David F. Karnosky

Growth responses of Populus tremuloides clones to interacting elevated carbon dioxide and tropospheric ozone

The Intergovernmental Panel of Climate Change (IPCC) has concluded that the greenhouse gases carbon dioxide (CO2) and tropospheric ozone (O3) are increasing concomitantly globally. Little is known about the effect of these interacting gases on growth, survival, and productivity of forest ecosystems. In this study we assess the effects of three successive years of exposure to combinations of elevated CO2 and O3 on growth responses in a five trembling aspen (Populus tremuloides) clonal mixture in a regenerating stand. The experiment is located in Rhinelander, Wisconsin, USA (45 degrees N 89 degrees W) and employs free air carbon dioxide and ozone enrichment (FACE) technology. The aspen stand was exposed to a factorial combination of four treatments consisting of elevated CO2 (560 ppm), elevated O3 (episodic exposure-90 microl l(-1) hour(-1)), a combination of elevated CO2 and O3, and ambient control in 30 m treatment rings with three replications. Our overall results showed that our three growth parameters including height, diameter and volume were increased by elevated CO2, decreased by elevated O3, and were not significantly different from the ambient control under elevated CO2 + O3. However, there were significant clonal differences in the responses; all five clones exhibited increased growth with elevated CO2, one clone showed an increase with elevated O3, and two clones showed an increase over the control with elevated CO2 + O3, two clones showed a decrease, and one was not significantly different from the control. Notably. there was a significant increase in current terminal shoot dieback with elevated CO2 during the 1999-2000 dormant season. Dieback was especially prominent in two of the five clones, and was attributed to those clones growing longer into the autumnal season where they were subject to frost. Our results show that elevated O3 negates expected positive growth effects of elevated CO2 in Populus tremuloides in the field, and suggest that future climate model predictions should take into account the offsetting effects of elevated O3 on CO2 enrichment when estimating future growth of trembling aspen stands.

Next generation of elevated [CO2] experiments with crops: A critical investment for feeding the future world

A rising global population and demand for protein-rich diets are increasing pressure to maximize agricultural productivity. Rising atmospheric [CO(2)] is altering global temperature and precipitation patterns, which challenges agricultural productivity. While rising [CO(2)] provides a unique opportunity to increase the productivity of C(3) crops, average yield stimulation observed to date is well below potential gains. Thus, there is room for improving productivity. However, only a fraction of available germplasm of crops has been tested for CO(2) responsiveness. Yield is a complex phenotypic trait determined by the interactions of a genotype with the environment. Selection of promising genotypes and characterization of response mechanisms will only be effective if crop improvement and systems biology approaches are closely linked to production environments, that is, on the farm within major growing regions. Free air CO(2) enrichment (FACE) experiments can provide the platform upon which to conduct genetic screening and elucidate the inheritance and mechanisms that underlie genotypic differences in productivity under elevated [CO(2)]. We propose a new generation of large-scale, low-cost per unit area FACE experiments to identify the most CO(2)-responsive genotypes and provide starting lines for future breeding programmes. This is necessary if we are to realize the potential for yield gains in the future.

Consequences of elevated carbon dioxide and ozone for foliar chemical composition and dynamics in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera).

Atmospheric chemical composition affects foliar chemical composition, which in turn influences the dynamics of both herbivory and decomposition in ecosystems. We assessed the independent and interactive effects of CO2 and O3 fumigation on foliar chemistry of quaking aspen (Populus tremuloides) and paper birch (Betula papyrifera) at a Free-Air CO2 Enrichment (FACE) facility in northern Wisconsin. Leaf samples were collected at five time periods during a single growing season, and analyzed for nitrogen. starch and condensed tannin concentrations, nitrogen resorption efficiencies (NREs), and C:N ratios. Enriched CO2 reduced foliar nitrogen concentrations in aspen and birch; O3 only marginally reduced nitrogen concentrations. NREs were unaffected by pollution treatment in aspen, declined with 03 exposure in birch, and this decline was ameliorated by enriched CO2. C:N ratios of abscised leaves increased in response to enriched CO2 in both tree species. O3 did not significantly alter C:N ratios in aspen, although values tended to be higher in + CO2 + O3 leaves. For birch, O3 decreased C:N ratios under ambient CO2 and increased C:N ratios under elevated CO2. Thus, under the combined pollutants, the C:N ratios of both aspen and birch leaves were elevated above the averaged responses to the individual and independent trace gas treatments. Starch concentrations were largely unresponsive to CO2 and O3 treatments in aspen. but increased in response to elevated CO2 in birch. Levels of condensed tannins were negligibly affected by CO2 and O3 treatments in aspen, but increased in response to enriched CO2 in birch. Results from this work suggest that changes in foliar chemical composition elicited by enriched CO2 are likely to impact herbivory and decomposition, whereas the effects of O3 are likely to be minor, except in cases where they influence plant response to CO2.

Agrobacterium rhizogenes-mediated genetic transformation and regeneration of a conifer:Larix decidua

SummaryGene transfer and plant regeneration systems have been developed for European larch (Larix decidua Mill.) in our laboratory. Aseptically germinated young seedlings were hypocotyl wound-inoculated withAgrobacterium rhizogenes strains 11325 containing a wild-type Ri (root-inducing) plasmid. Swollen stems appeared at infected wounds followed by either abundant hairy roots or adventitious shoot buds that developed within 3 to 4 wk after inoculation. No symptoms were seen on wounded but uninoculated seedlings. We demonstrated agrobacteria attached to larch cells by examination of scanning electron micrographs. Subsequently, calli derived from symptomatic tissues exhibited phytohormone autotrophic growth. Adventitious buds were elongated and rooted in vitro before being transferred to the greenhouse where the transformed whole plants grew normally. Transformants tested positive for opine production and transformation was further confirmed by Southern blot analysis with larch genomic DNAs isolated from both proliferated calli and needle tissue of transgenic plants.

Impacts of elevated atmospheric CO2 on forest trees and forest ecosystems: knowledge gaps

Atmospheric CO(2) is rising rapidly, and options for slowing the CO(2) rise are politically charged as they largely require reductions in industrial CO(2) emissions for most developed countries. As forests cover some 43% of the Earths surface, account for some 70% of terrestrial net primary production (NPP), and are being bartered for carbon mitigation, it is critically important that we continue to reduce the uncertainties about the impacts of elevated atmospheric CO(2) on forest tree growth, productivity, and forest ecosystem function. In this paper, I review knowledge gaps and research needs on the effects of elevated atmospheric CO(2) on forest above- and below-ground growth and productivity, carbon sequestration, nutrient cycling, water relations, wood quality, phenology, community dynamics and biodiversity, antioxidants and stress tolerance, interactions with air pollutants, heterotrophic interactions, and ecosystem functioning. Finally, I discuss research needs regarding modeling of the impacts of elevated atmospheric CO(2) on forests.Even though there has been a tremendous amount of research done with elevated CO(2) and forest trees, it remains difficult to predict future forest growth and productivity under elevated atmospheric CO(2). Likewise, it is not easy to predict how forest ecosystem processes will respond to enriched CO(2). The more we study the impacts of increasing CO(2), the more we realize that tree and forest responses are yet largely uncertain due to differences in responsiveness by species, genotype, and functional group, and the complex interactions of elevated atmospheric CO(2) with soil fertility, drought, pests, and co-occurring atmospheric pollutants such as nitrogen deposition and O(3). Furthermore, it is impossible to predict ecosystem-level responses based on short-term studies of young trees grown without interacting stresses and in small spaces without the element of competition. Long-term studies using free-air CO(2) enrichment (FACE) technologies or forest stands around natural CO(2) vents are needed to increase the knowledge base on forest ecosystem responses to elevated atmospheric CO(2). In addition, new experimental protocols need to continue to be developed that will allow for mature trees to be examined in natural ecosystems. These studies should be closely linked to modeling efforts so that the inference capacity from these expensive and long-term studies can be maximized.

Isoprene emission from terrestrial ecosystems in response to global change: minding the gap between models and observations

Coupled surface–atmosphere models are being used with increased frequency to make predictions of tropospheric chemistry on a ‘future’ earth characterized by a warmer climate and elevated atmospheric CO2 concentration. One of the key inputs to these models is the emission of isoprene from forest ecosystems. Most models in current use rely on a scheme by which global change is coupled to changes in terrestrial net primary productivity (NPP) which, in turn, is coupled to changes in the magnitude of isoprene emissions. In this study, we conducted measurements of isoprene emissions at three prominent global change experiments in the United States. Our results showed that growth in an atmosphere of elevated CO2 inhibited the emission of isoprene at levels that completely compensate for possible increases in emission due to increases in aboveground NPP. Exposure to a prolonged drought caused leaves to increase their isoprene emissions despite reductions in photosynthesis, and presumably NPP. Thus, the current generation of models intended to predict the response of isoprene emission to future global change probably contain large errors. A framework is offered as a foundation for constructing new isoprene emission models based on the responses of leaf biochemistry to future climate change and elevated atmospheric CO2 concentrations.

Respiratory Oxygen Uptake Is Not Decreased by an Instantaneous Elevation of [CO2], But Is Increased with Long-Term Growth in the Field at Elevated [CO2]

Averaged across many previous investigations, doubling the CO2 concentration ([CO2]) has frequently been reported to cause an instantaneous reduction of leaf dark respiration measured as CO2 efflux. No known mechanism accounts for this effect, and four recent studies have shown that the measurement of respiratory CO2 efflux is prone to experimental artifacts that could account for the reported response. Here, these artifacts are avoided by use of a high-resolution dual channel oxygen analyzer within an open gas exchange system to measure respiratory O2 uptake in normal air. Leaf O2 uptake was determined in response to instantaneous elevation of [CO2] in nine contrasting species and to long-term elevation in seven species from four field experiments. Over six hundred separate measurements of respiration failed to reveal any decrease in respiratory O2 uptake with an instantaneous increase in [CO2]. Respiration was found insensitive not only to doubling [CO2], but also to a 5-fold increase and to decrease to zero. Using a wide range of species and conditions, we confirm earlier reports that inhibition of respiration by instantaneous elevation of [CO2] is likely an experimental artifact. Instead of the expected decrease in respiration per unit leaf area in response to long-term growth in the field at elevated [CO2], there was a significant increase of 11% and 7% on an area and mass basis, respectively, averaged across all experiments. The findings suggest that leaf dark respiration will increase not decrease as atmospheric [CO2] rises.

Effects of elevated CO2 and O3 on aspen clones varying in O3 sensitivity: can CO2 ameliorate the harmful effects of O3?

To determine whether elevated CO2 reduces or exacerbates the detrimental effects of O3 on aspen (Populus tremuloides Michx.). aspen clones 216 and 271 (O3 tolerant), and 259 (O3 sensitive) were exposed to ambient levels of CO2 and O3 or elevated levels of CO2, O3, or CO2 + O3 in the FACTS II (Aspen FACE) experiment, and physiological and molecular responses were measured and compared. Clone 259. the most O3-sensitive clone, showed the greatest amount of visible foliar symptoms as well as significant decreases in chlorophyll, carotenoid, starch, and ribulose-1, 5-bisphosphate carboxylase/oxygenase (Rubisco) concentrations and transcription levels for the Rubisco small subunit. Generally, the constitutive (basic) transcript levels for phenylalanine ammonialyase (PAL) and chalcone synthase (CHS) and the average antioxidant activities were lower for the ozone sensitive clone 259 as compared to the more tolerant 216 and 271 clones. A significant decrease in chlorophyll a, b and total (a + b) concentrations in CO2, O3, and CO2 + O3 plants was observed for all clones. Carotenoid concentrations were also significantly lower in all clones; however. CHS transcript levels were not significantly affected, suggesting a possible degradation of carotenoid pigments in O3-stressed plants. Antioxidant activities and PAL and 1-aminocyclopropane-l-carboxylic acid (ACC)-oxidase transcript levels showed a general increase in all O3 treated clones, while remaining low in CO2 and CO2 + O3 plants (although not all differences were significant). Our results suggest that the ascorbate-glutathione and phenylpropanoid pathways were activated under ozone stress and suppressed during exposure to elevated CO2. Although CO2 + O2 treatment resulted in a slight reduction of O3-induced leaf injury, it did not appear to ameliorate all of the harmful affects of O3 and, in fact. may have contributed to an increase in chloroplast damage in all three aspen clones.

Impacts of global change on diseases of agricultural crops and forest trees

The fourth assessment report of the Intergovernmental Panel on Climate Change projects rising levels of greenhouse gas and global temperature. The well-known dependence of plant diseases on weather has long been exploited for predicting epidemics and to time applications of control measures for tactical disease management. Fingerprints of inter-annual climatic variation on pathogens have recently been shown in literature linking pathogen abundance to atmospheric composition. Past reviews have dealt with impacts of changing atmospheric composition and climate on diseases, regional or country-wide assessments of climate change impacts and impacts on specific disease/pathogen or pathogen groups. All agree on paucity of knowledge prompting a need to generate new empirical data on host‐pathogen biology under a changing climate. Focused on experimental research, the purpose of this review is to summarize published and unpublished studies on plant pathogens and diseases in free-air CO2 enrichment (FACE) facilities and open top chambers and other current non-FACE research to offer a summary of future research needs and opportunities. Critical review of recent literature on the influence of elevated CO2 and O3 on agriculture and forestry species forms a major part of the treatise. Summaries of unpublished or ongoing experimental research on plant pathogens from FACE studies are included as a catalogue of work in this neglected area. The catalogue and knowledge gaps are intended as a resource for workers initiating research in this area as well as the general scientific community grappling with the design and scope of next generation of FACE facilities.

Isoprene emission rates under elevated CO2 and O3 in two field-grown aspen clones differing in their sensitivity to O3.

Isoprene is the most important nonmethane hydrocarbon emitted by plants. The role of isoprene in the plant is not entirely understood but there is evidence that it might have a protective role against different oxidative stresses originating from heat shock and/or exposure to ozone (O(3)). Thus, plants under stress conditions might benefit by constitutively high or by higher stress-induced isoprene emission rates. In this study, measurements are presented of isoprene emission from aspen (Populus tremuloides) trees grown in the field for several years under elevated CO(2) and O(3). Two aspen clones were investigated: the O(3)-tolerant 271 and the O(3)-sensitive 42E. Isoprene emission decreased significantly both under elevated CO(2) and under elevated O(3) in the O(3)-sensitive clone, but only slightly in the O(3)-tolerant clone. This study demonstrates that long-term-adapted plants are not able to respond to O(3) stress by increasing their isoprene emission rates. However, O(3)-tolerant clones have the capacity to maintain higher amounts of isoprene emission. It is suggested that tolerance to O(3) is explained by a combination of different factors; while the reduction of O(3) uptake is likely to be the most important, the capacity to maintain higher amounts of isoprene is an important factor in strengthening this character.