Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests.

Three young northern temperate forest communities in the north-central United States were exposed to factorial combinations of elevated carbon dioxide (CO2 ) and tropospheric ozone (O3 ) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO2 enhanced ecosystem C content by 11%, whereas elevated O3 decreased ecosystem C content by 9%.

The transcriptome of Populus in elevated CO reveals increased anthocyanin biosynthesis during delayed autumnal senescence.

*The delay in autumnal senescence that has occurred in recent decades has been linked to rising temperatures. Here, we suggest that increasing atmospheric CO2 may partly account for delayed autumnal senescence and for the first time, through transcriptome analysis, identify gene expression changes associated with this delay. *Using a plantation of Populus x euramericana grown in elevated [CO2] (e[CO2]) with free-air CO2 enrichment (FACE) technology, we investigated the molecular and biochemical basis of this response.

Advances in understanding ozone impact on forest trees: messages from novel phytotron and free-air fumigation studies.

Recent evidence from novel phytotron and free-air ozone (O3) fumigation experiments in Europe and America on forest tree species is highlighted in relation to previous chamber studies. Differences in O3 sensitivity between pioneer and climax species are examined and viewed for trees growing at the harsh alpine timberline ecotone. As O3 apparently counteracts positive effects of elevated CO2 and mitigates productivity increases, response is governed by genotype, competitors, and ontogeny rather than species per se.

Challenges in elevated CO2 experiments on forests.

Current forest Free Air CO(2) Enrichment (FACE) experiments are reaching completion. Therefore, it is time to define the scientific goals and priorities of future experimental facilities. In this opinion article, we discuss the following three overarching issues (i) What are the most urgent scientific questions and how can they be addressed? (ii) What forest ecosystems should be investigated? (iii) Which other climate change factors should be coupled with elevated CO(2) concentrations in future experiments to better predict the effects of climate change? Plantations and natural forests can have conflicting purposes for high productivity and environmental protection.

Differential response of aspen and birch trees to heat stress under elevated carbon dioxide.

The effect of high temperature on photosynthesis of isoprene-emitting (aspen) and non-isoprene-emitting (birch) trees were measured under elevated CO(2) and ambient conditions. Aspen trees tolerated heat better than birch trees and elevated CO(2) protected photosynthesis of both species against moderate heat stress. Elevated CO(2) increased carboxylation capacity, photosynthetic electron transport capacity, and triose phosphate use in both birch and aspen trees.

Will photosynthetic capacity of aspen trees acclimate after long-term exposure to elevated CO2 and O3?

Photosynthetic acclimation under elevated carbon dioxide (CO(2)) and/or ozone (O(3)) has been the topic of discussion in many papers recently. We examined whether or not aspen plants grown under elevated CO(2) and/or O(3) will acclimate after 11 years of exposure at the Aspen Face site in Rhinelander, WI, USA. We studied diurnal patterns of instantaneous photosynthetic measurements as well as A/C(i) measurements monthly during the 2004-2008 growing seasons.

Gene expression responses of paper birch (Betula papyrifera) to elevated CO2 and O3 during leaf maturation and senescence.

Gene expression responses of paper birch (Betula papyrifera) leaves to elevated concentrations of CO(2) and O(3) were studied with microarray analyses from three time points during the summer of 2004 at Aspen FACE. Microarray data were analyzed with clustering techniques, self-organizing maps, K-means clustering and Sammon's mappings, to detect similar gene expression patterns within sampling times and treatments. Most of the alterations in gene expression were caused by O(3), alone or in combination with CO(2).

DNA damage in Populus tremuloides clones exposed to elevated O3.

The effects of elevated concentrations of atmospheric tropospheric ozone (O(3)) on DNA damage in five trembling aspen (Populus tremuloides Michx.) clones growing in a free-air enrichment experiment in the presence and absence of elevated concentrations of carbon dioxide (CO(2)) were examined. Growing season mean hourly O(3) concentrations were 36.

Elevated CO2 response of photosynthesis depends on ozone concentration in aspen.

The effect of elevated CO(2) and O(3) on apparent quantum yield (varphi), maximum photosynthesis (P(max)), carboxylation efficiency (V(cmax)) and electron transport capacity (J(max)) at different canopy locations was studied in two aspen (Populus tremuloides) clones of contrasting O(3) tolerance. Local light climate at every leaf was characterized as fraction of above-canopy photosynthetic photon flux density (%PPFD). Elevated CO(2) alone did not affect varphi or P(max), and increased J(max) in the O(3)-sensitive, but not in the O(3)-tolerant clone.

Response and potential of agroforestry crops under global change.

The use of agroforestry crops is a promising tool for reducing atmospheric carbon dioxide concentration through fossil fuel substitution. In particular, plantations characterised by high yields such as short rotation forestry (SRF) are becoming popular worldwide for biomass production and their role acknowledged in the Kyoto Protocol. While their contribution to climate change mitigation is being investigated, the impact of climate change itself on growth and productivity of these plantations needs particular attention, since their management might need to be modified accordingly.

Diurnal changes in photosynthetic parameters of Populus tremuloides, modulated by elevated concentrations of CO2 and/or O3 and daily climatic variation.

The diurnal changes in light-saturated photosynthesis (Pn) under elevated CO(2) and/or O(3) in relation to stomatal conductance (g(s)), water potential, intercellular [CO(2)], leaf temperature and vapour-pressure difference between leaf and air (VPD(L)) were studied at the Aspen FACE site. Two aspen (Populus tremuloides Michx.) clones differing in their sensitivity to ozone were measured.

Analysis of a Farquhar-von Caemmerer-Berry leaf-level photosynthetic rate model for Populus tremuloides in the context of modeling and measurement limitations.

The balance of mechanistic detail with mathematical simplicity contributes to the broad use of the Farquhar, von Caemmerer and Berry (FvCB) photosynthetic rate model. Here the FvCB model was coupled with a stomatal conductance model to form an [A,g(s)] model, and parameterized for mature Populus tremuloides leaves under varying CO(2) and temperature levels. Data were selected to be within typical forest light, CO(2) and temperature ranges, reducing artifacts associated with data collected at extreme values.

Leaf size and surface characteristics of Betula papyrifera exposed to elevated CO2 and O3.

Betula papyrifera trees were exposed to elevated concentrations of CO(2) (1.4 x ambient), O(3) (1.2 x ambient) or CO(2) + O(3) at the Aspen Free-air CO(2) Enrichment Experiment.

Spring leaf flush in aspen (Populus tremuloides) clones is altered by long-term growth at elevated carbon dioxide and elevated ozone concentration.

Early spring leaf out is important to the success of deciduous trees competing for light and space in dense forest plantation canopies. In this study, we investigated spring leaf flush and how long-term growth at elevated carbon dioxide concentration ([CO(2)]) and elevated ozone concentration ([O(3)]) altered leaf area index development in a closed Populus tremuloides (aspen) canopy. This work was done at the Aspen FACE experiment where aspen clones have been grown since 1997 in conditions simulating the [CO(2)] and [O(3)] predicted for approximately 2050.

Transcriptomic comparison in the leaves of two aspen genotypes having similar carbon assimilation rates but different partitioning patterns under elevated [CO2].

This study compared the leaf transcription profiles, physiological characteristics and primary metabolites of two Populus tremuloides genotypes (clones 216 and 271) known to differ in their responses to long-term elevated [CO2] (e[CO2]) at the Aspen free-air CO2 enrichment site near Rhinelander, WI, USA. The physiological responses of these clones were similar in terms of photosynthesis, stomatal conductance and leaf area index under e[CO2], yet very different in terms of growth enhancement (0-10% in clone 216; 40-50% in clone 271). Although few genes responded to long-term exposure to e[CO2], the transcriptional activity of leaf e[CO2]-responsive genes was distinctly different between the clones, differentially impacting multiple pathways during both early and late growing seasons.

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.

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.

Effects of decadal exposure to interacting elevated CO2 and/or O3 on paper birch (Betula papyrifera) reproduction.

We studied the effects of long-term exposure (nine years) of birch (Betula papyrifera) trees to elevated CO(2) and/or O(3) on reproduction and seedling development at the Aspen FACE (Free-Air Carbon Dioxide Enrichment) site in Rhinelander, WI. We found that elevated CO(2) increased both the number of trees that flowered and the quantity of flowers (260% increase in male flower production), increased seed weight, germination rate, and seedling vigor. Elevated O(3) also increased flowering but decreased seed weight and germination rate.

Wood properties of trembling aspen and paper birch after 5 years of exposure to elevated concentrations of CO(2) and O(3).

We investigated the interactive effects of elevated concentrations of carbon dioxide ([CO(2)]) and ozone ([O(3)]) on radial growth, wood chemistry and structure of five 5-year-old trembling aspen (Populus tremuloides Michx.) clones and the wood chemistry of paper birch (Betula papyrifera Marsh.).

Carbon gain and bud physiology in Populus tremuloides and Betula papyrifera grown under long-term exposure to elevated concentrations of CO2 and O3.

Paper birch (Betula papyrifera Marsh.) and three trembling aspen clones (Populus tremuloides Michx.) were studied to determine if alterations in carbon gain in response to an elevated concentration of CO(2) ([CO(2)]) or O(3) ([O(3)]) or a combination of both affected bud size and carbohydrate composition in autumn, and early leaf development in the following spring.

Air quality in natural areas: interface between the public, science and regulation.

Natural areas are important interfaces between air quality, the public, science and regulation. In the United States and Canada, national parks received over 315million visits during 2004. Many natural areas have been experiencing decreased visibility, increased ozone (O(3)) levels and elevated nitrogen deposition.

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.

Impacts of elevated atmospheric CO2 and O3 on paper birch (Betula papyrifera): reproductive fitness.

Atmospheric CO2 and tropospheric O3 are rising in many regions of the world. Little is known about how these two commonly co-occurring gases will affect reproductive fitness of important forest tree species. Here, we report on the long-term effects of CO2 and O3 for paper birch seedlings exposed for nearly their entire life history at the Aspen FACE (Free Air Carbon Dioxide Enrichment) site in Rhinelander, WI.

Impacts of air pollution and climate change on forest ecosystems--emerging research needs.

Outcomes from the 22nd meeting for Specialists in Air Pollution Effects on Forest Ecosystems "Forests under Anthropogenic Pressure--Effects of Air Pollution, Climate Change and Urban Development", September 10-16, 2006, Riverside, CA, are summarized. Tropospheric or ground-level ozone (O3) is still the phytotoxic air pollutant of major interest. Challenging issues are how to make O3 standards or critical levels more biologically based and at the same time practical for wide use; quantification of plant detoxification processes in flux modeling; inclusion of multiple environmental stresses in critical load determinations; new concept development for nitrogen saturation; interactions between air pollution, climate, and forest pests; effects of forest fire on air quality; the capacity of forests to sequester carbon under changing climatic conditions and coexposure to elevated levels of air pollutants; enhanced linkage between molecular biology, biochemistry, physiology, and morphological traits.

Micropropagation and germplasm conservation of Central Park Splendor Chinese elm (Ulmus parvifolia Jacq. 'A/Ross Central Park') trees.

Micropropagation offers opportunities to propagate, preserve and ship tree germplasm. It also reduces the risk of moving pathogens and insects with the germplasm due to built-in pathogen detection capabilities of aseptic cultures. For the past few decades, our laboratory has been involved in a project to preserve and restore a large, cold hardy, and historically important Chinese elm (Ulmus parvifolia Jacq.