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.
Extensive evidence shows that increasing carbon dioxide concentration ([CO 2 ]) stimulates, and increasing temperature decreases, both net photosynthetic carbon assimilation (A) and biomass production for C 3 plants. However the [CO 2 ]-induced stimulation in A is projected to increase further with warmer temperature. While the influence of increasing temperature and [CO 2 ], 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 [CO 2 ] (approximately 585 mmol mol 21) 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 CO 2 Enrichment experiment at the Soybean Free Air CO 2 Enrichment facility. Measurements of A, stomatal conductance, and intercellular [CO 2 ] 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 [CO 2 ] and led in turn to lower yields. Increasing both [CO 2 ] and temperature stimulated A relative to elevated [CO 2 ] 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 [CO 2 ] was elevated. All treatments lowered the harvest index for soybean, although the effect of elevated [CO 2 ] 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 [CO 2 ] to mitigate the influence of rising temperatures on photosynthesis, growth, and yields of C 3 crops.
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.
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 largerscale 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.
HighlightExpression of cyanobacterial FBP/SBPase in soybean prevents yield depression in a free air CO2 enrichment experiment under simulated future climate conditions with combination of elevated CO2 and elevated temperature.
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.
Rosenthala, David M.; Ruiz-Vera, Ursula M.; Siebers, Matthew H.; Gray, Sharon B.; Bernacchi, Carl J.; and Ort, Donald R., "Biochemical acclimation, stomatal limitation and precipitationpatterns underlie decreases in photosynthetic stimulation of soybean(Glycine max) at elevated [CO 2 Contents lists available at ScienceDirect Plant Science j o u r n a l h o m e p a g e : w w w . e l s e v i e r . c o m / l o c a t e / p l a n t s c i (1) the acclimation of two biochemical parameters that frequently limit photosynthesis (A), the maximum carboxylation capacity of Rubisco (V c,max ) and the maximum potential linear electron flux through photosystem II (J max ), (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 [CO 2 ] was consistent over two growing seasons with respect to V c,max and J max . However, elevated temperature significantly decreased J max contributing to lower photosynthetic stimulation by elevated CO 2 . Large seasonal differences in precipitation altered soil moisture availability modulating the complex effects of elevated temperature and CO 2 on biochemical and structural properties related to A. Elevated temperature also reduced the benefit of elevated [CO 2 ] by eliminating decreases in stomatal limitation at elevated [CO 2 ]. These results highlight the critical importance of considering multiple environmental factors (i.e. temperature, moisture, [CO 2 ]) when trying to predict plant productivity in the context of climate change.
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.
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