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Growth Responses of Yellow-Poplar (Liriodendron Tulipifera L.) Exposed

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Growth Responses of Yellow-Poplar (Liriodendron Tulipifera L.) Exposed Plant, Cell and Environment (2002) 25, 1527- 1537 Growth responses of yellow-popl~r(Li~i~de~d~~~ ~~lipife~~ L.) exposed to 5 years of O3 alone or Gornbi~edwith elevated C02 J. REBBECK & A. J. SCHERZER Northeastern Research Station, USDA Forest Service, 359 Main Road, Delaware, OH 43015, USA ABSTRACT INTRODUCTION Field-grown yellow-poplar (Liriodendron tulipifera L.) Current projections suggest that forests within the eastern were fumigated from May to October in 1992-96 within United States will continue to be exposed to increasing open-top chambers to determine the impact of ozone (0,) levels of tropospheric O3 (Chameides et al. 1994) and ambi- alone or combined with elevated carbon dioxide (CO,) on ent C02(Watson et at. 1990). Rural regions of eastern USA sapling growth. Tkeatments were replicated three times and typically average summer daytime levels in excess of included charcoal-filtered air (CF); 1x ambient ozone 0.05 p.p.m. O3(Lefohn, Edwards & Adams 1994) with occa- (1x 03);1.5 x ambient ozone (1.5 x 03);1.5 x ambient sional O3 episodes above 0.100 p.p.m. Tropospheric O3has ozone plus 350 p.p.m. carbon dioxide (1-5 x O3+ CO,) (tar- long been known to be phytotoxic (Middleton 1956) and get of 700p.p.m. CO,); and open-air chamberless plot causes the greatest amount of damage on vegetation of any (OA). After five seasons, the total cumulative O3 exposure gaseous pollutant by reducing growth and productivity of (SU~~=sumof hourly O3 concentrations during the many plant species through reductions in photosynthesis, study) ranged from 145 (CF) to 861 (14 x 0,) pq.m. x h accelerated leaf senescence and decreased root growth (parts per million hour). Ozone had no statistically signifi- (Bortier, Ceulemans & de Temmerman 2000; Rudorff, Mul- cant effect on yellow-poplar growth or biomass, even chi & Lee 2000). Elevated COz concentrations generally though total root biomass was reduced by 13'/0 in the increase plant growth through increased carbon assimila- 14 x 0,-exposed saplings relative to CF controls. Although tion, biomass and leaf area (Ceulemans & Mousseau. 1994; exposure to 1*5~0,+CO,had a stimulatory effect on Sax, Ellsworth & Heath 1998). It is hypothesized that plants yearly basal area growth increment after two seasons, sig- growing in elevated COzmay be protected from O3damage nificant increases in shoot and root biomass (-60% by reducing O3uptake because elevated C02often reduces increase relative to all others) were not detected until the stomata1 conductance (Eamus & Jarvis 1989; Nousseau & fifth season. After five seasons, the yearly basal area growth Saugier 1992). Elevated C02 may also provide substrates increment of saplings exposed to 1.5 x O3+ C02-air for detoxification and repair processes against O3 damage increased by 41'%0relative to all others. Based on this multi- (Rudorff et al. 2000). year study, it appears that chronic O3 effects on yellow- Empirical information on the combined effects of the 0, poplar growth are limited and slow to manifest, and are and C02 on woody plants is limited (Allen 1990; Olszyk consistent with previous studies that show yellow-poplar et al. 2000). In some studies, elevated C02offsets the neg- growth is not highly responsive to O3exposure. In addition, ative effects of O3 in deciduous species (Volin, Reich & these results show that enriched C02may ameliorate the Givnish 1998; Broadmeadow & Jackson 2000), whereas in negative effects of elevated O3 on yellow-poplar shoot others no ameliorative effects are detected (Kull etal, growth and root biomass under field conditions. 1996). Multi-year field fumigations are needed to accu- rately assess the chronic response of tree species to the Key-words: elevated O3 + elevated CO,; hardwood trees; combination of elevated C02and 03.This can be achieved scaling O3 responses. by exposing the same population of trees to gaseous pollut- anis during their development from seedlings to saplings or mature trees (Lee & Jarvis 1995; Retzlaff, Williams & DeJong 1997; Rey & Jarvis 1997). These studies are very costly to run and only a few have been undertaken, result- ing in a paucity of empirical .data CHI the response of sap- lings and , mature trees to gaseous pollutants (Karnosky Correspondence: Joanne Rebbeck. Fax: +I: 740-368-0252;e-mail: et al. 1999; Isebrands et at. 2001). These types of studies can [email protected] serve as an important linkage to extrapolate seedling pol- @ 2002 Blackwell Publishing Ltd 1527 1528 J. Rebbeck & A. J. Scherzer lutant response data to larger and more structurally com- In 1992, methyl [1-[(butylamino)carbonyl]-1H-benzimi- plex older trees. dazol-2-yl] carbamate [Benomyl 50Em(14.7 mg active ingre- In the current study, we investigated how field-planted dient L-I); DuPont, Wilmington, DE, USA] was applied yellow-poplar (Liriodendron tulipifera L.) seedlings grown biweekly from mid-August to midSeptember as a prophy- under less than optimal conditions (unmanipulated soil lactic to control powdery mildew, a serious foliar disease of moisture and fertility) responded to chronic levels of ele- yellow-poplar in central Ohio that can kill young seedlings. vated 0, either alone or combined with elevated CO, over As observed in previous studies, Benomyl did not protect five seasons. The effects of elevated 0, on yellow-poplar, the seedlings from oxidant injury (Rebbeck 1996a; Loats & an ecologically and economically important hardwood spe- Rebbeck 1999). No other applications of pesticides were cies in eastern USA forests, have been well studied in pot- made during the study. In 1993and 1994, sporadic outbreaks ted seedlings under environmentally controlled conditions of aphids were observed with a fairly uniform distribution showing both inhibitions and stimulations in growth and in all plots. Insecticidal soap [Safero (39 mL L-I); Gardens physiological processes (Chappelka, Chevone & Seiler Alive, Lawrenceburg, IN, USA] was applied biweekly to 1988; Jensen & Patton 1990; Roberts 1990; Cannon, Rob- suppress aphid populations. Biological control measures erts & Bargar 1993; Rebbeck & Loats 1997; Loats & were subsequently adopted and commercially reared lady- Rebbeck 1999). Long considered an 0,-sensitive species, bugs (Hippodamia sp.) were released biweekly from mid- yellow-poplar is used as a bioindicator of 0, in eastern US July until late August during each season. forests (Davis & Skelly 1992). Our objectives were to deter- mine: (1) if the responses of field-grown saplings to multi- Pollutant exposure and experimental design year O3exposures were comparable with those of scedlings in shorter-term 0, exposures under more controlled condi- The experiment was a randomized block design with three tions; and (2) if doubling ambient CO, concentrations replicates of the following treatments: (1) charcoal-filtered would ameliorate negative 0, growth effects. air (CF); (2) 1 x ambient 03(1 x 03); (3) 1.5~ambient 0, (1.5 x 0,); (4) 1.5 x ambient O3 plus 350 p.p.m. CO, above MATERIALS AND METHODS ambient (target CO, concentration was 700p.p.m) Seedling culture and site characteristics (1.5 x 0, + CO,); and (5) open-air chamberless plot (OA). Due to the prohibitive costs associated with supplying twice In 1991, a field plantation of yellow-poplar was established ambient CO, to three additional open-top chambers, we did at the Northeastern Research Station's Forestry Sciences not characterize the effects of elevated C02alone on yel- Laboratory in Delaware, Ohio, USA (latitude 40'21' N lon- low-poplar growth. Gases were dispensed 24 h per day gitude 83'04' W), by clearing a 26 m by 73 m area in a 20- from mid-May until mid-October in 1992-96. The polyvi- year-old-abandoned American elm (Ulmus americana L) nyl-chloride film panels were removed each season when plantation. The glacial till soil was primarily of the Blunt gas exposures were terminated. In spring 1994, standard Series with pockets of the Morley Series. Soil textures were OTCs (3 m in diameter, 2.4 m high) were extended to a determined to be primarily clay loams. Soils were chemi- height of 4.6 m. The mean volumetric airflow within cham- cally analysed prior to planting to a depth of 25 cm and bers was 47 m3 min-I ? 10% and was provided by a single were considered adequate in total N (24-7 pg g-I NH,-N blower. In spring 1995, standard OTCs were replaced with and 8.2 pg g-I NO,-N). Mean soil pH was 6.39 and CEC was larger open-top chambers (4-6 m in diameter, 9.2 m high) 13.9 meqI100 g dry soil. Extractable ion concentrations similar to those developed by Heagle et al. (1989), but dou- were 7.4 pg g-I P; 103.5 pg g-I K; 1909 pg g-I Ca; and bled in height. Air was blown into the bottom of each 217.6 pg g-I Mg. Soil chemical properties were measured chamber via two 2HP, 91.4-cm-diameter fans (Penn Venti- annually; no major changes in concentrations were lator Inc., Philadelphia, PA, USA), each housed in a galva- observed throughout the study. nized sheet-metal box (123 cm wide, 123 cm high, 123 cm One-year-old bareroot stock originating from seeds long). Unfiltered ambient air entered each chamber after collected in south-eastern Tennessee was obtained from a passing through particulate filters and connecting plastic private nursery in western Pennsylvania, USA. Approxi- ducts at approximately 6.8 m3s-I. mately 250 uniform bare-rooted seedlings were planted Ozone was generated from vaporized liquid oxygen with 2.1 m by 2.1 m apart in July 1991 to serve as buffer trees an electric spark discharge 0, generator (OREC Model between the plots. In May 1992 within each chamber or 03V10-0; OREC, Glendale, AZ, USA) and was dispensed plot, 12 randomly chosen seedlings were planted 47 cm into the 1x0, and 1.5 x O3 OTC's through Teflon tubing apart in a circular pattern approximately 1.8 m in diameter when ambient levels exceeded 0.03 p.p.m.
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