Stomatal regulation in a changing climate: a field study using Free Air Temperature Increase (FATI) and Free Air CO2 Enrichment (FACE)

This study investigates effects of climate warming (+ 2.5°C ubove ambient) and elevated CO₂ concentration (600 μmol mol^?1) on the stomatal functioning and the water relations of Lolium perenne, using Free Air Temperature Increase (FATI) and Free Air CO₂ Enrichment (FACE). Compared to growth at ambient temperature, whole-season temperature increase reduced leaf stomatal conductance, but only at the top of the canopy (-14.6 and -8.8% at ambient and elevated CO₂, respectively). However, because higher canopy temperature raised the leaf-to-air vapour pressure difference, leaf transpiration rate increased (+28% at ambient and +48% at elevated CO₂) and instantaneous leaf water use efficiency, derived from short-term measurements of assimilation and transpiration rate, declined (-11% at ambient and -13% at elevated CO₂). Nevertheless, at the stand level, growth at + 2.5°C reduced transpiration due to fewer tillers per plant and a smaller leaf area per tiller. This sparser vegetation was also more closely coupled to the atmosphere and maintained a drier internal microclimate. To assess whether the stomatal behaviour observed in this experiment could be explained by prevailing concepts of stomatal functioning, three models were applied (Cowan 1977; Ball, Woodrow & Berry 1987; Leuning 1995). The latter model accounted for the highest proportion of variability in the data (58%) and was insensitive to CO₂ and temperature regime, which suggests that the principles of stomatal regulation are not affected by changes in CO₂ or climate.     

Influence of High Temperature on End-of-Season Tundra CO2 Exchange

The high-arctic terrestrial environment is generally recognized as one of the world's most sensitive areas with regard to global warming. In this study, we examined the influence of an isolated warm period on net ecosystem carbon dioxide (CO₂) exchange at high latitude during autumn. Using the Free Air Temperature Increase (FATI) technique, we manipulated air, soil, and vegetation temperatures in late August in a tundra site at Zackenberg in the National Park of North and East Greenland (74°N 21°W). The consequences for gross canopy photosynthesis, canopy respiration, and belowground respiration of increasing these temperatures by approximately 2.5°C were determined with closed dynamic CO₂ exchange systems. Under current temperatures, the ecosystem acted as a net CO₂ source, releasing 19 g CO₂-C m⁻² over the 14-day study period. Warm soils and senescing vegetation in autumn were unequivocally responsible for this efflux. Heating enhanced CO₂ efflux to 29 g CO₂-C m⁻². This effect was attributed to a 39% increase in belowground respiration, which was the main component of the carbon (C) budget. Gross photosynthesis, on the other hand, was not affected significantly by the simulated warming. Although the aftereffects of an isolated warm period on the C balance in early winter could be significant, simulations with a simple C budget model suggest that soil carbon pools are not affected to a great extent by such a climatic disturbance. The influence on atmospheric carbon, however, appears to be significant. Received 9 June 2000; accepted 20 December 2000.     

Evergreen leaf respiration acclimates to long-term nocturnal warming under field conditions

Acclimation of plant respiration rates (R) to climate warming is highly variable and many results appear contradictory. We tested the recently suggested hypotheses that pre‐existing, long‐lived leaves should exhibit a relatively limited ability for R to acclimate to climate warming, and that acclimation would occur via changes in the short‐term temperature sensitivity of respiration. Seedlings of a subalpine, evergreen tree species (Eucalyptus pauciflora) were grown under naturally fluctuating conditions within its natural distribution. We used a free air temperature increase (FATI) system of infra‐red ceramic lamps to raise night‐time leaf temperatures by 0.3±0.1, 1.3±0.1, and 2.2±0.1 °C above ambient for 1 year. Light‐saturated assimilation rates and plant growth did not change with nocturnal FATI treatments. Leaf R measured at prevailing temperatures did not differ between FATI treatments. Within each FATI treatment, nocturnal leaf R was highly sensitive to artificial temperature changes within minutes, and also correlated strongly with natural nocturnal and seasonal temperature variation. The corresponding values of Q₁₀ of R varied according to time scale of measurements, but did not vary between FATI treatments. Instead, acclimation of R to nocturnal FATI occurred through changes in the base rate of respiration.     

Future atmospheric CO2 leads to delayed autumnal senescence

Growing seasons are getting longer, a phenomenon partially explained by increasing global temperatures. Recent reports suggest that a strong correlation exists between warming and advances in spring phenology but that a weaker correlation is evident between warming and autumnal events implying that other factors may be influencing the timing of autumnal phenology. Using freely rooted, field‐grown Populus in two Free Air CO₂ Enrichment Experiments (AspenFACE and PopFACE), we present evidence from two continents and over 2 years that increasing atmospheric CO₂ acts directly to delay autumnal leaf coloration and leaf fall. In an atmosphere enriched in CO₂ (by ～45% of the current atmospheric concentration to 550 ppm) the end of season decline in canopy normalized difference vegetation index (NDVI) – a commonly used global index for vegetation greenness – was significantly delayed, indicating a greener autumnal canopy, relative to that in ambient CO₂. This was supported by a significant delay in the decline of autumnal canopy leaf area index in elevated as compared with ambient CO₂, and a significantly smaller decline in end of season leaf chlorophyll content. Leaf level photosynthetic activity and carbon uptake in elevated CO₂ during the senescence period was also enhanced compared with ambient CO₂. The findings reveal a direct effect of rising atmospheric CO₂, independent of temperature in delaying autumnal senescence for Populus, an important deciduous forest tree with implications for forest productivity and adaptation to a future high CO₂ world.     

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.     

Growth in elevated CO2 protects photosynthesis against high-temperature damage

We present evidence that plant growth at elevated atmospheric CO₂ increases the high‐temperature tolerance of photosynthesis in a wide variety of plant species under both greenhouse and field conditions. We grew plants at ambient CO₂ (~ 360 μ mol mol ^{? 1}) and elevated CO₂ (550–1000 μ mol mol ^{? 1}) in three separate growth facilities, including the Nevada Desert Free‐Air Carbon Dioxide Enrichment (FACE) facility. Excised leaves from both the ambient and elevated CO₂ treatments were exposed to temperatures ranging from 28 to 48 °C. In more than half the species examined (4 of 7, 3 of 5, and 3 of 5 species in the three facilities), leaves from elevated CO₂‐grown plants maintained PSII efficiency (F_v/F_m) to significantly higher temperatures than ambient‐grown leaves. This enhanced PSII thermotolerance was found in both woody and herbaceous species and in both monocots and dicots. Detailed experiments conducted with Cucumis sativus showed that the greater F_v/F_m in elevated versus ambient CO₂‐grown leaves following heat stress was due to both a higher F_m and a lower F_o, and that F_v/F_m differences between elevated and ambient CO₂‐grown leaves persisted for at least 20 h following heat shock. Cucumis sativus leaves from elevated CO₂‐grown plants had a critical temperature for the rapid rise in F_o that averaged 2·9 °C higher than leaves from ambient CO₂‐grown plants, and maintained a higher maximal rate of net CO₂ assimilation following heat shock. Given that photosynthesis is considered to be the physiological process most sensitive to high‐temperature damage and that rising atmospheric CO₂ content will drive temperature increases in many already stressful environments, this CO₂‐induced increase in plant high‐temperature tolerance may have a substantial impact on both the productivity and distribution of many plant species in the 21st century.     

Physiological ecology of desert biocrust moss following 10 years exposure to elevated CO₂: evidence for enhanced photosynthetic thermotolerance

In arid regions, biomes particularly responsive to climate change, mosses play an important biogeochemical role as key components of biocrusts. Using the biocrust moss Syntrichia caninervis collected from the Nevada Desert Free Air CO₂ Enrichment Facility, we examined the physiological effects of 10 years of exposure to elevated CO₂, and the effect of high temperature events on the photosynthetic performance of moss grown in CO₂‐enriched air. Moss exposed to elevated CO₂ exhibited a 46% decrease in chlorophyll, a 20% increase in carbon and no difference in either nitrogen content or photosynthetic performance. However, when subjected to high temperatures (35–40°C), mosses from the elevated CO₂ environment showed higher photosynthetic performance and photosystem II (PSII) efficiency compared to those grown in ambient conditions, potentially reflective of a shift in nitrogen allocation to components that offer a higher resistance of PSII to heat stress. This result suggests that mosses may respond to climate change in markedly different ways than vascular plants, and observed CO₂‐induced photosynthetic thermotolerance in S. caninervis will likely have consequences for future desert biogeochemistry.     

Response of soil surface CO2 flux in a boreal forest to ecosystem warming

Soil surface carbon dioxide (CO₂) flux (R_S) was measured for 2 years at the Boreal Soil and Air Warming Experiment site near Thompson, MB, Canada. The experimental design was a complete random block design that consisted of four replicate blocks, with each block containing a 15 m × 15 m control and heated plot. Black spruce [Picea mariana (Mill.) BSP] was the overstory species and Epilobium angustifolium was the dominant understory. Soil temperature was maintained (～5 °C) above the control soil temperature using electric cables inside water filled polyethylene tubing for each heated plot. Air inside a 7.3‐m‐diameter chamber, centered in the soil warming plot, contained approximately nine black spruce trees was heated ～5 °C above control ambient air temperature allowing for the testing of soil‐only warming and soil+air warming. Soil surface CO₂ flux (R_S) was positively correlated (P < 0.0001) to soil temperature at 10 cm depth. Soil surface CO₂ flux (R_S) was 24% greater in the soil‐only warming than the control in 2004, but was only 11% greater in 2005, while R_S in the soil+air warming treatments was 31% less than the control in 2004 and 23% less in 2005. Live fine root mass (< 2 mm diameter) was less in the heated than control treatments in 2004 and statistically less (P < 0.01) in 2005. Similar root mass between the two heated treatments suggests that different heating methods (soil‐only vs. soil+air warming) can affect the rate of decomposition.     

Infrared heater arrays for warming ecosystem field plots

There is a need for methodology to warm open‐field plots in order to study the likely effects of global warming on ecosystems in the future. Herein, we describe the development of arrays of more powerful and efficient infrared heaters with ceramic heating elements. By tilting the heaters at 45° from horizontal and combining six of them in a hexagonal array, good uniformity of warming was achieved across 3‐m‐diameter plots. Moreover, there do not appear to be obstacles (other than financial) to scaling to larger plots. The efficiency [η_h (%); thermal radiation out per electrical energy in] of these heaters was higher than that of the heaters used in most previous infrared heater experiments and can be described by: η_h= 10 + 25exp(? 0.17 u), where u is wind speed at 2 m height (m s^{? 1}). Graphs are presented to estimate operating costs from degrees of warming, two types of plant canopy, and site windiness. Four such arrays were deployed over plots of grass at Haibei, Qinghai, China and another at Cheyenne, Wyoming, USA, along with corresponding reference plots with dummy heaters. Proportional integral derivative systems with infrared thermometers to sense canopy temperatures of the heated and reference plots were used to control the heater outputs. Over month‐long periods at both sites, about 75% of canopy temperature observations were within 0.5 °C of the set‐point temperature differences between heated and reference plots. Electrical power consumption per 3‐m‐diameter plot averaged 58 and 80 kW h day^{? 1} for Haibei and Cheyenne, respectively. However, the desired temperature differences were set lower at Haibei (1.2 °C daytime, 1.7 °C night) than Cheyenne (1.5 °C daytime, 3.0 °C night), and Cheyenne is a windier site. Thus, we conclude that these hexagonal arrays of ceramic infrared heaters can be a successful temperature free‐air‐controlled enhancement (T‐FACE) system for warming ecosystem field plots.     

Functional ecology of shrub seedlings after a natural recruitment event at the Nevada Desert FACE Facility

Seedling recruitment is an important determinant of community structure in desert ecosystems. Positive photosynthetic growth and water balance responses to increasing atmospheric carbon dioxide (CO₂) concentrations ([CO₂]) are predicted to be substantial in desert plants, suggesting that recruitment could be stimulated. However, to date no studies have addressed the response of perennial plant recruitment in natural populations of desert shrubs exposed to elevated [CO₂]. In April 1997, we employed Free‐Air Carbon Dioxide Enrichment (FACE) in order to increase atmospheric [CO₂] in an undisturbed Mojave Desert ecosystem from ambient (～～ 370 µmol mol^?1) to elevated CO₂ (～～ 550 µmol mol^?1). From 1997 to 2001 we seasonally examined survival, growth, gas exchange and water potential responses of Larrea tridentata and Ambrosia dumosa seedlings that germinated in Fall, 1997. Recruitment densities were not influenced by [CO₂] in either species, although a two‐fold higher adult Ambrosia density under elevated [CO₂] resulted in two‐fold higher seedling density (0.87 vs 0.40 seedlings m^?2). Mortality was greatest for both species during the first summer (1998), despite above‐average rainfall during the previous Winter–Spring. A significant [CO₂] × time interaction revealed that early survival was greater under elevated CO₂, whereas a significant species time interaction revealed that overall survival was greater for Ambrosia (28%) than for Larrea (15%), regardless of [CO₂]. Microsite (understorey or interspace) alone had no significant influence on survival. Significant species, microsite and species × microsite effects on growth (seedling height, stem diameter and canopy size) were found, but elevated CO₂ had minimal impact on these parameters. Photosynthetic rates (A_sat) for both species were higher at elevated [CO₂] during certain seasons, but not consistently so. These results suggest that increased atmospheric [CO₂] may enhance carbon (C) assimilation and survival of aridland perennial shrubs during favourable growing conditions, but that it may not counteract the effects of prolonged drought on mortality.     

COMPARATIVE CACTUS ARCHITECTURE AND PAR INTERCEPTION

Because CO₂ uptake by cacti can be limited by low levels of photosynthetically active radiation (PAR) and because plant form affects PAR interception, various cactus forms were studied using a computer model, field measurements, and laboratory phototropic studies. Model predictions indicated that CO₂ uptake by individual stems at an equinox was greatest when the stems were vertical, but at the summer and the winter solstice CO₂ uptake was greatest for stems tilted 30° away from the equator. Stem tilting depended on form and taxonomic group; four barrel cacti in Ferocactus and in Copiapoa and four cylindropuntias in Opuntia tilted toward a horizontal light beam by an average of 18°, 48°, and 52°, respectively, after growth periods of 1 to 4 yr. In contrast, three columnar species showed no significant phototropic response, perhaps because structural stability requires their massive stems to be erect. Field plants of the dense, multiple-stemmed shrub Opuntia echinocarpa had stems which tended to radiate outward from the plant base, and, although this would not influence the total PAR intercepted, it would result in a more uniform PAR distribution and hence higher CO₂ uptake. For O. echinocarpa and the even denser, mound-forming Echinocereus engelmannii, PAR and chlorophyll decreased approximately exponentially with depth into the canopy. The canopies of O. echinocarpa and other cylindropuntias did not extend to the ground; in certain species, such truncation apparently resulted from a combination of very low PAR levels just below the lower canopy edge and the light-dependent growth responses of individual stems. In addition, although the canopy surfaces of O. echinocarpa and O. acanthocarpa tilted toward the equator by about 30°, the canopies of other cylindropuntias tilted less or not at all; the computer model predicted that a 30° tilt would decrease interstem shading, increase daily PAR, and increase nocturnal CO₂ uptake by as much as 54, 26, and 24%, respectively. Not only can the shape of cacti be affected by PAR, but also shape influences PAR interception and hence CO₂ uptake.     

Characteristics of free air carbon dioxide enrichment of a northern temperate mature forest

In 2017, the Birmingham Institute of Forest Research (BIFoR) began to conduct Free Air Carbon Dioxide Enrichment (FACE) within a mature broadleaf deciduous forest situated in the United Kingdom. BIFoR FACE employs large‐scale infrastructure, in the form of lattice towers, forming ‘arrays’ which encircle a forest plot of ~30 m diameter. BIFoR FACE consists of three treatment arrays to elevate local CO₂ concentrations (e[CO₂]) by +150 µmol/mol. In practice, acceptable operational enrichment (ambient [CO₂] + e[CO₂]) is ±20% of the set point 1‐min average target. There are a further three arrays that replicate the infrastructure and deliver ambient air as paired controls for the treatment arrays. For the first growing season with e[CO₂] (April to November 2017), [CO₂] measurements in treatment and control arrays show that the target concentration was successfully delivered, that is: +147 ± 21 µmol/mol (mean ± SD) or 98 ± 14% of set point enrichment target. e[CO₂] treatment was accomplished for 97.7% of the scheduled operation time, with the remaining time lost due to engineering faults (0.6% of the time), CO₂ supply issues (0.6%) or adverse weather conditions (1.1%). CO₂ demand in the facility was driven predominantly by wind speed and the formation of the deciduous canopy. Deviations greater than 10% from the ambient baseline CO₂ occurred <1% of the time in control arrays. Incidences of cross‐contamination >80 µmol/mol (i.e. >53% of the treatment increment) into control arrays accounted for <0.1% of the enrichment period. The median [CO₂] values in reconstructed three‐dimensional [CO₂] fields show enrichment somewhat lower than the target but still well above ambient. The data presented here provide confidence in the facility setup and can be used to guide future next‐generation forest FACE facilities built into tall and complex forest stands.     

Temporal dynamics and spatial variability in the enhancement of canopy leaf area under elevated atmospheric CO2

Increased canopy leaf area (L) may lead to higher forest productivity and alter processes such as species dynamics and ecosystem mass and energy fluxes. Few CO₂ enrichment studies have been conducted in closed canopy forests and none have shown a sustained enhancement of L. We reconstructed 8 years (1996–2003) of L at Duke's Free Air CO₂ Enrichment experiment to determine the effects of elevated atmospheric CO₂ concentration ([CO₂]) on L before and after canopy closure in a pine forest with a hardwood component, focusing on interactions with temporal variation in water availability and spatial variation in nitrogen (N) supply. The dynamics of L were reconstructed using data on leaf litterfall mass and specific leaf area for hardwoods, and needle litterfall mass and specific leaf area combined with needle elongation rates, and fascicle and shoot counts for pines. The dynamics of pine L production and senescence were unaffected by elevated [CO₂], although L senescence for hardwoods was slowed. Elevated [CO₂] enhanced pine L and the total canopy L (combined pine and hardwood species; P<0.050); on average, enhancement following canopy closure was ～16% and 14% respectively. However, variation in pine L and its response to elevated [CO₂] was not random. Each year pine L under ambient and elevated [CO₂] was spatially correlated to the variability in site nitrogen availability (e.g. r²=0.94 and 0.87 in 2001, when L was highest before declining due to droughts and storms), with the [CO₂]‐induced enhancement increasing with N (P=0.061). Incorporating data on N beyond the range of native fertility, achieved through N fertilization, indicated that pine L had reached the site maximum under elevated [CO₂] where native N was highest. Thus closed canopy pine forests may be able to increase leaf area under elevated [CO₂] in moderate fertility sites, but are unable to respond to [CO₂] in both infertile sites (insufficient resources) and sites having high levels of fertility (maximum utilization of resources). The total canopy L, representing the combined L of pine and hardwood species, was constant across the N gradient under both ambient and elevated [CO₂], generating a constant enhancement of canopy L. Thus, in mixed species stands, L of canopy hardwoods which developed on lower fertility sites (～3 g N inputs m^?2 yr^?1) may be sufficiently enhanced under elevated [CO₂] to compensate for the lack of response in pine L, and generate an appreciable response of total canopy L (～14%).     

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 (g_s). If maize photosynthesis is not stimulated in elevated [CO₂], then reduced g_s 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.     

Modification of the response of photosynthetic productivity to rising temperature by atmospheric CO2 concentrations: Has its importance been underestimated?

Abstract. Climate change will include correlated increases in temperature and atmospheric CO₂ concentration (C_a). Rising temperatures will increase the ratio of photorespiratory loss of carbon to photosynthetic gain, whilst rising C_a will have an opposing effect. The mechanism of these effects at the level of carboxylation in C₃ photosynthesis are quantitatively well understood and provide a basis for models of the response of leaf and canopy carbon exchange to climate change. The principles of such a model are referred to here and used to quantitatively examine the implications of concurrent increase in temperature and C_a. Simulations of leaf photosynthesis show the increase, with elevation of C_a from 350 to 650 μmol mol^-1, in light saturated rates of CO₂ uptake (A_sat) and maximum quantum yields (φ) to rise with temperature. An increase in C_a from 350 to 650 μmol mol^-1 can increase A_sat by 20% at 10°C and by 105% at 35°C, and can raise the temperature optimum of A_sat by 5°C. This pattern of change agrees closely with experimental data. At the canopy level, simulations also suggest a strong interaction of increased temperature and CO₂ concentration. Predictions are compared with the findings of long-term field studies. The principles used here suggest that elevated C_a will alter both the magnitude of the response of leaf and canopy carbon gain to rising temperature, and sometimes, the direction of response. Findings question the value of models for predicting plant production in response to climate change which ignore the direct effects of rising C_a and the modifications that rising C_a imposes on the temperature response of net CO₂ exchange.     

Increased temperatures may safeguard the nutritional quality of crops under future elevated CO2 concentrations

Iron (Fe) and zinc (Zn) deficiencies are a global human health problem that may worsen by the growth of crops at elevated atmospheric CO₂ concentration (eCO₂). However, climate change will also involve higher temperature, but it is unclear how the combined effect of eCO₂ and higher temperature will affect the nutritional quality of food crops. To begin to address this question, we grew soybean (Glycine max) in a Temperature by Free‐Air CO₂ Enrichment (T‐FACE) experiment in 2014 and 2015 under ambient (400 μmol mol^?1) and elevated (600 μmol mol^?1) CO₂ concentrations, and under ambient and elevated temperatures (+2.7°C day and +3.4°C at night). In our study, eCO₂ significantly decreased Fe concentration in soybean seeds in both seasons (?8.7 and ?7.7%) and Zn concentration in one season (?8.9%), while higher temperature (at ambient CO₂ concentration) had the opposite effect. The combination of eCO₂ with elevated temperature generally restored seed Fe and Zn concentrations to levels obtained under ambient CO₂ and temperature conditions, suggesting that the potential threat to human nutrition by increasing CO₂ concentration may not be realized. In general, seed Fe concentration was negatively correlated with yield, suggesting inherent limitations to increasing seed Fe. In addition, we confirm our previous report that the concentration of seed storage products and several minerals varies with node position at which the seeds developed. Overall, these results demonstrate the complexity of predicting climate change effects on food and nutritional security when various environmental parameters change in an interactive manner.     

Photosynthesis, light and nitrogen relationships in a young deciduous forest canopy under open-air CO2 enrichment

Leaf photosynthesis (Ps), nitrogen (N) and light environment were measured on Populus tremuloides trees in a developing canopy under free‐air CO₂ enrichment in Wisconsin, USA. After 2 years of growth, the trees averaged 1·5 and 1·6 m tall under ambient and elevated CO₂, respectively, at the beginning of the study period in 1999. They grew to 2·6 and 2·9 m, respectively, by the end of the 1999 growing season. Daily integrated photon flux from cloud‐free days (PPFD_day,sat) around the lowermost branches was 16·8 ± 0·8 and 8·7 ± 0·2% of values at the top for the ambient and elevated CO₂ canopies, respectively. Elevated CO₂ significantly decreased leaf N on a mass, but not on an area, basis. N per unit leaf area was related linearly to PPFD_day,sat throughout the canopies, and elevated CO₂ did not affect that relationship. Leaf Ps light‐response curves responded differently to elevated CO₂, depending upon canopy position. Elevated CO₂ increased Ps_sat only in the upper (unshaded) canopy, whereas characteristics that would favour photosynthesis in shade were unaffected by elevated CO₂. Consequently, estimated daily integrated Ps on cloud‐free days (Ps_day,sat) was stimulated by elevated CO₂ only in the upper canopy. Ps_day,sat of the lowermost branches was actually lower with elevated CO₂ because of the darker light environment. The lack of CO₂ stimulation at the mid‐ and lower canopy was probably related to significant down‐regulation of photosynthetic capacity; there was no down‐regulation of Ps in the upper canopy. The relationship between Ps_day,sat and leaf N indicated that N was not optimally allocated within the canopy in a manner that would maximize whole‐canopy Ps or photosynthetic N use efficiency. Elevated CO₂ had no effect on the optimization of canopy N allocation.     

Some effects of air temperature and humidity on crop and leaf photosynthesis, transpiration and resistance to gas transfer

Measurements of carbon dioxide exchange and transpiration were made, at various air temperatures, on wheat and barley using a field enclosure system. From these were derived the stomatal and mesophyll resistances to carbon dioxide transfer. Optimum temperatures for net CO₂ uptake were about 24°C for wheat and barley. Above these optima, as temperature increased so net CO₂ uptake rates decreased, because of increasing stomatal and mesophyll resistances; transpiration rates decreased in wheat but were constant in barley. In laboratory growth cabinets, wheat plants were subjected to different regimes of temperature and humidity. Optimum temperature for net CO₂ uptake of individual leaves was 25°C. At constant humidity, a decline in net uptake rates above 25°C was associated with large increases in mesophyll resistance. At a constant 25°C, as the vapour pressure deficit (v.p.d. was increased above 1 k Pa (10 mb) v.p.d. the net uptake declined, with an increase in mesophyll resistance and a small increase in stomatal resistance. When the v.p.d. exceeded 1 k Pa at a temperature of 30°C, conditions that are experienced by field plants, then there were large increases in both mesophyll and stomatal resistances and the net uptake rates declined. Photo-respiration, as measured by CO₂ uptake in oxygen-free air, was independent of temperature, but both dark respiration and CO₂ compensation concentration increased with temperature.     

Rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) growth and nitrogen distribution under elevated CO<Subscript>2</Subscript> concentration and air temperature

A field experiment was conducted to investigate the effects of elevated atmospheric CO₂ concentration and temperature, singly and in combination, on grain yield and the distribution of nitrogen (N) in different rice organs. The rice ‘Wuyunjing 23’ was planted under four treatments: ambient CO₂ and temperature (ACT), elevated CO₂ (200 μmol mol^?1 higher than ambient CO₂) (EC), elevated temperature (1 °C above the ambient temperature) (ET), and the combination of elevated CO₂ and temperature (ECT) under T-FACE (temperature and CO₂ free air controlled enrichment) system. CO₂-induced increment and temperature-induced reduction in grain yield was 6.0 and 25.2% in 2013, and that was 9.8 and 10.8% in 2014, respectively. Dry matter (DM) production in different organs increased under EC at vegetative stage but decreased under ET at reproductive stage. The negative effects of temperature on grain yield and DM was weakened when combined with CO₂ enrichment. And the trends of decrease for yield and DM under ET and ECT in 2013 were more obvious than those in 2014 due to the annual temperature differences. Furthermore, ET led to greater distribution of N in root and stem but not for panicle than that under ACT. These mainly demonstrated that the rice production would be suffered varying degree of loss under global warming in future although the CO₂ enrichment could alleviate the effects of high temperature on rice growth.     

Physiological responses of Opuntia ficus-indica to growth temperature

The influences of various day/night air temperatures on net CO₂ uptake and nocturnal acid accumulation were determined for Opuntia ficus-indica, complementing previous studies on the water relations and responses to photosynthetically active radiation (PAR) for this widely cultivated cactus. As for other Crassulacean acid metabolism (CAM) plants, net nocturnal CO₂ uptake had a relatively low optimal temperature, ranging from 11°C for plants grown at day/night air temperatures of 10°C/0°C to 23°C at 45°C/35°C. Stomatal opening, which occurred essentially only at night and was measured by changes in water vapor conductance, progressively decreased as the measurement temperature was raised. The CO₂ residual conductance, which describes chlorenchyma properties, had a temperature optimum a few degrees higher than the optimum for net CO₂ uptake at all growth temperatures. Nocturnal CO₂ uptake and acid accumulation summed over the whole night were maximal for growth temperatures near 25°C/15°C, CO₂ uptake decreasing more rapidly than acid accumulation as the growth temperature was raised. At day/night air temperatures that led to substantial nocturnal acid accumulation (25°C/15°C.). 90% saturation of acid accumulation required a higher total daily PAR than at non-optimal growth temperatures (10°C/0°C and 35°C/25°C). Also, the optimal temperature of net CO₂ uptake shifted downward when the plants were under drought conditions at all three growth temperatures tested, possibly reflecting an increased fractional importance of respiration at the higher temperatures during drought. Thus, water status, ambient PAR, and growth temperatures must all be considered when predicting the temperature response of gas exchange for O. ficus-indica and presumably for other CAM plants.