Short- and long-term warming events on photosynthetic physiology, growth, and yields of field grown crops.

Global temperatures are rising from increasing concentrations of greenhouse gases in the atmosphere associated with anthropogenic activities. Global warming includes a warmer shift in mean temperatures as well as increases in the probability of extreme heating events, termed heat waves. Despite the ability of plants to cope with temporal variations in temperature, global warming is increasingly presenting challenges to agroecosystems. The impact of warming on crop species has direct consequences on food security, therefore understanding impacts and opportunities to adapt crops to global warming necessitates experimentation that allows for modification of growth environments to represent global warming scenarios. Published studies addressing crop responses to warming are extensive, however, in-field studies where growth temperature is manipulated to mimic global warming are limited. Here, we provide an overview of in-field heating techniques employed to understand crop responses to warmer growth environments. We then focus on key results associated with season-long warming, as expected with rising global mean temperatures, and with heat waves, as a consequence of increasing temperature variability and rising global mean temperatures. We then discuss the role of rising temperatures on atmospheric water vapor pressure deficit and potential implications for crop photosynthesis and productivity. Finally, we review strategies by which crop photosynthetic processes might be optimized to adapt crops to the increasing temperatures and frequencies of heat waves. Key findings from this review are that higher temperatures consistently reduce photosynthesis and yields of crops even as atmospheric carbon dioxide increases, yet potential strategies to minimize losses from high-temperature exist.

Evaluation of the effects of elevated CO2 concentrations on the growth of cassava storage roots by destructive harvests and ground penetrating radar scanning approaches.

Cassava (Manihot esculenta Crantz) production will need to be improved to meet future food demands in Sub-Saharan Africa. The selection of high-yielding cassava cultivars requires a better understanding of storage root development. Additionally, since future production will happen under increasing atmospheric CO2 concentrations ([CO2 ]), cultivar selection should include responsiveness to elevated [CO2 ]. Five farmer-preferred African cassava cultivars were grown for three and a half months in a Free Air CO2 Enrichment experiment in central Illinois. Compared to ambient [CO2 ] (~400 ppm), cassava storage roots grown under elevated [CO2 ] (~600 ppm) had a higher biomass with some cultivars having lower storage root water content. The elevated [CO2 ] stimulation in storage root biomass ranged from 33% to 86% across the five cultivars tested documenting the importance of this trait in developing new cultivars. In addition to the destructive harvests to obtain storage root parameters, we explored ground penetrating radar as a nondestructive method to determine storage root growth across the growing season.

Field-grown ictB tobacco transformants show no difference in photosynthetic efficiency for biomass relative to the wild type.

In this study, four tobacco transformants overexpressing the inorganic carbon transporter B gene (ictB) were screened for photosynthetic performance relative to the wild type (WT) in field-based conditions. The WT and transgenic tobacco plants were evaluated for photosynthetic performance to determine the maximum rate of carboxylation (Vc, max), maximum rate of electron transport (Jmax), the photosynthetic compensation point (Γ*), quantum yield of PSII (ΦPSII), and mesophyll conductance (gm). Additionally, all plants were harvested to compare differences in above-ground biomass. Overall, transformants did not perform better than the WT on photosynthesis-, biomass-, and leaf composition-related traits. This is in contrast to previous studies that have suggested significant increases in photosynthesis and yield with the overexpression of ictB, although not widely evaluated under field conditions. These findings suggest that the benefit of ictB is not universal and may only be seen under certain growth conditions. While there is certainly still potential benefit to utilizing ictB in the future, further effort must be concentrated on understanding the underlying function of the gene and in which environmental conditions it offers the greatest benefit to crop performance. As it stands at present, it is possible that ictB overexpression may be largely favorable in controlled environments, such as greenhouses.

High sink strength prevents photosynthetic down-regulation in cassava grown at elevated CO2 concentration.

Cassava has the potential to alleviate food insecurity in many tropical regions, yet few breeding efforts to increase yield have been made. Improved photosynthetic efficiency in cassava has the potential to increase yields, but cassava roots must have sufficient sink strength to prevent carbohydrates from accumulating in leaf tissue and suppressing photosynthesis. Here, we grew eight farmer-preferred African cassava cultivars under free-air CO2 enrichment (FACE) to evaluate the sink strength of cassava roots when photosynthesis increases due to elevated CO2 concentrations ([CO2]). Relative to the ambient treatments, elevated [CO2] treatments increased fresh (+27%) and dry (+37%) root biomass, which was driven by an increase in photosynthesis (+31%) and the absence of photosynthetic down-regulation over the growing season. Moreover, intrinsic water use efficiency improved under elevated [CO2] conditions, while leaf protein content and leaf and root cyanide concentrations were not affected. Overall, these results suggest that higher cassava yields can be expected as atmospheric [CO2] increases over the coming decades. However, there were cultivar differences in the partitioning of resources to roots versus above-grown biomass; thus, the particular responses of each cultivar must be considered when selecting candidates for improvement.

Canopy warming accelerates development in soybean and maize, offsetting the delay in soybean reproductive development by elevated CO2 concentrations.

Increases in atmospheric CO2 concentrations ([CO2 ]) and surface temperature are known to individually have effects on crop development and yield, but their interactive effects have not been adequately investigated under field conditions. We evaluated the impacts of elevated [CO2 ] with and without canopy warming as a function of development in soybean and maize using infrared heating arrays nested within free air CO2 enrichment plots over three growing seasons. Vegetative development accelerated in soybean with temperature plus elevated [CO2 ] resulting in higher node number. Reproductive development was delayed in soybean under elevated [CO2 ], but warming mitigated this delay. In maize, both vegetative and reproductive developments were accelerated by warming, whereas elevated [CO2 ] had no apparent effect on development. Treatment-induced changes in the leaf carbohydrates, dark respiration rate, morphological parameters, and environmental conditions accompanied the changes in plant development. We used two thermal models to investigate their ability to predict the observed development under warming and elevated [CO2 ]. Whereas the growing degree day model underestimated the thermal threshold to reach each developmental stage, the alternative process-based model used (ß function) was able to predict crop development under climate change conditions.

The Role of Sink Strength and Nitrogen Availability in the Down-Regulation of Photosynthetic Capacity in Field-Grown Nicotiana tabacum L. at Elevated CO2 Concentration.

Down-regulation of photosynthesis is among the most common responses observed in C3 plants grown under elevated atmospheric CO2 concentration ([CO2]). Down-regulation is often attributed to an insufficient capacity of sink organs to use or store the increased carbohydrate production that results from the stimulation of photosynthesis by elevated [CO2]. Down-regulation can be accentuated by inadequate nitrogen (N) supply, which may limit sink development. While there is strong evidence for down-regulation of photosynthesis at elevated [CO2] in enclosure studies most often involving potted plants, there is little evidence for this when [CO2] is elevated fully under open-air field treatment conditions. To assess the importance of sink strength on the down-regulation of photosynthesis and on the potential of N to mitigate this down-regulation under agriculturally relevant field conditions, two tobacco cultivars (Nicotiana tabacum L. cv. Petit Havana; cv. Mammoth) of strongly contrasting ability to produce the major sink of this crop, leaves, were grown under ambient and elevated [CO2] and with two different N additions in a free air [CO2] (FACE) facility. Photosynthetic down-regulation at elevated [CO2] reached only 9% in cv. Mammoth late in the season likely reflecting sustained sink strength of the rapidly growing plant whereas down-regulation in cv. Petit Havana reached 25%. Increased N supply partially mitigated down-regulation of photosynthesis in cv. Petit Havana and this mitigation was dependent on plant developmental stage. Overall, these field results were consistent with the hypothesis that sustained sink strength, that is the ability to utilize photosynthate, and adequate N supply will allow C3 crops in the field to maintain enhanced photosynthesis and therefore productivity as [CO2] continues to rise.

Expression of cyanobacterial FBP/SBPase in soybean prevents yield depression under future climate conditions.

Predictions suggest that current crop production needs to double by 2050 to meet global food and energy demands. Based on theory and experimental studies, overexpression of the photosynthetic enzyme sedoheptulose-1,7-bisphosphatase (SBPase) is expected to enhance C3 crop photosynthesis and yields. Here we test how expression of the cyanobacterial, bifunctional fructose-1,6/sedoheptulose-1,7-bisphosphatase (FBP/SBPase) affects carbon assimilation and seed yield (SY) in a major crop (soybean, Glycine max). For three growing seasons, wild-type (WT) and FBP/SBPase-expressing (FS) plants were grown in the field under ambient (400 µmol mol−1) and elevated (600 µmol mol−1) CO2 concentrations [CO2] and under ambient and elevated temperatures (+2.7 °C during daytime, +3.4 °C at night) at the SoyFACE research site. Across treatments, FS plants had significantly higher carbon assimilation (4­14%), Vc,max (5­8%), and Jmax (4­8%). Under ambient [CO2], elevated temperature led to significant reductions of SY of both genotypes by 19­31%. However, under elevated [CO2] and elevated temperature, FS plants maintained SY levels, while the WT showed significant reductions between 11% and 22% compared with plants under elevated [CO2] alone. These results show that the manipulation of the photosynthetic carbon reduction cycle can mitigate the effects of future high CO2 and high temperature environments on soybean yield.

Intensifying drought eliminates the expected benefits of elevated carbon dioxide for soybean.

Stimulation of C3 crop yield by rising concentrations of atmospheric carbon dioxide ([CO2]) is widely expected to counteract crop losses that are due to greater drought this century. But these expectations come from sparse field trials that have been biased towards mesic growth conditions. This eight-year study used precipitation manipulation and year-to-year variation in weather conditions at a unique open-air field facility to show that the stimulation of soybean yield by elevated [CO2] diminished to zero as drought intensified. Contrary to the prevalent expectation in the literature, rising [CO2] did not counteract the effect of strong drought on photosynthesis and yield because elevated [CO2] interacted with drought to modify stomatal function and canopy energy balance. This new insight from field experimentation under hot and dry conditions, which will become increasingly prevalent in the coming decades, highlights the likelihood of negative impacts from interacting global change factors on a key global commodity crop in its primary region of production.

Canopy warming caused photosynthetic acclimation and reduced seed yield in maize grown at ambient and elevated [CO2 ].

Rising atmospheric CO2 concentration ([CO2 ]) and attendant increases in growing season temperature are expected to be the most important global change factors impacting production agriculture. Although maize is the most highly produced crop worldwide, few studies have evaluated the interactive effects of elevated [CO2 ] and temperature on its photosynthetic physiology, agronomic traits or biomass, and seed yield under open field conditions. This study investigates the effects of rising [CO2 ] and warmer temperature, independently and in combination, on maize grown in the field throughout a full growing season. Free-air CO2 enrichment (FACE) technology was used to target atmospheric [CO2 ] to 200 µmol mol(-1) above ambient [CO2 ] and infrared heaters to target a plant canopy increase of 3.5 °C, with actual season mean heating of ~2.7 °C, mimicking conditions predicted by the second half of this century. Photosynthetic gas-exchange parameters, leaf nitrogen and carbon content, leaf water potential components, and developmental measurements were collected throughout the season, and biomass and yield were measured at the end of the growing season. As predicted for a C4 plant, elevated [CO2 ] did not stimulate photosynthesis, biomass, or yield. Canopy warming caused a large shift in aboveground allocation by stimulating season-long vegetative biomass and decreasing reproductive biomass accumulation at both CO2 concentrations, resulting in decreased harvest index. Warming caused a reduction in photosynthesis due to down-regulation of photosynthetic biochemical parameters and the decrease in the electron transport rate. The reduction in seed yield with warming was driven by reduced photosynthetic capacity and by a shift in aboveground carbon allocation away from reproduction. This field study portends that future warming will reduce yield in maize, and this will not be mitigated by higher atmospheric [CO2 ] unless appropriate adaptation traits can be introduced into future cultivars.

Biochemical acclimation, stomatal limitation and precipitation patterns underlie decreases in photosynthetic stimulation of soybean (Glycine max) at elevated [CO2] and temperatures under fully open air field conditions.

The net effect of elevated [CO2] and temperature on photosynthetic acclimation and plant productivity is poorly resolved. We assessed the effects of canopy warming and fully open air [CO2] enrichment on (1) the acclimation of two biochemical parameters that frequently limit photosynthesis (A), the maximum carboxylation capacity of Rubisco (Vc,max) and the maximum potential linear electron flux through photosystem II (Jmax), (2) the associated responses of leaf structural and chemical properties related to A, as well as (3) the stomatal limitation (l) imposed on A, for soybean over two growing seasons in a conventionally managed agricultural field in Illinois, USA. Acclimation to elevated [CO2] was consistent over two growing seasons with respect to Vc,max and Jmax. However, elevated temperature significantly decreased Jmax contributing to lower photosynthetic stimulation by elevated CO2. Large seasonal differences in precipitation altered soil moisture availability modulating the complex effects of elevated temperature and CO2 on biochemical and structural properties related to A. Elevated temperature also reduced the benefit of elevated [CO2] by eliminating decreases in stomatal limitation at elevated [CO2]. These results highlight the critical importance of considering multiple environmental factors (i.e. temperature, moisture, [CO2]) when trying to predict plant productivity in the context of climate change.

Modelling C3 photosynthesis from the chloroplast to the ecosystem.

Globally, photosynthesis accounts for the largest flux of CO2 from the atmosphere into ecosystems and is the driving process for terrestrial ecosystem function. The importance of accurate predictions of photosynthesis over a range of plant growth conditions led to the development of a C3 photosynthesis model by Farquhar, von Caemmerer & Berry that has become increasingly important as society places greater pressures on vegetation. The photosynthesis model has played a major role in defining the path towards scientific understanding of photosynthetic carbon uptake and the role of photosynthesis on regulating the earth's climate and biogeochemical systems. In this review, we summarize the photosynthesis model, including its continued development and applications. We also review the implications these developments have on quantifying photosynthesis at a wide range of spatial and temporal scales, and discuss the model's role in determining photosynthetic responses to changes in environmental conditions. Finally, the review includes a discussion of the larger-scale modelling and remote-sensing applications that rely on the leaf photosynthesis model and are likely to open new scientific avenues to address the increasing challenges to plant productivity over the next century.

Global warming can negate the expected CO2 stimulation in photosynthesis and productivity for soybean grown in the Midwestern United States.

Extensive evidence shows that increasing carbon dioxide concentration ([CO2]) stimulates, and increasing temperature decreases, both net photosynthetic carbon assimilation (A) and biomass production for C3 plants. However the [CO2]-induced stimulation in A is projected to increase further with warmer temperature. While the influence of increasing temperature and [CO2], independent of each other, on A and biomass production have been widely investigated, the interaction between these two major global changes has not been tested on field-grown crops. Here, the interactive effect of both elevated [CO2] (approximately 585 µmol mol(-1)) and temperature (+3.5°C) on soybean (Glycine max) A, biomass, and yield were tested over two growing seasons in the Temperature by Free-Air CO2 Enrichment experiment at the Soybean Free Air CO2 Enrichment facility. Measurements of A, stomatal conductance, and intercellular [CO2] were collected along with meteorological, water potential, and growth data. Elevated temperatures caused lower A, which was largely attributed to declines in stomatal conductance and intercellular [CO2] and led in turn to lower yields. Increasing both [CO2] and temperature stimulated A relative to elevated [CO2] alone on only two sampling days during 2009 and on no days in 2011. In 2011, the warmer of the two years, there were no observed increases in yield in the elevated temperature plots regardless of whether [CO2] was elevated. All treatments lowered the harvest index for soybean, although the effect of elevated [CO2] in 2011 was not statistically significant. These results provide a better understanding of the physiological responses of soybean to future climate change conditions and suggest that the potential is limited for elevated [CO2] to mitigate the influence of rising temperatures on photosynthesis, growth, and yields of C3 crops.

Future carbon dioxide concentration decreases canopy evapotranspiration and soil water depletion by field-grown maize.

Maize, in rotation with soybean, forms the largest continuous ecosystem in temperate North America, therefore changes to the biosphere-atmosphere exchange of water vapor and energy of these crops are likely to have an impact on the Midwestern US climate and hydrological cycle. As a C4 crop, maize photosynthesis is already CO2 -saturated at current CO2 concentrations ([CO2 ]) and the primary response of maize to elevated [CO2 ] is decreased stomatal conductance (gs ). If maize photosynthesis is not stimulated in elevated [CO2 ], then reduced gs is not offset by greater canopy leaf area, which could potentially result in a greater ET reduction relative to that previously reported in soybean, a C3 species. The objective of this study is to quantify the impact of elevated [CO2 ] on canopy energy and water fluxes of maize (Zea mays). Maize was grown under ambient and elevated [CO2 ] (550 µmol mol(-1) during 2004 and 2006 and 585 µmol mol(-1) during 2010) using Free Air Concentration Enrichment (FACE) technology at the SoyFACE facility in Urbana, Illinois. Maize ET was determined using a residual energy balance approach based on measurements of sensible (H) and soil heat fluxes, and net radiation. Relative to control, elevated [CO2 ] decreased maize ET (7-11%; P < 0.01) along with lesser soil moisture depletion, while H increased (25-30 W m(-2) ; P < 0.01) along with higher canopy temperature (0.5-0.6 °C). This reduction in maize ET in elevated [CO2 ] is approximately half that previously reported for soybean. A partitioning analysis showed that transpiration contributed less to total ET for maize compared to soybean, indicating a smaller role of stomata in dictating the ET response to elevated [CO2 ]. Nonetheless, both maize and soybean had significantly decreased ET and increased H, highlighting the critical role of elevated [CO2 ] in altering future hydrology and climate of the region that is extensively cropped with these species.