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Research into crop yield and resilience has underpinned global food security, evident in yields tripling in the past 5 decades. The challenges that global agriculture now faces are not just to feed 10+ billion people within a generation, but to do so under a harsher, more variable, and less predictable climate, and in many cases with less water, more expensive inputs, and declining soil quality. The challenges of climate change are not simply to breed for a "hotter drier climate," but to enable resilience to floods and droughts and frosts and heat waves, possibly even within a single growing season. How well we prepare for the coming decades of climate variability will depend on our ability to modify current practices, innovate with novel breeding methods, and communicate and work with farming communities to ensure viability and profitability. Here we define how future climates will impact farming systems and growing seasons, thereby identifying the traits and practices needed and including exemplars being implemented and developed. Critically, this review will also consider societal perspectives and public engagement about emerging technologies for climate resilience, with participatory approaches presented as the best approach., (© American Society of Plant Biologists 2022. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Deployment of the Rht-B1b and Rht-D1b dwarfing genes helped facilitate the Green Revolution to increase wheat yields globally. Much is known of the influence of these genes on plant height and agronomic performance, but not of their effects on root architecture. We assessed 29 near-isogenic lines (NILs) representing 11 Green Revolution and alternative dwarfing genes across multiple genetic backgrounds for root architecture characteristics in controlled and field environments. Genetic background did not influence plant height, but had a small and significant (P<0.05) effect on root architecture. All dwarfing gene NILs were significantly (P<0.01) shorter compared with tall controls. The Green Revolution Rht-B1b and Rht-D1b sometimes had longer seedling roots but were not different from their respective tall controls for root depth in the field. The Rht8, Rht12, and Rht18 dwarfing gene NILs produced long seminal roots in seedling pouches, and a greater maximum rooting depth (MRD) and root penetration rate (RPR) in the field. Genotypic increases in MRD and RPR were strongly correlated with increased harvest index and grain yield, particularly in dry environments. Careful root phenotyping highlights the potential of novel dwarfing genes for wheat genetic improvement under water-limited conditions., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Key Message: Greater embryo size in a large and carefully phenotyped mapping population was genetically associated with a greater number of longer seminal roots to increase grain yield in droughted field environments. Breeding modification of root architecture is challenging in field environments owing to genetic and phenotypic complexity, and poor repeatability with root sampling. Seeds from a large mapping population varying in embryo size were harvested from a common glasshouse and standardised to a common size before assessing in rolled germination paper at 12 and 20 °C for seedling growth. Differences in genotype means were large and heritabilities high (h 2 = 0.55-0.93) indicating strong and repeatable genotypic differences for most root traits. Seminal roots 1 to 3 were produced on all seedlings, whereas growth of seminal roots 4, 5 and 6 was associated with differences in embryo size. Increases in seminal root number from 4 to 6 per plant were strongly, genetically correlated with increases in total seminal length (r g = 0.84, < 0.01). Multivariate analysis confirmed initiation and growth of seminal roots 1, 2 and 3, and of roots 4, 5 and 6 behaved as genetically independent (r Pg = 0.15 ns) cohorts. Tails representing extremes in seedling root length and number were associated with significant differences in grain yield of up to 35% in droughted field environments but were not different in irrigated environments. Increases in grain yield were linked to greater lengths of seminal roots 4, 5 and 6 and were largely independent of plant height or development. This is the first report on the genetic relationship of seedling root architecture and embryo size, and potential in selection of seminal root size for accessing deep-soil moisture in droughted environments., (© 2022. Crown.)

Six cycles of recurrent selection for early shoot vigour in wheat resulted in significant increases in leaf width and shoot biomass. Here, in replicated controlled-environment studies, the effect of early shoot vigour on root biomass, rhizosheath size, root hair length, and cell size in the roots and leaves was examined across different cycles of selection. Increased shoot vigour was associated with greater root biomass, larger rhizosheath size, and longer root hairs. Our findings demonstrate that rhizosheath size was a reliable surrogate for root hair length in this germplasm. Examination of the root epidermis revealed that the 'cell body' of the trichoblasts (hair-forming cells) and the atrichoblasts (non-hair-forming cells) decreased in size as shoot vigour increased. Therefore, in higher vigour germplasm, longer root hairs emerged from smaller trichoblasts so that total trichoblast volume (root hair plus cell body) was generally similar regardless of shoot vigour. Similarly, the sizes of the four main cell types on the leaf epidermis became progressively smaller as shoot vigour increased, which also increased stomatal density. The relationship between shoot vigour and root traits is considered, and the potential contribution of below-ground root traits to performance and competitiveness of high vigour germplasm is discussed., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The challenge in establishing an early-sown wheat crop in southern Australia is the need for consistently high seedling emergence when sowing deep in subsoil moisture (>10 cm) or into dry top-soil (4 cm). However, the latter is strongly reliant on a minimum soil water availability to ensure successful seedling emergence. This study aimed to: (1) evaluate 233 Australian and selected international wheat genotypes for consistently high seedling emergence under limited soil water availability when sown in 4 cm of top-soil in field and glasshouse (GH) studies; (2) ascertain genetic loci associated with phenotypic variation using a genome-wide association study (GWAS); and (3) compare across loci for traits controlling coleoptile characteristics, germination, dormancy, and pre-harvest sprouting. Despite significant ( P < 0.001) environment and genotype-by-environment interactions within and between field and GH experiments, eight genotypes that included five cultivars, two landraces, and one inbred line had consistently high seedling emergence (mean value > 85%) across nine environments. Moreover, 21 environment-specific quantitative trait loci (QTL) were detected in GWAS analysis on chromosomes 1B, 1D, 2B, 3A, 3B, 4A, 4B, 5B, 5D, and 7D, indicating complex genetic inheritance controlling seedling emergence. We aligned QTL for known traits and individual genes onto the reference genome of wheat and identified 16 QTL for seedling emergence in linkage disequilibrium with coleoptile length, width, and cross-sectional area, pre-harvest sprouting and dormancy, germination, seed longevity, and anthocyanin development. Therefore, it appears that seedling emergence is controlled by multifaceted networks of interrelated genes and traits regulated by different environmental cues., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Francki, Stainer, Walker, Rebetzke, Stefanova and French.)

Canopy ground cover (GC) is an important agronomic measure for evaluating crop establishment and early growth. This study evaluates the reliability of GC estimates, in the presence of varying light and dew on leaves, from three different ground-based sensors: (1) normalized difference vegetation index (NDVI) from the commercially available GreenSeeker®; (2) RGB images from a digital camera, where GC was determined as the portion of pixels from each image meeting a greenness criterion (i.e., (Green - Red)/(Green + Red) > 0); and (3) LiDAR using two separate approaches: (a) GC from LiDAR red reflectance (whereby red reflectance less than five was classified as vegetation) and (b) GC from LiDAR height (whereby height greater than 10 cm was classified as vegetation). Hourly measurements were made early in the season at two different growth stages (tillering and stem elongation), among wheat genotypes highly diverse for canopy characteristics. The active NDVI showed the least variation through time and was particularly stable, regardless of the available light or the presence of dew. In addition, between-sample-time Pearson correlations for NDVI were consistently high and significant ( P < 0.0001), ranging from 0.89 to 0.98. In comparison, GC from LiDAR and RGB showed greater variation across sampling times, and LiDAR red reflectance was strongly influenced by the presence of dew. Excluding times when the light was exceedingly low, correlations between GC from RGB and NDVI were consistently high (ranging from 0.79 to 0.92). The high reliability of the active NDVI sensor potentially affords a high degree of flexibility for users by enabling sampling across a broad range of acceptable light conditions., Competing Interests: The authors declare that there is no conflict of interest regarding the publication of this article., (Copyright © 2021 David M. Deery et al.)

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 [CO 2 ] (e[CO 2 ]) alone might limit the area where the reduced-tillering trait is advantageous, the most likely climate scenario of e[CO 2 ] 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., (© 2020 John Wiley & Sons Ltd.)

Highly repeatable, nondestructive, and high-throughput measures of above-ground biomass (AGB) and crop growth rate (CGR) are important for wheat improvement programs. This study evaluates the repeatability of destructive AGB and CGR measurements in comparison to two previously described methods for the estimation of AGB from LiDAR: 3D voxel index (3DVI) and 3D profile index (3DPI). Across three field experiments, contrasting in available water supply and comprising up to 98 wheat genotypes varying for canopy architecture, several concurrent measurements of LiDAR and AGB were made from jointing to anthesis. Phenotypic correlations at discrete events between AGB and the LiDAR-derived biomass indices were significant, ranging from 0.31 ( P < 0.05) to 0.86 ( P < 0.0001), providing confidence in the LiDAR indices as effective surrogates for AGB. The repeatability of the LiDAR biomass indices at discrete events was at least similar to and often higher than AGB, particularly under water limitation. The correlations between calculated CGR for AGB and the LiDAR indices were moderate to high and varied between experiments. However, across all experiments, the repeatabilities of the CGR derived from the LiDAR indices were appreciably greater than those for AGB, except for the 3DPI in the water-limited environment. In our experiments, the repeatability of either LiDAR index was consistently higher than that of AGB, both at discrete time points and when CGR was calculated. These findings provide promising support for the reliable use of ground-based LiDAR, as a surrogate measure of AGB and CGR, for screening germplasm in research and wheat breeding., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2020 David M. Deery et al.)

Simple and repeatable methods are needed to select for deep roots under field conditions. A large-scale field experiment was conducted to assess the association between canopy temperature (CT) measured by airborne thermography and rooting depth determined by the core-break method. Three wheat populations, C306×Westonia (CW), Hartog×Drysdale (HD), and Sundor×Songlen (SS), were grown on stored soil water in NSW Australia in 2017 (n=196-252). Cool and warm CT extremes ('tails') were cored after harvest (13-32% of each population). Rooting depth was significantly correlated with CT at late flowering (r= -0.25, -0.52, and -0.23 for CW, HD, and SS, respectively, P<0.05 hereafter), with normalized difference vegetation index (NDVI) at early grain filling (r=0.30-0.39), and with canopy height (r=0.23-0.48). The cool tails showed significantly deeper roots than the respective warm tails by 8.1 cm and 6.2 cm in CW and HD, and correspondingly, greater yields by an average 19% and 7%, respectively. This study highlighted that CT measured rapidly by airborne thermography or NDVI at early grain filling could be used to guide selection of lines with deeper roots to increase wheat yields. The remote measurement methods in this study were repeatable and high throughput, making them well suited to use in breeding programmes., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Water limits crop productivity, so selecting for a minimal yield gap in drier environments is critical to mitigate against climate change and land-use pressure. We investigated the responses of relative water content (RWC), stomatal conductance, chlorophyll content, and metabolites in flag leaves of commercial wheat (Triticum aestivum L.) cultivars to three drought treatments in the glasshouse and in field environments. We observed strong genetic associations between glasshouse-based RWC, metabolites, and yield gap-based drought tolerance (YDT; the ratio of yield in water-limited versus well-watered conditions) across 18 field environments spanning sites and seasons. Critically, RWC response to glasshouse drought was strongly associated with both YDT (r2=0.85, P<8E-6) and RWC under field drought (r2=0.77, P<0.05). Moreover, multiple regression analyses revealed that 98% of genetic YDT variance was explained by drought responses of four metabolites: serine, asparagine, methionine, and lysine (R2=0.98; P<0.01). Fitted coefficients suggested that, for given levels of serine and asparagine, stronger methionine and lysine accumulation was associated with higher YDT. Collectively, our results demonstrate that high-throughput, targeted metabolic phenotyping of glasshouse-grown plants may be an effective tool for selection of wheat cultivars with high field-derived YDT., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Background: Good establishment is important for rapid leaf area development in wheat crops. Poor establishment results in fewer, later-emerging plants, reduced leaf area and tiller number. In addition, poorly established crops suffer from increased soil moisture loss through evaporation and greater competition from weeds while fewer spikes are produced which can reduce grain yield. By protecting the emerging first leaf, the coleoptile is critical for achieving good establishment, and its length and interaction with soil physical properties determine the ability of a cultivar to emerge from depth., Results: Here we characterise a locus on chromosome 1AS, that increases coleoptile length in wheat, which we designate as Lcol-A1. We identified Lcol-A1 by bulked-segregant analysis and used a Halberd-derived population to fine map the gene to a 2 cM region, equivalent to 7 Mb on the IWGSC genome reference sequence of Chinese Spring (RefSeqv1.0). By sowing recently released cultivars and near-isogenic lines in the field at both conventional and deep sowing depths, we confirmed that Locl-A1 was associated with increased emergence from depth in the presence and absence of conventional dwarfing genes. Flanking markers IWB58229 and IWA710 were developed to assist breeders to select for long coleoptile wheats., Conclusions: Increased coleoptile length is sought in many global wheat production areas to improve crop emergence. The identification of the gene Lcol-A1, together with tools to allow wheat breeders to track the gene, will enable improvements to be made for this important trait.

Infrared canopy temperature (CT) is a well-established surrogate measure of stomatal conductance. There is ample evidence showing that genotypic variation in stomatal conductance is associated with grain yield in wheat. Our goal was to determine when CT repeatability is greatest (throughout the season and within the day) to guide CT deployment for research and wheat breeding. CT was measured continuously with ArduCrop wireless infrared thermometers from post-tillering to physiological maturity, and with airborne thermography on cloudless days from manned helicopter at multiple times before and after flowering. Our experiments in wheat, across two years contrasting for water availability, showed that repeatability for CT was greatest later in the season, during grain-filling, and usually in the afternoon. This was supported by the observation that repeatability for ArduCrop, and more so for airborne CT, was significantly associated ( P < 0.0001) with calculated clear-sky solar radiation and to a lesser degree, vapor pressure deficit. Adding vapor pressure deficit to a model comprising either clear-sky solar radiation or its determinants, day-of-year and hour-of-day, made little to no improvement to the coefficient of determination. Phenotypic correlations for airborne CT afternoon sampling events were consistently high between events in the same year, more so for the year when soil water was plentiful ( r = 0.7 to 0.9) than the year where soil water was limiting ( r = 0.4 to 0.9). Phenotypic correlations for afternoon airborne CT were moderate between years contrasting in soil water availability ( r = 0.1 to 0.5) and notably greater on two separate days following irrigation or rain in the drier year, ranging from r = 0.39 to 0.53 ( P < 0.0001) for the midday events. For ArduCrop CT the pattern of phenotypic correlations, within a given year, was similar for both years: phenotypic correlations were higher during the grain-filling months of October and November and for hours-of-day from 11 onwards. The lowest correlations comprised events from hours-of-day 8 and 9 across all months. The capacity for the airborne method to instantaneously sample CT on hundreds of plots is more suited to large field experiments than the static ArduCrop sensors which measure CT continuously on a single experimental plot at any given time. Our findings provide promising support for the reliable deployment of CT phenotyping for research and wheat breeding, whereby the high repeatability and high phenotypic correlations between afternoon sampling events during grain-filling could enable reliable screening of germplasm from only one or two sampling events.

Improved genetic, genomic and statistical technologies have increased the capacity to enrich breeding populations for key alleles underpinning adaptation and continued genetic gain. In turn, directed genomic selection together with increased heritability will reduce genetic variance to narrow the genetic base in many crop breeding programs. Diverse genetic resources (GR), including wild and weedy relatives, landraces and reconstituted synthetics, have potential to contribute novel alleles for key traits. Targeted trait identification may also identify genetic diversity in addressing new challenges including the need for modified root architecture, greater nutrient-use efficiency, and adaptation to warmer air and soil temperatures forecast with climate change. Yet while core collections and other GR sources have historically been invaluable for major gene control of disease and subsoil constraints, the mining of genetically (and phenotypically) complex traits in GR remains a significant challenge owing to reduced fertility, limited seed quantities and poor adaptation through linkage drag with undesirable alleles. High-throughput field phenomics (HTFP) offers the opportunity to capture phenotypically complex variation underpinning adaptation in traditional phenotypic selection or statistics-based breeding programs. Targeted HTFP will permit the reliable phenotyping of greater numbers of GR-derived breeding lines using smaller plot sizes and at earlier stages of population development to reduce the duration of breeding cycles and the loss of potentially important alleles with linkage drag. Two key opportunities are highlighted for use of HTFP in selection among GR-derived wheat breeding lines for greater biomass and stomatal conductance through canopy temperature., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Early vigour, or faster early leaf area development, has been considered an important trait for rainfed wheat in dryland regions such as Australia. However, early vigour is a genetically complex trait, and results from field experiments have been highly variable. Whether early vigour can lead to improved water use efficiency and crop yields is strongly dependent on climate and management conditions across the entire growing season. Here, we present a modelling framework for simulating the impact of early vigour on wheat growth and yield at eight sites representing the major climate types in Australia. On a typical soil with plant available water capacity (PAWC) of 147 mm, simulated yield increase with early vigour associated with larger seed size was on average 4% higher compared with normal vigour wheat. Early vigour through selection of doubled early leaf sizes could increase yield by 16%. Increase in yield was mainly from increase in biomass and grain number, and was reduced at sites with seasonal rainfall plus initial soil water <300 mm. Opportunities exists for development of early vigour wheat varieties for wetter sites. Soil PAWC could play a significant role in delivering the benefit of early vigour and would require particular attention., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

In spite of the increasing expectation for process-based crop modelling to capture genotype (G) by environment (E) by management (M) interactions to support breeding selections, it remains a challenge to use current crop models to accurately predict phenotypes from genotypes or from candidate genes. We use wheat as a target crop and the APSIM farming systems model (Holzworth et al., 2014) as an example to analyse the current status of process-based crop models with a major focus on need to improve simulation of specific eco-physiological processes and their linkage to underlying genetic controls. For challenging production environments in Australia, we examine the potential opportunities to capture physiological traits, and to integrate genetic and molecular approaches for future model development and applications. Model improvement will require both reducing the uncertainty in simulating key physiological processes and enhancing the capture of key observable traits and underlying genetic control of key physiological responses to environment. An approach consisting of three interactive stages is outlined to (i) improve modelling of crop physiology, (ii) develop linkage from model parameter to genotypes and further to loci or alleles, and (iii) further link to gene expression pathways. This helps to facilitate the integration of modelling, phenotyping, and functional gene detection and to effectively advance modelling of G×E×M interactions. While gene-based modelling is not always needed to simulate G×E×M, including well-understood gene effects can improve the estimation of genotype effects and prediction of phenotypes. Specific examples are given for enhanced modelling of wheat in the APSIM framework., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Abiotic stress tolerance traits are often complex and recalcitrant targets for conventional breeding improvement in many crop species. This study evaluated the potential of genomic selection to predict water-soluble carbohydrate concentration (WSCC), an important drought tolerance trait, in wheat under field conditions. A panel of 358 varieties and breeding lines constrained for maturity was evaluated under rainfed and irrigated treatments across two locations and two years. Whole-genome marker profiles and factor analytic mixed models were used to generate genomic estimated breeding values (GEBVs) for specific environments and environment groups. Additive genetic variance was smaller than residual genetic variance for WSCC, such that genotypic values were dominated by residual genetic effects rather than additive breeding values. As a result, GEBVs were not accurate predictors of genotypic values of the extant lines, but GEBVs should be reliable selection criteria to choose parents for intermating to produce new populations. The accuracy of GEBVs for untested lines was sufficient to increase predicted genetic gain from genomic selection per unit time compared to phenotypic selection if the breeding cycle is reduced by half by the use of GEBVs in off-season generations. Further, genomic prediction accuracy depended on having phenotypic data from environments with strong correlations with target production environments to build prediction models. By combining high-density marker genotypes, stress-managed field evaluations, and mixed models that model simultaneously covariances among genotypes and covariances of complex trait performance between pairs of environments, we were able to train models with good accuracy to facilitate genetic gain from genomic selection., (Copyright © 2018 Ovenden et al.)

Crop improvement efforts are targeting increased above-ground biomass and radiation-use efficiency as drivers for greater yield. Early ground cover and canopy height contribute to biomass production, but manual measurements of these traits, and in particular above-ground biomass, are slow and labor-intensive, more so when made at multiple developmental stages. These constraints limit the ability to capture these data in a temporal fashion, hampering insights that could be gained from multi-dimensional data. Here we demonstrate the capacity of Light Detection and Ranging (LiDAR), mounted on a lightweight, mobile, ground-based platform, for rapid multi-temporal and non-destructive estimation of canopy height, ground cover and above-ground biomass. Field validation of LiDAR measurements is presented. For canopy height, strong relationships with LiDAR ( r 2 of 0.99 and root mean square error of 0.017 m) were obtained. Ground cover was estimated from LiDAR using two methodologies: red reflectance image and canopy height. In contrast to NDVI, LiDAR was not affected by saturation at high ground cover, and the comparison of both LiDAR methodologies showed strong association ( r 2 = 0.92 and slope = 1.02) at ground cover above 0.8. For above-ground biomass, a dedicated field experiment was performed with destructive biomass sampled eight times across different developmental stages. Two methodologies are presented for the estimation of biomass from LiDAR: 3D voxel index (3DVI) and 3D profile index (3DPI). The parameters involved in the calculation of 3DVI and 3DPI were optimized for each sample event from tillering to maturity, as well as generalized for any developmental stage. Individual sample point predictions were strong while predictions across all eight sample events, provided the strongest association with biomass ( r 2 = 0.93 and r 2 = 0.92) for 3DPI and 3DVI, respectively. Given these results, we believe that application of this system will provide new opportunities to deliver improved genotypes and agronomic interventions via more efficient and reliable phenotyping of these important traits in large experiments.

Key Message: Water-soluble carbohydrate accumulation can be selected in wheat breeding programs with consideration of genetic × environmental interactions and relationships with other important characteristics such as relative maturity and nitrogen concentration, although the correlation between WSC traits and grain yield is low and inconsistent. The potential to increase the genetic capacity for water-soluble carbohydrate (WSC) accumulation is an opportunity to improve the drought tolerance capability of rainfed wheat varieties, particularly in environments where terminal drought is a significant constraint to wheat production. A population of elite breeding germplasm was characterized to investigate the potential for selection of improved WSC concentration and total amount in water deficit and well-watered environments. Accumulation of WSC involves complex interactions with other traits and the environment. For both WSC concentration (WSCC) and total WSC per area (WSCA), strong genotype × environment interactions were reflected in the clear grouping of experiments into well-watered and water deficit environment clusters. Genetic correlations between experiments were high within clusters. Heritability for WSCC was larger than for WSCA, and significant associations were observed in both well-watered and water deficit experiment clusters between the WSC traits and nitrogen concentration, tillering, grains per m 2 , and grain size. However, correlations between grain yield and WSCC or WSCA were weak and variable, suggesting that selection for these traits is not a better strategy for improving yield under drought than direct selection for yield.

Improving water-use efficiency by incorporating drought avoidance traits into new wheat varieties is an important objective for wheat breeding in water-limited environments. This study uses genome wide association studies (GWAS) to identify candidate loci for water-soluble carbohydrate accumulation-an important drought-avoidance characteristic in wheat. Phenotypes from a multi-environment trial with experiments differing in water availability and separate single nucleotide polymorphism (SNP) and diversity arrays technology (DArT) marker sets were used to perform the analyses. Significant associations for water-soluble carbohydrate accumulation were identified on chromosomes 1A, 1B, 1D, 2D, and 4A. Notably, these loci did not collocate with the major loci identified for relative maturity. Loci on chromosome 1D collocated with markers previously associated with the high molecular weight glutenin Glu-D1 locus. Genetic × environmental interactions impacted the results strongly, with significant associations for carbohydrate accumulation identified only in the water-deficit experiments. The markers associated with carbohydrate accumulation may be useful for marker-assisted selection of drought tolerance in wheat., (Copyright © 2017 Ovenden et al.)

Lower canopy temperature (CT), resulting from increased stomatal conductance, has been associated with increased yield in wheat. Historically, CT has been measured with hand-held infrared thermometers. Using the hand-held CT method on large field trials is problematic, mostly because measurements are confounded by temporal weather changes during the time required to measure all plots. The hand-held CT method is laborious and yet the resulting heritability low, thereby reducing confidence in selection in large scale breeding endeavors. We have developed a reliable and scalable crop phenotyping method for assessing CT in large field experiments. The method involves airborne thermography from a manned helicopter using a radiometrically-calibrated thermal camera. Thermal image data is acquired from large experiments in the order of seconds, thereby enabling simultaneous measurement of CT on potentially 1000s of plots. Effects of temporal weather variation when phenotyping large experiments using hand-held infrared thermometers are therefore reduced. The method is designed for cost-effective and large-scale use by the non-technical user and includes custom-developed software for data processing to obtain CT data on a single-plot basis for analysis. Broad-sense heritability was routinely >0.50, and as high as 0.79, for airborne thermography CT measured near anthesis on a wheat experiment comprising 768 plots of size 2 × 6 m. Image analysis based on the frequency distribution of temperature pixels to remove the possible influence of background soil did not improve broad-sense heritability. Total image acquisition and processing time was ca. 25 min and required only one person (excluding the helicopter pilot). The results indicate the potential to phenotype CT on large populations in genetics studies or for selection within a plant breeding program.

Increases in air and soil temperatures will impact cereal growth and reduce crop yields. Little is known about how increasing temperatures will impact seedling growth and crop establishment. Climate forecast models predict that by 2060, mean and maximum air temperatures in the Australian wheatbelt will increase by 2-4°C during the March-June sowing period, and particularly at lower latitudes. Concomitant increases in soil temperature will shorten coleoptile length to reduce crop establishment, particularly where deep sowing to access sub-surface moisture. Mean coleoptile length was reduced in commercial wheat (Triticum aestivum L.) germplasm with increasing soil temperature (106mm and 51mm at 15°C and 31°C, respectively). Coleoptile lengths of modern semidwarf varieties were significantly (P<0.01) shorter than those of older tall wheats at 15°C (95mm and 135mm) and 31°C (46mm and 70mm). A 12-parent diallel indicated large additive and small non-maternal genetic effects for coleoptile length at 15°C and 27°C. Large genotype rank changes for coleoptile length across temperatures (rs=0.37, P<0.05) contributed to smaller entry-mean heritabilities (0.41-0.67) to reduce confidence in selection for long-coleoptile genotypes across contrasting temperatures. General combining ability effects were strongly correlated across temperatures (rp=0.81, P < 0.01), indicating the potential of some donors in identification of progeny with consistently longer coleoptiles. Warmer soils in future will contribute to poor establishment and crop failure, particularly with deep-sown semidwarf wheat. Breeding long-coleoptile genotypes with improved performance will require targeted selection at warmer temperatures in populations incorporating novel sources of reduced height and greater coleoptile length.

Early vigour is an important physiological trait to improve establishment, water-use efficiency, and grain yield for wheat. Phenotyping large numbers of lines is challenging due to the fast growth and development of wheat seedlings. Here we developed a new photo-based workflow to monitor dynamically the growth and development of the wheat canopy of two wheat lines with a contrasting early vigour trait. Multiview images were taken using a 'vegetation stress' camera at 2 d intervals from emergence to the sixth leaf stage. Point clouds were extracted using the Multi-View Stereo and Structure From Motion (MVS-SFM) algorithm, and segmented into individual organs using the Octree method, with leaf midribs fitted using local polynomial function. Finally, phenotypic parameters were calculated from the reconstructed point cloud including: tiller and leaf number, plant height, Haun index, phyllochron, leaf length, angle, and leaf elongation rate. There was good agreement between the observed and estimated leaf length (RMSE=8.6mm, R (2)=0.98, n=322) across both lines. Significant contrasts of phenotyping parameters were observed between the two lines and were consistent with manual observations. The early vigour line had fewer tillers (2.4±0.6) and larger leaves (308.0±38.4mm and 17.1±2.7mm for leaf length and width, respectively). While the phyllochron of both lines was quite similar, the non-vigorous line had a greater Haun index (more leaves on the main stem) on any date, as the vigorous line had slower development of its first two leaves. The workflow presented in this study provides an efficient method to phenotype individual plants using a low-cost camera (an RGB camera is also suitable) and could be applied in phenotyping for applications in both simulation modelling and breeding. The rapidity and accuracy of this novel method can characterize the results of specific selection criteria (e.g. width of leaf three, number of tillers, rate of leaf appearance) that have been or can now be utilized to breed for early leaf growth and tillering in wheat., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Rhizosheaths comprise soil bound to roots, and in wheat (Triticum aestivum L.) rhizosheath size correlates with root hair length. The aims of this study were to determine the effect that a large rhizosheath has on the phosphorus (P) acquisition by wheat and to investigate the genetic control of rhizosheath size in wheat grown on acid soil.Near-isogenic wheat lines differing in rhizosheath size were evaluated on two acid soils. The soils were fertilized with mineral nutrients and included treatments with either low or high P. The same soils were treated with CaCO3 to raise the pH and detoxify Al(3+) Genotypic differences in rhizosheath size were apparent only when soil pH was low and Al(3+) was present. On acid soils, a large rhizosheath increased shoot biomass compared with a small rhizosheath regardless of P supply. At low P supply, increased shoot biomass could be attributed to a greater uptake of soil P, but at high P supply the increased biomass was due to some other factor. Generation means analysis indicated that rhizosheath size on acid soil was controlled by multiple, additive loci. Subsequently, a quantitative trait loci (QTL) analysis of an F6 population of recombinant inbred lines identified five major loci contributing to the phenotype together accounting for over 60% of the total genetic variance. One locus on chromosome 1D accounted for 34% of the genotypic variation. Genetic control of rhizosheath size appears to be relatively simple and markers based on the QTL provide valuable tools for marker assisted breeding., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Genotypic variation in ear morphology is linked to differences in photosynthetic potential to influence grain yield in winter cereals. Awns contribute to photosynthesis, particularly under water-limited conditions when canopy assimilation is restricted. We assessed performance of up to 45 backcross-derived, awned-awnletted NILs representing four diverse genetic backgrounds in 25 irrigated or rainfed, and droughted environments in Australia and Mexico. Mean environment grain yields were wide-ranging (1.38-7.93 t ha(-1)) with vegetative and maturity biomass, plant height, anthesis date, spike number, and harvest index all similar (P >0.05) for awned and awnletted NILs. Overall, grain yields of awned-awnletted sister-NILs were equivalent, irrespective of yield potential and genetic background. Awnletted wheats produced significantly more grains per unit area (+4%) and per spike (+5%) reflecting more fertile spikelets and grains in tertiary florets. Increases in grain number were compensated for by significant reductions in grain size (-5%) and increased frequency (+0.8%) of small, shrivelled grains ('screenings') to reduce seed-lot quality of awnletted NILs. Post-anthesis canopies of awnletted NILs were marginally warmer over all environments (+0.27 °C) but were not different and were sometimes cooler than awned NILs at cooler air temperatures. Awns develop early and represented up to 40% of total spikelet biomass prior to ear emergence. We hypothesize that the allocation of assimilate to large and rapidly developing awns decreases spikelet number and floret fertility to reduce grain number, particularly in distal florets. Individual grain size is increased to reduce screenings and to increase test weight and milling quality, particularly in droughted environments. Despite the average reduction in grain size, awnless lines could be identified that combined higher grain yield with larger grain size, increased grain protein concentration, and reduced screenings., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Genetic modification of shoot and root morphology has potential to improve water and nutrient uptake of wheat crops in rainfed environments. Near-isogenic lines (NILs) varying for a tillering inhibition (tin) gene and representing multiple genetic backgrounds were phenotyped in contrasting, controlled environments for shoot and root growth. Leaf area, shoot and root biomass were similar until tillering, whereupon reduced tillering in tin-containing NILs produced reductions of up to 60% in total leaf area and biomass, and increases in total root length of up to 120% and root biomass to 145%. Together, the root-to-shoot ratio increased two-fold with the tin gene. The influence of tin on shoot and root growth was greatest in the cv. Banks genetic background, particularly in the biculm-selected NIL, and was typically strongest in cooler environments. A separate de-tillering study confirmed greater root-to-shoot ratios with regular tiller removal in non-tin-containing genotypes. In validating these observations in a rainfed field study, the tin allele had a negligible effect on seedling growth but was associated with significantly (P<0.05) reduced tiller number (-37%), leaf area index (-26%), and spike number (-35%) to reduce plant biomass (-19%) at anthesis. Root biomass, root-to-shoot ratio at early stem elongation, and root depth at maturity were all increased in tin-containing NILs. Soil water use was slowed in tin-containing NILs, resulting in greater water availability, greater stomatal conductance, cooler canopy temperatures, and maintenance of green leaf area during grain-filling. Together these effects contributed to increases in harvest index and grain yield. In both the controlled and field environments, the tin gene was commonly associated with increased root length and biomass, but the significant influence of genetic background and environment suggests careful assessment of tin-containing progeny in selection for genotypic increases in root growth., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Quantitative trait loci (QTLs) for shoot biomass were identified in wheat grown on a soil high in total phosphorus (P) but low in plant-available P. The two populations screened included recombinant inbred lines (RILs) from Chuan-Mai 18/Vigour 18 and doubled-haploid lines from Kukri/Janz. Glasshouse-grown plants were harvested at the five-leaf stage. Seven QTLs for shoot biomass were identified in the RILs, with the largest on chromosome 7A accounting for 7.4% of the phenotypic variance. RILs from the upper tail had larger embryos than RILs from the lower tail. Tail lines were then grown in non-limiting P and the results indicated that early vigour and the capacity to access P contributed to the initial distribution. The influence of early vigour on P nutrition was examined further with advanced vigour lines (AVLs). The AVLs accumulated more shoot biomass, maintained lower shoot P concentrations, and showed greater P-acquisition efficiency than Vigour 18. Nine QTLs for shoot biomass were identified in the Kukri/Janz population. Two on chromosomes 4B and 4D accounted for 24.8% of the variance. Candidates underlying these QTLs are the Rht genes. We confirmed the influence of these genes using near-isogenic lines with different Rht alleles. The dwarf and semi-dwarf alleles affected shoot and root biomass at high and low P but not the efficiency of P acquisition. We conclude that early vigour contributed to the distributions in both populations. Early vigour can increase plant growth at suboptimal P and some sources can also improve the efficiency of P acquisition., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Increasing climate variability, particularly variability in the timing and amount of soil water, means that breeding wheat (Triticum aestivum L.) varieties with stable high grain yields is increasingly more challenging. Changing environmental conditions in water-limited rainfed environments will alter genotype ranking to reduce confidence in the identification of consistently higher yielding performers. Greater early vigour (EV) and transpiration efficiency (TE) are two physiological traits that have demonstrated benefits as breeding targets for efficient water-use in Mediterranean in-season water and monsoonal stored water environments, respectively. This Perspective discusses the hypothesis that combining higher TE and greater EV will broaden the adaptation and increase grain yields for wheats grown across most rainfed environments. We examine the physiology underpinning adaptation with greater EV and higher TE, as well as the challenges and potential benefits of deploying these traits in combination. We then discuss how these two traits interact with different environments and, in particular, the different wheat-growing regions of Australia. We conclude that the combination of these two traits is genetically and physiologically feasible, as well theoretically beneficial to average yield in most rainfed environments. Hence, we suggest a strategy for reliably managing the complex genetics underpinning EV and TE when phenotyping and selecting both traits in commercial wheat breeding programs.

Shoot and root system size influences N uptake in wheat (Triticum aestivum L.). Previously, we showed that four wheat genotypes with different biomass had similar N uptake at tillering. In the present study, we determined whether the similarity in N uptake in these genotypes was associated with genotypic differences in the affinity of the root system for NO3- uptake. Kinetic parameters of NO3- uptake were measured in hydroponic seedlings of vigorous and nonvigorous early growth wheat genotypes by exposing them to solutions with differing concentrations of K15NO3 for 15min. In the low concentration range, the high-affinity transport system of the nonvigorous cultivar Janz showed a higher maximum influx rate than the three vigorous lines and a higher affinity than two of the three vigorous lines. At high NO3- concentrations, where the low-affinity transport system was functional, the responsiveness of NO3- uptake to external concentrations was greater in Janz than in the vigorous lines. Both the high- and low-affinity transport systems were inducible. The genotypic variation in the kinetic parameters of NO3- uptake was large enough to offset differences in morphological traits and should be considered in efforts to improve N uptake. In a field trial, the growth and N uptake performance of the four wheat genotypes was investigated over the winter-spring growing season (June-November of 2010). The field trial showed that although early N uptake was disproportionately large relative to biomass accumulation, the differences in uptake at tillering can be changed by subsequent patterns of uptake.

Background: The flag leaf of a wheat (Triticum aestivum L.) plant rolls up into a cylinder in response to drought conditions and then unrolls when leaf water relations improve. This is a desirable trait for extending leaf area duration and improving grain size particularly under drought. But how do we quantify this phenotype so that different varieties of wheat or different treatments can be compared objectively since this phenotype can easily be confounded with inter-genotypic differences in root-water uptake and/or transpiration at the leaf level if using traditional methods?, Results: We present a new method to objectively test a range of lines/varieties/treatments for their propensity of leaves to roll. We have designed a repeatable protocol and defined an objective measure of leaf curvature called "rolled-upness" which minimises confounding factors in the assessment of leaf rolling in grass species. We induced leaf rolling by immersing leaf strips in an osmoticum of known osmotic pressure. Using micro-photographs of individual leaf cross-sections at equilibrium in the osmoticum, two approaches were used to quantify leaf rolling. The first was to use some properties of the convex hull of the leaf cross-section. The second was to use cubic smoothing splines to approximate the transverse leaf shape mathematically and then use a statistic derived from the splines for comparison. Both approaches resulted in objective measurements that could differentiate clearly between breeding lines and varieties contrasting genetically in their propensity for leaf rolling under water stress. The spline approach distinguished between upward and downward curvature and allowed detailed properties of the rolling to be examined, such as the position on the strip where maximum curvature occurs., Conclusions: A method applying smoothing splines to skeletonised images of transverse wheat leaf sections enabled objective measurements of inter-genotypic variation for hydronastic leaf rolling in wheat. Mean-curvature of the leaf cross-section was the measure selected to discriminate between genotypes, as it was straightforward to calculate and easily construed. The method has broad applicability and provides an avenue to genetically dissect the trait in cereals.

Seedling roots enable plant establishment. Their small phenotypes are measured routinely. Adult root systems are relevant to yield and efficiency, but phenotyping is challenging. Root length exceeds the volume of most pots. Field studies measure partial adult root systems through coring or use seedling roots as adult surrogates. Here, we phenotyped 79 diverse lines of the small grass model Brachypodium distachyon to adults in 50-cm-long tubes of soil with irrigation; a subset of 16 lines was droughted. Variation was large (total biomass, ×8; total root length [TRL], ×10; and root mass ratio, ×6), repeatable, and attributable to genetic factors (heritabilities ranged from approximately 50% for root growth to 82% for partitioning phenotypes). Lines were dissected into seed-borne tissues (stem and primary seminal axile roots) and stem-borne tissues (tillers and coleoptile and leaf node axile roots) plus branch roots. All lines developed one seminal root that varied, with branch roots, from 31% to 90% of TRL in the well-watered condition. With drought, 100% of TRL was seminal, regardless of line because nodal roots were almost always inhibited in drying topsoil. Irrigation stimulated nodal roots depending on genotype. Shoot size and tillers correlated positively with roots with irrigation, but partitioning depended on genotype and was plastic with drought. Adult root systems of B. distachyon have genetic variation to exploit to increase cereal yields through genes associated with partitioning among roots and their responsiveness to irrigation. Whole-plant phenotypes could enhance gain for droughted environments because root and shoot traits are coselected., (© 2015 American Society of Plant Biologists. All Rights Reserved.)

The breeding of wheat with greater early vigour has potential to increase water- and nutrient-use efficiency, as well as to improve weed competitiveness to raise crop yields profitably. Given that wheat is inherently conservative in its early growth, a sustained breeding effort was initiated to increase genetically seedling leaf area in developing novel high vigour germplasm. A recurrent selection programme was initiated by intercrossing a genetically diverse set of 28 vigorous wheat lines identified globally. These were intercrossed at random and S1:2 progeny with the largest leaf 1 and 2 widths were intermated to develop new populations for assessment of early growth. This procedure was repeated for up to 60 segregating families per cycle across six cycles over 15 years. Thirty random S1:2 progeny were retained from each cycle and seed-increased together to produce seed for early vigour assessment in multiple sowings. The most vigorous wheat seedlings were identified in later cycles, with some lines producing more than double the leaf area and biomass of elite commercial wheat varieties. Phenotypic selection for greater leaf width was associated with a realized significant (P<0.01) linear increase per seedling of 0.41 mm per cycle (+7.1%) for mean leaf width, and correlated linear increases in total leaf area and biomass of 4.48 cm(2) per cycle (+10.3%) and 10.8 mg per cycle (+5.3%), respectively. Genetic gains in widths of leaves 2 (+8.4%) and 3 (+11.5%) were significantly (P<0.01) greater than for leaf 1 (+5.3%). Selection for greater leaf width was associated with linear increases in coleoptile tiller leaf area, small curvilinear increases in leaf 1 length, and reductions in numbers of leaves and mainstem tillers. Genetic variances were large and heritabilities high for leaf width and total leaf area in each cycle, but reduced linearly in size with selection across cycles. Coupling diverse germplasm with a simple, inexpensive, and repeatable selection process has confirmed the value of recurrent selection in developing uniquely vigorous wheat germplasm for use as parents in commercial breeding., (© The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology.)