APSIM-based modeling approach to understand sorghum production environments in Mali.

Sorghum production system in the semi-arid region of Africa is characterized by low yields which are generally attributed to high rainfall variability, poor soil fertility, and biotic factors. Production constraints must be well understood and quantified to design effective sorghum-system improvements. This study uses the state-of-the-art in silico methods and focuses on characterizing the sorghum production regions in Mali for drought occurrence and its effects on sorghum productivity. For this purpose, we adapted the APSIM-sorghum module to reproduce two cultivated photoperiod-sensitive sorghum types across a latitude of major sorghum production regions in Western Africa. We used the simulation outputs to characterize drought stress scenarios. We identified three main drought scenarios: (i) no-stress; (ii) early pre-flowering drought stress; and (iii) drought stress onset around flowering. The frequency of drought stress scenarios experienced by the two sorghum types across rainfall zones and soil types differed. As expected, the early pre-flowering and flowering drought stress occurred more frequently in isohyets < 600 mm, for the photoperiod-sensitive, late-flowering sorghum type. In isohyets above 600 mm, the frequency of drought stress was very low for both cultivars. We quantified the consequences of these drought scenarios on grain and biomass productivity. The yields of the highly-photoperiod-sensitive sorghum type were quite stable across the higher rainfall zones > 600 mm, but was affected by the drought stress in the lower rainfall zones < 600 mm. Comparatively, the less photoperiod-sensitive cultivar had notable yield gain in the driest regions < 600 mm. The results suggest that, at least for the tested crop types, drought stress might not be the major constraint to sorghum production in isohyets > 600 mm. The findings from this study provide the entry point for further quantitative testing of the Genotype × Environment × Management options required to optimize sorghum production in Mali. Supplementary Information: The online version contains supplementary material available at 10.1007/s13593-023-00909-5.

Contrasting leaf-scale photosynthetic low-light response and its temperature dependency are key to differences in crop-scale radiation use efficiency.

Radiation use efficiency (RUE) is a key crop adaptation trait that quantifies the potential amount of aboveground biomass produced by the crop per unit of solar energy intercepted. But it is unclear why elite maize and grain sorghum hybrids differ in their RUE at the crop level. Here, we used a non-traditional top-down approach via canopy photosynthesis modelling to identify leaf-level photosynthetic traits that are key to differences in crop-level RUE. A novel photosynthetic response measurement was developed and coupled with use of a Bayesian model fitting procedure, incorporating a C4 leaf photosynthesis model, to infer cohesive sets of photosynthetic parameters by simultaneously fitting responses to CO2 , light, and temperature. Statistically significant differences between leaf photosynthetic parameters of elite maize and grain sorghum hybrids were found across a range of leaf temperatures, in particular for effects on the quantum yield of photosynthesis, but also for the maximum enzymatic activity of Rubisco and PEPc. Simulation of diurnal canopy photosynthesis predicted that the leaf-level photosynthetic low-light response and its temperature dependency are key drivers of the performance of crop-level RUE, generating testable hypotheses for further physiological analysis and bioengineering applications.

Two decades of harnessing standing genetic variation for physiological traits to improve drought tolerance in maize.

We review approaches to maize breeding for improved drought tolerance during flowering and grain filling in the central and western US corn belt and place our findings in the context of results from public breeding. Here we show that after two decades of dedicated breeding efforts, the rate of crop improvement under drought increased from 6.2 g m-2 year-1 to 7.5 g m-2 year-1, closing the genetic gain gap with respect to the 8.6 g m-2 year-1 observed under water-sufficient conditions. The improvement relative to the long-term genetic gain was possible by harnessing favourable alleles for physiological traits available in the reference population of genotypes. Experimentation in managed stress environments that maximized the genetic correlation with target environments was key for breeders to identify and select for these alleles. We also show that the embedding of physiological understanding within genomic selection methods via crop growth models can hasten genetic gain under drought. We estimate a prediction accuracy differential (Δr) above current prediction approaches of ~30% (Δr=0.11, r=0.38), which increases with increasing complexity of the trait environment system as estimated by Shannon information theory. We propose this framework to inform breeding strategies for drought stress across geographies and crops.

A cross-scale analysis to understand and quantify the effects of photosynthetic enhancement on crop growth and yield across environments.

Photosynthetic manipulation provides new opportunities for enhancing crop yield. However, understanding and quantifying the importance of individual and multiple manipulations on the seasonal biomass growth and yield performance of target crops across variable production environments is limited. Using a state-of-the-art cross-scale model in the APSIM platform we predicted the impact of altering photosynthesis on the enzyme-limited (Ac ) and electron transport-limited (Aj ) rates, seasonal dynamics in canopy photosynthesis, biomass growth, and yield formation via large multiyear-by-location crop growth simulations. A broad list of promising strategies to improve photosynthesis for C3 wheat and C4 sorghum were simulated. In the top decile of seasonal outcomes, yield gains were predicted to be modest, ranging between 0% and 8%, depending on the manipulation and crop type. We report how photosynthetic enhancement can affect the timing and severity of water and nitrogen stress on the growing crop, resulting in nonintuitive seasonal crop dynamics and yield outcomes. We predicted that strategies enhancing Ac alone generate more consistent but smaller yield gains across all water and nitrogen environments, Aj enhancement alone generates larger gains but is undesirable in more marginal environments. Large increases in both Ac and Aj generate the highest gains across all environments. Yield outcomes of the tested manipulation strategies were predicted and compared for realistic Australian wheat and sorghum production. This study uniquely unpacks complex cross-scale interactions between photosynthesis and seasonal crop dynamics and improves understanding and quantification of the potential impact of photosynthesis traits (or lack of it) for crop improvement research.

Genetic modification of PIN genes induces causal mechanisms of stay-green drought adaptation phenotype.

The stay-green trait is recognized as a key drought adaptation mechanism in cereals worldwide. Stay-green sorghum plants exhibit delayed senescence of leaves and stems, leading to prolonged growth, a reduced risk of lodging, and higher grain yield under end-of-season drought stress. More than 45 quantitative trait loci (QTL) associated with stay-green have been identified, including two major QTL (Stg1 and Stg2). However, the contributing genes that regulate functional stay-green are not known. Here we show that the PIN FORMED family of auxin efflux carrier genes induce some of the causal mechanisms driving the stay-green phenotype in sorghum, with SbPIN4 and SbPIN2 located in Stg1 and Stg2, respectively. We found that nine of 11 sorghum PIN genes aligned with known stay-green QTL. In transgenic studies, we demonstrated that PIN genes located within the Stg1 (SbPIN4), Stg2 (SbPIN2), and Stg3b (SbPIN1) QTL regions acted pleiotropically to modulate canopy development, root architecture, and panicle growth in sorghum, with SbPIN1, SbPIN2, and SbPIN4 differentially expressed in various organs relative to the non-stay-green control. The emergent consequence of such modifications in canopy and root architecture is a stay-green phenotype. Crop simulation modelling shows that the SbPIN2 phenotype can increase grain yield under drought.

Radiation use efficiency increased over a century of maize (Zea mays L.) breeding in the US corn belt.

In the absence of stress, crop growth depends on the amount of light intercepted by the canopy and the conversion efficiency [radiation use efficiency (RUE)]. This study tested the hypothesis that long-term genetic gain for grain yield was partly due to improved RUE. The hypothesis was tested using 30 elite maize hybrids commercialized in the US corn belt between 1930 and 2017. Crops grown under irrigation showed that pre-flowering crop growth increased at a rate of 0.11 g m-2 year-1, while light interception remained constant. Therefore, RUE increased at a rate of 0.0049 g MJ-1 year-1, translating into an average of 3 g m-2 year-1 of grain yield over 100 years of maize breeding. Considering that the harvest index has not changed for crops grown at optimal density for the hybrid, the cumulative RUE increase over the history of commercial maize breeding in the USA can account for ~32% of the documented yield trend for maize grown in the central US corn belt. The remaining RUE gap between this study and theoretical maximum values suggests that a yield improvement of a similar magnitude could be achieved by further increasing RUE.

Detecting Sorghum Plant and Head Features from Multispectral UAV Imagery.

In plant breeding, unmanned aerial vehicles (UAVs) carrying multispectral cameras have demonstrated increasing utility for high-throughput phenotyping (HTP) to aid the interpretation of genotype and environment effects on morphological, biochemical, and physiological traits. A key constraint remains the reduced resolution and quality extracted from "stitched" mosaics generated from UAV missions across large areas. This can be addressed by generating high-quality reflectance data from a single nadir image per plot. In this study, a pipeline was developed to derive reflectance data from raw multispectral UAV images that preserve the original high spatial and spectral resolutions and to use these for phenotyping applications. Sequential steps involved (i) imagery calibration, (ii) spectral band alignment, (iii) backward calculation, (iv) plot segmentation, and (v) application. Each step was designed and optimised to estimate the number of plants and count sorghum heads within each breeding plot. Using a derived nadir image of each plot, the coefficients of determination were 0.90 and 0.86 for estimates of the number of sorghum plants and heads, respectively. Furthermore, the reflectance information acquired from the different spectral bands showed appreciably high discriminative ability for sorghum head colours (i.e., red and white). Deployment of this pipeline allowed accurate segmentation of crop organs at the canopy level across many diverse field plots with minimal training needed from machine learning approaches.

Reproductive resilience but not root architecture underpins yield improvement under drought in maize.

Because plants capture water and nutrients through roots, it was proposed that changes in root systems architecture (RSA) might underpin the 3-fold increase in maize (Zea mays L.) grain yield over the last century. Here we show that both RSA and yield have changed with decades of maize breeding, but not the crop water uptake. Results from X-ray phenotyping in controlled environments showed that single cross (SX) hybrids have smaller root systems than double cross (DX) hybrids for root diameters between 2465 µm and 181µm (P<0.05). Soil water extraction measured under field conditions ranged between 2.6 mm d-1 and 2.9 mm d-1 but were not significantly different between SX and DX hybrids. Yield and yield components were higher for SX than DX hybrids across densities and irrigation (P<0.001). Taken together, the results suggest that changes in RSA were not the cause of increased water uptake but an adaptation to high-density stands used in modern agriculture. This adaptation may have contributed to shift in resource allocation to the ear and indirectly improved reproductive resilience. Advances in root physiology and phenotyping can create opportunities to maintain long-term genetic gain in maize, but a shift from ideotype to crop and production system thinking will be required.

In pursuit of a better world: crop improvement and the CGIAR.

The CGIAR crop improvement (CI) programs, unlike commercial CI programs, which are mainly geared to profit though meeting farmers' needs, are charged with meeting multiple objectives with target populations that include both farmers and the community at large. We compiled the opinions from >30 experts in the private and public sector on key strategies, methodologies, and activities that could the help CGIAR meet the challenges of providing farmers with improved varieties while simultaneously meeting the goals of: (i) nutrition, health, and food security; (ii) poverty reduction, livelihoods, and jobs; (iii) gender equality, youth, and inclusion; (iv) climate adaptation and mitigation; and (v) environmental health and biodiversity. We review the crop improvement processes starting with crop choice, moving through to breeding objectives, production of potential new varieties, selection, and finally adoption by farmers. The importance of multidisciplinary teams working towards common objectives is stressed as a key factor to success. The role of the distinct disciplines, actors, and their interactions throughout the process from crop choice through to adoption by farmers is discussed and illustrated.

Tackling G × E × M interactions to close on-farm yield-gaps: creating novel pathways for crop improvement by predicting contributions of genetics and management to crop productivity.

KEY MESSAGE: Climate change and Genotype-by-Environment-by-Management interactions together challenge our strategies for crop improvement. Research to advance prediction methods for breeding and agronomy is opening new opportunities to tackle these challenges and overcome on-farm crop productivity yield-gaps through design of responsive crop improvement strategies. Genotype-by-Environment-by-Management (G × E × M) interactions underpin many aspects of crop productivity. An important question for crop improvement is "How can breeders and agronomists effectively explore the diverse opportunities within the high dimensionality of the complex G × E × M factorial to achieve sustainable improvements in crop productivity?" Whenever G × E × M interactions make important contributions to attainment of crop productivity, we should consider how to design crop improvement strategies that can explore the potential space of G × E × M possibilities, reveal the interesting Genotype-Management (G-M) technology opportunities for the Target Population of Environments (TPE), and enable the practical exploitation of the associated improved levels of crop productivity under on-farm conditions. Climate change adds additional layers of complexity and uncertainty to this challenge, by introducing directional changes in the environmental dimension of the G × E × M factorial. These directional changes have the potential to create further conditional changes in the contributions of the genetic and management dimensions to future crop productivity. Therefore, in the presence of G × E × M interactions and climate change, the challenge for both breeders and agronomists is to co-design new G-M technologies for a non-stationary TPE. Understanding these conditional changes in crop productivity through the relevant sciences for each dimension, Genotype, Environment, and Management, creates opportunities to predict novel G-M technology combinations suitable to achieve sustainable crop productivity and global food security targets for the likely climate change scenarios. Here we consider critical foundations required for any prediction framework that aims to move us from the current unprepared state of describing G × E × M outcomes to a future responsive state equipped to predict the crop productivity consequences of G-M technology combinations for the range of environmental conditions expected for a complex, non-stationary TPE under the influences of climate change.

Towards a multiscale crop modelling framework for climate change adaptation assessment.

Predicting the consequences of manipulating genotype (G) and agronomic management (M) on agricultural ecosystem performances under future environmental (E) conditions remains a challenge. Crop modelling has the potential to enable society to assess the efficacy of G × M technologies to mitigate and adapt crop production systems to climate change. Despite recent achievements, dedicated research to develop and improve modelling capabilities from gene to global scales is needed to provide guidance on designing G × M adaptation strategies with full consideration of their impacts on both crop productivity and ecosystem sustainability under varying climatic conditions. Opportunities to advance the multiscale crop modelling framework include representing crop genetic traits, interfacing crop models with large-scale models, improving the representation of physiological responses to climate change and management practices, closing data gaps and harnessing multisource data to improve model predictability and enable identification of emergent relationships. A fundamental challenge in multiscale prediction is the balance between process details required to assess the intervention and predictability of the system at the scales feasible to measure the impact. An advanced multiscale crop modelling framework will enable a gene-to-farm design of resilient and sustainable crop production systems under a changing climate at regional-to-global scales.

Genotypic variation in whole-plant transpiration efficiency in sorghum only partly aligns with variation in stomatal conductance.

Water scarcity can limit sorghum (Sorghum bicolor (L.) Moench) production in dryland agriculture, but increased whole-plant transpiration efficiency (TEwp, biomass production per unit of water transpired) can enhance grain yield in such conditions. The objectives of this study were to quantify variation in TEwp for 27 sorghum genotypes and explore the linkages of this variation to responses of the underpinning leaf-level processes to environmental conditions. Individual plants were grown in large lysimeters in two well-watered experiments. Whole-plant transpiration per unit of green leaf area (TGLA) was monitored continuously and stomatal conductance and maximum photosynthetic capacity were measured during sunny conditions on recently expanded leaves. Leaf chlorophyll measurements of the upper five leaves of the main shoot were conducted during early grain filling. TEwp was determined at harvest. The results showed that diurnal patterns in TGLA were determined by vapour pressure deficit (VPD) and by the response of whole-plant conductance to radiation and VPD. Significant genotypic variation in the response of TGLA to VPD occurred and was related to genotypic differences in stomatal conductance. However, variation in TGLA explained only part of the variation in TEwp, with some of the residual variation explained by leaf chlorophyll readings, which were a reflection of photosynthetic capacity. Genotypes with different genetic background often differed in TEwp, TGLA and leaf chlorophyll, indicating potential differences in photosynthetic capacity among these groups. Observed differences in TEwp and its component traits can affect adaptation to drought stress.

Water Use Efficiency as a Constraint and Target for Improving the Resilience and Productivity of C3 and C4 Crops.

The ratio of plant carbon gain to water use, known as water use efficiency (WUE), has long been recognized as a key constraint on crop production and an important target for crop improvement. WUE is a physiologically and genetically complex trait that can be defined at a range of scales. Many component traits directly influence WUE, including photosynthesis, stomatal and mesophyll conductances, and canopy structure. Interactions of carbon and water relations with diverse aspects of the environment and crop development also modulate WUE. As a consequence, enhancing WUE by breeding or biotechnology has proven challenging but not impossible. This review aims to synthesize new knowledge of WUE arising from advances in phenotyping, modeling, physiology, genetics, and molecular biology in the context of classical theoretical principles. In addition, we discuss how rising atmospheric CO2 concentration has created and will continue to create opportunities for enhancing WUE by modifying the trade-off between photosynthesis and transpiration.

Quantifying impacts of enhancing photosynthesis on crop yield.

Enhancing photosynthesis is widely accepted as critical to advancing crop yield. However, yield consequences of photosynthetic manipulation are confounded by feedback effects arising from interactions with crop growth, development dynamics and the prevailing environment. Here, we developed a cross-scale modelling capability that connects leaf photosynthesis to crop yield in a manner that addresses the confounding factors. The model was validated using data on crop biomass and yield for wheat and sorghum from diverse field experiments. Consequences for yield were simulated for major photosynthetic enhancement targets related to leaf CO2 and light energy capture efficiencies, and for combinations of these targets. Predicted impacts showed marked variation and were dependent on the photosynthetic enhancement, crop type and environment, especially the degree of water limitation. The importance of interdependencies operating across scales of biological organization was highlighted, as was the need to increase understanding and modelling of the photosynthesis-stomatal conductance link to better quantify impacts of enhancing photosynthesis.

Simulating daily field crop canopy photosynthesis: an integrated software package.

Photosynthetic manipulation is seen as a promising avenue for advancing field crop productivity. However, progress is constrained by the lack of connection between leaf-level photosynthetic manipulation and crop performance. Here we report on the development of a model of diurnal canopy photosynthesis for well watered conditions by using biochemical models of C3 and C4 photosynthesis upscaled to the canopy level using the simple and robust sun-shade leaves representation of the canopy. The canopy model was integrated over the time course of the day for diurnal canopy photosynthesis simulation. Rationality analysis of the model showed that it simulated the expected responses in diurnal canopy photosynthesis and daily biomass accumulation to key environmental factors (i.e. radiation, temperature and CO2), canopy attributes (e.g. leaf area index and leaf angle) and canopy nitrogen status (i.e. specific leaf nitrogen and its profile through the canopy). This Diurnal Canopy Photosynthesis Simulator (DCaPS) was developed into a web-based application to enhance usability of the model. Applications of the DCaPS package for assessing likely canopy-level consequences of changes in photosynthetic properties and its implications for connecting photosynthesis with crop growth and development modelling are discussed.

Multi-Spectral Imaging from an Unmanned Aerial Vehicle Enables the Assessment of Seasonal Leaf Area Dynamics of Sorghum Breeding Lines.

Genetic improvement in sorghum breeding programs requires the assessment of adaptation traits in small-plot breeding trials across multiple environments. Many of these phenotypic assessments are made by manual measurement or visual scoring, both of which are time consuming and expensive. This limits trial size and the potential for genetic gain. In addition, these methods are typically restricted to point estimates of particular traits, such as leaf senescence or flowering and do not capture the dynamic nature of crop growth. In water-limited environments in particular, information on leaf area development over time would provide valuable insight into water use and adaptation to water scarcity during specific phenological stages of crop development. Current methods to estimate plant leaf area index (LAI) involve destructive sampling and are not practical in breeding. Unmanned aerial vehicles (UAV) and proximal-sensing technologies open new opportunities to assess these traits multiple times in large small-plot trials. We analyzed vegetation-specific crop indices obtained from a narrowband multi-spectral camera on board a UAV platform flown over a small pilot trial with 30 plots (10 genotypes randomized within 3 blocks). Due to variable emergence we were able to assess the utility of these vegetation indices to estimate canopy cover and LAI over a large range of plant densities. We found good correlations between the Normalized Difference Vegetation Index (NDVI) and the Enhanced Vegetation Index (EVI) with plant number per plot, canopy cover and LAI both during the vegetative growth phase (pre-anthesis) and at maximum canopy cover shortly after anthesis. We also analyzed the utility of time-sequence data to assess the senescence pattern of sorghum genotypes known as fast (senescent) or slow senescing (stay-green) types. The Normalized Difference Red Edge (NDRE) index which estimates leaf chlorophyll content was most useful in characterizing the leaf area dynamics/senescence patterns of contrasting genotypes. These methods to monitor dynamics of green and senesced leaf area are suitable for out-scaling to enhance phenotyping of additional crop canopy characteristics and likely crop yield responses among genotypes across large fields and multiple dates.

Connecting Biochemical Photosynthesis Models with Crop Models to Support Crop Improvement.

The next advance in field crop productivity will likely need to come from improving crop use efficiency of resources (e.g., light, water, and nitrogen), aspects of which are closely linked with overall crop photosynthetic efficiency. Progress in genetic manipulation of photosynthesis is confounded by uncertainties of consequences at crop level because of difficulties connecting across scales. Crop growth and development simulation models that integrate across biological levels of organization and use a gene-to-phenotype modeling approach may present a way forward. There has been a long history of development of crop models capable of simulating dynamics of crop physiological attributes. Many crop models incorporate canopy photosynthesis (source) as a key driver for crop growth, while others derive crop growth from the balance between source- and sink-limitations. Modeling leaf photosynthesis has progressed from empirical modeling via light response curves to a more mechanistic basis, having clearer links to the underlying biochemical processes of photosynthesis. Cross-scale modeling that connects models at the biochemical and crop levels and utilizes developments in upscaling leaf-level models to canopy models has the potential to bridge the gap between photosynthetic manipulation at the biochemical level and its consequences on crop productivity. Here we review approaches to this emerging cross-scale modeling framework and reinforce the need for connections across levels of modeling. Further, we propose strategies for connecting biochemical models of photosynthesis into the cross-scale modeling framework to support crop improvement through photosynthetic manipulation.

Hybrid variation for root system efficiency in maize: potential links to drought adaptation.

Water availability can limit maize (Zea mays L.) yields, and root traits may enhance drought adaptation if they can moderate temporal patterns of soil water extraction to favour grain filling. Root system efficiency (RSE), defined as transpiration per unit leaf area per unit of root mass, represents the functional mass allocation to roots to support water capture relative to the allocation to aerial mass that determines water demand. The aims of this study were to identify the presence of hybrid variation for RSE in maize, determine plant attributes that drive these differences and illustrate possible links of RSE to drought adaptation via associations with water extraction patterns. Individual plants for a range of maize hybrids were grown in large containers in shadehouses in Queensland, Australia. Leaf area, shoot and root mass, transpiration, root distribution and soil water were measured in all or selected experiments. Significant hybrid differences in RSE existed. High RSE was associated with reduced dry mass allocation to roots and more efficient water capture per unit of root mass. It was also weakly negatively associated with total plant dry mass, reducing preanthesis water use. This could increase grain yield under drought. RSE provides a conceptual physiological framework to identify traits for high-throughput phenotyping in breeding programs.

Soil water capture trends over 50 years of single-cross maize (Zea mays L.) breeding in the US corn-belt.

Breeders have successfully improved maize (Zea mays L.) grain yield for the conditions of the US corn-belt over the past 80 years, with the past 50 years utilizing single-cross hybrids. Long-term improvement for grain yield under water-limited conditions has also been reported. Grain yield under water-limited conditions depends on water use, water use efficiency, and harvest index. It has been hypothesized that long-term genetic gain for yield could be due, in part, to increased water capture from the soil. This hypothesis was tested using a set of elite single-cross hybrids that were released by DuPont Pioneer between 1963 and 2009. Eighteen hybrids were grown in the field during 2010 and 2011 growing seasons at Woodland, CA, USA. Crops grew predominantly on stored soil water and drought stress increased as the season progressed. Soil water content was measured to 300cm depth throughout the growing season. Significant water extraction occurred to a depth of 240-300cm and seasonal water use was calculated from the change in soil water over this rooting zone. Grain yield increased significantly with year of commercialization, but no such trend was observed for total water extraction. Therefore, the measured genetic gain for yield for the period represented by this set of hybrids must be related to either increased efficiency of water use or increased carbon partitioning to the grain, rather than increased soil water uptake.