Saman Seneweera

Increasing CO2 threatens human nutrition

Dietary deficiencies of zinc and iron are a substantial global public health problem. An estimated two billion people suffer these deficiencies, causing a loss of 63 million life-years annually. Most of these people depend on C3 grains and legumes as their primary dietary source of zinc and iron. Here we report that C3 grains and legumes have lower concentrations of zinc and iron when grown under field conditions at the elevated atmospheric CO2 concentration predicted for the middle of this century. C3 crops other than legumes also have lower concentrations of protein, whereas C4 crops seem to be less affected. Differences between cultivars of a single crop suggest that breeding for decreased sensitivity to atmospheric CO2 concentration could partly address these new challenges to global health.

Changes in source-sink relations during development influence photosynthetic acclimation of rice to free air CO2 enrichment (FACE)

Relationships between photosynthetic acclimation and changes in the balance between source-sink supply and demand of carbon (C) and nitrogen (N) were tested using rice (Oryza sativa L. cv. Akitakomachi). Plants were field-grown in northern Japan at ambient CO2 partial pressure [p(CO2)] or free air CO2 enrichment (FACE; p(CO2) ~ 26-32 Pa above ambient) with low, medium or high N supplies. Leaf CO2 assimilation rates (A) and biochemical parameters were measured at 32-36 (eighth leaf) and 76-80 (flag leaf) d after transplanting, representing stages with a contrasting balance between C and N supply and demand in sources and sinks. Acclimation due to FACE was pronounced in flag leaves at each N supply. This was not fully accounted for by reductions in leaf N concentrations, because A/N and Vcmax/N were lower in FACE-grown flag leaves. Acclimation did not occur in the eighth leaf, and A/N and Vcmax/N was not significantly increased in FACE-grown leaves. Soluble protein / sucrose and amino acid / sucrose concentrations decreased under FACE, whereas sucrose phosphate synthase protein levels increased. At flag leaf stage, there was a discrepancy between the demand and supply of N, which was resolved by enhanced leaf N remobilization, associated with the lower Rubisco concentrations under FACE. In contrast to the early growth stage, enhanced growth of rice plants was accompanied by increased plant N uptake in FACE. We conclude that photosynthetic acclimation in flag leaves occurs under FACE because there is a large demand for N for reproductive development, relative to supply of N from root uptake and remobilization from leaves.

Accelerated early growth of rice at elevated CO2 : Is it related to developmental changes in the shoot apex?

The influence of elevated CO2 on the development of the shoot apex and on subsequent vegetative growth and grain yield was investigated using rice (Oryza sativa L. cv Jarrah) grown in flooded soil at either 350 or 700 [mu]L CO2 L-1. At 8 d after planting (DAP), elevated CO2 increased the height and diameter of the apical dome and lengths of leaf primordia and tiller buds but had no effect on their numbers. By 16 DAP, there were five tiller buds in the apex at 700 [mu]L CO2 L-1 compared with only three tiller buds at 350 [mu]L CO2 L-1. These changes in development of the shoot apex at high CO2 were forerunners to faster development of the vegetative shoot at elevated CO2 between 11 and 26 DAP as evidenced by increases in the relative growth rates of the shoot and tillers. Accelerated development at high CO2 was responsible for the 42% increase in tiller number at the maximum tillering stage and the 57% enhancement of grain yield at the final harvest. The link between high CO2 effects on development during the first 15 DAP and final tiller number and grain yield was demonstrated by delaying exposure of plants to high CO2 for 15 d. The delay totally inhibited the tillering response to high CO2, and the increase in grain yield of 20% arose from a greater number of grains per panicle. Consequently, it can be concluded that accelerated development in the shoot apex early in development is crucial for obtaining maximum increases in grain yield at elevated atmospheric CO2 concentrations.

Diurnal Regulation of Leaf Blade Elongation in Rice by CO2 (Is it Related to Sucrose-Phosphate Synthase Activity?).

The relationship between leaf blade elongation rates (LER) and sucrose-phosphate synthase (SPS) activity was investigated at different times during ontogeny of rice (Oryza sativa L. cv Jarrah) grown in flooded soil at either 350 or 700 [mu]L CO2 L-1. High CO2 concentrations increased LER of expanding blades and in vivo activity (Vlimiting) SPS activity of expanded blades during the early vegetative stage (21 d after planting [DAP]), when tiller number was small and growing blades were strong carbohydrate sinks. Despite a constant light environment, there was a distinct diurnal pattern in LER, Vlimiting SPS activity, and concentration of soluble sugars, with an increase in the early part of the light period and a decrease later in the light period. The strong correlation (r = 0.65) between LER and Vlimiting SPS activity over the diurnal cycle indicated that SPS activity played an important role in controlling blade growth. The higher Vlimiting SPS activity at elevated CO2 at 21 DAP was caused by an increase in the activation state of the enzyme rather than an increase in Vmax. Fructose and glucose accumulated to a greater extent than sucrose at high CO2 and may have been utilized for synthesis of cell-wall components, contributing to higher specific leaf weight. By the mid-tillering stage (42 DAP), CO2 enrichment enhanced Vlimiting and Vmax activities of source blades. Nevertheless, LER was depressed by high CO2, probably because tillers were stronger carbohydrate sinks than growing blades.

Glyphosate resistance of C3 and C4 weeds under rising atmospheric CO2

The present paper reviews current knowledge on how changes of plant metabolism under elevated CO2 concentrations (e[CO2]) can affect the development of the glyphosate resistance of C3 and C4 weeds. Among the chemical herbicides, glyphosate, which is a non-selective and post-emergence herbicide, is currently the most widely used herbicide in global agriculture. As a consequence, glyphosate resistant weeds, particularly in major field crops, are a widespread problem and are becoming a significant challenge to future global food production. Of particular interest here it is known that the biochemical processes involved in photosynthetic pathways of C3 and C4 plants are different, which may have relevance to their competitive development under changing environmental conditions. It has already been shown that plant anatomical, morphological, and physiological changes under e[CO2] can be different, based on (i) the plant’s functional group, (ii) the available soil nutrients, and (iii) the governing water status. In this respect, C3 species are likely to have a major developmental advantage under a CO2 rich atmosphere, by being able to capitalize on the overall stimulatory effect of e[CO2]. For example, many tropical weed grass species fix CO2 from the atmosphere via the C4 photosynthetic pathway, which is a complex anatomical and biochemical variant of the C3 pathway. Thus, based on our current knowledge of CO2 fixing, it would appear obvious that the development of a glyphosate-resistant mechanism would be easier under an e[CO2] in C3 weeds which have a simpler photosynthetic pathway, than for C4 weeds. However, notwithstanding this logical argument, a better understanding of the biochemical, genetic, and molecular measures by which plants develop glyphosate resistance and how e[CO2] affects these measures will be important before attempting to innovate sustainable technology to manage the glyphosate-resistant evolution of weeds under e[CO2]. Such information will be of essential in managing weed control by herbicide use, and to thus ensure an increase in global food production in the event of increased atmospheric [CO2] levels.

Improving yield potential in crops under elevated CO2: Integrating the photosynthetic and nitrogen utilization efficiencies

Increasing crop productivity to meet burgeoning human food demand is challenging under changing environmental conditions. Since industrial revolution atmospheric CO2 levels have linearly increased. Developing crop varieties with increased utilization of CO2 for photosynthesis is an urgent requirement to cope with the irreversible rise of atmospheric CO2 and achieve higher food production. The primary effects of elevated CO2 levels in most crop plants, particularly C3 plants, include increased biomass accumulation, although initial stimulation of net photosynthesis rate is only temporal and plants fail to sustain the maximal stimulation, a phenomenon known as photosynthesis acclimation. Despite this acclimation, grain yield is known to marginally increase under elevated CO2. The yield potential of C3 crops is limited by their capacity to exploit sufficient carbon. The “C fertilization” through elevated CO2 levels could potentially be used for substantial yield increase. Rubisco is the rate-limiting enzyme in photosynthesis and its activity is largely affected by atmospheric CO2 and nitrogen availability. In addition, maintenance of the C/N ratio is pivotal for various growth and development processes in plants governing yield and seed quality. For maximizing the benefits of elevated CO2, raising plant nitrogen pools will be necessary as part of maintaining an optimal C/N balance. In this review, we discuss potential causes for the stagnation in yield increases under elevated CO2 levels and explore possibilities to overcome this limitation by improved photosynthetic capacity and enhanced nitrogen use efficiency. Opportunities of engineering nitrogen uptake, assimilatory, and responsive genes are also discussed that could ensure optimal nitrogen allocation toward expanding source and sink tissues. This might avert photosynthetic acclimation partially or completely and drive for improved crop production under elevated CO2 levels.

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.

Effects of elevated CO2 on plant growth and nutrient partitioning of rice (Oryza sativa L.) at rapid tillering and physiological maturity

Abstract This work investigates the relationship between plant growth, grain yield, nutrient acquisition and partitioning in rice (Oryza sativa L.) under elevated CO2. Plants were grown hydroponically in growth chambers with a 12-h photoperiod at either 370 or 700 µmol CO2 mol−1 concentration. Plant dry mass (DM), grain yield and macro- and micronutrient concentrations of vegetative organs and grains were determined. Elevated CO2 increased biomass at tillering, and this was largely due to an increase in root mass by 160%. Elevated CO2 had no effect on total nutrient uptake (N, P, K, Mg and Ca). However, nutrient partitioning among organs was significantly altered. N partitioning to leaf blades was significantly decreased, whereas the N partitioning into the leaf sheaths and roots was increased. Nutrient use efficiency of N, P, K, and Mg in all organs was significantly increased at elevated CO2. At harvest maturity, grain yield was increased by 27% at elevated CO2 while grain (protein) concentration was decreased by a similar magnitude (28%), suggesting that critical nutrient requirements for rice might need to be reassessed with global climate change.

Canopy development and hydraulic function in Eucalyptus tereticornis grown in drought in CO2-enriched atmospheres

We report on the relationship between growth, partitioning of shoot biomass and hydraulic development of Eucalyptus tereticornis Sm. grown in glasshouses for six months. Close coordination of stem vascular capacity and shoot architecture is vital for survival of eucalypts, especially as developing trees are increasingly subjected to spasmodic droughts and rising atmospheric CO2 levels. Trees were exposed to constant soil moisture deficits in 45 L pots (30-50% below field capacity), while atmospheric CO2 was raised to 700 μL CO2 L-1 in matched glasshouses using a hierarchical, multi-factorial design. Enrichment with CO2 stimulated shoot growth rates for 12-15 weeks in well-watered trees but after six months of CO2 enrichment, shoot biomasses were not significantly heavier (30% stimulation) in ambient conditions. By contrast, constant drought arrested shoot growth after 20 weeks under ambient conditions, whereas elevated CO2 sustained growth in drought and ultimately doubled the shoot biomass relative to ambient conditions. These growth responses were achieved through an enhancement of lateral branching up to 8-fold due to CO2 enrichment. In spite of larger transpiring canopies, CO2 enrichment also improved the daytime water status of leaves of droughted trees. Stem xylem development was highly regulated, with vessels per unit area and cross sectional area of xylem vessels in stems correlated inversely across all treatments. Furthermore, vessel numbers related to the numbers of leaves on lateral branches, broadly supporting predictions arising from Pipe Model Theory that the area of conducting tissue should correlate with leaf area. Diminished water use of trees in drought coincided with a population of narrower xylem vessels, constraining hydraulic capacity of stems. Commensurate with the positive effects of elevated CO2 on growth, development and leaf water relations of droughted trees, the capacity for long-distance water transport also increased.

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.