Does Elevated [CO2] Only Increase Root Growth in the Topsoil? A FACE Study with Lentil in a Semi-Arid Environment.

Atmospheric carbon dioxide concentrations [CO2] are increasing steadily. Some reports have shown that root growth in grain crops is mostly stimulated in the topsoil rather than evenly throughout the soil profile by e[CO2], which is not optimal for crops grown in semi-arid environments with strong reliance on stored water. An experiment was conducted during the 2014 and 2015 growing seasons with two lentil (Lens culinaris) genotypes grown under Free Air CO2 Enrichment (FACE) in which root growth was observed non-destructively with mini-rhizotrons approximately every 2-3 weeks. Root growth was not always statistically increased by e[CO2] and not consistently between depths and genotypes. In 2014, root growth in the top 15 cm of the soil profile (topsoil) was indeed increased by e[CO2], but increases at lower depths (30-45 cm) later in the season were greater than in the topsoil. In 2015, e[CO2] only increased root length in the topsoil for one genotype, potentially reflecting the lack of plant available soil water between 30-60 cm until recharged by irrigation during grain filling. Our limited data to compare responses to e[CO2] showed that root length increases in the topsoil were correlated with a lower yield response to e[CO2]. The increase in yield response was rather correlated with increases in root growth below 30 cm depth.

Assessing the evolution of wheat grain traits during the last 166 years using archived samples.

The current study focuses on yield and nutritional quality changes of wheat grain over the last 166 years. It is based on wheat grain quality analyses carried out on samples collected between 1850 and 2016. Samples were obtained from the Broadbalk Continuous Wheat Experiment (UK) and from herbaria from 16 different countries around the world. Our study showed that, together with an increase in carbohydrate content, an impoverishment of mineral composition and protein content occurred. The imbalance in carbohydrate/protein content was specially marked after the 1960's, coinciding with strong increases in ambient [CO2] and temperature and the introduction of progressively shorter straw varieties. The implications of altered crop physiology are discussed.

Elevated CO2 has concurrent effects on leaf and grain metabolism but minimal effects on yield in wheat.

While the general effect of CO2 enrichment on photosynthesis, stomatal conductance, N content, and yield has been documented, there is still some uncertainty as to whether there are interactive effects between CO2 enrichment and other factors, such as temperature, geographical location, water availability, and cultivar. In addition, the metabolic coordination between leaves and grains, which is crucial for crop responsiveness to elevated CO2, has never been examined closely. Here, we address these two aspects by multi-level analyses of data from several free-air CO2 enrichment experiments conducted in five different countries. There was little effect of elevated CO2 on yield (except in the USA), likely due to photosynthetic capacity acclimation, as reflected by protein profiles. In addition, there was a significant decrease in leaf amino acids (threonine) and macroelements (e.g. K) at elevated CO2, while other elements, such as Mg or S, increased. Despite the non-significant effect of CO2 enrichment on yield, grains appeared to be significantly depleted in N (as expected), but also in threonine, the S-containing amino acid methionine, and Mg. Overall, our results suggest a strong detrimental effect of CO2 enrichment on nutrient availability and remobilization from leaves to grains.

A reduced-tillering trait shows small but important yield gains in dryland wheat production.

Reducing the number of tillers per plant using a tiller inhibition (tin) gene has been considered as an important trait for wheat production in dryland environments. We used a spatial analysis approach with a daily time-step coupled radiation and transpiration efficiency model to simulate the impact of the reduced-tillering trait on wheat yield under different climate change scenarios across Australia's arable land. Our results show a small but consistent yield advantage of the reduced-tillering trait in the most water-limited environments both under current and likely future conditions. Our climate scenarios show that whilst elevated [CO2 ] (e[CO2 ]) alone might limit the area where the reduced-tillering trait is advantageous, the most likely climate scenario of e[CO2 ] combined with increased temperature and reduced rainfall consistently increased the area where restricted tillering has an advantage. Whilst long-term average yield advantages were small (ranged from 31 to 51 kg ha-1  year-1 ), across large dryland areas the value is large (potential cost-benefits ranged from Australian dollar 23 to 60 MIL/year). It seems therefore worthwhile to further explore this reduced-tillering trait in relation to a range of different environments and climates, because its benefits are likely to grow in future dry environments where wheat is grown around the world.

Carbon sink strength of nodules but not other organs modulates photosynthesis of faba bean (Vicia faba) grown under elevated [CO2 ] and different water supply.

Photosynthetic stimulation by elevated [CO2 ] (e[CO2 ]) may be limited by the capacity of sink organs to use photosynthates. In many legumes, N2 -fixing symbionts in root nodules provide an additional sink, so that legumes may be better able to profit from e[CO2 ]. However, drought not only constrains photosynthesis but also the size and activity of sinks, and little is known about the interaction of e[CO2 ] and drought on carbon sink strength of nodules and other organs. To compare carbon sink strength, faba bean was grown under ambient (400 ppm) or elevated (700 ppm) atmospheric [CO2 ] and subjected to well-watered or drought treatments, and then exposed to 13 C pulse-labelling using custom-built chambers to track the fate of new photosynthates. Drought decreased 13 C uptake and nodule sink strength, and this effect was even greater under e[CO2 ], and was associated with an accumulation of amino acids in nodules. This resulted in decreased N2 fixation, and increased accumulation of new photosynthates (13 C/sugars) in leaves, which in turn can feed back on photosynthesis. Our study suggests that nodule C sink activity is key to avoid sink limitation in legumes under e[CO2 ], and legumes may only be able to achieve greater C gain if nodule activity is maintained.

The water use dynamics of canola cultivars grown under elevated CO2 are linked to their leaf area development.

The 'CO2 fertilisation effect' is often predicted to be greater under drier than wetter conditions, mainly due to hypothesised early season water savings under elevated [CO2] (e[CO2]). However, water savings largely depend on the balance between CO2-induced improvement of leaf-level water use efficiency and CO2-stimulation of transpiring leaf area. The dynamics of water use during the growing season can therefore vary depending on leaf area development. Two canola (Brassica napus L.) cultivars of contrasting growth and vigour (vigorous hybrid cv. Hyola 50 and non-hybrid cv. Thumper) were grown under ambient [CO2] (a[CO2], ∼400 µmol mol-1) or e[CO2] (∼700 µmol mol-1) with two water treatments (well-watered and mild drought) in a glasshouse to investigate the interdependence of leaf area development and water use. Dynamics of water use during the growing season varied depending on [CO2] and cultivars. Early stimulation of leaf growth under e[CO2], which also depended on cultivar, overcompensated for the effect of increased leaf-level water use efficiency, so that weekly water use was greater and water depletion from soil greater under e[CO2] than a[CO2]. This result shows that the balance between leaf area and water use efficiency stimulation by e[CO2] can tip towards early depletion of available soil water, so that e[CO2] does not lead to water savings, and the 'CO2 fertilisation effect' is not greater under drier conditions.

Elevated [CO2] mitigates the effect of surface drought by stimulating root growth to access sub-soil water.

Through stimulation of root growth, increasing atmospheric CO2 concentration ([CO2]) may facilitate access of crops to sub-soil water, which could potentially prolong physiological activity in dryland environments, particularly because crops are more water use efficient under elevated [CO2] (e[CO2]). This study investigated the effect of drought in shallow soil versus sub-soil on agronomic and physiological responses of wheat to e[CO2] in a glasshouse experiment. Wheat (Triticum aestivum L. cv. Yitpi) was grown in split-columns with the top (0-30 cm) and bottom (31-60 cm; 'sub-soil') soil layer hydraulically separated by a wax-coated, root-penetrable layer under ambient [CO2] (a[CO2], ∼400 µmol mol-1) or e[CO2] (∼700 µmol mol-1) [CO2]. Drought was imposed from stem-elongation in either the top or bottom soil layer or both by withholding 33% of the irrigation, resulting in four water treatments (WW, WD, DW, DD; D = drought, W = well-watered, letters denote water treatment in top and bottom soil layer, respectively). Leaf gas exchange was measured weekly from stem-elongation until anthesis. Above-and belowground biomass, grain yield and yield components were evaluated at three developmental stages (stem-elongation, anthesis and maturity). Compared with a[CO2], net assimilation rate was higher and stomatal conductance was lower under e[CO2], resulting in greater intrinsic water use efficiency. Elevated [CO2] stimulated both above- and belowground biomass as well as grain yield, however, this stimulation was greater under well-watered (WW) than drought (DD) throughout the whole soil profile. Imposition of drought in either or both soil layers decreased aboveground biomass and grain yield under both [CO2] compared to the well-watered treatment. However, the greatest 'CO2 fertilisation effect' was observed when drought was imposed in the top soil layer only (DW), and this was associated with e[CO2]-stimulation of root growth especially in the well-watered bottom layer. We suggest that stimulation of belowground biomass under e[CO2] will allow better access to sub-soil water during grain filling period, when additional water is converted into additional yield with high efficiency in Mediterranean-type dryland agro-ecosystems. If sufficient water is available in the sub-soil, e[CO2] may help mitigating the effect of drying surface soil.

Water availability moderates N2 fixation benefit from elevated [CO2 ]: A 2-year free-air CO2 enrichment study on lentil (Lens culinaris MEDIK.) in a water limited agroecosystem.

Increased biomass and yield of plants grown under elevated [CO2 ] often corresponds to decreased grain N concentration ([N]), diminishing nutritional quality of crops. Legumes through their symbiotic N2 fixation may be better able to maintain biomass [N] and grain [N] under elevated [CO2 ], provided N2 fixation is stimulated by elevated [CO2 ] in line with growth and yield. In Mediterranean-type agroecosystems, N2 fixation may be impaired by drought, and it is unclear whether elevated [CO2 ] stimulation of N2 fixation can overcome this impact in dry years. To address this question, we grew lentil under two [CO2 ] (ambient ~400 ppm and elevated ~550 ppm) levels in a free-air CO2 enrichment facility over two growing seasons sharply contrasting in rainfall. Elevated [CO2 ] stimulated N2 fixation through greater nodule number (+27%), mass (+18%), and specific fixation activity (+17%), and this stimulation was greater in the high than in the low rainfall/dry season. Elevated [CO2 ] depressed grain [N] (-4%) in the dry season. In contrast, grain [N] increased (+3%) in the high rainfall season under elevated [CO2 ], as a consequence of greater post-flowering N2 fixation. Our results suggest that the benefit for N2 fixation from elevated [CO2 ] is high as long as there is enough soil water to continue N2 fixation during grain filling.

Benefits of increasing transpiration efficiency in wheat under elevated CO2 for rainfed regions.

Higher transpiration efficiency (TE) has been proposed as a mechanism to increase crop yields in dry environments where water availability usually limits yield. The application of a coupled radiation and TE simulation model shows wheat yield advantage of a high-TE cultivar (cv. Drysdale) over its almost identical low-TE parent line (Hartog), from about -7 to 558 kg/ha (mean 187 kg/ha) over the rainfed cropping region in Australia (221-1,351 mm annual rainfall), under the present-day climate. The smallest absolute yield response occurred in the more extreme drier and wetter areas of the wheat belt. However, under elevated CO2 conditions, the response of Drysdale was much greater overall, ranging from 51 to 886 kg/ha (mean 284 kg/ha) with the greatest response in the higher rainfall areas. Changes in simulated TE under elevated CO2 conditions are seen across Australia with notable increased areas of higher TE under a drier climate in Western Australia, Queensland and parts of New South Wales and Victoria. This improved efficiency is subtly deceptive, with highest yields not necessarily directly correlated with highest TE. Nevertheless, the advantage of Drysdale over Hartog is clear with the benefit of the trait advantage attributed to TE ranging from 102% to 118% (mean 109%). The potential annual cost-benefits of this increased genetic TE trait across the wheat growing areas of Australia (5 year average of area planted to wheat) totaled AUD 631 MIL (5-year average wheat price of AUD/260 t) with an average of 187 kg/ha under the present climate. The benefit to an individual farmer will depend on location but elevated CO2 raises this nation-wide benefit to AUD 796 MIL in a 2°C warmer climate, slightly lower (AUD 715 MIL) if rainfall is also reduced by 20%.

The relationship between transpiration and nutrient uptake in wheat changes under elevated atmospheric CO2.

The impact of elevated [CO2 ] (e[CO2 ]) on crops often includes a decrease in their nutrient concentrations where reduced transpiration-driven mass flow of nutrients has been suggested to play a role. We used two independent approaches, a free-air CO2 enrichment (FACE) experiment in the South Eastern wheat belt of Australia and a simulation study employing the agricultural production systems simulator (APSIM), to show that transpiration (mm) and nutrient uptake (g m-2 ) of nitrogen (N), potassium (K), sulfur (S), calcium (Ca), magnesium (Mg) and manganese (Mn) in wheat are correlated under e[CO2 ], but that nutrient uptake per unit water transpired is higher under e[CO2 ] than under ambient [CO2 ] (a[CO2 ]). This result suggests that transpiration-driven mass flow of nutrients contributes to decreases in nutrient concentrations under e[CO2 ], but cannot solely explain the overall decline.

The proportion of nitrate in leaf nitrogen, but not changes in root growth, are associated with decreased grain protein in wheat under elevated [CO2].

The atmospheric CO2 concentration ([CO2]) is increasing and predicted to reach ∼550ppm by 2050. Increasing [CO2] typically stimulates crop growth and yield, but decreases concentrations of nutrients, such as nitrogen ([N]), and therefore protein, in plant tissues and grains. Such changes in grain composition are expected to have negative implications for the nutritional and economic value of grains. This study addresses two mechanisms potentially accountable for the phenomenon of elevated [CO2]-induced decreases in [N]: N uptake per unit length of roots as well as inhibition of the assimilation of nitrate (NO3-) into protein are investigated and related to grain protein. We analysed two wheat cultivars from a similar genetic background but contrasting in agronomic features (Triticum aestivum L. cv. Scout and Yitpi). Plants were field-grown within the Australian Grains Free Air CO2 Enrichment (AGFACE) facility under two atmospheric [CO2] (ambient, ∼400ppm, and elevated, ∼550ppm) and two water treatments (rain-fed and well-watered). Aboveground dry weight (ADW) and root length (RL, captured by a mini-rhizotron root growth monitoring system), as well as [N] and NO3- concentrations ([NO3-]) were monitored throughout the growing season and related to grain protein at harvest. RL generally increased under e[CO2] and varied between water supply and cultivars. The ratio of total aboveground N (TN) taken up per RL was affected by CO2 treatment only later in the season and there was no significant correlation between TN/RL and grain protein concentration across cultivars and [CO2] treatments. In contrast, a greater percentage of N remained as unassimilated [NO3-] in the tissue of e[CO2] grown crops (expressed as the ratio of NO3- to total N) and this was significantly correlated with decreased grain protein. These findings suggest that e[CO2] directly affects the nitrate assimilation capacity of wheat with direct negative implications for grain quality.

Elevated atmospheric [CO2 ] can dramatically increase wheat yields in semi-arid environments and buffer against heat waves.

Wheat production will be impacted by increasing concentration of atmospheric CO2 [CO2 ], which is expected to rise from about 400 µmol mol(-1) in 2015 to 550 µmol mol(-1) by 2050. Changes to plant physiology and crop responses from elevated [CO2 ] (e[CO2 ]) are well documented for some environments, but field-level responses in dryland Mediterranean environments with terminal drought and heat waves are scarce. The Australian Grains Free Air CO2 Enrichment facility was established to compare wheat (Triticum aestivum) growth and yield under ambient (~370 µmol(-1) in 2007) and e[CO2 ] (550 µmol(-1) ) in semi-arid environments. Experiments were undertaken at two dryland sites (Horsham and Walpeup) across three years with two cultivars, two sowing times and two irrigation treatments. Mean yield stimulation due to e[CO2 ] was 24% at Horsham and 53% at Walpeup, with some treatment responses greater than 70%, depending on environment. Under supplemental irrigation, e[CO2 ] stimulated yields at Horsham by 37% compared to 13% under rainfed conditions, showing that water limited growth and yield response to e[CO2 ]. Heat wave effects were ameliorated under e[CO2 ] as shown by reductions of 31% and 54% in screenings and 10% and 12% larger kernels (Horsham and Walpeup). Greatest yield stimulations occurred in the e[CO2 ] late sowing and heat stressed treatments, when supplied with more water. There were no clear differences in cultivar response due to e[CO2 ]. Multiple regression showed that yield response to e[CO2 ] depended on temperatures and water availability before and after anthesis. Thus, timing of temperature and water and the crop's ability to translocate carbohydrates to the grain postanthesis were all important in determining the e[CO2 ] response. The large responses to e[CO2 ] under dryland conditions have not been previously reported and underscore the need for field level research to provide mechanistic understanding for adapting crops to a changing climate.

Expression patterns of C- and N-metabolism related genes in wheat are changed during senescence under elevated CO2 in dry-land agriculture.

Projected climatic impacts on crop yield and quality, and increased demands for production, require targeted research to optimise nutrition of crop plants. For wheat, post-anthesis carbon and nitrogen remobilisation from vegetative plant parts and translocation to grains directly affects grain carbon (C), nitrogen (N) and protein levels. We analysed the influence of increased atmospheric CO2 on the expression of genes involved in senescence, leaf carbohydrate and nitrogen metabolism and assimilate transport in wheat under field conditions (Australian Grains Free Air CO2 Enrichment; AGFACE) over a time course from anthesis to maturity, the key period for grain filling. Wheat grown under CO2 enrichment had lower N concentrations and a tendency towards greater C/N ratios. A general acceleration of the senescence process by elevated CO2 was not confirmed. The expression patterns of genes involved in carbohydrate metabolism, nitrate reduction and metabolite transport differed between CO2 treatments, and this CO2 effect was different between pre-senescence and during senescence. The results suggest up-regulation of N remobilisation and down-regulation of C remobilisation during senescence under elevated CO2, which is consistent with greater grain N-sink strength of developing grains.

Intraspecific variation in leaf growth of wheat (Triticum aestivum) under Australian Grain Free Air CO2 Enrichment (AGFACE): is it regulated through carbon and/or nitrogen supply?

Underlying physiological mechanisms of intraspecific variation in growth response to elevated CO2 concentration [CO2] were investigated using two spring wheat (Triticum aestivum L.) cultivars: Yitpi and H45. Leaf blade elongation rate (LER), leaf carbon (C), nitrogen (N) in the expanding leaf blade (ELB, sink) and photosynthesis (A) and C and N status in the last fully expanded leaf blade (LFELB, source) were measured. Plants were grown at ambient [CO2] (~384µmolmol-1) and elevated [CO2] (~550µmolmol-1) in the Australian Grains Free Air CO2 Enrichment facility. Elevated [CO2] increased leaf area and total dry mass production, respectively, by 42 and 53% for Yitpi compared with 2 and 13% for H45. Elevated [CO2] also stimulated the LER by 36% for Yitpi compared with 5% for H45. Yitpi showed a 99% increase in A at elevated [CO2] but no A stimulation was found for H45. There was a strong correlation (r2=0.807) between LER of the ELB and soluble carbohydrate concentration in LFELB. In ELB, the highest spatial N concentration was observed in the cell division zone, where N concentrations were 67.3 and 60.6mg g-1 for Yitpi compared with 51.1 and 39.2mg g-1 for H45 at ambient and elevated [CO2]. In contrast, C concentration increased only in the cell division and cell expansion zone of the ELB of Yitpi. These findings suggest that C supply from the source (LFELB) is cultivar dependent and well correlated with LER, leaf area expansion and whole-plant growth response to elevated [CO2].

Intraspecific variation in growth and yield response to elevated CO2 in wheat depends on the differences of leaf mass per unit area.

In order to investigate the underlying physiological mechanism of intraspecific variation in plant growth and yield response to elevated CO2 concentration [CO2], seven cultivars of spring wheat (Triticum aestivum L.) were grown at either ambient [CO2] (~384µmolmol-1) or elevated [CO2] (700µmolmol-1) in temperature controlled glasshouses. Grain yield increased under elevated [CO2] by an average of 38% across all seven cultivars, and this was correlated with increases in both spike number (productive tillers) (r=0.868) and aboveground biomass (r=0.942). Across all the cultivars, flag leaf photosynthesis rate (A) increased by an average of 57% at elevated [CO2]. The response of A to elevated [CO2] ranged from 31% (in cv. H45) to 75% (in cv. Silverstar). Only H45 showed A acclimation to elevated [CO2], which was characterised by lower maximum Rubisco carboxylation efficiency, maximum electron transport rate and leaf N concentration. Leaf level traits responsible for plant growth, such as leaf mass per unit area (LMA), carbon (C), N content on an area basis ([N]LA) and the C:N increased at elevated [CO2]. LMA stimulation ranged from 0% to 85% and was clearly associated with increased [N]LA. Both of these traits were positively correlated with grain yield, suggesting that differences in LMA play an important role in determining the grain yield response to elevated [CO2]. Thus increased LMA can be used as a new trait to select cultivars for a future [CO2]-rich atmosphere.

Will intra-specific differences in transpiration efficiency in wheat be maintained in a high CO2 world? A FACE study.

This study evaluates whether the target breeding trait of superior leaf level transpiration efficiency is still appropriate under increasing carbon dioxide levels of a future climate using a semi-arid cropping system as a model. Specifically, we investigated whether physiological traits governing leaf level transpiration efficiency, such as net assimilation rates (A(net)), stomatal conductance (g(s)) or stomatal sensitivity were affected differently between two Triticum aestivum L. cultivars differing in transpiration efficiency (cv. Drysdale, superior; cv. Hartog, low). Plants were grown under Free Air Carbon dioxide Enrichment (FACE, approximately 550 µmol mol⁻¹ or ambient CO2 concentrations (approximately 390 µmol mol⁻¹). Mean A(net) (approximately 15% increase) and gs (approximately 25% decrease) were less affected by elevated [CO2] than previously found in FACE-grown wheat (approximately 25% increase and approximately 32% decrease, respectively), potentially reflecting growth in a dry-land cropping system. In contrast to previous FACE studies, analyses of the Ball et al. model revealed an elevated [CO2] effect on the slope of the linear regression by 12% indicating a decrease in stomatal sensitivity to the combination of [CO2], photosynthesis rate and humidity. Differences between cultivars indicated greater transpiration efficiency for Drysdale with growth under elevated [CO2] potentially increasing the response of this trait. This knowledge adds valuable information for crop germplasm improvement for future climates.