NIMSS: View = Report

SAES-422 Multistate Research Activity Accomplishments Report=20 (termination)

Approved =

Project No. and Title:	NE10= 13=20 Mechanisms of Plant Responses to Ozone in the Northeastern US =
Period Covered:	10-2002 to 09-2007
Date of Report:	31-Mar-2008
Annual Meeting Dates:	08-May-2003 to 22-May-2007

Participants

Ainsworth, Lisa - USDA-ARS=20
Booker, Fitzgerald - USDA-ARS=20
Burkey, Kent - USDA-ARS=20
Bytnerowicz, Andrzej - US Forest Service=20
Carlson, John - Pennsylvania State University=20
Chappelka, Arthur - Auburn University=20
Chevone, Boris - Virginia Polytechnic Institute=20
Davison, Alan - Newcastle University=20
Decoteau, Dennis - Pennsylvania State University=20
Grantz, David - University of California at Riverside=20
Grulke, Nancy - US Forest Service=20
Karnosky, David - Michigan Technological University=20
King, John - North Carolina State University=20
Kohut, Robert - Cornell University=20
Krupa, Sagar - University of Minnesota=20
Lewis, Tim - US EPA=20
Long, Steve - University of Illinois=20
Manning, William - University of Massachusetts=20
Matyssek, Rainer - Technical University of Munich=20
McGrath, Margaret - Cornell University=20
Momen, Bahram - University of Maryland=20
Mulchi, Charles - University of Maryland=20
Muntifering, Russell - Auburn University=20
Neufeld, Howard - Appalachian State University=20
Percy, Kevin - Canadian Forest Service=20
Robinson, Michael - USDA-ARS=20
Sandermann, Heinrich - Ecotox=20
Skelly, John - Pennsylvania State University=20
Schaub, Marcus - Swiss Federal Research Institute WSL=20
Wiese, Cosima - College Misericordia=20
Zaleski, Rosemary - Exxon Mobil Biomedical Science=20
Zilinskas, Barbara - Rutgers University

Brief Summary of Minutes of Annual = Meeting

Annual=20 meeting dates and locations: May 8-9, 2003 (Raleigh, NC); May 20-22, = 2004=20 (Fresno, CA); May 19-20, 2005 (Asheville, NC); June 19-20, 2006 = (Champaign, IL);=20 May 21-22, 2007 (Rhinelander, WI).=20

CSREES Multistate Research Project Project No. NE-1013 Mechanisms of = Plant=20 Responses to Ozone in the Northeastern US=20

NE-1013 ANNUAL TECHNICAL COMMITTEE MEETING Raleigh, North Carolina = 8-9 May=20 2003 Minutes of the 2003 Annual Meeting=20

Meeting Attendees=20

Fitzgerald Booker, USDA-ARS, Raleigh, NC; Kent Burkey, USDA-ARS, = Raleigh, NC;=20 Art Chappelka, Auburn University, Auburn, AL; Boris Chevone, Virginia=20 Polytechnic Institute, Blacksburg, VA; Vahram Elagoz, University of=20 Massachusetts, Amherst, MA; Edwin Fiscus, USDA-ARS, Raleigh, North = Carolina;=20 David Grantz, Kearney Agricultural Center and UC-Riverside, Parlier, CA; = Sagar=20 Krupa, University of Minnesota, St. Paul, MN; Tim Lewis, US EPA, = Research=20 Triangle Park, NC; Bill Manning, University of Massachusetts, Amherst, = MA; Meg=20 McGrath, Cornell University, Riverhead, NY; Bahram Momen, University of=20 Maryland, College Park, MD; Charles Mulchi, University of Maryland, = College=20 Park, MD; Russell Muntifering, Auburn University, Auburn, AL; Howard = Neufeld,=20 Appalachian State University, Boone, NC; Mike Robinson, USDA-ARS, = Beltsville,=20 MD; Marcus Schaub, Swiss Federal Research Institute, Switzerland; Bob = Seem,=20 Cornell University, Ithaca, NY; John Skelly, The Pennsylvania State = University,=20 State College, PA;=20

The Technical Committee of NE-1013 was called to order by Chair F. = Booker at=20 8:30 AM on May 8, 2003 in Raleigh, NC.=20

Dr. Seem, Project Administrator, discussed final completion of the NE = 176=20 project. All members were requested to send a publication list from = annual=20 reports 1996 to 2002 to Dr. Seem who will use this information to = compile the=20 termination report.=20

Dr. Krupa raised concern about the lack of attendance of our CSREES=20 representative from Washington at the past two meetings. Dr. Seem stated = that=20 this was not uncommon due to funding issues and interest of the CSREES=20 representative. The possibility of recommending another individual from = CSREES=20 would be addressed by Dr. Seem. The comment was made that the current = annual=20 report should mention that about 50% of committee members in attendance = were=20 involved in writing or reviewing the new Ozone Criteria Document for = EPA. This=20 should be developed as an impact statement reflecting the outreach = component of=20 the Regional Project.=20

Station research reports were then presented by respective members = for the=20 remainder of May 8 and continued on May 9.=20

Dr. Burkey discussed the collaborative bean project utilizing = tolerant and=20 sensitive snapbeans. The experimental protocols for the bean project = were=20 presented to collaborating members and comments should be sent to Dr. = Burkey=20 prior to planting at the respective stations.=20

A discussion of the next meeting concerned issues of attendance of = the=20 European members. A suggestion was made to hold the Technical Committee = Meeting=20 in conjunction with the Air Pollution Workshop. No consensus was = reached. The=20 meeting site and date was to be decided by Dr. Grantz after = corresponding with=20 committee members. New European members to NE 1013 included M. Schwab, = J.=20 Jaeger. A. Davidson and H. Sandermann.=20

A motion was made and passed that two levels of membership to the = committee=20 be established: 1) full members and 2) corresponding members. = Corresponding=20 members would be those individuals interested in the research of the NE = 1013=20 Regional Project, but are unable to attend meetings on a regular basis.=20

Motion before the NE-1013 Technical Committee May 2003=20

Guidelines for NE-1013 membership and meeting attendance=20

In order to encourage attendance at NE-1013 Annual Meetings of the = Technical=20 Committee, and to provide opportunities for collaboration and project=20 coordination, and to provide a mechanism for maintaining current = membership=20 rolls, the following motion is proposed.=20

Attendance by members (or a representative of that member=19s = laboratory) of=20 one annual NE-1013 technical committee meeting within three consecutive = years is=20 encouraged. If attendance guidelines are not met, then the executive = committee=20 shall inquire into that member=19s commitment to the project either = directly or=20 through the administrative director. If a member cannot attend regular = meetings,=20 but would like to remain affiliated with the group, a member may be = classified=20 as an adjunct member who would be kept apprised of the group=19s = activities.=1D=20

This proposal is not meant to exclude or restrict membership to = NE-1013, nor=20 does it terminate membership if attendance guidelines are not met. It is = intended to foster participation in the group and to provide a mechanism = that=20 allows the Executive Committee to maintain a current membership roll.=20

Dr. Skelly requested that the committee write a letter to the PAES = director=20 stating the importance of maintaining an air pollution position at Penn = State=20 upon his retirement. The motion was made and passed that this be done = and that=20 Dr. Booker write the letter on behalf of the committee.=20

The meeting was adjourned at 12 N on May 9, 2003.=20

Respectfully submitted, Boris Chevone Secretary, NE 1013=20

CSREES Multistate Research Project Project No. NE-1013 Mechanisms of = Plant=20 Responses to Ozone in the Northeastern US NE-1013 ANNUAL TECHNICAL = COMMITTEE=20 MEETING Fresno, Parlier and Sequoia National Park, California 20-22 May = 2004=20

Minutes of the 2004 Annual Meeting=20

Meeting Attendees=20

Fitzgerald Booker, USDA-ARS, Raleigh, NC; Kent Burkey, USDA-ARS, = Raleigh, NC;=20 Andrzej Bytnerowicz, USDA Forest Service, Riverside, CA; Boris Chevone, = Virginia=20 Polytechnic Institute, Blacksburg, VA; Alan Davison, Newcastle = University,=20 Newcastle upon Tyne, U.K. Dennis Decoteau, Pennsylvania State = University, PA;=20 Annie Esperanza, US National Park Service, Sequoia National Park, CA; = David=20 Grantz, Kearney Agricultural Center and UC-Riverside, Parlier, CA; Nancy = Grulke=20 USDA Forest Service, Riverside, CA; Sagar Krupa, University of = Minnesota, St.=20 Paul, MN; Tim Lewis, US EPA, Research Triangle Park, NC; Bill Manning,=20 University of Massachusetts, Amherst, MA; Meg McGrath, Cornell = University,=20 Riverhead, NY; Bahram Momen, University of Maryland, College Park, MD; = Howard=20 Neufeld, Appalachian State University, Boone, NC; Heinrich Sandermann, = Institut=20 fuer Biochemische Pflanzenpathologie, Neuherberg, Germany; Anil = Shrestha,=20 Kearney Agricultural Center, Parlier, CA; Bob Seem, Cornell University, = Ithaca,=20 NY; Rosemary Zaleski, Exxon Mobil Biomedical Science, Annandale, NJ; = Barbara=20 Zilinskas, Rutgers University, New Brunswick, NJ;=20

SECRETARY'S REPORT=20

The NE-1013 Annual Technical Committee Meeting was called to order at = 8:15 AM=20 PST on May 20, 2004 by presiding chair F. Booker (USDA, NC). Welcoming = comments=20 were made by F. Booker and D. Grantz (Kearny Research Center, UC, CA), = the local=20 host, followed by introductions of the attending members. R. Seem, AES=20 administrative advisor, remarked that the new project had been approved = and is=20 now formally operational. Ray Knighton has been designated as the new = CSREES=20 advisor to the project in Washington, DC, but could not attend the = meeting due=20 to prior commitments.=20

Mr. Evan Shipp, meteorologist with the San Joaquin Valley, APCD, = presented a=20 talk on air quality in the valley. Violations of the 1 hr and 8 hr ozone = national air quality standard are common, with 160 ppb ozone typical for = 1 hr=20 high concentrations and 155 ppb for 8 hr concentrations. In the summer, = the 8 hr=20 standard is violated almost every day downwind from the = Fresno/Bakersfield metro=20 centers. Contributing to the high ozone concentrations are NOx, reactive = organic=20 compounds, high solar radiation, weak winds/strong inversions and = recirculation=20 of air masses. Following Mr. Shipps presentation, station reports = commenced.=20

At the conclusion of the station reports, the cooperative bean = project was=20 discussed. Stations involved in the project include NC, MD, NY, MA and = MN. S.=20 Krupa (MN) indicated that environmental and air quality data are = necessary for=20 modeling efforts. He would provide the other stations with a list of = minimum=20 data required. Another potential cooperative project was proposed = concerning the=20 role of ascorbate in ozone tolerance. This project is in the = developmental=20 stages at present.=20

Dennis Decoteau (PA), replacing J. Skelly, and Lew Ziska (USDA, MD),=20 replacing M. Robinson, were formally accepted as members. Steve Long = (IL) was=20 unanimously approved to join the project and a letter of invitation = would be=20 extended to him by F. Booker (USDA, NC). D. Decoteau was elected as = vice-chair=20 and B. Chevone (VA) agreed to remain as secretary for the next two = years.=20 Asheville, NC was approved as the site for the 2005 meeting and H. = Neufeld (NC)=20 and A. Chappelka (AL) would serve as local hosts. F. Booker (USDA, NC) = passed=20 the gavel to H. Neufeld (NC) as the incoming chair and H. Neufeld (NC) = adjourned=20 the meeting at noon on May 21, 2004. A field tour of the Kearny Research = Station=20 followed the formal meeting, with dinner at the Grantzs home. On May 22, = a tour=20 of ozone-impacted areas in the Sequoia National Park was led by Annie = Esperanza=20 (NPS) and Nancy Grulke (USDA/FS).=20

Respectfully submitted by, Boris Chevone Secretary, NE-1013 July 21, = 2004=20

REPORT OF THE SECRETARY NE-1013 Annual Technical Committee Meeting = May 19-20,=20 2005 Asheville, NC=20

Minutes of the 2005 Annual Meeting=20

Attendees of the NE1013 Technical Committee Meeting, 2005=20

Boris Chevone, VPI; Margaret Pippin, NASA Langely; Irene Ladd, NASA = Langely;=20 Jack Fishman, NASA Langely; Kirk Overmyer, NC, UNC-CH; Heinrich = Sandermann, GSF,=20 Germany; Bill Manning, MA; Kent Burkey, NC, USDA Raleigh; Margaret = McGrath, NY;=20 Don Davis, PA, PSU; Dennis Decoteau, PA; Fitz Booker, NC, USDA Raleigh; = Pat=20 Morgan, NC, USDA Raleigh; Stephanie Pilgrim, AL; Callie Nunley, AL; = Steve Long ,=20 IL; John Skelly, retired, PA; John Lin, AL; Russ Muntifering, AL; Alan = Davison,=20 UK, Newcastle, U.K.; David Grantz, CA; Ray Knighton, CSREES/USDA; Cosima = Wiese,=20 PA, College Misericordia;=20

The NE-1013 Annual Technical Committee Meeting was called to order at = 8:15 AM=20 EST on May 19, 2005 by presiding chair H. Neufeld (NC, Appalachian State = Univ.).=20 Welcoming comments were made by H. Neufeld, also the local host, = followed by=20 introductions of the attending members.=20

Bill Jackson, USDA-FS discussed ozone impacts in Class I wilderness = areas in=20 the southeastern U.S. High ozone concentrations occur in these areas but = are=20 dependent upon weather conditions. The concentrations are sufficient to = cause=20 foliar symptoms on milkweed and tulip poplar. Ten years of ozone data = across the=20 U.S. will soon be on the web from both high and low elevation sites. To = date,=20 the TREGRO model has shown that growth of red oak and red maple are not = affected=20 by current ambient ozone concentrations. The North Carolina Clean = Smokestack=20 Bill, reductions in emissions by TVA, the Clean Air Interstate Rule and=20 Knoxvilles attainment of the ozone NAAQS should contribute to a = reduction in=20 ambient ozone concentrations in nearby Class I areas.=20

Ms. Irene Ladd of the GLOBE outreach project, NASA Langley, then = presented=20 aspects of the program. One objective is to develop a common level of = knowledge=20 by the public of air pollution problems in the U.S. This education is = directed=20 toward developing an interest in young people to become air pollution=20 scientists. The program involves surface measurement of ozone using = ozone=20 sampling strips and planting ozone bio-indicator gardens.=20

Ms. Susan Sachs from the National Park Service discussed the ozone=20 bio-monitoring gardens in the Great Smoky Mts. There is more SOx and NOx = in the=20 Smokies than in any other National Parks. Three species have been = planted in the=20 indicator gardens and include crownbeard (Verbesina occidentalis), = cutleaf=20 coneflower (Rudbeckia laciniata) and tall milkweed (Asclepias exaltata). = Symptoms of stippling, chlorosis and necrosis are recorded weekly at = three=20 elevations and are animated over time to show the progression of foliar = injury=20 during the summer.=20

Following Ms. Sachs presentation, station reports commenced.=20

At the conclusion of the station reports, F. Booker (USDA, NC) = discussed the=20 NE1013 web page. The page has links to members of the project and to = other sites=20 showing ozone effects to vegetation. The URL is:=20 http://www.ncsu.edu/project/usda-ne-1013/.=20

Illinois was selected as the site of the next meeting and S. Long = would be=20 the local host.=20

The project renewal was then discussed and has to be submitted by = September=20 2006. The committee to develop the renewal consisted of D. Decoteau, = chair (PA),=20 S.Krupa (MN), B. Chevone (VA), A. Chappelka (AL) and D. Grantz (CA). A = draft of=20 the renewal is to be completed by the May Technical Committee Meeting, = 2006.=20

H. Neufeld formally closed the meeting at 12 noon, May 20, 2005.=20

Formally submitted by, Boris Chevone Secretary, NE-1013 July 28, 2005 =

REPORT OF THE SECRETARY NE-1013 Annual Technical Committee Meeting = June=20 19-20, 2006 Champaign, IL=20

Minutes of the 2006 Annual Meeting=20

Meeting Attendees=20

Ainsworth, Lisa - University of Illinois; Bernacchi, Carl - Illinois = State=20 Water Survey; Burkey, Kent - USDA ARS; Chappelka, Arthur - Auburn = University;=20 Chevone, Boris - Virginia Polytechnic Institute; Decoteau, Dennis - = Pennsylvania=20 State University; Knighton, Raymond - USDA CSREES; Krupa, Sagar - = University of=20 Minnesota; Leakey, Andrew - University of Illinois; Long, Steve - = University of=20 Illinois; Manning, William - University of Massachusetts; Morgan, = Patrick - USDA=20 ARS; Sandermann, Heinrich - Frieburg, Germany; Wittig, Victoria - = University of=20 Illinois; Zilinskas, Barbara - Rutgers University;=20

The NE-1013 Annual Technical Committee Meeting was called to order at = 8:15 AM=20 EST on June 19, 2006 by S. Long (IL) as presiding chair-elect D. = Decoteau (PA)=20 was delayed by inclement weather. Committee chair Howard Neufeld was = unable to=20 attend the meeting due to weather. Welcoming comments were made by S. = Long the=20 local host who mentioned the continuous crop plots started in the 1870s = at the=20 University of Illinois and the current SoyFACE project where a 20% yield = loss of=20 soybean has been observed under ambient ozone concentrations. = Introductions by=20 the attending members then followed. S. Krupa (MN) briefly discussed the = draft=20 renewal of NE1013 project. R. Knighton, the National Program Leader for = Air=20 Quality and the NE-1013 CSREES/USDA advisor, remarked that a major = program=20 concern was the contribution of agriculture to air quality. He further = commented=20 that the emission of ammonia, particulates and animal-produced reactive = VOCs in=20 relation to ozone production was an important consideration. Ammonium in = rainwater is increasing and the contribution from crop production is not = known.=20 Several committee members then raised a concern that the research focus = of=20 NE-1013 has been the effects of air quality (ozone) on crop production = and the=20 health of native vegetation. Most members expressed concern about = continuing in=20 the program if research emphasis shifted to a monitoring/modeling effort = from an=20 effects/mitigation effort.=20

Station reports were then presented. After the station reports, = discussion=20 ensued concerning the collaborative effort of the snapbean project. One = purpose=20 of the project was to determine the contribution of ambient ozone levels = to=20 yield loss of the sensitive cultivar compared to the tolerant one. This=20 information was deemed important to US EPA to consider when setting the = ozone=20 standard. A discussion then began on the renewal of NE-1013. Several = areas were=20 mentioned for inclusion in the renewal and the topics/coordinators were = natural=20 vegetation/A. Chappelka; water quality/air quality/B. Momen (MD); = mechanisms and=20 adaptation/S. Long and K. Burkey; education and outreach/D. Decoteau; = and=20 biomonitoring/W. Manning. R. Knighton suggested in the future that = station=20 reports, either oral or written, should be directed toward specific = objectives=20 of the existing proposal.=20

Potential new members included N. Grulke (USFS, CA) and D. Karnosky = (Michigan=20 Tech, MI). The meeting site for next year was selected as Rhinelander, = WI. D.=20 Decoteau adjourned the meeting at 12N on June 20, 2006.=20

Respectfully submitted, B. Chevone Secretary NE1013 Virginia Tech = Blacksburg,=20 VA=20

CSREES Multi-State Research Project NE-1013 Mechanisms of Plant = Response to=20 Ozone in the Northeastern US Holiday Inn Express, Rhinelander, WI May = 21-22,=20 2007=20

Minutes of the Meeting of the Technical Committee=20

Attendees: Lisa Ainsworth, USDA-ARS, Urbana, IL; Fitzgerald Booker, = USDA-ARS,=20 Raleigh, NC; Kent Burkey, USDA-ARS, Raleigh, NC; John Carlson, = Pennsylvania=20 State University; Dennis Decoteau, Pennsylvania State University; David = Grantz,=20 University of California, Riverside; Nancy Grulke, US Forest Service, = Riverside;=20 Dave Karnosky, Michigan Technological University; John King, NC State=20 University; Raymond Knighton, USDA-CSREES, Beltsville, MD; Mark Kubiske, = US=20 Forest Service, Rhinelander; Rainer Matyssek, Technical University of = Munich,=20 Germany; Margaret McGrath, Cornell University; Russ Muntifering, Auburn=20 University; Neil Nelson, US Forest Service, Rhinelander; Howard Neufeld, = Appalachian State University; Kevin Percy, Canadian Forest Service; = Heinrich=20 Sandermann, Ecotox, Germany; Cosima Wiese, College Misericordia; Barbara = Zilinskas, Rutgers University;=20

The meeting was called to order on May 21, 2007 at 9:00 a.m. by Fitz = Booker,=20 Chair of the Technical Committee, who introduced committee members and = presented=20 the history, objectives and collaborative projects of NE-1013. Ray = Knighton=20 informed the group that the renewal project NE-1030 had been approved = for a new=20 5-year period through September 30, 2012, and presented a number of = federal=20 budgetary items and funding opportunities that are pertinent to research = and=20 outreach activities of the Technical Committee. The committee was = invited to=20 submit information to Ray about ozone effects on specialty crops that he = could=20 take into consideration when formulating the RFPs for the new USDA = initiative on=20 specialty crops.=20

Station reports were then presented.=20

The following items were addressed during the business meeting:=20

1. NE-1030 was officially approved on 5/14/07.=20

2. David Grantz, Chair-Elect, will take over the position of Chair = after next=20 years 2008 meeting and will preside over the 2009 and 2010 meetings. A = new=20 Chair-Elect will need to be approved in 2008.=20

3. The US Forest Service has created the Paul Miller Clean Air Award, = a=20 national award for Forest Service employees to honor his memory. US = Forest=20 Service members on the Technical Committee were encouraged to consider=20 nominating deserving USFS individuals for this recognition. 4. NE-1013 = Annual=20 Reports for the period of June 2006 to May 2007 are due to F. Booker by = June 25,=20 2007. All committee members from land grant universities are required to = submit=20 annual reports, which basically contain the same information as annual = CRIS Form=20 AD-421, including publications. Fitz will summarize and submit along = with=20 meeting minutes. 5. An NE-1013 termination report is due for submission = to our=20 CSREES administrator no later than March 31, 2008. Station termination = reports=20 should be sent to Fitz by February 15, 2008. The termination report is = very=20 similar to the annual report except that accomplishments, impacts and=20 publications cover the entire span of the project. The following people = agreed=20 to help compose the termination report: Objective 1. (Describe the = spatial -=20 temporal variability of the adverse effects of ozone on crops and = forests - Meg=20 McGrath and Dennis Decoteau; Objective 2. (Assess the effects of ozone = on=20 structure, function and diversity of plant communities) - Russ = Muntifering and=20 Howie Neufeld; Objective 3. (Examine the joint effects of ozone with = other=20 growth regulating factors on crop and tree growth and productivity)- = Fitz Booker=20 and Dave Grantz; Objective 4. (Examine the molecular and physiological = basis of=20 ozone toxicity and tolerance in plants) - Barbara Zilinskas, Kent = Burkey;=20 Objective 5. (Develop numerical models to establish relationships = between=20 ambient O3 exposures and plant responses) - Sagar Krupa.=20

5. A report of the meeting is required by the EPA for funding that = has been=20 received. Howie Neufeld has $10,000 for travel. He needs official = receipts for=20 airplane, hotel and other expenses, except food receipts. He can handle = partial=20 reimbursement requests. 6. Kent Burkey provided an update on the snap = bean=20 project status. NJ, NC, PA, and NY will continue this research in 2007. = The=20 experiment may also be conducted in CA; however, the excessive heat = occurring=20 when ozone concentrations are highest may necessitate doing the work = during the=20 winter vegetable growing season when ozone levels are much lower. 7. Meg = McGrath=20 was elected next Secretary to replace Russ Muntifering, the current = Secretary.=20

8. Potential meeting locations were discussed. Next years meeting of = the new=20 NE-1030 project will be in the spring of 2008 at Auburn University, = hosted by=20 Art Chappelka and Russ Muntifering.=20

Dave Karnosky (MI), local host for the meeting and director of the = Aspen FACE=20 project, gave an overview of the Aspen FACE research, which is = attempting to see=20 how ozone alters the response of Northern forest ecosystems to elevated = carbon=20 dioxide. He discussed the variability displayed in growth responses to = these two=20 interacting gases, both interspecific and intraspecific. Later in the = program,=20 Dave led a field trip to the Aspen FACE project, which included a walk = through=20 of the four rings in the north replicate of the 12-ring experiment that = covers=20 some 20 ha on the USFS Harshaw Farm.=20

Respectfully submitted, R. Muntifering Secretary NE1013 Auburn = University=20 Auburn, AL=20

Accomplishments

The NE-1013 Project was = organized into=20 five distinct objectives. The Accomplishments of this successful = collaboration=20 are presented by Objective, below. The Impacts of these activities are = then=20 presented, integrated over all Objectives. At the conclusion of this = 5-year=20 project, the membership successfully prepared a renewal application, = approved as=20 NE-1030 for the 5-year period through September 30, 2012.=20

This project has been fundamental to increased understanding of ozone = effects=20 on plants. The collaborative effort has been the primary vehicle for = such=20 research in North America during this period. Meeting participants came = together=20 from 15 universities, two federal agencies and four foreign countries, = produced=20 120 peer-reviewed publications on ozone impacts, numerous book chapters, = and=20 four theses, and conducted a wide array of research and outreach = activities.=20 Several of the Technical Committee Members served as peer reviewers of = the US.=20 EPA 2006 Criteria Document for Ozone and Other Photochemical Oxidants. = Much of=20 the relevant new data regarding Welfare Effects and the associated = Secondary Air=20 Quality Standard for Ambient Ozone was developed by members of this = project.=20 Additionally, the project provided background briefing documents to the = USDA Air=20 Quality Task Force on Ozone Research and Vegetative Impacts and on Ozone = Effects=20 on Specialty Crops, to support preparation of the 2008 Farm Bill and to = use in=20 advising the Secretary of Agriculture regarding air quality policy. We = also=20 participated in reviews of the IPCC Fourth Assessment Report on Climate = Change.=20 The project has maintained a comprehensive web page since 2005, and = regularly=20 updates the site (http://www.ncsu.edu/project/usda-ne-1013/index.htm) = with=20 current information.=20

Objective 1. Describe the spatial - temporal variability of the = adverse=20 effects of ozone on crops and forests.=20

Adverse effects of ozone on crops and forests were investigated = through=20 experiments conducted with plants exposed to ambient ozone in various = locations=20 and time periods to achieve a diversity of ozone exposures. For example, = the=20 behavior of bean (Phaseolus vulgaris L.) selections R331 (tolerant to = ozone) and=20 S156 (susceptible to ozone) were compared in field plot evaluations over = several=20 planting dates, years and locations (states). Ambient ozone caused = severe injury=20 to leaves and defoliation in the ozone-sensitive snap bean cultivar, = S156,=20 across all of the tested locations and years. Total weight of bean pods=20 harvested for fresh-market consumption was 40 to 56% lower for S156 = compared=20 with the tolerant genotype when ozone concentrations were considered to = be=20 moderate to high. The R331 and S156 lines typically yielded similarly = under low=20 ozone concentrations, which usually occurred during the first or last = planting=20 dates at some of the sites. Overall, there was little spatial or = temporal=20 variability in the response of these bean lines to ambient ozone. Damage = occurred at ambient ozone levels in the sensitive line at a number of = locations=20 and over several years except when ozone concentrations were low.=20

In a similar type of study, biomass production of ozone-sensitive and = ozone-resistant clones of the commercial white clover line (Regal) were = compared=20 in New York. The sensitive clone was more severely injured and exhibited = up to=20 26% reduction in biomass production relative to the resistant clone when = ambient=20 ozone concentrations were high. The clones grew similarly when = concentrations=20 were low.=20

In contrast to the bean and clover studies, the amount of injury = observed on=20 grape foliage varied from year to year and was influenced by weather = conditions.=20 A study with Charbourcin grape in Pennsylvania found that ambient ozone = injury=20 included adaxial stipple and yellowing and defoliation of the older = leaves. The=20 drought of 2005 may have reduced foliar ozone injury compared with other = years=20 due to lowered ozone uptake. Also wet and cloudy conditions of 2003 and = 2004=20 contributed to reduced seasonal ambient ozone levels, which coincided = with less=20 injury. The Vidal variety of grape, which is considered tolerant to = ozone=20 injury, typically exhibited no foliar injury to ambient ozone levels.=20

Ambient ozone levels in the forests of western Virginia are = sufficient to=20 cause differential responses in ozone-sensitive and tolerant black = cherry trees.=20 Sensitive black cherry exhibited greater foliar injury symptoms and = lower=20 photosynthetic rates. These results demonstrate that even low ozone = levels (in=20 the 50 to 60 ppb range) can damage carbon assimilation processes in = sensitive=20 trees. Surprisingly, in Alabama, tree ring data from black cherry = indicated that=20 radial growth did not vary among ozone sensitivity groups during any = time period=20 analyzed.=20

In a study of native cutleaf coneflower (Rudbeckia laciniata) in the = Great=20 Smoky Mountain National Park, we found ozone injury to vary both = spatially and=20 temporally. These responses were not well correlated with ozone = concentrations.=20 In fact, injury was observed in relatively low-ozone years, indicating = that this=20 native plant can be quite sensitive to ozone, depending on other = environmental=20 conditions. Micro-site (seasonal rainfall or temperature patterns) and = genetic=20 factors influence plant sensitivity to ozone. In addition, it was found = that=20 sensitive coneflower developed injury earlier in the season than = insensitive=20 plants. Older leaf cohorts were more likely to exhibit the greatest = percent=20 injury by the end of the growing season. Also, leaf loss was more likely = for=20 older cohorts and lower leaf positions than in younger cohorts and upper = leaves,=20 respectively. Failure to take these factors into account can result in=20 underestimation of the effects of ozone on these plants.=20

Clearly, these results showed that medium to high ambient ozone=20 concentrations can damage vegetation and reduce yields, but that genetic = and=20 environmental factors strongly modulate the responses.=20

Objective 2. Assess the effects of ozone on structure, function and = diversity=20 of plant communities.=20

The overall goal of Objective 2 was to assess the effects of ozone on = structure, function and diversity of plant communities, notably forests = and=20 grasslands. Results have provided critically needed data on = physiological and=20 growth responses of mature trees and understory herbaceous species in = the field=20 under a range of ambient ozone exposure regimes. We determined changes = in tree=20 growth under ambient ozone conditions and elucidated underlying = mechanisms=20 responsible for differences in sensitivity of native wildflower species. = Also,=20 we characterized alterations in cell-wall constituents and secondary = metabolites=20 in ozone-exposed herbaceous vegetation that have implications to the = nutritional=20 ecology of economically important ruminant animals. Information needs in = these=20 areas have been identified as critical for the assessment of ozone = pollution=20 effects in natural ecosystems.=20

One of the fundamental differences identified by testing = responsiveness of=20 understory herbaceous species to rapid changes in light is that = ozone-sensitive=20 plants, such as cutleaf coneflower, have impaired stomatal functioning, = such as=20 non-responsiveness to changes in humidity or light compared to = ozone-insensitive=20 plants. It was found that ozone-sensitive plants fail to close their = stomata=20 under conditions where insensitive plants would, leading to reductions = in water=20 use efficiency and continued uptake of ozone by the plant, which = contributes to=20 ozone injury.=20

Ecosystem function can be altered by several processes, one of which = is=20 differential reproductive success. We found that reproductive effort of = selected=20 native plant species was affected by ozone. These effects can be = translated into=20 alterations in flowering patterns and abortion of seeds/fruits, and have = implications regarding establishment, survival, genetic stability and = vigor of=20 these species. Ozone reduces photosynthesis in native plants, especially = in=20 their older leaves, which translates into reduced starch reserves in = these=20 leaves and the underground rhizomes. Plants known to be sensitive suffer = greater=20 reductions in photosynthesis than those that are insensitive. = Sensitivity in one=20 herbaceous species was linked to extracellular levels of the antioxidant = ascorbic acid, suggesting that both uptake and biochemical = detoxification are=20 important mechanisms of response in these native plants.=20

Research under this objective has shown that, as a result of = accumulation of=20 secondary phenolic compounds, increased deposition and lignification of=20 cell-wall constituents and decreased in vitro digestibility in a number = of=20 ozone-sensitive plant species, predicted loss of forage nutritive = quality for=20 ruminant animals due to ozone injury can readily approach the same order = of=20 magnitude as that observed for biomass yield depression (ca. 5-15%). = These are=20 extremely important findings because total loss of consumable food value = (fractional reduction in yield =C3=97 fractional reduction in nutritive = quality for=20 ruminants) can be much more significant than biomass yield reductions = alone in=20 the assessment of the true economic impact of ozone on forages under = current and=20 future global-climate scenarios. Using ethylene diurea (EDU, an = antioxidant=20 protectant for plants) it was found that in purple coneflower (Echinacea = purpurea), EDU could ameliorate the deleterious effect of ozone on = nutritive=20 quality. Further testing is needed in this area to determine if this is = response=20 is evident among other plant species. Until recently, economic = assessment models=20 have included only the effects of yield depression.=20

Results of research under this objective also illustrate the value of = both=20 'real-world' and manipulative experiments, and the importance in=20 climate-change/nutritional ecology research of assessing effects of = co-exposure=20 to environmentally relevant levels of multiple air pollutants. For = example,=20 ground-level ozone, temperature and precipitation were observed to be = the most=20 important determinants of nutritive quality in a 5-year study with = alfalfa grown=20 under ambient concentrations of multiple air pollutants (ozone, nitrogen = oxides,=20 sulfur dioxide) and prevailing meteorological conditions, but their = relative=20 importance was largely dependent on yield such that ozone exposure was = the most=20 important determinant of quality in high-yielding but not in = low-yielding=20 alfalfa. We have observed in both intensively managed and semi-natural = systems=20 that adverse effects of ozone on forage quality may be amplified by high = soil=20 fertility or exposure to growth-stimulating levels of atmospheric N = deposition=20 compared with low soil fertility or growth-limiting levels of N = deposition,=20 respectively. We have discovered in our work with red (Trifolium = pratense) and=20 white (Trifolium repens) clover that, in contrast to recent reports of a = protective effect of elevated atmospheric carbon dioxide against yield = reduction=20 in plants under ozone stress, future increases in atmospheric carbon = dioxide=20 concentration might not be expected to ameliorate the negative impact of = elevated ozone on nutritive quality under global chemical-climate = scenarios=20 projected for the Northern Hemisphere through at least the first half of = the=20 21st Century.=20

Objective 3. Examine the joint effects of ozone with other growth = regulating=20 factors on crop and tree growth and productivity=20

Nutrition=20

The atmosphere has become a source of bioavailable N as well as = oxidizing=20 species such as ozone. Observations in a long-term experiment in a = sub-alpine=20 pasture in Switzerland examined the impact on pasture nutritive quality = of the=20 combined impacts of ozone and high N input. A pasture exposed to ozone = and N=20 deposition in Switzerland showed that nutritive quality was 7% lower for = elevated ozone treatments due to altered cell-wall chemical composition. = There=20 were long term changes in vegetative composition as well, with forbs = increasing=20 from 23 to 36%, and grasses and legumes both decreasing (from 68 to 60%, = and=20 from 9 to 3% of biomass, respectively).=20

Weeds and Invasive Vegetation=20

Competition relationships between crops and weeds can be altered by=20 concurrent exposure to ozone. However, knowledge of the ozone tolerance = of=20 individual species has not proven useful in predicting the outcome of=20 inter-specific plant competition in open top chambers studies in the San = Joaquin=20 Valley of California. For example, competition between Pima cotton and = the=20 globally significant C4 weed, yellow nutsedge (Cyperus esculentus L.) = was=20 affected substantially by ozone exposure. Cotton was generally more = sensitive to=20 ozone than nutsedge, and both competition from nutsedge and exposure to = ozone=20 reduced productivity of cotton. The two species inhibited the growth of = each=20 other to similar extents. At a high ozone concentration productivity of = cotton=20 was low, but the relative reduction by nutsedge competition at high = ozone was=20 similar to that at low ozone concentration. Growth of nutsedge in = competition=20 with cotton was greatest at high ozone, as the vigor of cotton declined. =

In contrast, similar competition studies with tomato, which is = somewhat more=20 ozone tolerant than cotton, led to differing results. Tomato became more = competitive at high ozone. Tomato growth was inhibited at the highest = ozone=20 concentration at all levels of nutsedge competition, while nutsedge was = less=20 affected. Nutsedge reduced tomato productivity under low and moderate = ozone=20 concentrations, but tomato was more competitive at high ozone = concentrations.=20 Nutsedge allocated greater resources to reproductive tubers at the = highest ozone=20 exposure which could make it even more invasive in future environments.=20

Horseweed is an economically important C4 weed that is becoming = increasingly=20 invasive in areas like the San Joaquin Valley. Glyphosate-resistant = horseweed=20 has almost completely replaced the wild-type, glyphosate-susceptible, = biotype.=20 Results of research conducted by this project suggest that ozone may be = a=20 significant contributor to this rapidly changing population structure. = Responses=20 to ozone are similar in glyphosate-sensitive and -resistant lines. = However,=20 additive effect of ozone and glyphosate is sufficient to drive the=20 glyphosate-sensitive population to extinction. The resistant biotype was = less=20 likely than the susceptible biotype to being driven out of the = population by the=20 combination of high ozone and glyphosate. Thus ozone may contribute to = the rapid=20 evolution of herbicide resistance and to rising crop production costs = due to=20 increased need for vegetation management, in addition to the well known = effect=20 of reducing yield directly. This is the first indication that = tropospheric ozone=20 may be a contributory factor in development of a serious agricultural = pest=20 through altered population structure.=20

Elevated Carbon Dioxide=20

Ozone is a component of the changing atmosphere, and will play a part = in=20 ongoing Global Change. A soybean-corn rotation open-air exposure = experiment=20 showed that for soybean, elevated carbon dioxide increased biomass and = seed=20 yield, elevated ozone decreased biomass and yield, and the deleterious = effects=20 of ozone were partially ameliorated by carbon dioxide when the two gas=20 treatments were combined. This and other studies have indicated that = protection=20 against ozone injury in many crops by elevated carbon dioxide can be = attributed=20 to reduced ozone uptake and possibly other factors, but there has been = little=20 direct testing of these hypotheses. Manipulation of ozone concentrations = and=20 estimates of plant ozone uptake indicated that equivalent ozone fluxes = that=20 suppressed net photosynthesis, growth, and yield at ambient = concentrations of=20 carbon dioxide were generally much less detrimental to plants treated=20 concurrently with elevated carbon dioxide. These responses appeared = unrelated to=20 effects on antioxidant metabolism. Plants treated with elevated carbon = dioxide=20 had higher rates of net photosynthesis due to higher intercellular = carbon=20 dioxide concentrations. Increased photoassimilation and decreased=20 photorespiration with elevated carbon dioxide would promote growth and = help=20 counter detrimental effects of ozone. Increasing concentrations of = atmospheric=20 carbon dioxide will likely ameliorate ozone damage to many crops due to = reduced=20 ozone uptake and increased carbon assimilation. Our study further = suggests that=20 elevated carbon dioxide may increase the threshold ozone flux for = biomass and=20 yield loss in soybean.=20

An experiment designed to test the effects of elevated carbon dioxide = and=20 ozone on soil carbon and nitrogen dynamics in a soybean-wheat no-till = system=20 using open-top chambers showed that elevated carbon dioxide increased = soybean=20 and wheat biomass production by 10 to 25% while ozone suppressed it by = 11 to=20 27%. In combination, elevated carbon dioxide ameliorated ozone effects = on=20 biomass. Treatment effects on biomass production dominated potential = impacts on=20 soil carbon dynamics as evidenced by litter levels in the treatment = plots and=20 minirhizotron images of root production.=20

Two winter-wheat cultivars, Gore and Susquehanna, were treated with = elevated=20 carbon dioxide and ozone individually and in combination. Elevated = carbon=20 dioxide resulted in stomatal closure and, under low ozone, induced = antioxidant=20 changes that could enhance defensive capacity for oxidative stress. = Yield under=20 the combination of elevated ozone and carbon dioxide tended to be = greater than=20 for elevated ozone alone. The results suggest that plant response to = ozone=20 depends upon a number of factors working together. The availability of = high=20 pools of antioxidants in wheat may further contribute to enhanced = oxidative=20 defense capabilities.=20

The AspenFACE experiment indicated that northern forests have the = capacity=20 for sustained growth stimulation due to elevated carbon dioxide, and = that=20 concurrent exposure to moderate levels of tropospheric ozone partially = or=20 totally compromises growth stimulation from elevated carbon dioxide. = Also,=20 elevated carbon dioxide and ozone have small effects on litter chemistry = and=20 specific rates of decomposition, while changes in litter inputs under = elevated=20 carbon dioxide and ozone will likely have large effects on soil organic = matter.=20

Objective 4. Examine the molecular and physiological basis of ozone = toxicity=20 and tolerance in plants.=20

Systemic effects of ozone=20

Ozone immediately affects leaf tissue; however, ozone can cause = systemic=20 effects on the whole plant. In snap beans, ozone treatment increased = shoot=20 respiration based on elevated metabolic heat rates measured with a=20 microcalorimeter. In both Pima cotton and muskmelon, ozone-induced = inhibition of=20 carbon assimilation and transport of carbohydrate from shoots to roots = did not=20 cause the expected reduction of root respiratory activity. As source = strength=20 and transport of carbohydrate to sink tissues declined, root respiration = per=20 unit fresh weight increased. The mechanism of these seemingly disparate=20 phenomena remains unknown. However, these findings suggest that the = hypothesis=20 that substrate control of root respiration, as it is modulated by ozone = impact=20 on shoot tissues, may not be supported.=20

In Pima cotton, exposure of the shoots to ozone led to genetic damage = in root=20 tips as visualized by an alkaline, single-cell electrophoretic assay of = damaged=20 DNA in isolated root tip cells. DNA damage increased with increasing = ozone=20 exposure. The results clearly indicate that the effects of ozone on = vegetation=20 are systemic, and suggest that translocated products of ozonation, or = other=20 signal transduction processes, are involved in reducing root = proliferation=20 following shoot exposure to ozone. Root growth is often inhibited by = ozone and=20 has been primarily attributed to reduced availability of carbon = resources needed=20 for growth. The finding that ozone induces genetic damage in root tips = adds a=20 new dimension to our understanding of the mechanisms of ozone toxicity.=20

Methyl jasmonate caused a suite of developmental changes reminiscent = of ozone=20 exposure. These preliminary experiments were in response to recent gene=20 expression studies that link jasmonate metabolism to ozone impacts. In = our=20 experiments, application of methyl jasmonate induced responses very = similar to=20 those of ozone. These similarities were sufficient to suggest that = ozone-induced=20 developmental or injury pathways were modulated by methyl jasmonate. = However,=20 application of methyl jasmonate in our experiments did not alter plant = responses=20 to ozone, indicating that ozone effects are propagated through a network = of=20 metabolic pathways and cellular processes that are only partly modulated = by=20 methly jasmonate. These and similar studies are important because they = help us=20 disect the biochemical and physiological mechanisms of ozone toxicity = which may=20 contribute to the engineering of ozone-tolerant crops.=20

Stomatal conductance in relation to ozone tolerance=20

Stomatal conductance was similar for ozone-sensitive (S156) and = tolerant=20 (R123) snap beans. While genetic variation was found for stomatal = density and=20 aperture on upper and lower leaf surfaces, these differences in stomatal = characteristics did not translate into differences in conductance rates, = suggesting that the observed genetic variation in ozone response is not = related=20 to differences in ozone uptake.=20

Ascorbic acid metabolism=20

Ascorbic acid (known also as vitamin C) is generally acknowledged to = play a=20 key role in plant response to oxidative stressors, including ozone. In=20 Arabidopsis, a putative F-Box gene, VCF1 (Vitamin C F-box 1), was = identified=20 that appears to negatively regulate gene expression in the = mannose-galactose=20 pathway of ascorbate biosynthesis. Plants in which VCF1 was inactivated = had=20 increased leaf ascorbate content and enhanced tolerance to ozone.=20

Photosynthesis in mature leaves of soybean was more sensitive to = ozone=20 exposure in the cultivar Forrest than in cultivar Essex. The ozone = tolerance of=20 Essex was associated with enhanced capacity to maintain ascorbate in the = reduced=20 form. The research provided further evidence that reduced ascorbate is = an=20 important antioxidant and the ascorbate-glutathione cycle plays a role = in=20 protecting the photosynthetic apparatus from ozone-induced damage.=20

Genetic variation in ozone response=20

Thirty soybean ancestors representing 92% of genes in modern U.S. and = Canadian cultivars were screened for ozone sensitivity. Two ancestors, = Fiskeby=20 III and Fiskeby 840-7-3, exhibited minimal foliar injury when exposed = for=20 six-days to 80 ppb ozone under greenhouse conditions and maintained = yield under=20 elevated ozone treatments during season-long exposures in open-top = chambers.=20 However, foliar injury in general was not a good predictor of seed yield = loss.=20 Specific ancestors exhibited low foliar injury with 25-30% yield loss = whereas=20 others were extensively injured with only 10% yield loss. Ozone effects = on seed=20 yield components were complex and included combinations of reduced seed = size and=20 reduced pod/seed number. These results suggest that screening of = germplasm for=20 ozone-tolerance based on foliar injury alone may not be the ideal or = appropriate=20 predictor of ozone effects on yield.=20

Antioxidants localized in the leaf apoplast and cell wall have the = potential=20 to scavenge ozone and ozone-derived reactive oxygen species thought to = be=20 involved in initiating foliar injury responses. The leaf apoplast from=20 ozone-sensitive and tolerant genotypes of soybean and tobacco contained = low=20 levels of ascorbic acid relative to total antioxidant capacity, evidence = that=20 soluble compounds other than ascorbate may contribute to ozone = scavenging=20 reactions. Soybean genotypes expressing differential ozone sensitivity = also=20 exhibited differences in composition of cell wall bound phenolic = compounds.=20 These results suggest that research on antioxidant mechanisms of ozone = tolerance=20 should be broadened to include other types of leaf chemistry in addition = to the=20 work being done on ascorbic acid.=20

G-proteins=20

The molecular signals that initiate ozone responses in plants are = thought to=20 originate in the leaf apoplast, but the mechanisms involved in sensing = and=20 propagating these signals are not known. Since GTPases (G-proteins) are = involved=20 in plant defense responses that share common features with ozone = responses,=20 Arabidopsis G-protein null mutants were tested for ozone sensitivity. = Mutants=20 with key deletions in the G-protein pathway did not exhibit the leaf = epinasty=20 observed in Columbia wild-type controls following ozone exposure, = evidence that=20 this particular phenotypic response to ozone is at least partially = G-protein=20 dependent. Biomass production, chlorophyll levels and photosynthesis = rates were=20 slightly lower in the G-protein null mutants following chronic ozone = exposure.=20 However, induction of peroxidase enzyme activity by ozone was similar in = both=20 G-protein mutants and wild-type controls. The results indicate that = stimulation=20 of peroxidase activity by ozone does not involve G-protein signaling = processes.=20 Ozone effects likely involve multiple pathways and cellular processes.=20 Suppression of the G-protein signaling pathway in Arabidopsis did not = markedly=20 increase its sensitivity to ozone, which suggests that other processes=20 compensated for the genomic changes or other modes of action are more = critical=20 in the etiology of ozone toxicity.=20

Objective 5. Develop numerical models to establish relationships = between=20 ambient ozone exposures and plant responses.=20

Alfalfa (Medicago sativa) cv. Beaver was grown under ambient = conditions at=20 multiple sites using the local grower cultivation practices. There were = two=20 harvests per growth season per plot (with 6 replicates). Since alfalfa=20 cultivation was on a five-year rotation cycle, each year a new seeding = was done=20 on separate study plots and harvested, starting with the year after = seeding and=20 during the following four years, with a total 78 harvests, but with = differing=20 age classes (2-5 years).=20

Alfalfa and total biomass (alfalfa + other plant species, weeds) = yields=20 varied between the two harvests at a given site during a given year and = between=20 sites and years, although there was no carry-over effect of stress from = one=20 harvest to another. Using the median yield value for all sites and years = combined, separately for alfalfa and for total biomass, data were = segregated=20 into two statistically differing classes: low=1Dand high. Alfalfa yields = for low=20 ranged from 85 to 17% of the median and from 117 to 218% for high. = Similarly for=20 total biomass yield values for low=1Dranged from 83 to 32% of the median = and from=20 117 to 197% for high. The low=1Dand high=1Dwere treated separately in = the yield=20 modeling of the alfalfa and the total biomass. The air quality and=20 meteorological variables were defined separately for each of three = growth stages=20 (time series) for use in the yield models. The independent variables in = each=20 stage were: ozone (median and 95th percentile hourly concentrations), = sulfur=20 dioxide and oxides of nitrogen exposure integrals (concentration x = exposure=20 duration) and temperature, relative humidity and global solar radiation, = and=20 precipitation depth totals. Mallows Critical Point Best regression was = used to=20 select the best yield models for further application. The initial = analyses were=20 done for alfalfa and total biomass separately, but with all yields, = sites and=20 years combined, the results giving very satisfactory values for both = adjusted=20 coefficient of variation (68-76%) and its significance p (0.000). Hourly = ozone=20 concentrations, the median and equal to or greater than the 95th = percentile=20 values were selected as important predictors of both alfalfa and total = biomass=20 yields. In addition to ozone, some of the other important predictors of = yield=20 included sulfur dioxide, oxides of nitrogen, temperature, global = radiation and=20 relative humidity.=20

Overall the models could account for 68 to 76% of the R2 variability = in the=20 yields of alfalfa or the total biomass respectively. Air quality (ozone = + sulfur=20 dioxide + oxides of nitrogen) influenced about 50% of the total = variation in all=20 of the alfalfa yields combined, with ozone accounting for one-half of = it. The=20 remaining 50% was due to variations in the climate and parameters that = were not=20 measured. Similar results were also obtained in the case of the total = biomass.=20 These results have been disseminated to the scientific community through = USDA-NE1013 Annual Technical Committee Meetings and currently three = journal=20 articles are in preparation.=20

Impacts

Ambient ozone concentrations during the growing season in all = locations=20 evaluated in this project strongly suppressed yields of = ozone-sensitive=20 snapbean genotypes. This implies that ambient ozone likely suppresses = yields=20 of ozone-sensitive crops in many regions of the U.S. This is supported = by the=20 findings of ozone-injury within open plots or in the semi-controlled = exposures=20 (open-top chambers or FACE systems) during summer seasons and provides = further=20 evidence that ozone is the cause of significant losses to many = agriculturally=20 important plants.=20
The information generated by this project is especially useful in = the=20 formulation of the National Ambient Air Quality Standard (NAAQS) for = ozone,=20 although the need is far from fulfilled. During the most recent review = of the=20 primary and secondary NAAQS for tropospheric ozone, it was concluded = that more=20 information on ozone-induced foliar injury and relationships to crop = growth=20 and productivity effects was needed.=20
Results of our studies have also been used by: 1) federal resource = managers in developing guidelines for protection of wilderness areas = such as=20 the National Parks; 2) federal agencies responsible for environmental = impact=20 assessment of expanded coal and oil industries in the Western US; and, = 3)=20 state agencies in the Southeastern US involved in transboundary = pollution=20 issues. Our findings about decreased nutritive quality and related = indirect=20 effects due to ozone are also under consideration for use in revising = extant=20 Critical Levels regulations in Europe.=20
Using a multivariate, multipoint statistical model, we = demonstrated that=20 ambient air quality contributed to 50% of alfalfa yield losses in an = Alberta,=20 Canada locale, and ozone was the most important air pollutant reducing = biomass. This study characterized the impacts of ambient ozone on a = crop yield=20 under ambient conditions. More importantly, it represents the first = attempt to=20 identify and separate the individual effects of ambient ozone on crop=20 productivity in the presence of other air pollutants and climatic = variables=20 that affect plant growth under field conditions.=20
Education and outreach activities performed by members of the = NE-1013=20 project include operation of the Air Quality Learning and = Demonstration Center=20 (PA), development of a website and web-based teaching modules (PA, = NC), site=20 tours (NC, PA, CA, NY), college course instruction (PSU, ASU, U Minn), = Master=20 Gardener sessions (NY), presentations to commercial growers and = extension=20 agents (CA, NY), and responses to local and national media about = ambient ozone=20 impacts on vegetation and ecosystem health.

Publications

2007=20

Booker, FL, KO Burkey, WA Pursley and AS Heagle. 2007. Elevated = carbon=20 dioxide and ozone effects on peanut. I. Gas-exchange, biomass, and leaf=20 chemistry. Crop Science 47:1475-1487.=20

Burkey, KO, FL Booker, WA Pursley and AS Heagle. 2007. Elevated = carbon=20 dioxide and ozone effects on peanut. II. Seed yield and quality. Crop = Science=20 47:1488-1497.=20

Calfapietra, C, AE Wiberley, TG Falbel, AR Linskey, G = Scarascia-Mugnozza, DF=20 Karnosky, F Loreto, and TD Sharkey. 2007. Isoprene synthase expression = and=20 protein levels are reduced under elevated O3 but not under elevated CO2 = (FACE)=20 in field-grown aspen trees. Plant Cell Environment 30:654-661.=20

Chen, X, C Tu, M Burton, D Watson, KO Burkey and S Hu. 2007. Plant = nitrogen=20 acquisition and interactions under elevated CO2: impact of endophytes = and=20 mycorrhizae. Global Change Biology. 13: 1238-1249.=20

Cheng, FY, KO Burkey, JM Robinson and FL Booker. 2007. Leaf = extracellular=20 ascorbate in relation to O3 tolerance of two soybean cultivars. = Environmental=20 Pollution 150:355-362.=20

Dubois, J.-J.B., EL Fiscus, FL Booker, MD Flowers and CD Reid. 2007.=20 Optimizing the statistical estimation of the parameters of the = Farquhar-von=20 Caemmerer-Berry model of photosynthesis. New Phytologist 176:402-414.=20

Fiscus, EL, FL Booker, J-JB Dubois, TR Rufty, JW Burton and WA = Pursley. 2007.=20 CO2 enhancement effects in container- versus ground-grown soybeans at = equal=20 planting densities. Crop Science 47:2486-2494.=20

Flowers, MD, EL Fiscus, KO Burkey, FL Booker and J-J Dubois. 2007.=20 Photosynthesis, chlorophyll fluorescence, and yield of snap bean = (Phaseolus=20 vulgaris L.) genotypes differing in sensitivity to ozone. Environmental = and=20 Experimental Botany 61:190-198.=20

Grantz, D.A., A. Shrestha, and H-B. Vu. 2008. Early vigor and ozone = response=20 in horseweed (Conyza canadensis) biotypes differing in glyphosate = resistance.=20 Weed Science 56:224=13230.=20

Holmes, WE, DR Zak, KS Pregitzer, and JS King. 2006. Elevated CO2 and = O3=20 alter soil nitrogen transformations beneath trembling aspen, paper = birch, and=20 sugar maple. Ecosystems 9:1354-1363.=20

Karnosky, DF, JM Skelly, KE Percy, and AH Chappelka. 2007. = Perspectives=20 regarding 50 years of research on effects of tropospheric ozone air = pollution on=20 U.S. Forests. Environmental Pollution 147:489-506.=20

Karnosky, DF, H Werner, T Holopainen, K Percy, T Oksanen, E Oksanen, = C=20 Heerdt, P Fabian, J Nagy, W Heilman, R Cox, N Nelson, and R Matyssek. = 2007.=20 Free-air exposure systems to scale up ozone research to mature trees. = Plant=20 Biology 9:181-190.=20

Kubiske, ME, VS Quinn, PE Marquardt, and DF Karnosky. 2007. Effects = of=20 elevated CO2 and/or O3 on intra- and interspecific competitive ability = of aspen.=20 Plant Biology 9:342-355.=20

Lin, JC, M. Nosal, RB Muntifering, and SV Krupa. 2007. Alfalfa = nutritive=20 quality for ruminant livestock as influenced by ambient air quality in=20 west-central Alberta. Environmental Pollution 149:99-103.=20

Lin, JC, K Nadarajah, M Volk, RB Muntifering and J Fuhrer. 2007. = Nutritive=20 quality of a species-rich, extensively managed pasture exposed to = elevated ozone=20 in a free-air fumigation system. Journal of Animal Science 90 (Suppl. = 1): 36.=20

Liu, L., J.S. King, and C.P. Giardina. 2007. Effects of elevated = atmospheric=20 CO2 and tropospheric O3 on nutrient dynamics: decomposition of leaf = litter in=20 trembling aspen and paper birch communities. Plant Soil, 299: 65-82.=20

Oncley, S.P., Foken, T., Vogt, R., Kohsiek, W., DeBruin,H.A.R., = Bernhofer,=20 C., Christen, A., van Gorsel, E., Grantz, D., Feigenwinter, C., Lehner, = I.,=20 Liebethal, D., Liu, H., Mauder, M., Pitacco, A., Ribeiro, L., and = Weidinger, T.=20 2007. The Energy Balance Experiment EBEX-2000. Part I: Overview and = energy=20 balance. Boundary Layer Meteorology 123: 1-28.=20

Percy, KE, M Nosal, W Heilman, T Dann, AH Legge, J Sober, and DF = Karnosky.=20 2007. New exposure-based metric approach for evaluating O3 risk to North = American aspen forests. Environmental Pollution 147:554-566.=20

Pregitzer, K.S., D.R. Zak, W.M. Loya, J.S. King, and A.J. Burton. = 2007. The=20 contribution of root systems to biogeochemical cycles in a changing = world. In Z.=20 Cardon and J. Whitbeck (eds) The rhizosphere-an ecological perspective.=20 Elsevier, Boston, pp. 155-178.=20

Zak D.R, W.E. Holmes, K.S. Pregitzer, J.S. King, D.S. Ellsworth, and = M.E.=20 Kubiske. 2007. Belowground competition and the response of developing = forest=20 communities to atmospheric CO2 and O3. Global Change Biology, 13: = 2230-2238.=20

2006=20

Burkey, K.O., H.S. Neufeld, L. Souza, A.H. Chappelka, and A.W. = Davison. 2006.=20 Seasonal profiles of leaf ascorbic acid in ozone-sensitive wildflowers.=20 Environmental Pollution. 143:427-434.=20

Bender, J., R. Muntifering, J. Lin and H. Weigel. 2006. Growth and = nutritive=20 quality of Poa pratensis as influenced by ozone and competition. = Environmental=20 Pollution 142: 109-115.=20

Decoteau, DR., J Ferdinand, J Savage, D Stevenson, and D Davis. 2006. = Advanced teacher training on air pollution effects on plants at the Air = Quality=20 Learning and Demonstration Center at the Arboretum at Penn State. = HortScience=20 41:1003.=20

Elagoz, V, S Han and WJ Manning. 2006. Acquired changes in stomatal=20 characteristics in response to ozone during plant growth and leaf = development of=20 bush beans (Phaseolus vulgaris L.) indicate phenotypic plasticity. = Environmental=20 Pollution 140:395-405.=20

Grulke, NE, HS Neufeld, AW Davison, M Roberts, AH Chappelka. 2006. = Stomatal=20 behavior of ozone-sensitive and -insensitive coneflowers (Rudbeckia = laciniata=20 var. digitata) in Great Smoky Mountains National Park. New Phytologist=20 173:100-109.=20

Grantz, D. and A. Shrestha. 2006. Tropospheric ozone and = interspecific=20 competition between yellow nutsedge and Pima cotton. Crop Science = 46:1879-1889.=20

Grantz, D., S. Gunn and H.-B. Vu. 2006. O3 impacts on plant = development: a=20 meta-analysis of root/shoot allocation and growth. Plant, Cell and = Environment=20 29:1193-1209.=20

Holmes, W.E., D.R. Zak, K.S. Pregitzer, and J.S. King. 2006. Elevated = CO2 and=20 O3 alter soil nitrogen transformations beneath trembling aspen, paper = birch, and=20 sugar maple. Ecosystems, 9:1354-1363.=20

Lewis, J., S. Ditchkoff, J. Lin, R. Muntifering and A.H. Chappelka. = 2006.=20 Nutritive quality of big bluestem (Andropogon gerardii) and eastern = gamagrass=20 (Tripsacum dactyloides) exposed to tropospheric ozone. Rangeland Ecol. = Mgmt.=20 59:267-274.=20

Long, S.P., E.A. Ainsworth, A.D.B. Leakey, A.D.B., J. Nosberger and = D.R. Ort.=20 2006. Food for thought: Lower-than-expected crop yield stimulation with = rising=20 CO2 concentrations. Science 312:1918-1921.=20

Muntifering, R.B., A.H. Chappelka, J.C. Lin, D.F. Karnosky and G.L. = Somers.=20 2006. Chemical composition and digestibility of Trifolium exposed to = elevated=20 ozone and carbon dioxide in a free-air (FACE) fumigation system. = Functional=20 Ecology 20: 269-275.=20

Muntifering, R.B., W.J. Manning, J.C. Lin and G.B. Robinson. 2006. = Short-term=20 exposure to ozone altered nutritive quality of alfalfa (Medicago sativa = L.)=20 under controlled exposure conditions. Environmental Pollution 140: 1-3.=20

Neufeld, H.S., A.H. Chappelka, K.O. Burkey, and A.W. Davison. 2006. = Reduced=20 ability of the SPAD meter to measure chlorophyll concentrations in = cutleaf=20 coneflower leaves exhibiting visible foliar injury from ozone. = Photosynthesis=20 Research 87: 281-286.=20

Souza, L., H.S. Neufeld, A.H. Chappelka, K.O. Burkey, and A.W. = Davison. 2006.=20 Seasonal development of ozone-induced foliar injury on tall milkweed = (Asclepias=20 exaltata) in Great Smoky Mountains National Park. Environmental = Pollution.=20 141:175-183.=20

Tu, C., F.L. Booker, D.M. Watson, X. Chen, T.W. Rufty, W. Shi and S. = Hu.=20 2006. Mycorrhizal mediation of plant N acquisition and residue = decomposition:=20 impact of mineral N inputs. Global Change Biology 12:793-803.=20

Weiser, G, WJ Manning, M Tausz and A Bytnerowicz. 2006. Evidence for=20 potential effects of ozone on Pinus cembra L. at mountain sites in = Europe: an=20 overview. Environmental Pollution 139:53-58.=20

2005=20

Ainsworth EA, SP Long. 2005. What have we learned from 15 years of = free-air=20 CO2 enrichment (FACE)? A meta-analytic review of the responses of=20 photosynthesis, canopy properties and plant production to rising CO2. = New=20 Phytologist 165:351-372.=20

Bernacchi CJ, PB Morgan, DR Ort, SP Long. 2005. The growth of soybean = under=20 free air [CO2] enrichment (FACE) stimulates photosynthesis while = decreasing in=20 vivo Rubisco capacity. Planta 220:434-446.=20

Booker, FL, EL Fiscus. 2005. Role of ozone flux and antioxidants in = the=20 suppression of ozone injury by elevated carbon dioxide in soybean. = Journal of=20 Experimental Botany 56:2139-2151.=20

Booker, FL, JE Miller, EL Fiscus, WA Pursley, LA Stefanski. 2005. = Comparative=20 responses of container- versus ground-grown soybean to elevated CO2 and = O3. Crop=20 Science 45:883-895.=20

Booker, FL, SA Prior, HA Torbert, EL Fiscus, WA Pursley, S Hu. 2005.=20 Decomposition of soybean grown under elevated concentrations of CO2 and = O3.=20 Global Change Biology 11:685-698.=20

Burkey, KO, JE Miller, EL Fiscus. 2005. Assessment of ambient ozone = effects=20 on vegetation using snap bean as a bioindicator species. Journal of=20 Environmental Quality 34:1081-1086.=20

Chapman, J.A., J.S. King, K.S. Pregitzer, and D.R. Zak. 2005. Effects = of=20 elevated concentrations of atmospheric CO2 and tropospheric O3 on = decomposition=20 of tree fine roots. Tree Physiology, 25:1501-1510.=20

Fiscus, EL, FL Booker, KO Burkey. 2005. Crop responses to ozone: = uptake,=20 modes of action, carbon assimilation and partitioning. Plant, Cell and=20 Environment 28:997-1011.=20

Grantz, DA. 2005. Ozone impacts on plants. In: P Dwivedi and RS = Dwivedi=20 (eds.). Physiology of Abiotic Stress in Plants. Oxford IBH Publishing = Co., New=20 Delhi.=20

Grantz, DA and A Shrestha. 2005. Ozone reduces crop yields and alters = competition with weeds such as yellow nutsedge. California Agriculture=20 59:137-143.=20

Hu, S, J Wu, KO Burkey, MK Firestone. 2005. Plant and microbial N = acquisition=20 under elevated atmospheric CO2 in two mesocosm experiments with annual = grasses.=20 Global Change Biology 11:213-223.=20

Hughes, NM, HS Neufeld, KO Burkey. 2005. Functional role of = anthocyanins in=20 high-light winter leaves of the evergreen herb Galax urceolata. New = Phytologist=20 168:575-587.=20

Long SP, Ainsworth EA, Leakey ADB, Morgan PB. 2005. Global food = insecurity.=20 Treatment of major food crops with elevated carbon dioxide or ozone = under=20 large-scale fully open-air conditions suggests recent models may have=20 overestimated future yields. Philosophical Transactions of the Royal = Society of=20 London B 360: 2011-2020.=20

Morgan, PB, GA Bollero, RL Nelson, FG Dohleman, SP Long. 2005. = Smaller than=20 predicted increase in aboveground net primary production and yield of=20 field-grown soybean under fully open-air [CO2] elevation. Global Change = Biology=20 11:1856-1865.=20

Sanz, J., R.B. Muntifering, V. Bermejo, B.S. Gimeno and S. Elvira. = 2005.=20 Ozone and increased nitrogen supply effects on the yield and nutritive = quality=20 of Trifolium subterraneum. Atmospheric Environment 39:5899-5907.=20

Schaub, M, JM Skelly, JW Zhang, JA Ferdinand, JE Savage, RE = Stevenson, DD=20 Davis, KC Steiner. 2005. Physiological and foliar symptom response in = the crowns=20 of Prunus serotina, Fraxinus americana and Acer rubrum canopy trees to = ambient=20 ozone under forest conditions. Environmental Pollution 133:553-567.=20

Shrestha, A and DA Grantz. 2005. Ozone impacts on competition between = tomato=20 and yellow nutsedge: Above- and below-ground effects. Crop Science = 45:1587-1595.=20

Wittig, VE, CJ Bernacchi, X-G Zhu, C. Calfapietra, R Ceulemans, P = Deangelis,=20 B Gielen, F Miglietta, PB Morgan, SP Long. 2005. Gross primary = production is=20 stimulated for three Populus species grown under free-air CO2 enrichment = from=20 planting through canopy closure. Global Change Biology 11:644-656.=20

2004=20

Booker, FL. 2004. Influence of ozone on ribonuclease activity in = wheat=20 (Triticum aestivum L.) leaves. Physiologia Plantarum 120:249-255.=20

Booker, FL, KO Burkey, K Overmyer, AM Jones. 2004. Differential = responses of=20 G-protein Arabidopsis thaliana mutants to ozone. New Phytologist = 162:633-641.=20

Booker, FL, EL Fiscus, JE Miller. 2004. Combined effects of elevated=20 atmospheric carbon dioxide and ozone on soybean whole-plant water use.=20 Environmental Management 33:S355-S362.=20

Bytnerowicz, A, B Godzik, K Grodzinska, W Fr=05czek, R Musselman, W = Manning, O=20 Badea, F Popescu, P Fleischer. 2004. Ambient ozone in forests of the = Central and=20 Eastern European mountains. Environmental Pollution 130: 5-16.=20

Estes, BL, SA Enebak, AH Chappelka. 2004. Loblolly pine seedling = growth after=20 inoculation with plant growth-promoting rhizobacteria and ozone = exposure.=20 Canadian Journal of Forest Research 34:1410-1416.=20

Finkelstein, PL, AW Davison, HS Neufeld, TP Meyers, AH Chappelka. = 2004.=20 Sub-canopy deposition of ozone in a stand of cutleaf coneflower. = Environmental=20 Pollution 131:295-303.=20

Grantz, DA and MJ Sanz. 2004. Common co-occurrence of citriculture = and ozone=20 air pollution: Potential for yield reductions. Proceedings 10th = International=20 Society of Citriculture Congress. Paper No. 97.=20

Grantz, DA and AK Murray. 2004. Effect of ozone on phloem transport = in=20 cotton. Proceedings of the 2004 Beltwide Cotton Conferences, San = Antonio, TX.=20 January 2004. Pp. 2144-2149.=20

Grantz, DA and A Shrestha. 2004. Ozone affects competition between = cotton and=20 nutsedge. Proceedings of the 2004 Beltwide Cotton Conferences, San = Antonio, TX.=20 January 2004. Pp. 2877-2882.=20

Gravano, E, F Bussotti, RJ Strasser, M Schaub, K Novak, JM Skelly, C = Tani.=20 2004. Ozone symptoms in leaves of woody plants in open-top chambers:=20 ultrastructural and physiological characteristics. Physiologia Plantarum = 121:620-633.=20

Gr=C3=BCnhage, L, SV Krupa, AH Legge, HJ J=C3=A4ger. 2004. Ambient = flux-based critical=20 values of ozone for protecting vegetation: differing spatial scales and=20 uncertainties in risk assessment. Atmospheric Environment 38:2433-2437.=20

Karnosky, D.F., Percy, K.E, Chappelka, A.H. and Krupa, S. (2004). Air = pollution and global change impacts on forest ecosystems: monitoring and = research needs. In Air Pollution, Global Change and Forests in the New=20 Millenium. eds. Karnosky, D.F., Percy, K.E., Simpson, C. and Chappelka, = A.H.=20 Elsevier Science, Amsterdam, The Netherlands. pp 447-459.=20

Krupa, SV, R Muntifering, AH Chappelka. 2004. Effects of ozone on = plant=20 nutritive quality characteristics for ruminant animals. The Botanica = 54:1-12.=20

Long SP, EA Ainsworth, A Rogers, DR Ort. 2004. Rising atmospheric = carbon=20 dioxide: plants FACE the future. Annual Review of Plant Biology = 55:591-628.=20

Lorence, A, BI Chevone, P Mendes, CL Nessler. 2004. Myo-inositol = oxygenase=20 offers a possible entry point into plant ascorbate biosynthesis. Plant=20 Physiology 134:1200-1205.=20

Manning, WJ, CJ Bergweiler. 2004. Assessing plant response to ambient = ozone:=20 growth of young apple trees in open-top chambers and corresponding = ambient air=20 plots. Environmental Pollution 132:503-508.=20

Manning, WJ, B Godzik. 2004. Bioindicator plants for ambient ozone in = Central=20 and Eastern Europe. Environmental Pollution 130:33-39.=20

Morgan PB, CJ Bernacchi, DR Ort, SP Long. 2004. An in vivo analysis = of the=20 effect of season-long open-air elevation of ozone to anticipated 2050 = levels on=20 photosynthesis in soybean. Plant Physiology 135:2348-2357.=20

Murray, AK and DA Grantz. 2004. Carbohydrate composition of cotton = aphid=20 honeydew. Proceedings of the 2004 Beltwide Cotton Conferences, San = Antonio, TX.=20 January 2004. Pp. 1590-1593.=20

Niyogi, D, H-I Chang, VK Saxena, T Holt, K Alapaty, FL Booker, F = Chen, KJ=20 Davis, B Holben, T Matsui, T Meyers, WC Oechel, RA Pielke, Sr, R Wells, = K=20 Wilson, Y Xue. 2004. Direct observations of the effects of aerosol = loading on=20 net ecosystem CO2 exchanges over different landscapes. Geophysical = Research=20 Letters 31: L20506, doi: 10.1029/2004GL020915.=20

Percy, K.E., Legge, A.H. and Krupa, S.V. (2004). Tropospheric ozone: = A=20 continuing threat to global forests? In Air Pollution, Global Change and = Forests=20 in the New Millennium. eds. Karnosky, D.F., Percy, K.E., Simpson, C. and = Chappelka, A.H. Elsevier Science, Amsterdam, The Netherlands. pp 85-118. =

Robinson, JM, J Lydon, CA Murphy, R Rowland, R Smith. 2004. Effect of = Pseudomonas syringae pv. tagetis infection on sunflower leaf = photosynthesis and=20 ascorbic acid relations. International Journal of Plant Science. = 165:263-271.=20

Robinson, JM, RC Sicher Jr. 2004. Antioxidant levels decline in = primary=20 leaves of barley during growth at ambient and elevated carbon dioxide = levels.=20 International Journal Of Plant Science. 165:965-972.=20

Sandermann, H. (ed). 2004. Molecular Ecotoxicology of Plants. Berlin: = Springer. 241 pp.=20

Sanz, J, RB Muntifering, BS Gimeno, V Bermejo. 2004. Nutritive = quality and=20 growth of Trifolium subterrraneum are modulated by ambient ozone = concentrations=20 and nitrogen fertilization. In: Garcia Criado, B, Garcia Cuidad, B, = Vasquez de=20 Aldana, A, Zabagogeazcoa, I. (eds.). Proceedings Spanish Society for = Pasture=20 Studies, Salamanca, Spain. 10-14 May. Pp. 197-201.=20

Schaub, M, JM Skelly, KC Steiner, DD Davis, SP Pennypacker, J.Zhang, = JA=20 Ferdinand, JE Savage, RE Stevenson. 2004. Physiological and foliar = responses of=20 Prunum serotina, Fraxinus americana and Acer rubrum seedlings to varying = soil=20 moisture and ozone. Environmental Pollution 124:307-320.=20

Skelly JM, DD Davis, DR Decoteau. 2004. Development of an air quality = learning and demonstration center at the arboretum at Penn State. = HortScience=20 39:810.=20

Wei, C, JM Skelly, SP Pennypacker, JA Ferdinand, JE Savage, RE = Stevenson, DD=20 Davis. 2004. Responses of hybrid poplar clones and red maple seedlings = to=20 ambient ozone under differing light within a mixed hardwood forest.=20 Environmental Pollution 130:199-214.=20

Wei, C, JM Skelly, SP Pennypacker, JA Ferdinand, JE Savage, RE = Stevenson, DD=20 Davis. 2004. Influence of light fleck and low light on foliar injury and = physiological responses of two hybrid poplar clones to ozone. = Environmental=20 Pollution 130:215-227.=20

2003=20

Burkey, KO, G Eason, EL Fiscus. 2003. Factors that affect leaf = extracellular=20 ascorbic acid content and redox status. Physiologia Plantarum 117:51-57. =

Bussotti, F, M Schaub, A Cozzi, N Krauchi, M Ferretti, K Novak, JM = Skelly.=20 2003. Assessment of ozone visible symptoms in the field: perspectives of = quality=20 control. Environmental Pollution 125:81-89.=20

Bytnerowicz, A, M Arbaugh, R Alonso (eds.). 2003. Ozone Air Pollution = in the=20 Sierra Nevada: Distribution and Effects on Forests. Elsevier, = Developments in=20 Environmental Science 2, Amsterdam, 402 pp, ISBN 0 08 044193 9.=20

Chappelka, AH, HS Neufeld, AW Davison, GL Somers, JR Renfro. 2003. = Ozone=20 injury on cutleaf coneflower (Rudbeckia laciniata) and crown-beard = (Verbesina=20 occidentalis) in Great Smoky Mountains National Park. Environmental = Pollution=20 125:53-59.=20

Davison, AW, HS Neufeld, AH Chappelka, K Wolff, PL Finkelstein. 2003. = Interpreting spatial variation in ozone symptoms shown by cutleaf = coneflower,=20 Rudbeckia laciniata L. Environmental Pollution:61-70.=20

Fuhrer, J, FL Booker. 2003. Ecological issues related to ozone: = agricultural=20 issues. Environment International 29:141-154.=20

Grantz, DA, JHB Garner, DW Johnson. 2003. Ecological effects of = particulate=20 matter. Environment International 29:213-239.=20

Grantz, D.A., V Silva, M Toyota, N Ott. 2003. Ozone increases root=20 respiration but decreases leaf CO2 assimilation in cotton and melon. = Journal of=20 Experimental Botany 54:2374-2384.=20

Gr=C3=BCnhage, L., Krupa, S. V., Legge, A. H., and J=C3=A4ger, H.-J. = (2003). Ambient=20 flux-based critical values of ozone for protecting vegetation: differing = spatial=20 scales and uncertainties in risk assessment. In Proc. UN-ECE Conf. = Ozone:=20 Critical Levels II, Gothenburg, Sweden, 19-23 November 2002. IVL, = Gothenburg,=20 Sweden. pp 19-34.=20

Grimmig, B, MN Gonzalez-Perez, G Leubner-Metzger, R Vogeli-Lange, F = Meins, R=20 Hain, J Penuelas, B Heidenreich, C Langebartels, D Ernst, H Sandermann. = 2003.=20 Ozone-induced gene expression occurs via ethylene-dependent and = -independent=20 signalling. Plant Molecular Biology 51:599-607.=20

Holmes, W.E., D.R. Zak, K.S. Pregitzer, and J.S. King. 2003. Soil = nitrogen=20 transformations under Populus tremuloides, Betula papyrifera, and Acer = saccharum=20 following 3 years exposure to elevated CO2 and O3. Global Change Biology = 9:1743-1750.=20

Krupa, SV. 2003. Joint effects of elevated levels of ultraviolet-B = radiation,=20 carbon dioxide and ozone on plants. Photochemistry and Photobiology = 78:535-542.=20

Krupa, SV, M Nosal, JA Ferdinand, RE Stevenson, JM Skelly. 2003. A=20 multi-variate statistical model integrating passive sampler and = meteorology data=20 to predict the frequency distributions of hourly ambient ozone (O3)=20 concentrations. Environmental Pollution 124, 173-178.=20

Loya, W.M., K.S. Pregitzer, N.J. Karberg, J.S. King, and = C.P.Giardina. 2003.=20 Reduction of soil carbon formation by tropospheric ozone under increased = carbon=20 dioxide levels. Nature 425:705-707.=20

Manning, WJ, RB Flagler, MA Frenkel. 2003. Assessing plant response = to=20 ambient ozone: growth of ozone-sensitive loblolly pine seedlings treated = with=20 ethylenediurea or sodium erythorbate. Environmental Pollution 126:73-81. =

Manning, WJ. 2003. Detecting plant effects is necessary to give = biological=20 significance to ambient ozone monitoring data and predictive ozone = standards.=20 Environmental Pollution 126:375-379.=20

Morgan PB, EA Ainsworth, SP Long. 2003. How does elevated ozone = impact=20 soybean? A meta-analysis of photosynthesis, growth and yield. Plant, = Cell and=20 Environment 26:1317-1328.=20

Novak, K, JM Skelly, M Schaub, N Krauchi, C Hug, W Landolt, P = Bleuler. 2003.=20 Ozone air pollution and foliar injury development on native plants of=20 Switzerland. Environmental Pollution 125:41-52.=20

Orendovici, T, JM Skelly, JA Ferdinand, JE Savage, MJ Sanz, GC Smith. = 2003.=20 Response of native plants of northeastern United States and southern = Spain to=20 ozone exposures; determining exposure/response relationships. = Environmental=20 Pollution 125:31-40.=20

Powell, MC, RB Muntifering, JC Lin, AH Chappelka. 2003. Yield and = nutritive=20 quality of sericea lespedeza (Lespedeza cuneata) and little bluestem=20 (Schizachyrium scoparium) exposed to ground-level ozone. Environmental = Pollution=20 122:313-322.=20

Radzio, JA, A Lorence, BI Chevone, CL Nessler. 2003. = L-Gulono-1,4-lactone=20 oxidase expression rescues vitamin C-deficient Arabidopsis (vtc) = mutants. Plant=20 Molecular Biology 53:837-844.=20

Schaub, M, JM Skelly, KC Steiner, DD Davis, SP Pennypacker, J Zhang, = JA=20 Ferdinand, JE Savage, RE Stevenson. 2003. Physiological and foliar = injury=20 responses of Prunus serotina, Fraxinus americana, and Acer rubrum = seedlings to=20 varying soil moisture and ozone. Environmental Pollution 124:307-320.=20

Watkinson, JI, AA Sioson, C Vasquez-Robinet, . . . BI Chevone . . . = 2003.=20 Photosynthetic acclimation is reflected in specific patterns of gene = expression=20 in drought-stressed loblolly pine. Plant Physiology 133:1702-1716.=20

Yuska, DE, JM Skelly, JA Ferdinand, RE Stevenson, JE Savage, JD = Mulik, A=20 Hines. 2003. Use of bioindicators and passive sampling devices to = evaluate=20 ambient ozone concentrations in north central Pennsylvania. = Environmental=20 Pollution 125:71-80.=20

2002=20

Barbo, DN, AH Chappelka, GL Somers, MS Miller-Goodman, K Stolte. = 2002. Ozone=20 impacts on loblolly pine (Pinus taeda L.) grown in a competitive = environment.=20 Environmental Pollution 116:27-36.=20

Burkey, KO, G Eason. 2002. Ozone tolerance in snap bean is associated = with=20 elevated ascorbic acid in the leaf apoplast. Physiologia Plantarum 114: = 387-394.=20

Bytnerowicz, A, B Godzik, W Frczek, K Grodziska, M Krywult, O Badea, = P=20 Baranok, O Blum, M Cerny, S Godzik, B Mankovska, W Manning, P Moravcik, = R=20 Musselman, J Oszlanyi, D Postelnicu, J Szdzuj, M Varsavova, M Zota. = 2002.=20 Distribution of ozone and other air pollutants in forests of the = Carpathian=20 Mountains in central Europe. Environmental Pollution 116:3-25.=20

Chappelka, AH. 2002. Reproductive development of blackberry (Rubus=20 cuneifolius), as influenced by ozone. New Phytologist 155: 249-255. = Elagoz, V,=20 WJ. Manning. 2002. Bean plants and ozone: morphology matters. = Environmental=20 Pollution 120: 521-524.=20

Elagoz, V, WJ. Manning. 2002. Bean plants and ozone: morphology = matters.=20 Environmental Pollution 120: 521-524.=20

Fiscus, EL, FL Booker. 2002. Growth of Arabidopsis flavonoid mutant = is=20 challenged by radiation longer than the UV-B band. Environmental and=20 Experimental Botany 48:213-224.=20

Fiscus, EL, JE Miller, FL Booker, AS Heagle, CD Reid. 2002. The = impact of=20 ozone and other limitations on the crop productivity response to CO2. = Technology=20 8: 181-192.=20

Grantz, D. A., J. H. B. Garner, and D. W. Johnson. 2002. Ecological = effects=20 of particulate matter. Environment International 29: 213-239.=20

Heagle, A.S., J.E. Miller, K.O. Burkey, G.P. Eason, and W.A. Pursley. = 2002.=20 Growth and yield responses of Phaseolus vulgaris to mixtures of carbon = dioxide=20 and ozone. Journal of Environmental Quality 31:2008-2014.=20

Heath, L.S., N. Ramakrishnan, R.R. Sederoff, R.W. Whetten, B.I. = Chevone, C.A.=20 Struble, V.Y. Jouenne, D. Chen, L. van-Zyl and R. Grene. 2002. Studying = the=20 functional genomics of stress responses in loblolly pine with the = Expresso=20 microarray experiment management system. Comparative and Functional = Genomics=20 3:226-243.=20

Krupa, SV. 2002. Sampling and physico-chemical analysis of = precipitation.=20 Environmental Pollution 120:565-594.=20

Krupa, S.V. and J.F. Moncrief. 2002. An integrative analysis of the = role of=20 atmospheric deposition and land management practices on nitrogen in the = US=20 agricultural sector. Environmental Pollution 118:273-283.=20

Lee, JC, KC Steiner, JW Zhang, JM Skelly. 2002. Heritability of ozone = sensitivity in open-pollinated families of black cherry (Prunus serotina = Ehrh.).=20 Forest Science 48:111-117.=20

Long SP , Naidu SL .2002. Effects of oxidants at the biochemical, = cell and=20 physiological levels, with particular reference to ozone. In: Air = Pollution and=20 Plant Life (eds Bell JNB ,Treshow M), pp. 69-88. John Wiley & Sons, = Ltd.,=20 West Sussex.=20

Manning, W. J. B. Godzik and R. Musselman. 2002. Potential = bioindicator plant=20 species for ambient ozone in forested mountain areas of central Europe.=20 Environmental Pollution 119: 283-290.=20

Novak, K.J. 2002. Ozone air pollution and foliar injury development = on native=20 plants of Switzerland. MS Thesis, The Pennsylvania State University, = University=20 Park, PA. 92 pp.=20

Orendovici, T. 2002. Response of native plants of northeastern United = States=20 of America and southeastern Spain to ozone exposures: Determining=20 exposure/response relationships. MS Thesis, The Pennsylvania State = University,=20 University Park, PA. 137 pp.=20

Wei, C. 2002. Influence of sunflecks on hybrid poplar, black cherry, = and red=20 maple response to ozone. Doctoral dissertation, The Pennsylvania State=20 University, University Park, PA. 125 pp.=20

West, D.H., A.H. Chappelka, K.M. Tilt, H.G. Ponder and J.D. Williams. = 2002.=20 Effect of tree shelters on growth and gas exchange of four tree species = under=20 field and nursery conditions. Journal of Environmental Horticulture 20: = 96-100.=20

Wohlgemuth, H, K Mittelstrass, S Kschieschan, J Bender, HJ Weigel, K=20 Overmyer, J Kangasjarvi, H Sandermann, C Langebartels. 2002. Activation = of an=20 oxidative burst is a general feature of sensitive plants exposed to the = air=20 pollutant ozone. Plant, Cell and Environment 25:717-726.=20

Yuska, D.E. 2002. Use of bioindicators and passive sampling devices = to=20 evaluate ambient ozone concentrations in north central Pennsylvania. MS = Thesis,=20 The Pennsylvania State University, University Park, PA. 70 pp.=20

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SAES-422 Multistate Research Activity Accomplishments Report=20 (termination)

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