Marta S. Lopes

Partitioning of assimilates to deeper roots is associated with cooler canopies and increased yield under drought in wheat

Dehydration avoidance through cooler canopy temperature (CT) has been shown to explain over 60% yield variation in a random progeny derived from a Seri/Babax cross. A near ‘isomorphic’ subset of Seri/Babax progeny and parents encompassing a restricted range of height and phenology were used for detailed characterisation of drought-adaptive trait expression under contrasting water regimes. Under drought, five of the six progeny out yielded the best parent Babax by up to 35%. The main physiological attributes associated with drought adaptation were increased root dry weight at depth, transpiration rate – evidenced by grain carbon isotope discrimination (Δ13C) – grain filling duration and decreased CT during grain filling. Furthermore, increased root mass at depth was associated with reduced levels of stem water soluble carbohydrates (WSC) when comparing genotypes. It is concluded that differences in rooting depth expressed among iso-morphic wheat sister lines explains superior adaptation to drought. These effects can be detected in season using remote sensing. In addition, the data suggest that accumulation of stem carbohydrates and deep rooting may be two alternative strategies for adapting to drought stress, the latter being beneficial where water is available at depth.

Enhancing drought tolerance in C4 crops

Adaptation to abiotic stresses is a quantitative trait controlled by many different genes. Enhancing the tolerance of crop plants to abiotic stresses such as drought has therefore proved to be somewhat elusive in terms of plant breeding. While many C(4) species have significant agronomic importance, most of the research effort on improving drought tolerance has focused on maize. Ideally, drought tolerance has to be achieved without penalties in yield potential. Possibilities for success in this regard are highlighted by studies on maize hybrids performed over the last 70 years that have demonstrated that yield potential and enhanced stress tolerance are associated traits. However, while our understanding of the molecular mechanisms that enable plants to tolerate drought has increased considerably in recent years, there have been relatively few applications of DNA marker technologies in practical C(4) breeding programmes for improved stress tolerance. Moreover, until recently, targeted approaches to drought tolerance have concentrated largely on shoot parameters, particularly those associated with photosynthesis and stay green phenotypes, rather than on root traits such as soil moisture capture for transpiration, root architecture, and improvement of effective use of water. These root traits are now increasingly considered as important targets for yield improvement in C(4) plants under drought stress. Similarly, the molecular mechanisms underpinning heterosis have considerable potential for exploitation in enhancing drought stress tolerance. While current evidence points to the crucial importance of root traits in drought tolerance in C(4) plants, shoot traits may also be important in maintaining high yields during drought.

Stay-green in spring wheat can be determined by spectral reflectance measurements (normalized difference vegetation index) independently from phenology

The green area displayed by a crop is a good indicator of its photosynthetic capacity, while chlorophyll retention or ‘stay-green’ is regarded as a key indicator of stress adaptation. Remote-sensing methods were tested to estimate these parameters in diverse wheat genotypes under different growing conditions. Two wheat populations (a diverse set of 294 advanced lines and a recombinant inbred line population of 169 sister lines derived from the cross between Seri and Babax) were grown in Mexico under three environments: drought, heat, and heat combined with drought. In the two populations studied here, a moderate heritable expression of stay-green was found–when the normalized difference vegetation index (NDVI) at physiological maturity was estimated using the regression of NDVI over time from the mid-stages of grain-filling to physiological maturity–and for the rate of senescence during the same period. Under heat and heat combined with drought environments, stay-green calculated as NDVI at physiological maturity and the rate of senescence, showed positive and negative correlations with yield, respectively. Moreover, stay-green calculated as an estimation of NDVI at physiological maturity and the rate of senescence regressed on degree days give an independent measurement of stay-green without the confounding effect of phenology. On average, in both populations under heat and heat combined with drought environments CTgf and stay-green variables accounted for around 30% of yield variability in multiple regression analysis. It is concluded that stay-green traits may provide cumulative effects, together with other traits, to improve adaptation under stress further.

Exploiting genetic diversity from landraces in wheat breeding for adaptation to climate change

Climate change has generated unpredictability in the timing and amount of rain, as well as extreme heat and cold spells that have affected grain yields worldwide and threaten food security. Sources of specific adaptation related to drought and heat, as well as associated breeding of genetic traits, will contribute to maintaining grain yields in dry and warm years. Increased crop photosynthesis and biomass have been achieved particularly through disease resistance and healthy leaves. Similarly, sources of drought and heat adaptation through extended photosynthesis and increased biomass would also greatly benefit crop improvement. Wheat landraces have been cultivated for thousands of years under the most extreme environmental conditions. They have also been cultivated in lower input farming systems for which adaptation traits, particularly those that increase the duration of photosynthesis, have been conserved. Landraces are a valuable source of genetic diversity and specific adaptation to local environmental conditions according to their place of origin. Evidence supports the hypothesis that landraces can provide sources of increased biomass and thousand kernel weight, both important traits for adaptation to tolerate drought and heat. Evaluation of wheat landraces stored in gene banks with highly beneficial untapped diversity and sources of stress adaptation, once characterized, should also be used for wheat improvement. Unified development of databases and promotion of data sharing among physiologists, pathologists, wheat quality scientists, national programmes, and breeders will greatly benefit wheat improvement for adaptation to climate change worldwide.

Integration of phenotyping and genetic platforms for a better understanding of wheat performance under drought

Identifying markers for physiological traits of proven value in breeding, especially ones that are consistent across environments with different patterns of stress, strengthens the toolkit to increase confidence in the value and delivery from physiological breeding. To identify markers relevant to drought adaptation, this review will highlight the importance of development and implementation of robust and repeatable phenotyping that is relevant to the different target drought types, and practical examples of managed environment facilities in Australia and Mexico are given. These facilities can be used as models to: (i) improve reliability and consistency of environments and genetic responses to the environment at a global scale; (ii) improve the capacity to deliver quantitative trait loci (QTLs) as user-friendly markers for enriching populations; and (iii) illustrate the use of populations with a narrow range of variation for phenology allowing the identification of QTLs for drought-adaptive traits. However, the importance of further optimizing phenology and plant height at a global scale is highlighted. Finally, the impact of physiological trait-based crossing is demonstrated and supports the need for urgent development of robust genetic markers.

Association mapping and nucleotide sequence variation in five drought tolerance candidate genes in spring wheat

Functional markers are needed for key genes involved in drought tolerance to improve selection for crop yield under moisture stress conditions. The objectives of this study were to (i) characterize five drought tolerance candidate genes, namely dehydration responsive element binding 1A (DREB1A), enhanced response to abscisic acid (ERA1‐B and ERA1‐D), and fructan 1‐exohydrolase (1‐FEH‐A and 1‐FEH‐B), in wheat (Triticum aestivum L.) for nucleotide and haplotype diversity, Tajimas D value, and linkage disequilibrium (LD) and (ii) associate within‐gene single nucleotide polymorphisms (SNPs) with phenotypic traits in a spring wheat association mapping panel (n = 126). Field trials were grown under contrasting moisture regimes in Greeley, CO, and Melkassa, Ethiopia, in 2010 and 2011. Genome‐specific amplification and DNA sequence analysis of the genes identified SNPs and revealed differences in nucleotide and haplotype diversity, Tajimas D, and patterns of LD. DREB1A showed associations (false discovery rate adjusted probability value = 0.1) with normalized difference vegetation index, heading date, biomass, and spikelet number. Both ERA1‐A and ERA1‐B were associated with harvest index, flag leaf width, and leaf senescence. 1‐FEH‐A was associated with grain yield, and 1‐FEH‐B was associated with thousand kernel weight and test weight. If validated in relevant genetic backgrounds, the identified marker–trait associations may be applied to functional marker‐assisted selection.

Acclimation to high CO2 in maize is related to water status and dependent on leaf rank

The responses of C(3) plants to rising atmospheric CO(2) levels are considered to be largely dependent on effects exerted through altered photosynthesis. In contrast, the nature of the responses of C(4) plants to high CO(2) remains controversial because of the absence of CO(2) -dependent effects on photosynthesis. In this study, the effects of atmospheric CO(2) availability on the transcriptome, proteome and metabolome profiles of two ranks of source leaves in maize (Zea mays L.) were studied in plants grown under ambient CO(2) conditions (350 +/- 20 µL L(-1) CO(2) ) or with CO(2) enrichment (700 +/- 20 µL L(-1) CO(2) ). Growth at high CO(2) had no effect on photosynthesis, photorespiration, leaf C/N ratios or anthocyanin contents. However, leaf transpiration rates, carbohydrate metabolism and protein carbonyl accumulation were altered at high CO(2) in a leaf-rank specific manner. Although no significant CO(2) -dependent changes in the leaf transcriptome were observed, qPCR analysis revealed that the abundance of transcripts encoding a Bowman-Birk protease inhibitor and a serpin were changed by the growth CO(2) level in a leaf rank specific manner. Moreover, CO(2) -dependent changes in the leaf proteome were most evident in the oldest source leaves. Small changes in water status may be responsible for the observed responses to high CO(2,) particularly in the older leaf ranks.

Modelling and genetic dissection of staygreen under heat stress

Key messageStaygreen traits are associated with heat tolerance in bread wheat. QTL for staygreen and related traits were identified across the genome co-located with agronomic and physiological traits associated to plant performance under heat stress.AbstractPlant chlorophyll retention—staygreen—is considered a valuable trait under heat stress. Five experiments with the Seri/Babax wheat mapping population were sown in Mexico under hot-irrigated environments. Normalized difference vegetation index (NDVI) during plant growth was measured regularly and modelled to capture the dynamics of plant greenness decay, including staygreen (Stg) at physiological maturity which was estimated by regression of NDVI during grainfilling. The rate of senescence, the percentage of plant greenness decay, and the area under the curve were also estimated based on NDVI measurements. While Stg and the best fitted curve were highly environment dependent, both traits showed strong (positive for Stg) correlations with yield, grainfilling rates, and extended grainfilling periods, while associations with kernel number and kernel weight were weak. Stg expression was largely dependent on rate of senescence which was related to the pattern of the greenness decay curve and the initial NDVI. QTL analyses revealed a total of 44 loci across environments linked to Stg and related traits, distributed across the genome, with the strongest and most repeatable effects detected on chromosomes 1B, 2A, 2B, 4A, 4B and 7D. Of these, some were common with regions controlling phenology but independent regions were also identified. The co-location of QTL for Stg and performance traits in this study confirms that the staygreen phenotype is a useful trait for productivity enhancement in hot-irrigated environments.

Genomic Prediction with Pedigree and Genotype × Environment Interaction in Spring Wheat Grown in South and West Asia, North Africa, and Mexico

Developing genomic selection (GS) models is an important step in applying GS to accelerate the rate of genetic gain in grain yield in plant breeding. In this study, seven genomic prediction models under two cross-validation (CV) scenarios were tested on 287 advanced elite spring wheat lines phenotyped for grain yield (GY), thousand-grain weight (GW), grain number (GN), and thermal time for flowering (TTF) in 18 international environments (year-location combinations) in major wheat-producing countries in 2010 and 2011. Prediction models with genomic and pedigree information included main effects and interaction with environments. Two random CV schemes were applied to predict a subset of lines that were not observed in any of the 18 environments (CV1), and a subset of lines that were not observed in a set of the environments, but were observed in other environments (CV2). Genomic prediction models, including genotype × environment (G×E) interaction, had the highest average prediction ability under the CV1 scenario for GY (0.31), GN (0.32), GW (0.45), and TTF (0.27). For CV2, the average prediction ability of the model including the interaction terms was generally high for GY (0.38), GN (0.43), GW (0.63), and TTF (0.53). Wheat lines in site-year combinations in Mexico and India had relatively high prediction ability for GY and GW. Results indicated that prediction ability of lines not observed in certain environments could be relatively high for genomic selection when predicting G×E interaction in multi-environment trials.

Traits associated with winter wheat grain yield in Central and West Asia

Improved adaptation of winter wheat (Triticum aestivum L.) to drought and heat may be influenced by days to heading, plant height, biomass, canopy temperature (CT) at grain filling, and rate of senescence. This study shows that, under supplemental irrigation or rainfed conditions, days to heading and plant height together explain up to 68% of grain yield (GY) variation, and these associations were further confirmed in several locations across West and Central Asia. Days to heading can be slightly reduced below that of check line Karahan to further improve GY while avoiding the effect of late frosts. Plant height has been decreased in recent germplasm, but further reductions below that of check line Karahan could still improve GY in a wide range of environments. However, in Iranian sites, taller genotypes showed better adaptation with higher biomass and increased reserves for grain filling. Canopy temperature and rate senescence were not associated with GY. A normalized difference vegetation index, used to estimate biomass (Feekes stages 4-5), had intermediate heritability across environments and correlated positively with GY under low plant density and should be explored further as a tool for early selection.