Uncovering hidden genetic variation in photosynthesis of field-grown maize under ozone pollution.

Ozone is the most damaging air pollutant to crops, currently reducing Midwest US maize production by up to 10%, yet there has been very little effort to adapt germplasm for ozone tolerance. Ozone enters plants through stomata, reacts to form reactive oxygen species in the apoplast and ultimately decreases photosynthetic C gain. In this study, 10 diverse inbred parents were crossed in a half-diallel design to create 45 F1 hybrids, which were tested for ozone response in the field using free air concentration enrichment (FACE). Ozone stress increased the heritability of photosynthetic traits and altered genetic correlations among traits. Hybrids from parents Hp301 and NC338 showed greater sensitivity to ozone stress, and disrupted relationships among photosynthetic traits. The physiological responses underlying sensitivity to ozone differed in hybrids from the two parents, suggesting multiple mechanisms of response to oxidative stress. FACE technology was essential to this evaluation because genetic variation in photosynthesis under elevated ozone was not predictable based on performance at ambient ozone. These findings suggest that selection under elevated ozone is needed to identify deleterious alleles in the world's largest commodity crop.

Diversity in stomatal function is integral to modelling plant carbon and water fluxes.

Stomatal pores on leaf surfaces respond to environmental and physiological signals to regulate leaf gas exchange. Mathematical models can predict stomatal conductance (g s), with one parameter (m or g l) reflecting the sensitivity of g s to the photosynthetic rate (A), atmospheric carbon dioxide concentration and atmospheric humidity, and a second parameter (g 0) representing the minimum g s. Such models are solved iteratively with a photosynthesis model to form the core of many models of crop or ecosystem carbon and water fluxes. For three decades, g s models have frequently been used assuming fixed parameter values for m or g 1 and g 0 across species and major plant functional types. This study of temperate tree species reveals significant interspecific variation in stomatal function. Applying species-specific parameterizations substantially reduced error in model predictions of g s by 34 to 64% and A by 52 to 60% and resulted in significant correlation between modelled and measured values. This work challenges the long-held assumption of fixed parameter values and, in doing so, suggests an approach for reducing modelling error across a wide range of ecological and agricultural applications.

A physiological and biophysical model of coppice willow (Salix spp.) production yields for the contiguous USA in current and future climate scenarios.

High-performance computing has facilitated development of biomass production models that capture the key mechanisms underlying production at high spatial and temporal resolution. Direct responses to increasing [CO2 ] and temperature are important to long-lived emerging woody bioenergy crops. Fast-growing willow (Salix spp.) within short rotation coppice (SRC) has considerable potential as a renewable biomass source, but performance over wider environmental conditions and under climate change is uncertain. We extended the bioenergy crop modeling platform, BioCro, to SRC willow by adding coppicing and C3 photosynthesis subroutines, and modifying subroutines for perennation, allocation, morphology, phenology and development. Parameterization with measurements of leaf photosynthesis, allocation and phenology gave agreement of modeled with measured yield across 23 sites in Europe and North America. Predictions for the continental USA suggest yields of ≥17 Mg ha(-1) year(-1) in a 4 year rotation. Rising temperature decreased predicted yields, an effect partially ameliorated by rising [CO2 ]. This model, based on over 100 equations describing the physiological and biophysical mechanisms underlying production, provides a new framework for utilizing mechanism of plant responses to the environment, including future climates. As an open-source tool, it is made available here as a community resource for further application, improvement and adaptation.

C3 and C4 plant responses to increased temperatures and altered monsoonal precipitation in a cool desert on the Colorado Plateau, USA.

Dryland ecosystems represent >40% of the terrestrial landscape and support over two billion people; consequently, it is vital to understand how drylands will respond to climatic change. However, while arid and semiarid ecosystems commonly experience extremely hot and dry conditions, our understanding of how further temperature increases or altered precipitation will affect dryland plant communities remains poor. To address this question, we assessed plant physiology and growth at a long-term (7-year) climate experiment on the Colorado Plateau, USA, where the community is a mix of shallow-rooted C3 and C4 grasses and deep-rooted C4 shrubs. The experiment maintained elevated-temperature treatments (+2 or +4 °C) in combination with altered summer monsoonal precipitation (+small frequent precipitation events or +large infrequent events). Increased temperature negatively affected photosynthesis and growth of the C3 and C4 grasses, but effects varied in their timing: +4 °C treatments negatively affected the C3 grass early in the growing season of both years, while the negative effects of temperature on the C4 grass were seen in the +2 and +4 °C treatments, but only during the late growing season of the drier year. Increased summer precipitation did not affect photosynthesis or biomass for any species, either in the year the precipitation was applied or the following year. Although previous research suggests dryland plants, and C4 grasses in particular, may respond positively to elevated temperature, our findings from a cool desert show marked declines in C3 and C4 photosynthesis and growth, with temperature effects dependent on the degree of warming and growing-season precipitation.

The effect of heat waves, elevated [CO2 ] and low soil water availability on northern red oak (Quercus rubra L.) seedlings.

The frequency and intensity of heat waves are predicted to increase. This study investigates whether heat waves would have the same impact as a constant increase in temperature with the same heat sum, and whether there would be any interactive effects of elevated [CO2 ] and soil moisture content. We grew Quercus rubra seedlings in treatment chambers maintained at either ambient or elevated [CO2 ] (380 or 700 µmol CO2 mol(-1) ) with temperature treatments of ambient, ambient +3 °C, moderate heat wave (+6 °C every other week) or severe heat wave (+12 °C every fourth week) temperatures. Averaged over a 4-week period, and the entire growing season, the three elevated temperature treatments had the same average temperature and heat sum. Half the seedlings were watered to a soil water content near field capacity, half to about 50% of this value. Foliar gas exchange measurements were performed morning and afternoon (9:00 and 15:00 hours) before, during and after an applied heat wave in August 2010. Biomass accumulation was measured after five heat wave cycles. Under ambient [CO2 ] and well-watered conditions, biomass accumulation was highest in the +3 °C treatment, intermediate in the +6 °C heat wave and lowest in the +12 °C heat wave treatment. This response was mitigated by elevated [CO2 ]. Low soil moisture significantly decreased net photosynthesis (Anet ) and biomass in all [CO2 ] and temperature treatments. The +12 °C heat wave reduced afternoon Anet by 23% in ambient [CO2 ]. Although this reduction was relatively greater under elevated [CO2 ], Anet values during this heat wave were still 34% higher than under ambient [CO2 ]. We concluded that heat waves affected biomass growth differently than the same amount of heat applied uniformly over the growing season, and that the plant response to heat waves also depends on [CO2 ] and soil moisture conditions.

The effect of induced heat waves on Pinus taeda and Quercus rubra seedlings in ambient and elevated CO2 atmospheres.

Here, we investigated the effect of different heat-wave intensities applied at two atmospheric CO2 concentrations ([CO2]) on seedlings of two tree species, loblolly pine (Pinus taeda) and northern red oak (Quercus rubra). Seedlings were assigned to treatment combinations of two levels of [CO2] (380 or 700 µmol mol(-1)) and four levels of air temperature (ambient, ambient +3°C, or 7-d heat waves consisting of a biweekly +6°C heat wave, or a monthly +12°C heat wave). Treatments were maintained throughout the growing season, thus receiving equal heat sums. We measured gas exchange and fluorescence parameters before, during and after a mid-summer heat wave. The +12°C heat wave, significantly reduced net photosynthesis (Anet) in both species and [CO2] treatments but this effect was diminished in elevated [CO2]. The decrease in Anet was accompanied by a decrease in Fv'/Fm' in P. taeda and ΦPSII in Q. rubra. Our findings suggest that, if soil moisture is adequate, trees will experience negative effects in photosynthetic performance only with the occurrence of extreme heat waves. As elevated [CO2] diminished these negative effects, the future climate may not be as detrimental to plant communities as previously assumed.

Effects of predicted future and current atmospheric temperature and [CO2] and high and low soil moisture on gas exchange and growth of Pinus taeda seedlings at cool and warm sites in the species range.

Predicted future changes in air temperature and atmospheric CO(2) concentration ([CO(2)]), coupled with altered precipitation, are expected to substantially affect tree growth. Effects on growth may vary considerably across a species range, as temperatures vary from sub-optimal to supra-optimal for growth. We performed an experiment simultaneously at two locations in the current range of loblolly pine, a cool site and a warm site, to examine the effect of future climate conditions on growth of loblolly pine seedlings in contrasting regions of the species range. At both sites 1-year-old loblolly pine seedlings were grown in current (local ambient temperature and [CO(2)]) and predicted future atmospheric conditions (ambient +2 °C temperature and 700 µmol mol(-1) [CO(2)]). Additionally, high and low soil moisture treatments were applied within each atmospheric treatment at each site by altering the amount of water provided to the seedlings. Averaged across water treatments, photosynthesis (A(net)) was 31% greater at the cool site and 34% greater at the warm site in elevated temperature and [CO(2)] compared with ambient temperature. Biomass accumulation was also stimulated by 38% at the cool site and by 24% at the warm site in that treatment. These results suggest that a temperature increase of 2 °C coupled with an increase in [CO(2)] (predicted future climate) will create conditions favorable for growth of this species. Reduced soil moisture decreased growth in both current and predicted atmospheric conditions. Biomass accumulation and A(net) were reduced by ∼39 and 17%, respectively, in the low water treatment. These results suggest that any benefit of future atmospheric conditions may be negated if soil moisture is reduced by altered precipitation patterns.

Higher growth temperatures decreased net carbon assimilation and biomass accumulation of northern red oak seedlings near the southern limit of the species range.

If an increase in temperature will limit the growth of a species, it will be in the warmest portion of the species distribution. Therefore, in this study we examined the effects of elevated temperature on net carbon assimilation and biomass production of northern red oak (Quercus rubra L.) seedlings grown near the southern limit of the species distribution. Seedlings were grown in chambers in elevated CO(2) (700 µmol mol(-1)) at three temperature conditions, ambient (tracking diurnal and seasonal variation in outdoor temperature), ambient +3 °C and ambient +6 °C, which produced mean growing season temperatures of 23, 26 and 29 °C, respectively. A group of seedlings was also grown in ambient [CO(2)] and ambient temperature as a check of the growth response to elevated [CO(2)]. Net photosynthesis and leaf respiration, photosynthetic capacity (V(cmax), J(max) and triose phosphate utilization (TPU)) and chlorophyll fluorescence, as well as seedling height, diameter and biomass, were measured during one growing season. Higher growth temperatures reduced net photosynthesis, increased respiration and reduced height, diameter and biomass production. Maximum net photosynthesis at saturating [CO(2)] and maximum rate of electron transport (J(max)) were lowest throughout the growing season in seedlings grown in the highest temperature regime. These parameters were also lower in June, but not in July or September, in seedlings grown at +3 °C above ambient, compared with those grown in ambient temperature, indicating no impairment of photosynthetic capacity with a moderate increase in air temperature. An unusual and potentially important observation was that foliar respiration did not acclimate to growth temperature, resulting in substantially higher leaf respiration at the higher growth temperatures. Lower net carbon assimilation was correlated with lower growth at higher temperatures. Total biomass at the end of the growing season decreased in direct proportion to the increase in growth temperature, declining by 6% per 1 °C increase in mean growing season temperature. Our observations suggest that increases in air temperature above current ambient conditions will be detrimental to Q. rubra seedlings growing near the southern limit of the species range.

Close coupling of whole-plant respiration to net photosynthesis and carbohydrates.

We studied the effect of changes in net photosynthesis (A(net)) on respiration, soluble sugars and carbohydrates in Populus deltoides Bartr. ex Marsh. saplings under controlled environmental conditions by making daily measurements of leaf respiration (R(d)), stem CO(2) efflux and root CO(2) efflux at a constant temperature in growth chambers. After a pretreatment period, one of three treatments was applied for 5 to 7 days: (1) increased atmospheric CO(2) concentration; (2) decreased photoperiod and photosynthetic photon flux (PPF); or (3) continuous darkness. Increased A(net) in response to elevated CO(2) concentration resulted in a sustained increase in whole-plant respiration, with R(d) increasing 46%, stem CO(2) efflux increasing 130% and root CO(2) efflux increasing 16%. Elevated CO(2) concentration also caused a significant increase in leaf soluble sugars. Decreasing photoperiod and PPF or complete darkness caused a rapid decrease in respiration throughout the saplings. In the low light treatment, R(d) decreased 40%, stem CO(2) efflux decreased 78%, root CO(2) efflux decreased 74% and significant decreases in leaf and root soluble sugar and leaf nonstructural carbohydrate concentrations were observed. Continuous darkness resulted in a 70% decrease in R(d), a 65% decrease in stem CO(2) efflux, a 73% decrease in root CO(2) efflux and significant decreases in leaf and root soluble sugar and root carbohydrate concentrations. In all treatments, changes in respiration rates in all tissues occurred within hours of treatment application. In addition, a diurnal pattern in root CO(2) efflux was observed throughout the experiment under constant environmental conditions. The observed rapid changes in whole-plant respiration following treatment application and the diurnal patterns in root CO(2) efflux suggest that growth and maintenance respiration in the saplings was strongly dependent on newly acquired carbohydrates.