Improving Rice Zinc Biofortification Success Rates Through Genetic and Crop Management Approaches in a Changing Environment.

Though rice is the predominant source of energy and micronutrients for more than half of the world population, it does not provide enough zinc (Zn) to match human nutritional requirements. Moreover, climate change, particularly rising atmospheric carbon dioxide concentration, reduces the grain Zn concentration. Therefore, rice biofortification has been recognized as a key target to increase the grain Zn concentration to address global Zn malnutrition. Major bottlenecks for Zn biofortification in rice are identified as low Zn uptake, transport and loading into the grain; however, environmental and genetic contributions to grain Zn accumulation in rice have not been fully explored. In this review, we critically analyze the key genetic, physiological and environmental factors that determine Zn uptake, transport and utilization in rice. We also explore the genetic diversity of rice germplasm to develop new genetic tools for Zn biofortification. Lastly, we discuss the strategic use of Zn fertilizer for developing biofortified rice.

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.

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].

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.

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.

Next generation of elevated [CO2] experiments with crops: a critical investment for feeding the future world.

A rising global population and demand for protein-rich diets are increasing pressure to maximize agricultural productivity. Rising atmospheric [CO(2)] is altering global temperature and precipitation patterns, which challenges agricultural productivity. While rising [CO(2)] provides a unique opportunity to increase the productivity of C(3) crops, average yield stimulation observed to date is well below potential gains. Thus, there is room for improving productivity. However, only a fraction of available germplasm of crops has been tested for CO(2) responsiveness. Yield is a complex phenotypic trait determined by the interactions of a genotype with the environment. Selection of promising genotypes and characterization of response mechanisms will only be effective if crop improvement and systems biology approaches are closely linked to production environments, that is, on the farm within major growing regions. Free air CO(2) enrichment (FACE) experiments can provide the platform upon which to conduct genetic screening and elucidate the inheritance and mechanisms that underlie genotypic differences in productivity under elevated [CO(2)]. We propose a new generation of large-scale, low-cost per unit area FACE experiments to identify the most CO(2)-responsive genotypes and provide starting lines for future breeding programmes. This is necessary if we are to realize the potential for yield gains in the future.

Physiological mechanisms of tolerance to high boron concentration in Brassica rapa.

Tolerance to high boron concentration in Brassica rapa was primarily due to low net boron uptake by the roots. However, in the two tolerant genotypes, 39-43% of boron uptake was retained in the tap roots, which limited boron accumulation in the leaves, and also contributed to boron tolerance. In the sensitive genotype, 99% of the increase in boron uptake caused by high soil boron accumulated in the leaves, particularly in the leaf margins. Despite higher transpiration rates, lower net boron uptake occurred in the tolerant genotypes. This result cannot be explained by passive boron uptake alone. Active boron efflux was probably responsible for differences in net boron uptake among tolerant and sensitive genotypes. Boron concentration was much lower in the cell walls than in the cell sap of leaves, indicating that storage of boron in the cell walls was not a tolerance mechanism. Despite high boron concentrations in the leaf symplasm, rates of photosynthesis, transpiration and growth were almost unaffected in the tolerant genotypes. The results demonstrate that boron tolerance in Brassica rapa involves boron exclusion at the root level, boron partitioning away from leaves and, as boron accumulates in leaves despite the first two mechanisms, boron tolerance of the leaf tissue itself.