  radish  oc-
curring after  exposures   to  mixtures  of
SO2/O3  were  greater than reductions from
SO2 alone.

             2. Genetic Factors

  Plant sensitivity to  SO2  can be considered
as  a  function of morphological  and  bio-
chemical  characteristics controlled  by  the
genetic plasticity  of the  plants  within a
population. Thus, some plants are more sensi-
tive  than others to  pollution stress. Both
inter-  and intraspecific differences  in sensi-
tivity  occur. For example, sensitivity varia-
tions within  species have been demonstrated
with spruce66 and  white pine.6'7   In this
regard, SO2  acts as a   selection  pressure
mechanism.   The  more   resistant  variants
within  a species continue normal growth and
existence, but  under this  SO2  pressure,  the
more  sensitive types  weaken and may not
survive within the polluted area.
  Shapiro,  Servis, and Welcher,68 based on
experiments  conducted  with  isolated DNA
and  bacteria,  using  sodium   bisulfite,  have
suggested  that SO2 in the atmosphere may
constitute a genetic hazard. At present, how-
ever, there is no  basis for extrapolating from
such experiments to living  organisms that are
structurally  and  functionally  more complex.

          3. Stage of Development

   The growth stage or phase  of development
at  which plants  are  exposed  to SO2 affects

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their susceptibility and  the yield losses asso-
ciated  with  injury.  Wells69  reported that
barley  yields were slightly reduced  by SO2
exposures when plants were from 10 to 60 cm
high, but that yields were reduced by 20 to
30 percent if plants were exposed during early
grain  development. For wheat, Brisley and
Jones7 °  demonstrated  greater yield  reduc-
tions  with exposure  in the  early stages  of
growth than in later stages. Thomas71 demon-
strated that injury to cereals at tillering could
be sustained with little loss in yield, but that
yield was reduced much more by injury after
culms had formed. In  several plant species,
van Haut5 3  reported a  "critical development
stage"  in which  there was a high probability
that leaf injury would result in reduced yield.
This critical  stage occurred with bean shortly
before flowering and during pod growth; with
radish, at'the  young seedling stage and again
as the roots began to increase in size; and with
oats just before panicle  emergence, at flower-
ing, and during flower opening.
  The leaves of most plants are more sensitive
to SO2  just after maximum  expansion has
occurred. Developing and older leaves tend to
be more  resistant.7 >4 3 >5 3 Variation in suscep-
tibility  between species has  been observed.
Van Haut5 3 found  that needles of pine and
larch are very sensitive to  SO2 before growth
has been completed.
    E.  PROBLEMS OF DIAGNOSIS AND
     ASSESSMENT OF THE ECONOMIC
      IMPACT OF SULFUR DIOXIDE

  The plant is a product of its environment.
Every environmental factor, favorable or un-
favorable, produces a response in the .plant.
Sulfur dioxide interacts  with other environ-
mental factors such as the climate, soil, biota
(insects, man,  and  microorganisms), and the
genetic constitution of the  plant to produce
responses  within the plant. Injury  produced
by  SO2  may  not only  be  modified  or ob-
scured by these  other environmental factors,
but the plant may develop injuries from these
other factors that  are difficult or impossible
to  distinguish from those  caused  by  SO2.
Ornamental   and  agronomic  crops  grown
under special management  practices must be
carefully  examined  before  attributing poor
growth to SO2. Many  bacterial, viral, and
fungal diseases,  as well as insect infestation,
can  produce  symptoms  in  plants  that  are
quite similar to  those produced by  SO2. To
aid in making  definitive diagnoses of  SO2
effects  upon  vegetation,  injuries must be
observed in the field and supported by labora-
tory  studies  using  different levels  of the
pollutant.  Laboratory   and   field  chamber
studies are essential  if qualitative and quanti-
tative  models  of   pollutant  effects  upon
vegetation are to be developed.  Since  it is
impossible to  include all parameters, labora-
tory  and  field chamber  studies  do  not
simulate ambient field conditions.
   The question  that must be answered in the
assessment  of  SO2 damage  to  plants  is
whether or not  the plant has been so altered
by the pollutant as  to significantly influence
its growth, survival, yield, or use.  In cases
where leaf injury impairs the use of the plant
for food,  as in the case of cabbage or lettuce,
or for  ornamental  purposes,  assessment  is
relatively   straightforward. However,  where
the marketable  product  is not influenced by
appearance, assessment of economic damage
is  more difficult. Hill and Thomas72 showed
that   the  yield  of  alfalfa  was  reduced in
proportion to the area of the leaf destroyed.
The  economic impact of leaf injury to fruit
trees  is extremely difficult to assess because
the effect upon the fruit is not known.
   In  discussing  the effect of air pollution on
vegetation, Guderian, van Haut,  and Strat-
mann7 3 have  suggested a method of making
the distinction between the terms injury and
damage in  cases  where  the effects  of air
pollution  on  vegetation  is  concerned. Ac-
cordingly, injury is defined as any identifiable
and  measurable  response of  a  plant  to air
pollution. Damage resulting from air pollution
injury is   defined  as any  identifiable  and
measureable adverse effect upon the desired
or intended use of  a plant or of a product
derived from the plant.  Thus, in using these
terms, leaf necrosis of alfalfa is a symptom of
injury; however, any assessment  of damage
requires a judgment that the injury affects the
yield or use of the plant.
   There are instances, also, where the esthetic
or sentimental value  of the plants is impaired

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by SO2. Examples of these are the vegetation
growing on a hillside and a tree planted by a
particular  college  or  university  class  as  a
memoir of their college  days.  In the first
instance, leaf damage or the death of the
plants  detracts from the view. In the second,
the tree is largely of sentimental value, and its
demise  is  extremely  difficult  to  assess in
dollars and cents.

 F. EFFECTS OF LOWER ORGANISMS

  The effects of SO2  on nonvascular plants
and on plant  pathogens have  been  studied by
many  investigators.  The  majority of these
studies  have  considered  the  incidence  of
specific organisms within areas influenced by
SO2  emissions.  The  absence of  species of
lichens and  bryophytes has  been  correlated
with the presence of  low  concentrations of
SO2-74"79  Lichens have  been used  in the
recognition and monitoring of SO2,80"82 and
qualitative  scales  for  estimating  SO2  con-
centrations have been developed on the  basis
of  sensitivity  differences  among  species.
Skye83  found  that  the  diversity  of species
was  reduced  in areas with  an  annual  SO2
concentration of approximately 39.3  jug/m3
(0.015 ppm) (determinations were averaged
over 4-week  periods).  Gilbert74 found that
several  species  of  bryophytes  and lichens
disappeared  when  winter averages (October-
April)  exceeded  52 ng/m3  (0.02 ppm). In a
lichen  transplant  study, death  of the  test
species occurred within 29 days at locations
with the highest average SO2 concentrations.
The  SO2  concentration, determined  inter-
mittently,   averaged   230   Mg/m3   (0.087
ppm).84
  The extreme sensitivity of lichens to SO2
appears to be due to  the breakdown of the
algal component.  Rao and LeBlanc85  have
shown that SO2  absorption by lichens causes
the  decomposition   of   chlorophyll  a  to
phaeophytin  a.  Experimentally, chlorophyll
breakdown occurred when the lichens  were
exposed to concentrations of 13,100 Mg/m3
(5 ppm) for 24 hours.
  The importance of long-term average con-
centrations  versus  many shorter terms of
higher  concentrations on the  reaction in
lichen populations awaits critical study.
   The effects of SO2 have also been inves-
tigated  in  relation  to  the  occurrence of
various   biotic   plant  diseases.  Koeck86
observed  the absence  of mildew on oak in
areas near SO2 sources, while the disease was
widespread  in  areas   distant  from  these
sources.  Scheffer and  Hedgcock87  observed
that the incidence of  several rust and other
fungal diseases was low in areas influenced by
SO2, but root rot caused by Armillaria mellea
and  bark  beetle infestations  were  more
prevalent on declining  trees affected by SO2.
This  relationship is  characteristic  of these
secondary pathogens.   For soil pathogens,
population  increases  may be related  to in-
creases  in  soil  acidity  in addition to  the
presence  of  more  susceptible   hosts.88
Saunders8 9  demonstrated  that SO2 reduced
the  incidence and severity  of  the fungus
Diplocarpon rosae, causing blackspot of roses.
Results  from field  studies suggested  that
average daily concentrations above 105 Mg/m3
(0.04 ppm) SO2 nearly eliminated the black-
spot disease.

   In sample areas near  a  Sudbury, Ontario,
smelter complex, Linzon90 noted that fewer
white pine trees were affected by blister rust,
heart rot, and insect infestation. In contrast,
the  occurrence  of bark  abnormalities  was
higher  on  white  pine  near  the smelter
complex than in other research areas. These
abnormalities  consisted  of a rough bark or
canker condition and a purple bark  condition
that  appeared   as  an  unnaturally  purplish
color.

   In southern Poland,91 acidification of tree
bark was correlated  with air pollution by
SO2. The relationship  of this phenomenon to
the  growth  and   development  of  bark
organisms has not been studied.

   In  severely  injured  conifer  stands,
Boesener9 2  found higher populations of bark
breeding insects  than in stands that exhibited
lower amounts of injury. Boesener indicated
that the  high insect populations accelerated
tree decline.  In another  insect population
study,  Przybylski93   observed  increases in
aphid  populations  in  areas  near  an  SO2
source. He  concluded  that this  increase  may
be related to reductions in aphid predators.
 8

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         G. ACID PRECIPITATION

   The oxidation and the solution of SO2  in
water  has  increased the acidity of precipita-
tion in several areas of the world. Based on
pH values dating from 1955, Oden94  reported
an  increase in  acidity of precipitation over a
12-year period in  Sweden. The lowest single
value,  a  pH of 2.8, was recorded in 1967. A
similar trend has been reported in the north-
eastern United States. Within this region, the
lowest  annual  average  pH   of  4.03  was
recorded at the Hubbard Brook Experimental
Forest,  New  Hampshire.95   At   Hubbard
Brook, the lowest single pH  was 3.0. Such
trends have been related to SO2 and to some
extent to oxides of nitrogen (NOX) emissions
from industrial complexes within these areas.
In addition, particulate matter emitted from
combustion processes also contributes to in-
creased acidity.96'97
   Values associated  with nonindustrialized
areas are also lower than the  neutral  pH of
7.0, but  these values  are  related to the
conversion of  normal  atmospheric  CO2 to
carbonic acid rather than to the stronger acids
resulting from  reactions with  SO2.  In  these
nonindustrial  areas, pH values between 4.9
and6.898'101 have been recorded.
   The acidic  precipitation has resulted in
increased   acidity  of  soils,   rivers,   and
lakes.7 ,7! ,94,! °2  Several researchers  have
related increased soil and water acidification
to   ground   level   concentrations  of
SO2.7'102'103  Increases in soil acidity can
affect  the availability of plant  nutrients and
change the species composition of soil micro-
organisms,  with possible concomitant reduc-
tions in  the  rates of mineralization  and
decomposition  processes.94'104'105  Changes
in these  processes  can affect the growth and
development of higher  plants.  Although soil
acidification  did  not  appear  to  affect de-
composition  processes   in  an  arid indus-
trialized  region  of   Czechoslovakia,  the
number of aerobic  bacteria and actinomycetes
was  reduced in research plots  near  the  pol-
lutant  source,  while   increases  in fungal
populations were recorded.79 Fungi  are more
tolerant of acidic conditions.
   Oden94  indicated  that effects  on  plant
growth are related to  the content  of basic
compounds in the soil. In this regard, soils of
basic composition, such as arable soils,  are
more resistant  to pH  change. Calculations
have revealed that acidification of these soils
will  require  a time  period  of 125  to  1000
years.   This  period could  increase  with
weathering  and  application of lime to soils.
For forest soils, which tend to be more acidic,
this time period is only 30 to 50 years. Sandy
soils are affected most by the acidification
reaction;  whereas,  soils  of  limestone and
basalts are affected  least because weathering
of these materials effectively neutralizes  the
acidic effect.
  Conclusive evidence involving the  effect of
soil  acidification on  forest productivity  has
not been  presented; however, several reports
have  described   possible  influences.105   In
Norway and Sweden, the amount of calcium
in upper  soil  zones was  related to  forest
productivity. This relation  was based on  the
conversion of calcium carbonate (CaCO3) to a
more soluble calcium sulfate (CaSO^) com-
pound by the action of sulfuric acid (.I2SO4)
from precipitation.  Compounds  of calcium
were then removed from the soil by leaching
and  run-off. Growth effects were estimated
from the  relationships of acid deposition  to
calcium removal and the effect of reduced
quantities of calcium on forest growth. Based
on  this information, an annual  decrease  in
growth  of  about  1  percent per year was
determined.
  Other  investigations  in   Sweden   have
compared tree growth in  areas affected  by
acidification  with relatively   unaffected
regions.  Comparisons  of  the  two   areas
indicated  that  tree  growth  was reduced  in
affected areas. In this case, the annual growth
reduction  amounted to approximately  0.3
percent per year. If the 1965 to 1970 levels of
sulfur emissions  remain constant, a reduction
in forest growth of between 10 and 15 percent
has been estimated for the year 2000. * °5
  The  effect  of acidic  precipitation  on
herbaceous  plants has  also  been  studied.
Cohen and  Ruston106  demonstrated reduc-
tions in the growth of timothy grown in pots
when plants were irrigated with acid rain and
H2SO4  solutions  at concentrations  between
10 and 320 ppm. This was within the range of
normal acidity levels of 5 to 100 ppm H2SO4 .

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  Leaching  of nutrients from plant foliage
has also  been associated with the increased
acidity   of  precipitation.   In  the  Ore
Mountains, Materna107 found that sulfur in
precipitation contributed 39  kg sulfur per
hectare (ha) per year to soil of an open area,
whereas soil under a  forest canopy  received
133 kg/ha-yr. With leaf fall, an additional 10
to 20 kg/ha-yr was supplied.  In addition to
increases in  soil acidity,  leaching from plant
foilage may  contribute to reductions in plant
nutrients, decreases in growth, and changes in
foliar microflora.
  The effect of increasing acidic precipitation
in the northeastern United States  on vegeta-
tion,  streams,  and the soil  has  not  been
adequately studied.

H. MISCELLANEOUS   ASPECTS  OF
   SULFUR  DIOXIDE   EFFECTS   ON
   VEGETATION

   Although  the   aspects  discussed  in  this
section do not impinge directly on air quality
standards, they should be  considered in any
control activity. These include possible usage
of vegetation as a sink for SO2 and the effects
of sulfuric acid aerosols.

    1. Vegetation as a Sulfur Dioxide Sink

   The maintenance of protective vegetational
areas,   or  green  belts,  near   industrial
complexes has been a recent  topic  in urban
and regional planning.1 ° 8 >* ° 9  The concept of
SO2 removal by vegetation results  from as-
similation of SO2  by plant foliage and  the
deflection of polluted air masses above vegeta-
tional areas.108'110 Martin and Barber's110
studies demonstrated  a maximum SO2 reduc-
tion  of  157 jug/m3-hr (0.06  ppm/hr)  by
hawthorne hedge. Variations in uptake were
associated with  the physiological  activity of
the plants and the environmental  conditions
affecting  the plants.  In areas containing  a
large number of emission sources, Wentzel1 ° 9
indicated that vegetation belts  offered only
limited protection. Lampadius111 found only
slight  differences  in  SO2  concentrations
within forest stands,  forest edges, and  clear-
ings. He  concluded that removal of SO2  by
forest vegetation was  of minor importance.
Although  vegetation may reduce the level of
SO2  in some instances, there  is no evidence
that  it will have a major impact on ambient
SO2  concentrations.

       2.  Effects of Sulfuric Acid Mist
               on Vegetation

   Thomas,  Hendricks,  and Hill5 2  discussed
experiments in which plants were treated with
sulfuric acid  aerosols  at  concentrations of
78,600 to 170,300  Mg/m3  (30 to  65 ppm).
Sulfuric acid droplets settled' on dry leaves
without causing  injury, but  when  the  leaf
surface was  wet,  a spotted  type  of injury
developed. Middleton, Darley, and  Brewer51
and  Thomas, Hendricks, and Hill5 2  reported
that this  type of injury occurred in the Los
Angeles  area during periods  of heavy  air
pollution  accompanied  by fog  when  the
surface of the  leaf may be wet. Injury may
also occur in the absence of fog near combus-
tion effluents containing sulfur oxides when
the  dew  point of the gas effluent  results  in
acid droplet formation.
   The sequence of symptom  development is
one  in which the exposed surface, usually the
upper surface, shows the initial necrosis. The
pH of moisture on the leaf surface may be less
than 3.0. Cellular  collapse and many small
spots develop progressively through the upper
epidermis, mesophyll, and lower epidermis of
the leaf, leaving scorched areas. No glazing or
bleaching accompanies  this injury,  and  leaf
areas  covered  by  exposed leaves  show  no
marking.  In  the Los Angeles  area, injury of
Swiss  chard  and  beets was  more  nearly
typical of all plant species  examined. Alfalfa
also  developed  a  spotted  injury  pattern.
Spinach, being more uniformly wetted by fog,
developed a more diffuse type' of injury.

   1.  EFFECTS ON BIOMASS  AND YIELD

   Although evidence has been presented (Sec-
tion B.3.d)  that shows a growth reduction  in
the  absence of visible injury,  the early litera-
ture supports the view  that visible injury is
closely correlated with  yield   and/or growth
reductions.  Thomas3 and  Katz9 concluded
that growth effects do not occur until at least
10

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 5 percent of the  foliage is visibly injured.
 Yield reduction from acute SO2  injury was
 found to  be equivalent to the removal of the
 same amount of leaf tissue. Light to moderate
 defoliation of cotton from SO2  exposure had
 no detrimental  effect on  fiber grade, staple
 length,   or   ginning   percentage.112
 Guderian113 found that  the order of sensi-
 tivity, as determined by  leaf  necrosis  and
 yield, was often  different when  comparing
 several grass and  forage  species  and  native
 plants common  to open fields. For example,
 based on leaf necrosis, alfalfa ranked high
 with  regard  to sensitivity;  however, when
 yield was considered, this plant ranked in the
 resistant   category.   Guderian  and   Strat-
 mann114 J11 s  found that growth and yield of
 potato were progressively reduced with in-
 creased pollution intensity. In addition, seed
 tubers "obtained from heavily polluted areas
 gave significantly lower yields in the following
 year than tubers of the same weight obtained
 from  crops   grown  in  control   areas.
 Guderian113 reported  changes  in the com-
 position  of plant societies after exposure to
 SO2.
   Many  studies have shown that  the  reduc-
 tion in  crop yield from exposure to  S02  is
 proportional to  the percentage  of leaf area
 destroyed.8>6 9,70,72,116  "j^is relationship  is
 adequately expressed in the equation:
              y - a - bx
(5-1)
where:   y  = yield  expressed  as percentage
              of control
         a  - a constant, approximately 100
              percent
         b  = slope  of yield/leaf-destruction
              curve
         x  = percentage of  leaf area  de-
              stroyed
Hill and Thomas7 2 exposed  field alfalfa plots
to 2620 to  13,100 Mg/m3 (1 to 5 ppm) of
SO2   for  a  single   exposure  or  multiple
exposures of 1  to 2 hours during the growth
of the crop. The equations developed  show:

   1.  Single fumigation at early, medium, or
       late stage, representing either 25, 50,
       or 80 percent  of the growth period of
       crop:
                   >> = 99.5 -0.30.x
                   n =96
                    r = 0.64 ±0.06
                  Sy = 7.4 percent
                                     (5-2)
             2.  Double   fumigation  at  early  and
                medium, early and  late,  or  medium
                and  late stages in the growth  of the
                crop:
                   y = 95.5-0.49.x
                   n = 34
                    r = 0.79 ±0.07
                     = 8.2 percent
                                     (5-3)
    3.  Triple fumigation at early, medium,
       and  late stages in the growth of the
       crop:
          y =96.6-0.75*
          n =12                      (5-4)
           r = 0.98 ±0.014
         Sy- 4.1 percent

where:   n = number of plots fumigated
         r = correlation coefficient
       Sy = standard deviation  of individ-
              ual yields from the regression
              line

Similar results  were obtained by  clipping a
percentage   of   leaf  tissue  equal  to  that
damaged   by  SO2  from  a  group  of  test
plants.72  An equation similar to that  above
was developed for alfalfa using exposures of 1
to 600 hours and from 262 to  7860
(0.1 to 3  ppm) of SO,-28
                   y = 99 - 0.37*
                   n = 103
                    r = 0.48
                     = 8.8 percent
                                     (5-5)
           Results from barley,69 wheat,70 and  cot-
         ton1 16  studies differ from alfalfa in that the
         production of grain  and cotton is a measure
         of the yield and not of the vegetative growth.
         The stage of development of the plant when
         leaf destruction occurs is very important, with
         the most important stage  of  growth being
         near the time of blossom  and  fruit  develop-
         ment. Examples for barley,6 9 using Equation
         5-1, show:
                                                                                       11

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  1. Early stage, less than 25 cm in height:

          y = 98 - 0.06*
          n = 18                     (5-
          r=0.13
         Sy = l2.2 percent
   2.  Heading out stage:

          y = 98 - 0.40*
          « = 60
          A-= 0.74
         •S  = 10.2 percent
(5-7)
Data  from experiments  dealing with  other
crops were used in the same basic equation
(5-1). Controlled  additions of SO2  in these
field  experiments  have  shown  correlations
between visible leaf injury and the  ultimate
crop yield.
  The  most   comprehensive  growth-yield
experiments conducted in the vicinity of an
SO2  source were  carried out  in Biersdorf,
Germany.114*115,117,118 A wide variety of
plants, including  cereals, vegetables,  trees,
forage and fruit crops, were studied at five
locations at differing distances from a single
pollution source (Table 5-1).! 14 >'15 >l 18 The
tests  were run  over  the 7-month  growing
seasons of 1959 to 1960. Foliar symptoms
were observed at all locations, and growth and
yield reductions were determined by compar-
ison with a control site free of SO2. A review
of maximum  SO2 concentrations  and the
percent of time that measureable SO2  levels
existed suggests that many  short-term high-
concentration  episodes were  responsible for
the injury and growth reductions  that oc-
curred. The 30-minute  averages are indicative
of the highest values that might produce an
effect in the time period used.
  Growth  suppression  and injury  develop-
ment  of   white   pines  were  reported  by
Linzon90,119  based on data from field plots
that  were  located up to a distance of 25 miles
from   a  smelter  complex   near  Sudbury,
Ontario,  Canada.  Results  from this forest
ecosystem  study  are   shown  in   Table
5-2.''9 >' 20 Average SO2 concentrations over
a 10-year period for 6-month growing seasons
are  shown for each of  three  experimental
locations.   The   frequency   of  30-minute
average concentrations above three values are
shown  to  provide  an  indication  of  the
maximum concentrations that might produce
injury and growth effects. The net change in
tree volume is used as the measure of growth.
  Linzon90 noted that persistent high con-
centrations  of SO2  produced  certain  well
defined  growth effects on white pine.  How-
ever, there was a sharp improvement in the
growth  of  white  pine when  SO2   levels
dropped below 655 Mg/m3 (0.25 ppm).
  In a comprehensive study of the smelter at
Trail,  British  Columbia,  the  growth  of
Douglas fir, yellow pine, and lodgepole pine
was adversely affected for a distance of 12 to
18   miles   from  the  smelter.9  Daessler,
Kaestner,   and  Ranft121  have   presented
evidence   showing  growth  reductions  for
several conifer and  deciduous tree species
growing near a zinc smelter. Chlorotic dwarf
of white pine  has been studied in  the United
States,122  but few  investigators  have  con-
sidered growth effect on other tree species. In
Germany, the  growth of European  beech and
larch was reduced in  areas influenced by SO2
emissions.13
  Growth suppression in the absence of foliar
injury   (Section  B.3.d)  for   ambient  air
exposures42 and  for controlled greenhouse
         exposures
                  4 0 ,4 1
               are presented in Table 5-3.
         Reduced  growth of several plants exposed in
         field chambers  to  know^1  concentrations of
         SO2 over specific time pe/iods are also shown
         in Table 5-3.113
           The  results  from  tobacco at 262 Mg/m3
         (0.1 ppm)SO2 and radish at 131 Mg/m3 (0.05
         ppm)  SO2  showed  a  reduction  in  certain
         growth parameters for these two species when
         grown  under  conditions of maximum sensi-
         tivity to  SO2.  The conditions under which
         these results were  obtained  would probably
         never  be duplicated under  ambient condi-
         tions.   These  controlled  studies  were   well
         conceived and reflect the  best growth  data
         available from more recent studies.

            ]. DOSE-INJURY RELATIONSHIP OF
             SULFUR DIOXIDE TO VEGETATION
                         RESPONSE

           The  interrelations  of time and  concentra-
         tion (dose) as they affect injury to plants are
12

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Table 5-1.  SULFUR DIOXIDE CONCENTRATIONS AND ASSOCIATED
 VEGETATIONAL EFFECTS IN RESEARCH AREAS AT BIERSDORF,
                                  GERMANY
Average
concentration,3
Mg/m3 (ppm)
7959 7960
338(0.129) 388(0.145)
183(0.070) 272(0.104)
Maximum
concentration,''
Jjg/m3 (ppm)
7959 7960
14,148(5.4) 17,292(6.6)
9432(3.6) 17,030(6.5)
Species

Wheat, rye, oats
Rape, alfalfa,
red clover
Potato, beet,
spinach
Tomato, carrot
Wheat, rye, oats
Rape
Alfalfa, red clover
Effects0

1
1,2
1,2
1
1
2
1,2
Potato, beets,
  spinach
Carrot
Apple, sweet cherry,
  plum, current,
                                                                              1,2



123(0.047) 134(0.051)










45(0.017) 66(0.025)







37(0.014) 26(0.010)

gooseberry
Pedunculate oak, red
beech, larch
6288(2.4) 5764(2.2) Wheat, rye, oats
Rape
Potato, beet
Alfalfa, red clover
Spinach
Apple, current,
gooseberry
Sweet cherry, plum
Current
Pedunculate oak, red
beech, spruce, larch
3406(1.3) 4978(1.9) Winter wheat
Potato
Spinach
Apple
Current
Gooseberry
Pedunculate oak, red
beech, spruce, larch
2096 (0.8) 4454 ( 1 .7) Spinach
Gooseberry
1,2

3
1
2
1
1,2
2

1,2
3
1,3

3
1
1
2
2
1,3
1,2;

3
2
1,3
,3








,3









,3




"Average concentrations for 7-month growing season (4/1-10/31) determined by multiplying the percent of time
 that measurable concentrations were found by average concentrations during this tune period Values reflect re-
 sults from five stations radiating from a single source.
''Maximum concentrations bated on 30-mmute averages
'Plant responses based on 1959 and 1960 growing season
 I * Reduction in yield
 2 * Reduction in quality
 3 • Reduction in growth (shoot height, diameter of stem, and/or foliage dry weight).
                                                                                          13

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        Table 5-2.  CHANGES IN NET TREE VOLUME OF EASTERN WHITE PINE
                    ASSOCIATED WITH SULFUR DIOXIDE CONCENTRATIONS
                      NEAR A SMELTER COMPLEX IN SUDBURY, CANADA
   Average
concentration,3
 Mg/m3 (ppm)
                                Concentration
                            frequencies, percent'3
655Mg/m3    1310/xg/m3    2620
(0.25 ppm)    (0.50 ppm)    (1.0 ppm)
                                                              Net average annual gain or
                                                              loss in total tree volume0
118(0.045)

45(0.017)

21 (0.008)

5.92

0.98

0.33

2.36

0.11

0.01

0.38

0.01

0.00

Tree volume reduced 1 .3% over
10-yr period.
Tree volume reduced 0.6%
over 10-yr period.
Tree volume increased 1 .6%
over 10-yr period^
aAverage concentrations for 6-month growing season (5/1-10/31) over a 10-year period (1954-1963). Values are
 from three stations radiating from a group of three major S02 sources.

^Concentration frequencies based on the percentage of 1 /2-hour average concentrations above the respective SC>2
 values over the 10-year period.

cWhite pine sampling areas were located several miles from the air monitoring sites, but were within the same con-
 centration isopleths.

^Increases in tree volume were measured at white pine sampling areas located near the SC>2 monitoring station far-
 thest from the three sources.
 essential elements of air quality criteria. There
 are  insufficient  data  in  the  literature  to
 develop equations capable of defining effects
 of chronic injury, or the reduction of growth,
 yield, or quality of plant material. There have
 been  several attempts  to develop  rational
 models   to   express   time-concentration-
 response results of  plants to acute exposures
 from  SO2 •  Several empirical relationships
 have been proposed that give some insight as
 to what may happen  under a given set  of
 circumstances.

         1.  Mathematical Equations

   The  first  dose-response  relationship  for
 SO2   was  developed  by O'Gara123   under
 growth  conditions that  produced  maximum
 sensitivity  in the plant studied. The equation
 was developed from exposures of alfalfa over
 a relatively  short period of  time  with the
 production  of   acute  injury.  Thomas and
 Hill124  modified the  O'Gara  equation for
                                              alfalfa,  but the generalized equation can be
                                              shown as:
                                                            t(c - a) = ,
                                                                   (5-8)
                                              where:   t -
                                                       c =
                                                       a =
                                           time, hours
                                           concentration  of   pollutant
                                           when it is above the threshold,
                                           ppm
                                           threshold  concentration below
                                           which no  injury occurs regard-
                                           less  of length of exposures,
                                           ppm
                                      b -   constant

                             The parameters a and b are dependent on the
                             species and variety of plant and the degree of
                             injury. The equation can be rearranged to:
                                                            c =-r
                                                                   (5-9)
                                              The plot of c versus \jt is a straight line. The
                                              parameter  a is the intercept for \/t = 0,  or
 14

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Species
Table 53.  GROWTH REDUCTION IN VEGETATION EXPOSED TO SULFUR
                DIOXIDE FOR LONG AND SHORT TIME PERIODS

                     Concentration3  Exposure                           Refer-
                          3 (ppm)      time        Effect   Conditions    ence
Tobacco
   (Nicotiana tabacum L.
   "BelW3")

Radish
   (Raphanus sativus L.
   "Cherry Bell")

Ryegrass
   (Lolium perenne L.
   "Aberystwyth S23")
Timothy
  (Phleum pratense L.)
                       262    (0.1)   8hr/day,
                                     5 days/wk,
                                     (4 wks)

                       131    (0.05)  8hr/day,
                                     5 days/wk,
                                     (5 wks)

                     <262  «0.1)b  63 days
                             <262  (<0.1)c  77 days
                     2489    (0.95)   8 hr
Alsike clover
  (Trifolium hybridum L.)     2489     (0.95)   8 hr
Crimson clover
  (Trifolium incarnatum L.)    2489     (0.95)   8 hr
Red clover
  (Trifolium pratense L.)
                     2489     (0.95)  12hr
Italian rye
  (Lolium multiflorum Lmk.)   2489
                             (0.95)  12hr
Mixtures of:
  T. pratense
      and
  L. multiflorum
  Vetch (Vicia saliva L. and
  V.faba L.), pea (Pisum
  arvense L.) and lupine
  (Lupinus lentens L.)
                     2489    (0.95)  12hr
                      996    (0.38)  48 hr
Reduced  Greenhouse    41
growth    exposure
          chambers

Reduced  Greenhouse    40
growth    exposure
          chambers

Reduced  Ambient       42
growth    air
          greenhouse
Reduced                 42
growth

Reduced  Field         113
growth    exposure
          chambers

Reduced  Field         113
growth    exposure
          chambers

Reduced  Field         113
growth    exposure
          chambers

Reduced  Field         113
growth    exposure
          chambers

Reduced  Field         113
growth    exposure
          chambers

Reduced  Field         113
growth    exposure
Growth    chambers
not
affected
Reduced  Field         113
growth    exposure
for all     chambers
species
a Average concentrations over the reported time periods. Inaccuracies associated with instrumentation result in
 deviations as great as ± 10 percent.
b Except 2 days at concentrations of 262 to 524 M8/m^ (0-1 to 0.2 ppm).
c Except 3 days at concentrations of 262 to 524/ug/m3 (0.1 to 0.2 ppm).
                                                                                  15

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when t is infinitely  large.  Thus a  could  be
considered  the  threshold  concentration for
injury.
   The O'Gara equation (5-8) could also  be
written:
          t = b
                  \
                c - a
                       (5-10)
Zahn125  proposed  a  function  that  he sug-
gested  would  fit  experimental  data  over
longer time periods better than the O'Gara
equation. The equation expressed in the same
form as Equation 5-10 above is:
                Q.SC
             c (c - a}
                                     (5-11)
The threshold  value  a was given as 0.1 for
alfalfa; b was called  a dimensional resistance
factor  that  incorporates  the  influence  of
environmental   conditions.  Comparing  the
three  equations (5-8, 5-10,  5-11), the  time
required for threshold injury for alfalfa at an
SO2 concentration of 1048 Mg/m3  (0.4 ppm)
would  be 13,  6,  or  10 hours. At  higher
concentrations  of SO2, there are only minor
differences of time.
   Guderian,  van   Haut,  and  Stratmann73
recognized that the O'Gara equation did not
give the  best  fit  to  their observations for
either short- or long-term exposures. This led
to the development of an exponential equa-
tion of the form:
               = Ke
                    -b(c-a)
                       (5-12)
where:  t
        K
        e
        b
        c -
        a  =
time, hours
vegetation lifetime, hours
base of the natural logarithm
biological   complex   factor
(which includes the influences
of environmental factors)
concentration  of  pollutant
when it is above the threshold,
ppm
injury threshold, ppm
These parameters vary with species, environ-
mental conditions, and degree of injury.
   In the midtime ranges of 0.5 to 12 hours,
all  of  the equations  fit the available  data
reasonably well; however,  the  exponential
form  (Equation 5-12) fits over a wider range
of time.  These equations relate a given time
and concentration to a specific percentage of
injury.  They  have  been  developed  using
experimental data from  a limited number of
plant  species.
  These  two-dimensional models are limited
in their  application  since they  do  not  in-
corporate  the  relationships  of  the  many
factors that affect plant response to SO2. A
multivariate model is needed if these relation-
ships  are  to  be  considered.  Wolozin and
Landau126  proposed a nonlinear  function
incorporating all relevant factors that affect a
plant's response. They  suggest that in any
multivariate analysis the following factors be
considered: differing SO2 levels, duration and
frequency of  such  levels, relative humidity,
temperature,  diurnal  pattern  of SO2 con-
centrations,  species  of plant,  and  stage  of
plant growth.

            2. Dose-Injury Data

   Since  useful mathematical models are  not
available, an  extensive summary  of time-
concentration-response data  found  in  the
literature is necessary. A discussion of growth
effects was presented in Section I,  and  the
data were summarized in Tables 5-1, 5-2, and
5-3.   This  section   will be   limited  to  a
discussion of acute  effects that occur over a
relatively short time span,  results  of field
exposures where identifiable injury is present,
and results  of experiments utilizing mixtures
of pollutants. In most cases (except Table 5-7,
which shows the response of white pine and
radish to low mixture concentrations over a
period of several weeks), the responses noted
are the  result  of  acute exposures  to  the
toxicant in question.
   A study comparing spruce forests in a high
and   in   a  medium  pollution  area  was
conducted in Czechoslovakia.4 6 Results are in
terms of a  relative  determination  of foliar
injury to spruce. Four-month growing season
averages  and 30-minute maximum concentra-
tions  of SO2  are  reported.  Although  the
injury results, presented in Table 5-4, are not
easily quantified, the injury observed was of
the acute type. Materna, Jirgle, and Kucera46
 16

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          Table 54. SEASONAL AVERAGE SULFUR DIOXIDE CONCENTRATIONS
                          ASSOCIATED WITH FOLIAR INJURY TO SPRUCE
                             IN TWO LOCATIONS IN CZECHOSLOVAKIA
      Average
 concentrations,3 >
   Mg/m3  (ppm)
    Maximum
concentrations,'3 >c
      3  (ppm)
Foliar injury
    68 (0.026)
    943 (0.36)
   Severe
    47(0.018)
    812(0.31)
   Moderate
 a Average concentrations for 4 months of the growing season (6/1/66 - 9/30/66) determined from day and night
  monthly averages.

 bMonitoring instruments functioned with an error less than 10 percent only when concentrations were above
  150Mg/m3.
 c Maximum concentrations based on highest 30-minute average.
state   that   their  monitoring  instruments
functioned with an error less than 10 percent
only  when  concentrations  were above  150
Mg/rn3.

  Many  experiments have related  time  and
concentration to a response in plants sensitive
to SO2. It is reasonable that experiments and
field observations relating to short-time acute
threshold responses be reviewed  for inclusion
in a table of plant responses. Results from
three   investigations  are shown  in  Table
5.5 70,127,128 The resuits Of tne white pine
study127  are included because they  report
effects at much  lower  concentrations than
noted  before.  It should  be noted  that the
procedures  followed   in  making  the plant
grafts and the measurement techniques used
make the results in the reference of question-
able value.12 7

  Table   5-6  lists  agricultural  and  forest
species  growing  in  the  area of  Sudbury,
Ontario,  Canada.129  Shown are the minimal
average  concentrations for which injury  was
observed  after  exposures  of 1,  2,  4,  and  8
hours.  These field  observations relate to the
total pollution load over  the 8-hour averaging
period. The  average injury was 10 percent on
the leaves affected.12 9
         The  interaction of SO2  with other pol-
      lutants  was briefly discussed in Section D. 1 .f.
      Only interactions between SO2  and 03 and
      SO2  and  NO2   have  been  studied.  These
      results,  presented in Table 5-7,40'6 1"6S  are
      not  based  on  extensive  studies  but  are
      preliminary. They point out some conflicting
      reports, which need  in-depth  study. It is
      apparent  that, under  certain conditions and
      with  given levels of gases in the gas mixtures,
      some plants  can  be more severely affected
      than  by individual pollutants. However, there
      are   cases   where  plants  are  apparently
      protected  by  pollutant mixtures.  Neverthe-
      less, the potential for damage at low concen-
      trations of pollutant mixtures exists.
         Using the  threshold  concentrations from
      Tables  5-5,  5-6,  and  5-7 and  information
      involving   effects   that   relate   time  and
      concentration   over  short  time   periods,
      projected  SO2 concentrations causing  injury
      to  three susceptible groupings of plants were
      developed (Table 5-8). This table  was taken
      with  minor  changes   from  Heggestad and
      Heck.130  Table  5-9 gives a complete  list of
      plants  that have  been  studied in relation to
      SO2. The  plants in Table 5-9 are categorized
      using the sensitivity scale used in  Table 5-8.
      Within  each susceptibility grouping, the plants
      are listed alphabetically by family.
                                                                                       17

-------
        Table 5-5.  CONCENTRATIONS OF SULFUR DIOXIDE CAUSING  INJURY
                                         TO SENSITIVE VEGETATION3
Concentration'1
Species jug/m3 (ppm)
White pine
(Pinus strobus L.)









Alfalfa
(Medicago saliva L.)



Broccoli
(Brassica oleracea var.
botrytis L.)



Apple
(Malus sp. "Manks
Codlin")


Pear
Prunus sp,
"Legipont"
"Conference"

Mountain ash
(Sorbus aucuparia L.)




131

131

131

262

262


1310

1310


655

1310

1310

1258




1258

1336


1415

2175


(0.05)

(0.05)

(0.05)

(0.10)

(0.10)


(0.5)

<0.5)


(0.25)

(0.5)

(0.5)

(0.48)




(0.48)

(0.51)


(0.54)

(0.83)

Exposure
time, hr

1

2

3

1

2.5


4

4


4

4

4

6




6

6


3

3

Effect'

Needle injury
rating of 3
Needle injury
rating of 5
Needle injury
rating of 8
Needle injury
rating of 5
Needle injury
rating of 8

5% leaf
injury
19% leaf
injury

6% leaf
injury
4% leaf
injury
None

Leaf injury
rating of 6



Leaf injury
rating of 4
Leaf injury
rating of 5

Leaf injury
rating of 3
Leaf injury
rating of 7
Conditions

Branch
exposure
chamber
in greenhouse







Greenhouse
exposure
chambers


Same





Branch
exposure
chambers in
natural stands

Same




Same



Refer-
ence

127










70




70





128




128




128



        'The vegetation was observed or exposed when growing under environmental conditions that made it most sensi-
          tive to SO2.
        bAverage concentrations over the reported time periods. Inaccuracies associated with instrumentation result in de-
          viations a* great as ±10 percent.
        cThe effects are reported differently in each reference. Their definition is briefly described.
           I. Reference 127  The needle injury rating is based on a 1 Io8 scale with 7 as no injury and 8 as 2 to 3 cm of
             tip necrosis.
           2. Reference 70:  The values reflect the average percentage foliar injury on the three most severely injured leaves.
           3. Reference 128:  The leaf injury rating is based on a 0 to 10 scale with 0 as no injury and 10 as the entire leaf
             surface injured.
18

-------
     Table 5-6. SULFUR DIOXIDE CONCENTRATIONS CAUSING INJURY
                     TO AGRICULTURAL AND FOREST SPECIESa
   Species
Buckwheat
  (Fagopyrum sp.)
Barley
  (Hordeum vulgare L.)
Red clover
  (Trifolium pratense, L.)
Radish
  (Raphanus sativus, L.)
Oats
  (A vena sativa, L.)
Peas
  (Pisum sativum, L.)
Rhubarb
  (Rheum rhaponticum, L.)
Timothy
  (Phleum pratense, L.)
Swiss chard
  (Beta vulgaris var. cicla, L.)
Beans
  (Phaseolus sp.)
Beets
  (Beta vulgaris, L.)
Turnips
  (Brassica rapa, L.)
Carrots
  (Daucus carota, L.)
Cucumbers
  (Cucumis sativa, L.)
Lettuce
  (Lactuca sativa, L.)
                                           Maximum average concentrations^*
    1 hr,         2 hr,         4 hr,          8 hr,
     3  (ppm) jug/m3 (ppm)  ;ug/m3 (ppm) jug/m3  (ppm)
     Agricultural

1467   (0.56)  1022   (0.39)   681  (0.26)   393   (0.15)

1651   (0.63)  1153   (0.44)   629  (0.24)   314   (0.12)

1834   (0.70)  1205   (0.46)   707  (0.27)   367   (0.14)

1991   (0.76)  1415   (0.54)   760  (0.29)   367   (0.14)

1651   (0.63)  1546   (0.59)   891  (0.34)   445   (0.17)

1651   (0.63)  1546   (0.59)   891  (0.34)   445   (0.17)

1651   (0.63)  1546   (0.59)   891  (0.34)   445   (0.17)

1729   (0.66)  1415   (0.54)  1048  (0.40)   550   (0.21)

2306   (0.88)  1677   (0.64)  1074  (0.42)   707   (0.27)

1205   (0.46)  1179   (0.45)  1127  (0.43)   550   (0.21)

3432   (1.31)  2017   (0.77)  1179  (0.45)   603   (0.23)

3432   (1.31)  2017   (0.77)  1179  (0.45)   603   (0.23)

2830   (1.08)  2070   (0.79)  1310  (0.50)   655   (0.25)

2830   (1.08)  2070   (0.79)  1310  (0.50)   655   (0.25)

1677   (0.64)  1467   (0.56)  1126  (0.43)   996   (0.38)

1677   (0.64)  1467   (0.56)  1126  (0.43)   996   (0.38)
Tomatoes
  (Lycopersicon esculcntum, Mill.)
Potatoes                        1677   (0.64)  1467  (0.56)  1126  (0.43)  996   (0.38)
  (Solanum tuberosum, L.)
Raspberry                       1939   (0.74)  1651  (0.63)  1389  (0.53) 1022   (0.39)
  (Rubus idaeus,  L.)
Celery                          2279   (0.87)  1939  (0.74)  1441  (0.55)  760   (0.29)
  (Apium graveolens, L.)
Spinach                        3511   (1.34)  2384  (0.91)  1310  (0.50)  891   (0.34)
  (Spinacea oleracea, L.)
                                                                                  19

-------
           Table 56.  SULFUR DIOXIDE CONCENTRATIONS CAUSING INJURY
                     TO AGRICULTURAL AND FOREST SPECIES3 (Continued)
Maximum average concentrations15
Species
Ihr,
Mg/m3 (ppm) /ug
2 hr, 4 hr,
;/m3 (ppm) Mg/m3 (ppm) Mg/
8hr,
m3 (ppm)
Forest
 Cabbage                         2463   (0.94)  2332  (0.89) 1834  (0.70)  1179   (0.45)
   (Brassica oleracea, L.)
 Corn                             	-     	
   (Zea mays, L.)c
 Bracken fern                     1179   (0.45)   891  (0.34)   625  (0.25)   550   (0.21)
   (Pteridium aquilinum, L.)
 Large tooth aspen                1729   (0.66)  1126  (0.43)   969  (0.37)   524   (0.20)
   (Populus grandidentata Michx)
 Willow                          1074   (0.41)   996  (0.38)   865  (0.33)   786   (0.30)
   (Salix sp.)
 Trembling aspen                 1100   (0.42)  1022  (0.39)   681  (0.26)   341   (0.13)
   (Populus tremuloides Michx)
 Jack pine                        1362   (0.52)  1153  (0.44)   760  (0.29)   524   (0.20)
   (Pinus banksiana Lamb.)
 White pine                      1179   (0.45)   917  (0.35)   655  (0.25)   550   (0.21)
   (Pinus strobus L.)
 Alder                           1205   (0.46)  1126  (0.43) 1126  (0.43)   550   (0.21)
   (Alnus sp.)
 Red pine                        2043   (0.78)  1809  (0.69) 1153  (0.44)   786   (0.30)
   (Pinus resinosa Ait)
 Balsam poplar                   2149   (0.82)  1703  (0.65) 1179  (0.45)   681   (0.26)
   (Populus balsamifera L.)
 Austrian pine                    1729   (0.66)  1179  (0.45) 1153  (0.44)   865   (0.33)
   (Pinus nigra Arnold)
 Witch hazel                     2987   (1.14)  1965  (0.75) 1179  (0.45)   603   (0.23)
   (Hamametis virginiana, L.)
 Red oak                         2332   (0.89)  2149  (0.82) 1598  (0.61)  1074   (0.41)
   (Quercus sp.)
 Sugar maple                     2149   (0.82)  1703  (0.65) 1624  (0.62)  1205   (0.46)
   (Acer saccharum Marsh.)
 White spruce                    2279   (0.87)  2070  (0.79) 1834  (0.70)  1310   (0.50)
   (Picea glauca  (Moench) (Voss)
 Cedar                            --     .-     -	-
     f
   (Thuja occidentalis, L.)c


 aThe vegetation was observed when growing under environmental conditions that made it most sensitive to SO2.

 ''Average concentrations over the reported time periods. Inaccuracies associated with instrumentation result in
  deviations as great as ±10 percent.

 cNever injured near recorder stations.
20

-------
     Table 5-7. VEGETATIONAL RESPONSE TO SULFUR DIOXIDE IN COMBINATION
                        WITH EITHER OZONE OR NITROGEN DIOXIDE3
   Species
    Concentration
       ratio, b
    Mg/m3 (ppm)
             Exposure
                time
         Effect,0
         percent
          foliar
          injury
         Refer-
         ence
Tomato
  (Lycopersicon esculentum Mill.)
Alfalfa
  (Medicago sativa L.)
Broccoli
  (Brassica oleracea var. botrytis L.)
Cabbage
  (Brassica oleracea var. capitata L.)
Spinach
  (Spinacia oleracea L.)

Tobacco
  (Nicotiana tabacum L.)
  BelW3
  Bel W3
  BelB
  BelB
                                   Sulfur dioxide/ozone
 262/196
 655/490
1310/980
2620/1960
(0.1/0.1)
(0.25/0.1)
(0.5/0.1)
(1.0/0.1)
1310/98    (0.5/0.05)
 262/196   (0.1/0.1)
 655/196   (0.25/0.1)
1310/196   (0.5/0.1)
2620/196   (1.0/0.1)
4hr
4hr
4hr
4hr
               4hr
               4hr
               4hr
               4hr
               4hr
-25d
- 3
  4
-33d
          -17
           19d
           21d
           55d
1310/98
262/196
655/196
1310/196
(0.5/0.05)
(0.1/0.1)
(0.25/0.1)
(0.5/0.1)
4hr
4hr
4hr
4hr
17d
34d
11
14
1310/98
262/196
655/196
1310/196
2620/196
(0.5/0.05)
(0.1/0.1)
(0.25/0.1)
(0.5/0.1)
(1.0/0.1)
4hr
4hr
4hr
4hr
4hr
4
12
14
47d
-42d
          -48d
70
           70
                                                70
                                               70
           70
655/59
655/98
1310/98
262/196
685/196
1310/196
655/59
655/98
(0.25/0.03)
(0.25/0.05)
(0.5/0.05)
(0.1/0.1)
(0.25/0.1)
(0.5/0.1)
(0.25/0.03)
(0.25/0.05)
2hr
4hr
4hr
4hr
4hr
4hr
2hr
4hr
15d
16d
55d
8
75d
73d
9d
3d
67
70




67
70
                                                                                  21

-------
    Table 5-7. VEGETATIONAL RESPONSE TO SULFUR DIOXIDE IN COMBINATION
                 WITH EITHER OZONE OR NITROGEN DIOXIDE3 (Continued)
Species
Concentration
ratio, b
jug/m3 (ppm)
Exposure
time
Effect,c
percent
foliar
injury
Refer-
ence
Bromegrass
  (Bromus inermis L.)

Radish
  (Raphanus sativus L.)
White pine
  (Pinus strobus L.)
Radish
  (Raphanus sativus L.)
Bean
  (Phaseolus vulgaris L.)
Oats
  (A vena saliva L.)
2620/196   (1.0/0.1)
4hr
-6 Id
1310/98
262/196
655/196
1310/196
262/196
131/98
131/98
(0.5/0.05)
(0.1/0.1)
(0.25/0.1)
(0.5/0.1)
(0.1/0.1)
(0.05/0.05)
(0.05/0.05)
4hr
4hr
4hr
4hr
4 to 8 wk,
5 days/wk,
4 to 8 hr/day
10 to 30 days,
12 hr/day
5 wk,
5 days/wk,
8 hr/day
6
31d
22d
45d
9
Trace -ex ten-
sive necrosis
Reduced
growth
70
                                              70
Sulfur dioxide/nitrogen dioxide


 131/188   (0.05/0.1)       4hr        0
 262/188   (0.1/0.1)        4hr       11
 262/282   (0.1/0.15)       4 hr       24
 655/282   (0.25/0.15)      4 hr        4
 524/376   (0.2/0.2)        4 hr       16
655/94
131/188
262/188
262/282
655/282
524/376
(0.25/0.05)
(0.05/0.1)
(0.1/0.1)
(0.1/0.15)
(0.25/0.15)
(0.2/0.2)
4hr
4hr
4hr
4hr
4hr
4hr
3
0
27
12
0
10
                                              68
                                                                              69
                                              40
                    71
                                              71
22

-------
      Table 57.  VEGETATIONAL RESPONSE TO SULFUR DIOXIDE IN COMBINATION
                      WITH EITHER OZONE OR NITROGEN DIOXIDE3 (Continued)

Species


Concentration
ratio,b
/ig/m3 (ppm)

Exposure
time

Fffprt c
percent
foliar
injury

Refer-
ence

Radish
   (Raphanus sativtts L.)
262/188   (01/0.1)         4hr         27
262/282   (0.1/0.15)        4 hr         24
655/282   (0.25/0.15)      4 hr          4
131/470   (0.05/0.25)      4 hr         13
71
 Soybean
   (Glycine max L.)
Tomato
131/94
524/94
655/94
262/188
262/282
655/282
524/376
131/470
ntum Mill.) 655/94
131/188
262/188
262/282
655/282
131/470
(0.05/0.05)
(0.2/0.05)
(0.25/0.05)
(0.1/0.1)
(0.1/0.15)
(0.25/0.15)
(0.2/0.2)
(0.05/0.25)
(0.25/0.05)
(0.05/0.1)
(0.1/0.1)
(0.1/0.15)
(0.25/0.15)
(0.05/0.25)
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
4hr
2
6
7
35
20
1
9
2
1
0
1
17
0
0
71







71





aThe vegetation was grown under greenhouse conditions such that the plants were most sensitive to the pollutants
 and pollutant mixtures.

b Average concentration of pollutants. Inaccuracies associated with instrumentation result in deviations as great as
 ±10 percent.

c The effects are reported differently in each reference. Their definition is briefly described:
   1.  References 67, 68, and 70: Percentages are expressed as the difference between the percent of foliar injury
      from the SO2/03 mixture and the additive percent injury of the single gas exposures. Minus signs indicate
      that injury from the mixture was less than the additive injury from single gas treatments.
   2.  Reference 69:  Descriptive only.
   3.  Reference 40:  Growth reductions from the mixture were either less than additive or equal to the additive
      effects of single gas treatments.
   4.  Reference 71:  Foliar injury from the S02/NO2 mixtures occurred at pollution levels below the threshold
      injury concentration for SO2 (0.5 ppm) or NO2 (2.0 ppm) when used alone.

dPercentage differences are significant at the 0.05 level.
                                                                                               23

-------
             Table 58. PROJECTED SULFUR DIOXIDE CONCENTRATIONS
                         THAT WILL PRODUCE THRESHOLD INJURY TO
                         VEGETATION FOR SHORT TERM EXPOSURES3

                             Concentration producing injury in three
                                 susceptibility groups of plants
Time,
hours
0.5
1.0
2.0
4.0
8.0
Sensitive,
Mg/m3 (pPm)
2620
1310
655
262
131
to
to
to
to
to
10,480
7860
5240
2620
1310
(1.0
(0.5
(0.25
(0.1
(0.05
to 4.0)
to 3.0)
to 2.0)
to 1.0)
to 0.5)
Intermediate,
Mg/m3 (ppm)
9170 to
6550 to
3930 to
1310to
524 to
3 1 ,440
26,200
19,650
13,100
6550
(3.5 to
(2.5 to
(1.5 to
(0.5 to
(0.2 to
12)
10)
7.5)
5)
2.5)
Resistant,
Mg/m3 (ppm)
^26,200
^20,960
^15,720
^10,480
^ 5240
(^10)
(^ 8)
£ 6)
(^ 4)
£ 2)
a Values were developed from subjective evaluations of injury reported in the literature where both time and con-
 centration were considered. The concentrations and times shown for each susceptibility grouping are reasonable
 only when the plants are growing under the most sensitive environmental conditions and stage of plant maturity.
24

-------
           Table 59. LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                             BY SENSITIVITY TO SULFUR DIOXIDE3

Species                                                                       Reference

                                        Sensitive

Aceraceae
     Maple (Acer pseudoplatanus L.)

Amaranthaceae
     Pigweed (Amaranthus retroflexus L.)

Begoniaceae
     Begonia (Begonia sp.)

Bignoniaceae
     Catalpa (Catalpa sp.)

Carophyllaceae
     Bouncing bet (Saponaria officinalis L.)
     Sweet William (Dianthus baratus L.)

Chenopodiaceae
     Beet (Beta vulgaris L.)
     Lamb's quarters (Chenopodium album L.)
     Spinach (Spinacia oleracea L.)
     Swiss chard (Beta vulgaris var. cicla L.)

Convolvulaceae
     Sweet potato (Ipomea batata L.)

Compositae
     Aster (Aster sp.)
     Bachelor's buttons (Centaurea cyanus L.)
     Cocklebur (Xanthium sp.)
     Cosmos (Bidens sp.)
     Dandelion (Taraxacum officinale Weber)
     Endive (Cichorium endivia L.)
     Marigold (Tagetes sp.)
     Prickly lettuce (Lactuca scariola L.)
     Ragweed (Ambrosia sp.)
     Zinnia (Zinnia sp.)

Cruciferae
     Broccoli (Brassica oleracea var. botrytis L.)
     Brussels sprouts (Brassica oleracea var. gemmifera L.)
     Cabbage (Brassica oleracea var. capitata L.)
     Kale (Brassica oleracea var.acephala  DC.)
     Mustard, black (Brassica sp.)
     Mustard, hedge (Sisymbrium sp.)
     Nasturtium (Nasturtium  sp.)
128


131


 28


 28
 28
 28
 28
131
 28,50
 28,50
 28,47
 28
 28
 28,131
 28,132,50
 28,131
 50,47
 28
 28
 28
 28
 70
 28
 70
 28
 28
 28
 28
      25

-------
           Table 5-9. LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                       BY SENSITIVITY TO SULFUR DIOXIDE3 (Continued)
Species
   Reference
 Cruciferae (continued)
     Radish (Raphanus sativus L.)
     Turnip (Brassica rapa L.)

 Curcurbitaceae
     Cucumber (Cucumis sativus L.)
     Pumpkin (Cucurbita pepo L.)
     Squash (Cucurbita maxima Duchesne)

 Euphorbia ceae
     Rubber (Hevea brasiliensis Muell.)

 Fagaceae
     Beech (Fagus silvatica L.)

 Gramineae
     Barley (Hordeum vulgare L.)
     Bentgrass (Agrostis palustris Huds.)
     Bluegrass (Poa annua. L.)
     Bromegrass (Bromus sp.)
     Fescue, red (Festuca rubra L.)
     Junegrass (Poa pratensis L.)
     Oats (Avena sativa L.)
     Orchardgrass (Dactylis glomerata L.)
     Rye (Secale cereale L.)
     Ryegrass (Lolium sp.)
     Wheat (Triticum aestivum L.)

Iridaceae
     Gladiolus (Gladiolus Sp.)
     Iris (Iris sp.)

Labiatae
     Coleus (Coleus blumei Benth.)

 Leguminosae
     Alfalfa (Medicago sativa L.)
     Bean (Phaseolus  vulgaris L.)
     Bean, lima (Phaseolus lunatus L.)
     Pea (Pisum sativum L.)
     Sweet clover (Meliotus sp.)
     Sweet pea (Lathyrus odoratus L.)
     Soybean (Glycine max.  Merr.)
     Vetch (Vicia sp.)
    70,50
    28
   131
    28
    28
   133
   131
    28,47
   132
   132
    70
   132
    28
     9,50,47
    28
    28,50,47
    28
    28
   131
   131,28
   131
 70,89,50,47
 51,134
 70,28
 28
 28,47
 28,50
 70
131
26

-------
           Table 5-9.  LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                       BY SENSITIVITY TO SULFUR DIOXIDE3 (Continued)
Species
 Reference
Liliaceae
     Leek (Allium porrum L.)
     Onion (Allium cepa L.)

Malvaceae
     Cotton (Gossypium hirsutum L.)
     Hollyhock (Althaea sp.)
     Mallow (Malva sp.)

Nyctaginaceae
     Four o'clock ( Mirabilis jalapa L.)

Pinaceae
     Larch (Larix sp.)
     White pine (Pinus strobus L.)

Plantaginaceae*
     Plantain (Plantago sp.)

Polygonaceae
     Buckwheat (Fagopyrum sp.)

     Rhubarb (Rheum rhaponticum L.)
     Smartweed (Polygonum sp.)
     Sorrel (Rumex sp.)

Rosaceae
     Apple (Malus sp.)
     Apricot (Prunus sp.)
     European cherry (Prunus padus L.)
     Mountain ash (Sorbus aucuparia L.)
     Pacific ninebark (Physocarpus capitatus (Pursh) Ktze.)
     Peach (Prunus sp.)
     Pear (Pyrus sp.)
     Prune (Prunus sp.)
     Saskatton seniceberry (Amelianchier alnifolia Nutt.)

Saxifragaceae
     Gooseberry (Ribes sp.)
     Hydrangea (Hydrangea sp.)

Solanaceae
     Eggplant (Solanum melongena L.)
     Nightshade (Solanum sp.)
     Petunia (Petunia sp.)
     Tobacco (Nicotiana tabacum  L.)
 28
 70
 28,131
 28
 28
 28
 89,129
129
 28
129,131,
 50,47
 28
 28,47
 128,28,131
 131,128
 128
 87
 28
 128
 128
 131
 87
 28
 28

 28,47
 28
135
70,136
                                                                                  27

-------
           Table 5-9.  LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                       BY SENSITIVITY TO SULFUR DIOXIDE3 (Continued)
Species
Reference
Ulmaceae
     Chinese elm (Ulmus parvifolia Jacq.)
     Elm {Ulmus sp.)

Umbellifereae
     Carrot (Daucus carota L.)
     Celery (Apiwn graveolens L.)
     Parsley (Petroselinum crispum Nym.)
     Parsnip (Pastinaca sp.)

Vitaceae
     Wild grape (Vitis labrusca L.)
137
 28
 28
131
 28
 28
131
                                    Intermediate
Aceraceae
     Box elder (Acer negundo L.)
     Maple (Acer sp.)
     Norway maple (Acer platanoides L.)

Anacardiaceae
     Sumac (Rhus sp.)

Asclepiadaceae
     Milkweed (Asclepias sp.)

Begoniaceae
     Begonia (Begonia sp.)

Betulaceae
     California hazel (Corylus califomica (A.D.C.) Rose.)

Cannaceae
     Canna (Canna sp.)

Caprifoliaceae
     Columbia snowberry (Symphoricarpos rivularis Suks.)
     Honeysuckle (Lonicera sp.)
     Snowball (Viburnum sp.)

Compositae
     Chrysanthemum (Chrysanthemum sp.)

Cruciferae
     Horse-radish (Armoracia rusticana Gaertn., B Bay. and Scherb)
     Shepherd's purse (Capsella bursa-pastora L.)
 28
 28
137
 28


 28


 138


 87


 28
 87
 28
 28
 28
 28
 28
28

-------
           Table 5-9. LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                       BY SENSITIVITY TO SULFUR DIOXIDE3 (Continued)
Species
Reference
Cucurbitaceae
     Cucumber (Cucumis sativa L.)

Euphoribiaceae
     Castor bean (Ricinus communis L.)

Fagaceae
     Pin oak (Quercus palustris L.)

Ginkgoaceae
     Ginkgo (Ginkgo sp.)
 28
 47,28
137
137
Gramineae
     Kentucky bluegrass (foa pratensis L.)
     Salt grass (Spartina sp.)

Iridaceae
     Gladiolus (Gladiolus sp.)
137
 28


 28
Labiatae
     Salvia (Salvia sp.)

Leguminosae
     Wisteria (Wisteria sp.)

Liliaceae
     Onion (A Ilium cepa L.)

Malvaceae
     Hibiscus (Hibiscus sp.)

Oleaceae
     Lilac (Syringa vulgaris L.)

Pinaceae
     Douglas fir (Pseudotsuga taxifolia Brit.)
     Fir (Abies sp.)
     Pine, lodgepole (Pinus.contorta Dougl.)
     Pine, ponderosa (Pinus ponderosa Law)
     Pine, western white (Pinus monticola Dougl.)

Portulacaceae
     Purslane (Portulaca sp.)
 47


 28


 28


 28


 28
 87
  9
 87
  9
 87
 28
                                                                                   29

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           Table 5-9.  LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                      BY SENSITIVITY TO SULFUR DIOXIDE3 (Continued)
Species
Reference
Roseaceae
    Plum (Prunus sp.)
    Rose (Rosa sp.)
    Sweet cherry (Prunus avium L.)

Salicaceae
    Poplar (Populus sp.)

Saxifragaceae
    Mock-orange (Philadelphus sp.)
    Mock-orange, Lewis (Philadelphus lewisii Pursh.)

Scrophulariaceae
    Snapdragon (Antirrhinum sp.)

Solanoceae
    Potato, Irish (Solarium tuberosum L.)
    Tobacco (Nicotiana tabacum L.)
 28
 28
 28
 28
 28
 87
139
 28
140
Vitaceae
     Virginia creeper (Parthenocissus quinquefolia Planch.)
 28
                                       Resistant
 Caryophyllaceae
     Dianthus (Dianthus sp.)

 Cucurbitaceae
     Cantaloupe (Cucumis melo L.)

 Ericaceae
     Rhododendron (Rhododendron sp.)

 Fagaceae
     Oak (Quercus sp.)
     Oak, live (Quercus virginiana Mill.)
     Oak, pin (Quercus palustris L.)
     Oak, white (Quercus alba L.)

 Gramineae
     Corn (Zea  mays L.)
 139


  28


 139
 141
  28
 137
 141
 129
30

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            Table 5-9.  LISTS OF PLANTS IN THREE SUSCEPTIBILITY GROUPS
                        BY SENSITIVITY TO SULFUR DIOXIDE3 (Continued)

 Species                                                                        Reference
 Liliaceae
     Lily (Lilium speciosum Thunb.)                                              47

 Oleaceae
     Privet (Ligustrum sp.)                                                       28

 Orchidaceae
     Orchid (Cattleya sp.)                                                       50,139,47
            (Cymbidium sp.)                                                    50,141,47
            (Odontoglossum sp.)                                                50,141,47
            (Oncidium sp.)                                                      50,141,47

 Pinaceae
     Arb'or-vitae (Thuja sp.)                                                      28
     Cedar (Thuja occidentalis L.)                                                129

 Rubiaceae
     Gardenia (Gardenia sp.)                                                    139

 Rutaceae
     Citrus (Citrus sp.)                                                           28

 Umbellifereae
     Celery (Apium graveolens L.)                                                 28
aPlants were placed into the three susceptibility groups as defined in Table 5-7. The time-concentration data were
 obtained for each plant by Checking the appropriate reference.
                                                                                     31

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              K.  SUMMARY

  Plant species and  varieties  vary in sensi-
tivity  to  SO2. This  is  the  result of  the
interaction  of  environmental  and genetic
factors   that  influence  plant   response.
Temperature,  humidity, light,  other air pol-
lutants, edaphic conditions, the stage of plant
growth and the selective  pressures between
and within species all interact in affecting the
sensitivity  of plants  to  injury  from  sulfur
dioxide.   (Section  D.I,  2,  and  3.)  Since
ambient air is composed of many pollutants,
interaction  with  other  pollutants  must  be
considered in analyzing the effects of SO2  on
vegetation. In this regard, adverse foliar and
growth effects from pollutant  mixtures may
be of  a larger magnitude than effects from
exposures to SO2 alone. (Section  D.l.e.)
  The response of a  given variety or species
of plants to a specific air pollutant cannot be
predetermined  on the  basis of the  known
response of related plants to  the same pol-
lutant. Neither can the response  be predeter-
mined by a  given response  of  a  plant  to
similar  doses  of  different  pollutants. The
interplay   of  genetic   susceptibility  and
environmental influences  must be considered
for each plant and  pollutant. (Section D.)
  The  responses  of vegetation  to  sulfur
dioxide may  be classified into  two general
categories — visible effects and subtle effects.
Visible  effects   are   visually   identifiable
pigmented or  necrotic  foliar  patterns that
result  from major physiological  disturbances
to plant cells. Subtle effects  are those that
produce measurable growth  or physiological
changes in  plants but do not cause visible
injury. (Section B.)
   Visible  effects can be subdivided into acute
and  chronic injury. Acute injury  is severe
injury that  occurs within a few hours after
exposure to SO2  and is characterized by the
collapse of cells with the subsequent develop-
ment of  necrotic patterns.  It is associated
with high, short-term SO2 concentrations. In
broad-leaved  plants,  it  is  characterized  by
white  or  brown  interveinal  and  marginal
necrosis of  the leaf.  Red to brown colored
necrotic   lesions  occur  in  conifers.  This
necrotic response usually involves the needle
tip, but other portions of the needle may also
be affected. Acute injury patterns are general-
ly more characteristic of a specific pollutant
than chronic injury patterns. (Section B.I.a.)
  Chronic injury  results in light to severe
injury  that  develops  from  exposure over an
extended time period.  It is associated  with
long-term exposures where the pollutant con-
centration is  sufficiently  high  to produce
some   cell  destruction  or   disruption.
Symptoms are usually  in  the form of  leaf
chlorisis, but necrotic areas may also develop.
Foliar  injury  is followed by leaf abscission,
and  the  response  may  resemble  normal
senescence.  Acute and  chronic injuries may
develop on  the same leaves. (Section B.l.b.)
  Subtle  effects   implies   that   SO2   can
interfere  with  physiological   and/or   bio-
chemical  processes,  and  with plant growth
and yield without attendant development of
visible  symptoms. Processes that have  been
studied  include   photosynthesis,  stomatal
behavior, chemical composition,  and reduc-
tions in growth and yield. (Section B.2)
  The  term  physiological  effects includes
both subtle and visible effects. Physiological
changes in plants precede the  visible expres-
sions  of injury; however, visible  injury may
not  occur  at  all.  Changes  in  the  plant
processes, enzyme systems, and chemical com-
position  may  result  in growth and  yield
reductions in the absence  of  visible injury.
(Section B.3.)
  The mechanism by which plants are injured
by SO2 is not understood. Acute injury does
not occur if the rate  of SO2 absorption does
not  exceed  the  capacity  of  the  plant to
oxidize  sulfite to  sulfate  ions.  Under  long-
term  SO2 stress,  sulfate  thus formed may
accumulate, with the subsequent development
of chronic injury symptoms. (Section C.)
  In  assessing SO2  damage to  plants, the
most  significant question  is whether or not
the plant has been so altered by the pollutant
that its growth, survival, yield, or  use has been
impaired.
  Except in those instances where damage to
the plant  foliage  results in decrease in the
value  of the  product,  economic damage is
extremely difficult to assess. (Section E.)
  Growth and/or yield reductions may occur
without visible injury to  plants. (Section
B.S.d) Laboratory studies demonstrated  that
32

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reduced  root weights of  radishes occurred
with  exposure  to  SO2  at  concentrations of
131  to 160  Mg/m3  (0.05  to 0.06 ppm), 40
hr/week for 5 weeks. Reduction in the growth
of tobacco occurred with  exposure to SO2
concentrations  of  262 Mg/m3  (0.1 ppm), 40
hr/week  for  4  weeks. (Section B.S.d., Table
5-3) The conditions under  which the studies
were  conducted,  however, would  probably
seldom, if ever, be reached  in the ambient air.
More studies are  needed before a definitive
statement can be made.
  Most reports,  however,   have  considered
that  visible injury is required for reductions in
growth and yield.  Many studies have shown
that   the  reduction  in  crop  yield  from
exposure  to  SO2  is  proportional to  the
percentage   of   leaf  area   destroyed.  The
relationship between the percentage of  leaf
destroyed and  reduction in crop yield  has
been  expressed  in  the equation y = a -  bx,
where y  = the yield expressed as the per-
centage of the control and x - the percentage
of leaf area destroyed. The constant a is about
100   percent, and b  is the  slope of  the
yield/leaf-destruction curve. (Section I, Equa-
tion 5-1.)
  Foliar  injury of agronomic crops and trees
was reported  at SO2 concentrations of 1074
to 1650 Mg/m3  (0.41 to 0.63 ppm) for 1-hour
exposure periods when these exposure periods
were  within 8-hour time periods with average
concentrations of  314 to 786 Mg/m3 (0.12 to
0.30  ppm).  Concentrations of 314 Mg/m3
(0.12 ppm) injured barley after 8-hour periods.
(Tables 5-5  and   5-6.) In  Germany, growth
reductions  of  several  forage plants were
demonstrated after field exposures  of 2489
Mg/m3 (0.95  ppm) SO2  for 8 hours.  (Table
5-3.) For Italian rye, growth was also affected
at this concentration in exposure periods of
12 hours, but when this species was grown in
combination with red clover, growth was  not
affected  by  SO2   exposure. (Table  5-3.) In
contrast,   reduced  growth   of  rye  grass
occurred  when  average daily SO2 concentra-
tions were less than 262 Mg/m3  (0.1 ppm) for
about 96 percent of the experimental periods
and no greater than 524 Mg/m3  (0.2  ppm)
during the remaining periods of time. (Section
B.3.d, Table 5-3.)
  Growth, yield, and quality effects have also
been related to growing season average emis-
sions   from  single  sources.  In  Germany,
reductions  in these parameters were demon-
strated for spinach and gooseberry at growing
season averages of 26 to 37 Mg/m3 (0.010 to
0.014  ppm)  SO2.   A  larger  number  of
agronomic species were affected at averages of
45 to 66 Mg/m3 (0.017 to 0.025 ppm) SO2.
In this study, the effects were associated with
maximum 30-minute values of 2096 to 4978
Mg/m3  (0.8 to 1.9 ppm) SO2. (Table 5-1.)

  Laboratory and field chamber studies are
essential if qualitative and quantitative models
of pollutant effects upon vegetation are to be
developed.  Since it is impossible to include all
parameters,  laboratory  and  field  chamber
studies do  not  simulate ambient field condi-
tions. The following results were  obtained
through field chamber studies. Several forage
plants exhibited growth reductions after field
exposures of 2489 Mg/m3  (0.95 ppm) SO2 for
8 hours.  (Table 5-3.)  Injury  to the foliage of
varieties of apple and pear trees occurred after
6-hour exposures to  1258 Mg/m3 (0.48 ppm)
SO2 ; however,  foliar  injury  of mountain ash
occurred  after exposure to 1415 Mg/m3 (0.54
ppm) SO2 for 3 hours. (Table 5-5.)

  The interrelations  of  time and  the con-
centration of a pollutant are extremely impor-
tant in determining the amount of injury that
will be produced by a given pollutant. Several
attempts have been made  to develop rational
models  that   express  time-concentration-
response  results of plants to acute exposures
of SO2. (Section J.I.)

  Since  ambient air contains many  pol-
lutants,  interaction   with other  pollutants
must be considered in analyzing the effects of
SO2  on  vegetation.  In this regard, adverse
foliar  and  growth  effects from  pollutant
mixtures may  be of a larger magnitude than
effects from single SO2  exposures.  (Section
J.2.) Foliar injury of three  of six agronomic
crops  (alfalfa,   broccoli,  and  radish)  was
greater  after   4-hour  exposures  involving
SO2/O3 mixtures of 262/196 Mg/m3 (0.1/0.1
ppm) for each pollutant than for ozone alone.
No injury was observed after exposure to SO2
                                                                                     33

-------
alone. (Table 5-7.) In addition, growth reduc-
tions of radish, occurring after exposures to
SO2/O3 mixtures of 131/94/xg/m3 (0.05/0.05
ppm)  8 hr/day, 5  days/week for 5 weeks,
were greater than reductions from single SO2
exposures. (Section B.l.f.)
  Foliar injury to  four of five  agronomic
crops  (beans,  oats,  radish, and soybeans)
developed  after   4-hour  exposures  to
SO2/NO2   mixtures  of   262/188  Mg/m3
(0.1/0.1 ppm) of each pollutant.  The con-
centration  used was  below the injury thres-
hold for each of the gases. (Table 5-7.)
  Lichens  and  bryophytes are very sensitive
to the  presence of  SO2. Lichens have been
used  in the recognition and monitoring of
SO2. The presence of several fungal pathogens
has  been  reduced  in SO2  polluted  areas.
(Section F.)
  Another  effect   of  SO2  involves  the
acidification of precipitation. The  oxidation
and  solution  of SO2  in  precipitation has
increased the  acidity of soil  and water in
many  parts of the  world. This increase in
acidity may reduce populations of micro-
organisms  and  affect  the  process  of de-
composition   and  mineralization.   Acid
precipitation  may  also  contribute  to the
leaching of nutrients from  plant foliage and
from the soil.  (Section G.) Further studies of
the effects of acid rainfall in the United States
are  needed.
  Sulfuric acid  mists may  occur when heavy
air  pollution is accompanied by fog.  These
mists result in  necrotic spots, usually on the
upper  surface,  which  may  then develop
progressively through to the lower epidermis.
(Section H.2.)
  Reduced growth  of white pine occurred
with average SO2 concentrations of 45 Mg/m3
(0.017  ppm) associated with peak 30-minute
maximums of 3249 jug/m3  (1-24 ppm) during
growing seasons over a 10-year period. (Table
5-2.)
  Since  short-term   concentrations  are
probably  more  important  than  long-term
averages in the development of vegetational
injury,  growing season or annual averages as
well as the maximum concentrations must be
shown  if  they are  to  have  any value in
determining  causal  relationships.  In  this
regard, there is a need for the development of
mathematical  equations  to  express  relation-
ships  between   short-term  concentrations,
long-term averages, and vegetational  response
to sulfur dioxide. (Section J.I.)

            L. CONCLUSIONS

   The final chapter of Air Quality Criteria for
Sulfur Oxides  includes  summaries  of the
preceding chapters  of that document and
conclusions based upon them. The summary
of vegetation effects presented in that chapter
(Chapter  10,   Section  A.6)   no   longer
represents  the  best information  currently
available, and the reader is referred instead to
the  preceding section  of  this  report. The
conclusions   related to  vegetation   in  that
document (Chapter  10, Section  B.4) also are
superseded by those presented in this section,
as is the brief  statement in  the  "Resume"
(Chapter 10,  Section C).
   The conclusions  that  follow are derived
from a  careful  evaluation  by the  Environ-
mental Protection Agency of the foreign and
American studies cited herein. They represent
the Agency's best judgment of the effects that
may occur when various levels of pollution
are reached in the atmosphere. The data from
which the conclusions were derived, and the
qualifications that should  be considered in
using the data,  are identified by section or
table reference in each case.
   In applying the guidelines presented in the
following  paragraphs,  factors  other  than
pollutant concentration  that affect  a plant's
response to pollution, including the sensitivity
of the given variety or species to  the pol-
lutant,  duration of exposure,  temperature,
humidity, interaction  with  other  pollutants,
edaphic  conditions, and  state of plant de-
velopment, should be  kept in  mind.  Since
short-term  concentrations'are probably more
important  than  long-term  averages in the
development  of vegetational injury, maximum
concentrations as well  as growing season or
annual averages must be  specified in evalua-
tion of  long-term exposures.  In this regard,
there  is  a   need  for  the  development of
mathematical equations that express relation-
ships  between   short-term  concentrations,
long-term averages, and vegetation response to
sulfur dioxide.
34

-------
   For plants such as maple trees, spinach, and
sweet potatoes that are sensitive  to sulfur
dioxide, damage or reduction in growth or
yield may result from short-term exposures as
low as 131 to  1316 ME/m3 (0.05 to 0.5 ppm)
over periods of 8 hours or 2620 to 10,480
Aig/m3  (1.0 to 4.0 ppm)  over periods  of !/2
hour.  More resistant plants such as oak trees
and corn may  require exposures of over 5240
Mg/m3 (2 ppm) for the 8-hour period or over
26,000  Mg/m3 (10  ppm) for the  '/--hour
period. (Section J.2, Tables 5-8 and 5-9.)
   Growing season average concentrations as
low as 26 to 66 ng/m3 (0.010 to 0.025 ppm)
have been reported to affect  a large number
of agronomic  species. These  averages  were
associated with maximum 30-minute values of
2096  to  4978  /ag/m3  (0.8  to 1.9  ppm).
(Section 1, Table 5-1.)
   Foliar and  growth  affects  of mixtures of
SO2  with  other  pollutants  may  be greater
than the effects of SO2 alone. Mixtures of
both SO2 and ozone  and SO2 and nitrogen
dioxide  have been found to produce greater
effects than either pollutant alone.

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                                                                                    35

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106. Cohen,  J. B.  and A. G. Ruston.  Smoke:
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107. Materna, J. Zvysovani odolnosti Drevin
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108. Lanphear,  F.  O.   Urban  Vegetation:
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109. Wentzel,  K.   F.   Pestebni  zmirnujici
     prostredky proti imisim.  [Sylvicultural
     Measures for  Reducing  the  Effects  of
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     prumyslovych  exhalaci   na  lesni  hos-
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     vakia, October 11-14, 1966. Sponsored
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110. Martin,   A.  and F.  R.  Barber.  Some
     Measurements of Loss of Atmospheric
     Sulphur  Dioxide  near Foliage.  Atmos.
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111. Lampadius, F. Die Bedeutung der SO2-
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    Forstlichen Rauchschadentherapie. [The
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    Forests in Relation to the Control of Air
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    77(2):   503-511,   1968.   (Typescript
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112. Davis,  C.  R., G. W. Morgan, and D.  R.
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    Agron. J. 57: 250-251, 1965.
113. Guderian,  R.   Reaktionen  von Pflan-
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    auf  Schwefeldioxideirwirkungen.  [Re-
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    missions- und  Bodennutzungsschutz  des
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114. Guderian,  R.  and   H.   Stratmann.
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    [Field  Experiments  to Determine  the
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115. Guderian, R. and H. Stratmann. Freilan-
    dversuche zur  Ermittlung von Schwefel-
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    [Field   Experiments   for   Determining
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    tion.]   Forschungsberichte  des  Landes
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116. Brisley, H. R., C.  R. Davis, and J.  A.
    Booth.  Sulfur  Dioxide Fumigation  of
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    1959.
                                                                                    41

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117. Guderian,   R.   Zur  Methodik   der
    Ermittlung  von   SO2 -Toleranzgrenzen
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    script Trans.)
118. Stratmann,  H.  Freilandversuche   zur
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    the  Effects of SO2  on Vegetation.  II.
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    Ground Level  Concentrations.]
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    in-Westfalen. Essen,  W. Germany.  No.
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119. Linzon,  S.  N. Economic  Effects  of
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120. Dreisinger, B. R. Sulphur Dioxide Levels
    and the Effects of the Gas on Vegetation
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121. Daessler,  H. G., W.  Kaestner,  and  H.
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    akuten Einflussbereich einer Zinkhuette
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    77(2):   145-158,   1968.   (English
    Abstract.) Also in Forestry Abstracts.
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124. Thomas,  M.  D.  and  G.  R.  Hill, Jr.
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125. Zahn,  R.   Untersuchurigen ueber  die
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                                                                                    43

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 BIBLIOGRAPHIC DATA
 SHEET
1. Report No.
  EPA-R3-73-030
                                                                      3. Recipient's Accession No.
4. Title and Subtitle
  EFFECTS OF  SULFUR OXIDES  IN THE ATMOSPHERE ON  VEGETATION;
  Revised Chapter 5 for Air Quality  Criteria for Sulfur Oxides
                                                 5- Report Date
                                                 September 1973
                                                 6.
7. Author(s)
                                                 8- Performing Organization Kept.
                                                   No.
9. Performing Organization Name and Address
 National Environmental  Research Center
 Office of  Research and  Development
 U.S. Environmental Protection Agency
 Research Triangle Park, North Carolina 27711
                                                 10. Project/Task/Work Unit No.
                                                 11. Contract/Grant No.
 12. Sponsoring Organization Name and Address
                                                 13. Type of Report & Period
                                                    Covered
                                                                       14.
 15. Supplementary Notes
 16. Abstracts
 Limitations in  the criteria for secondary standards  in the publication  "Air Quality
 Criteria for  Sulfur Oxides," which became apparent  since the  adoption of  Air Quality
 Standards, prompted review and revision  of Chapter  5, "Effects  of Sulfur  Oxides in the
 Atmosphere on Vegetation."  This document presents  the revision to Chapter 5, and also
 includes revised portions  of Chapter  10,"Summary  and Conclusions," that relate to
 effects on vegetation.  The document,  based primarily on a thorough review of available
 literature, summarizes current scientific knowledge  of air pollution effects by sulfur
 oxides upon vegetation and also points up the major  deficiencies in that  knowledge.
 17. Key Words and Document Analysis.  17o. Descriptors
 Air  pollution
 Pollution
 Vegetation*
 Sulfur dioxide*
 Air  quality criteria*
 Agriculture*
17b. Identifiers/Open-Ended Terms

 *Air  pollution effects
17c. COSATI Field/Group
18. Availability Statement
                                     19. Security Class (This
                                        Report)
                                          UNCLASSIFIED
                                                          20. Security Class (This
                                                             Page
                                                                UNCLASSIFIED
21. No. of Pages

    54
                                                                                 22. Price
FORM NTIS-33 
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       (e.g.,  date of issue, date of approval, date of preparation.


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      from the performing organization.

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  14*  Sponsoring Agency  Code.   Leave  blank.

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      Translation of  ...   Presented at conference of  ... To bt published in ...  Supersedes  . . .       Supplements . . •

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      If thp report contains a significant  bibliography or literature survey, mention u here.

  17.  Key Words and Document  Analysis,  (a).  Descriptors.  Select from the Thesaurus of  Engineering and Scientific Terms the
      proper authon?,ed terms that identify the major concept  of the research and  are sufficiently  specific and piecise to be used
      as index entries for cataloging.
      (b).  Identifiers and Open-Ended Yermj.  Use identifiers for project names, code names, equipment designators,  etc.  Use
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FORM NTIS-35 (REV. 3-72)                                                                                   USCOMM-DC

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