Crop/Plant Modeling Supports Plant Breeding: II. Guidance of Functional Plant Phenotyping for Trait Discovery.

Observable morphological traits are widely employed in plant phenotyping for breeding use, which are often the external phenotypes driven by a chain of functional actions in plants. Identifying and phenotyping inherently functional traits for crop improvement toward high yields or adaptation to harsh environments remains a major challenge. Prediction of whole-plant performance in functional-structural plant models (FSPMs) is driven by plant growth algorithms based on organ scale wrapped up with micro-environments. In particular, the models are flexible for scaling down or up through specific functions at the organ nexus, allowing the prediction of crop system behaviors from the genome to the field. As such, by virtue of FSPMs, model parameters that determine organogenesis, development, biomass production, allocation, and morphogenesis from a molecular to the whole plant level can be profiled systematically and made readily available for phenotyping. FSPMs can provide rich functional traits representing biological regulatory mechanisms at various scales in a dynamic system, e.g., Rubisco carboxylation rate, mesophyll conductance, specific leaf nitrogen, radiation use efficiency, and source-sink ratio apart from morphological traits. High-throughput phenotyping such traits is also discussed, which provides an unprecedented opportunity to evolve FSPMs. This will accelerate the co-evolution of FSPMs and plant phenomics, and thus improving breeding efficiency. To expand the great promise of FSPMs in crop science, FSPMs still need more effort in multiscale, mechanistic, reproductive organ, and root system modeling. In summary, this study demonstrates that FSPMs are invaluable tools in guiding functional trait phenotyping at various scales and can thus provide abundant functional targets for phenotyping toward crop improvement.

Crop/Plant Modeling Supports Plant Breeding: I. Optimization of Environmental Factors in Accelerating Crop Growth and Development for Speed Breeding.

The environmental conditions in customered speed breeding practice are, to some extent, empirical and, thus, can be further optimized. Crop and plant models have been developed as powerful tools in predicting growth and development under various environments for extensive crop species. To improve speed breeding, crop models can be used to predict the phenotypes resulted from genotype by environment by management at the population level, while plant models can be used to examine 3-dimensional plant architectural development by microenvironments at the organ level. By justifying the simulations via numerous virtual trials using models in testing genotype × environment × management, an optimized combination of environmental factors in achieving desired plant phenotypes can be quickly determined. Artificial intelligence in assisting for optimization is also discussed. We admit that the appropriate modifications on modeling algorithms or adding new modules may be necessary in optimizing speed breeding for specific uses. Overall, this review demonstrates that crop and plant models are promising tools in providing the optimized combinations of environment factors in advancing crop growth and development for speed breeding.

Securing maize reproductive success under drought stress by harnessing CO2 fertilization for greater productivity.

Securing maize grain yield is crucial to meet food and energy needs for the future growing population, especially under frequent drought events and elevated CO2 (eCO2) due to climate change. To maximize the kernel setting rate under drought stress is a key strategy in battling against the negative impacts. Firstly, we summarize the major limitations to leaf source and kernel sink in maize under drought stress, and identified that loss in grain yield is mainly attributed to reduced kernel set. Reproductive drought tolerance can be realized by collective contribution with a greater assimilate import into ear, more available sugars for ovary and silk use, and higher capacity to remobilize assimilate reserve. As such, utilization of CO2 fertilization by improved photosynthesis and greater reserve remobilization is a key strategy for coping with drought stress under climate change condition. We propose that optimizing planting methods and mining natural genetic variation still need to be done continuously, meanwhile, by virtue of advanced genetic engineering and plant phenomics tools, the breeding program of higher photosynthetic efficiency maize varieties adapted to eCO2 can be accelerated. Consequently, stabilizing maize production under drought stress can be achieved by securing reproductive success by harnessing CO2 fertilization.

Effects of mulching types on the yield and water utilization by broomcorn millet (Panicum miliaceum): Results of a study in the Loess Plateau, China.

Mulching practices have been widely adopted to improve rainfed crop productivity. However, the major resources including water, heat, and light that influenced the yield of broomcorn millet in different dryland regions have rarely been explored. A three-season field experiment with three mulching practices i.e. traditional planting with non-mulching (TP), ridge-furrow mulching system (RF), and plastic film mulching (PFM) was conducted in three semi-arid regions in the Loess Plateau, China, i.e. Guyuan city (GY), Huining county (HN), and Yulin city (YL) between 2020 and 2022 to investigate the impacts of mulching regimes on soil hydrothermal conditions, agronomic characteristics, leaf photosynthetic properties, broomcorn millet yield, and water use efficiency (WUE). Results showed that both PFM and RF treatments increased soil temperature and moisture, and enhanced dry matter accumulation by promoting leaf photosynthetic capacity and chlorophyll content, thereby improving broomcorn millet yield and WUE. PFM and RF increased the average broomcorn millet yield by 15.08% and 24.86% at GY site, 20.86% and 25.61% at HN site, and 15.75% and 25.57% at YL site, respectively, and increased the average WUE by 16.31% and 27.48% at GY site, 23.21% and 28.80% at HN site, 15.55% and 28.57% at YL site, respectively. Partial least squares path modeling analysis revealed that soil moisture was an important environmental factor in determining broomcorn millet yield. Overall, RF practice can be taken to improve the management of agricultural climate factors and maximize yield, thereby promoting the sustainable development of dryland agriculture in the Loess Plateau.

Improving wheat seedling quality through deep ploughing and soil compaction at sowing in lime concretion black soil.

The straw incorporation in lime concretion black soil compromises the emergence and quality of winter wheat seedlings in Huaibei Plain, China, lowering the potential of wheat productivity. To overcome the disadvantage, a two-year field experiment was conducted in 2017-18 and 2018-19 to investigate the effects of different tillage modes on seedling emergence and subsequent seedling growth, and final grain yield (GY) in winter wheat. The modes are rotary tillage with compaction after sowing (RCT), rotary tillage after deep ploughing (PT) and rotary tillage after deep ploughing with compaction after sowing (PCT), with the traditional rotary tillage (RT) method as the control. Compared to RT, greater soil moisture content (SMC) at the seedling stage was observed in deep ploughing or compaction treatment, and the highest SMC was achieved in PCT; the time of reaching the maximum number of seedlings was 1 d sooner in RCT or PT, and 3 d in PCT; the seedling number in RCT, PT and PCT was significantly increased by 32.6%, 34.5% and 61.5% respectively. The population size, shoot and root growth of winter wheat in ploughing mode was significantly enhanced than that of rotary treatment at the over-wintering stage; compared to no compaction after sowing, plant growth in compaction treatments was significantly promoted with greater plant population size and height of seedlings. At harvest, GY in RCT, PT and PCT was significantly improved by 5.87%, 10.8% and 16.4%, respectively, compared to RT and the highest GY was achieved in PCT by up to 8, 350.1 kg ha-1 due to the increased spike number. In conclusion, the seedling quality in the straw incorporation practice was improved through rotary after deep ploughing and compaction after sowing for lime concretion black soil in Huaibei Plain, China or a similar soil type.

Effects of Spraying KH2PO4 on Flag Leaf Physiological Characteristics and Grain Yield and Quality under Heat Stress during the Filling Period in Winter Wheat.

As one of the most important wheat-producing areas in China, wheat is prone to heat stress during the grain filling period in the Huang-Huai-Hai Plain (3HP), which lowers yields and degrades the grain quality of wheat. To assess the effects of spraying potassium dihydrogen phosphate (KH2PO4) on the physiological traits in flag leaves and grain yield (GY) and quality under heat stress during the filling period, we conducted a two-year field experiment in the winter wheat growing seasons of 2020-2022. In this study, spraying water combined with heat stress (HT), 0.3% KH2PO4 (KDP), and 0.3% KH2PO4 combined with heat stress (PHT) were designed, and spraying water alone was used as a control (CK). The dates for the spraying were the third and eleventh day after anthesis, and a plastic film shed was used to impose heat stress on the wheat plants during the grain filling period. The results showed that spraying KH2PO4 significantly improved the chlorophyll content and net photosynthesis rate (Pn) in flag leaves compared with the non-sprayed treatments. Compared with CK, the Pn in HT decreased by 8.97% after heat stress, while Pn in PHT decreased by 7.44% compared to that of KDP. The activities of superoxide dismutase, catalase, and peroxidase in flag leaves were significantly reduced when the wheat was subjected to heat stress, while malonaldehyde content increased, and the enzyme activities were significantly enhanced when KH2PO4 was sprayed. Heat stress significantly decreased the contribution rate of dry matter accumulation (DM) after anthesis of wheat to grain (CRAA), whereas spraying KH2PO4 significantly increased the CRAA and harvest index. At maturity, the DM in CK was significantly higher than that in HT, KDP was significantly higher than PHT, and KDP had the highest DM. Compared with CK, the GY in KDP significantly increased by 9.85% over the two years, while the GY in HT decreased by 11.44% compared with that of CK, and the GY in PHT decreased by 6.31% compared to that of KDP. Spraying KH2PO4 after anthesis primarily helped GY by maintaining a high thousand grain weight to lessen the negative effects of heat stress on wheat. Moreover, heat stress significantly reduced protein concentration, wet gluten content, dough development time, and hardness index in grains of mature, while spraying KH2PO4 maintained a sufficient grain quality under the conditions of achieving higher yields. Overall, spraying KH2PO4 after anthesis could enhance the heat stress resistance of wheat and maintain the photosynthetic capacity of flag leaves, ensuring the dry matter production and reducing the negative effects on grain yield and quality in the 3HP.

Characteristics of historical precipitation for winter wheat cropping in the semi-arid and semi-humid area.

Winter wheat (Triticum aestivum L.) is one of major crops in the area along Huai river, China where it is a semi-arid and semi-humid region with sufficient precipitation for an entire season, but with uneven distribution within various growth stages. The instability of precipitation is an important factor in limiting wheat production potential under climate change. Therefore, it is essential to characterise the precipitation associated with different crop developmental stages. Based on climate data from 1999 to 2020 in six representative meteorological stations, we characterised the historical precipitation relating to seven key growth stages in winter wheat. There is no clear trend of interannual variation of precipitation for wheat season, with an average of precipitation of 414.4 ± 121.2 mm. In terms of the distribution of precipitation grade within a season, light rain was dominant. Continuous rain occurred frequently during the pre-winter seedling and overwintering stages. The critical period of water demand, such as jointing and booting, has less precipitation. The fluctuation range of precipitation in sowing, heading-filling and maturation stages is large, which means that there is flood and drought at times. In conclusion, these findings provide a foundation for instructing winter wheat cropping in confronting with waterlogging and drought risk due to uneven precipitation in 'Yanhuai' region, China.

Physiology and Molecular Breeding in Sustaining Wheat Grain Setting and Quality under Spring Cold Stress.

Spring cold stress (SCS) compromises the reproductive growth of wheat, being a major constraint in achieving high grain yield and quality in winter wheat. To sustain wheat productivity in SCS conditions, breeding cultivars conferring cold tolerance is key. In this review, we examine how grain setting and quality traits are affected by SCS, which may occur at the pre-anthesis stage. We have investigated the physiological and molecular mechanisms involved in floret and spikelet SCS tolerance. It includes the protective enzymes scavenging reactive oxygen species (ROS), hormonal adjustment, and carbohydrate metabolism. Lastly, we explored quantitative trait loci (QTLs) that regulate SCS for identifying candidate genes for breeding. The existing cultivars for SCS tolerance were primarily bred on agronomic and morphophysiological traits and lacked in molecular investigations. Therefore, breeding novel wheat cultivars based on QTLs and associated genes underlying the fundamental resistance mechanism is urgently needed to sustain grain setting and quality under SCS.

Overcoming Reproductive Compromise Under Heat Stress in Wheat: Physiological and Genetic Regulation, and Breeding Strategy.

The reproductive compromise under heat stress is a major obstacle to achieve high grain yield and quality in wheat worldwide. Securing reproductive success is the key solution to sustain wheat productivity by understanding the physiological mechanism and molecular basis in conferring heat tolerance and utilizing the candidate gene resources for breeding. In this study, we examined the performance on both carbon supply source (as leaf photosynthetic rate) and carbon sink intake (as grain yields and quality) in wheat under heat stress varying with timing, duration, and intensity, and we further surveyed physiological processes from source to sink and the associated genetic basis in regulating reproductive thermotolerance; in addition, we summarized the quantitative trait loci (QTLs) and genes identified for heat stress tolerance associated with reproductive stages. Discovery of novel genes for thermotolerance is made more efficient via the combination of transcriptomics, proteomics, metabolomics, and phenomics. Gene editing of specific genes for novel varieties governing heat tolerance is also discussed.

Sugar conundrum in plant-pathogen interactions: roles of invertase and sugar transporters depend on pathosystems.

It has been increasingly recognized that CWIN (cell wall invertase) and sugar transporters including STP (sugar transport protein) and SWEET (sugar will eventually be exported transporters) play important roles in plant-pathogen interactions. However, the information available in the literature comes from diverse systems and often yields contradictory findings and conclusions. To solve this puzzle, we provide here a comprehensive assessment of the topic. Our analyses revealed that the regulation of plant-microbe interactions by CWIN, SWEET, and STP is conditioned by the specific pathosystems involved. The roles of CWINs in plant resistance are largely determined by the lifestyle of pathogens (biotrophs versus necrotrophs or hemibiotrophs), possibly through CWIN-mediated salicylic acid or jasmonic acid signaling and programmed cell death pathways. The up-regulation of SWEETs and STPs may enhance or reduce plant resistance, depending on the cellular sites from which pathogens acquire sugars from the host cells. Finally, plants employ unique mechanisms to defend against viral infection, in part through a sugar-based regulation of plasmodesmatal development or aperture. Our appraisal further calls for attention to be paid to the involvement of microbial sugar metabolism and transport in plant-pathogen interactions, which is an integrated but overlooked component of such interactions.

Combating Dual Challenges in Maize Under High Planting Density: Stem Lodging and Kernel Abortion.

High plant density is considered a proficient approach to increase maize production in countries with limited agricultural land; however, this creates a high risk of stem lodging and kernel abortion by reducing the ratio of biomass to the development of the stem and ear. Stem lodging and kernel abortion are major constraints in maize yield production for high plant density cropping; therefore, it is very important to overcome stem lodging and kernel abortion in maize. In this review, we discuss various morphophysiological and genetic characteristics of maize that may reduce the risk of stem lodging and kernel abortion, with a focus on carbohydrate metabolism and partitioning in maize. These characteristics illustrate a strong relationship between stem lodging resistance and kernel abortion. Previous studies have focused on targeting lignin and cellulose accumulation to improve lodging resistance. Nonetheless, a critical analysis of the literature showed that considering sugar metabolism and examining its effects on lodging resistance and kernel abortion in maize may provide considerable results to improve maize productivity. A constructive summary of management approaches that could be used to efficiently control the effects of stem lodging and kernel abortion is also included. The preferred management choice is based on the genotype of maize; nevertheless, various genetic and physiological approaches can control stem lodging and kernel abortion. However, plant growth regulators and nutrient application can also help reduce the risk for stem lodging and kernel abortion in maize.

Functional-Structural Plant Models Mission in Advancing Crop Science: Opportunities and Prospects.

Functional-structural plant models (FSPMs) have been evolving for over 2 decades and their future development, to some extent, depends on the value of potential applications in crop science. To date, stabilizing crop production by identifying valuable traits for novel cultivars adapted to adverse environments is topical in crop science. Thus, this study will examine how FSPMs are able to address new challenges in crop science for sustainable crop production. FSPMs developed to simulate organogenesis, morphogenesis, and physiological activities under various environments and are amenable to downscale to the tissue, cellular, and molecular level or upscale to the whole plant and ecological level. In a modeling framework with independent and interactive modules, advanced algorithms provide morphophysiological details at various scales. FSPMs are shown to be able to: (i) provide crop ideotypes efficiently for optimizing the resource distribution and use for greater productivity and less disease risk, (ii) guide molecular design breeding via linking molecular basis to plant phenotypes as well as enrich crop models with an additional architectural dimension to assist breeding, and (iii) interact with plant phenotyping for molecular breeding in embracing three-dimensional (3D) architectural traits. This study illustrates that FSPMs have great prospects in speeding up precision breeding for specific environments due to the capacity for guiding and integrating ideotypes, phenotyping, molecular design, and linking molecular basis to target phenotypes. Consequently, the promising great applications of FSPMs in crop science will, in turn, accelerate their evolution and vice versa.

Targeting salt stress coping mechanisms for stress tolerance in Brassica: A research perspective.

Brassica genus comprises numerous cultivated brassica species with various economic importance. Salt stress is an overwhelming problem causing serious losses in Brassica species (e.g. B. napus, B. rapa, B. oleracea, B. juncea) growth and grain yield production by inducing ionic and ROS toxicity. Given that a significant variation exists in salt tolerance level in Brassica genus, Brassica species exhibited numerous salt tolerance mechanisms which were either overlooked or given less importance to improve and understand innate salt stress tolerance mechanism in Brassica species. In this review, we tried to highlight the importance and recent findings relating to some overlooked and potential mechanisms such as role of neurotransmitters, and role of cytosolic Ca2+ and ROS as signaling elements to enhance salt stress tolerance. Studies revealed that salt tolerant brassica species retained more K+ in leaf mesophyll which confers overall salinity tolerance in salt tolerance brassica species. Neurotransmitter such as melatonin, dopamiane and eATP regulates K+ and Ca2+ permeable ion channels and plays a very crucial role in ionic homeostasis under salinity stress in brassica. At the end, the numerous possible salt stress agronomic strategies were also discussed to mitigate the severity of the salt stress in Brassica species.

Early nodulin 93 protein gene: essential for induction of somatic embryogenesis in oil palm.

KEY MESSAGE: Transcript profiling during the early induction phase of oil palm tissue culture and RNAi studies in a model somatic embryogenesis system showed that EgENOD93 expression is essential for somatic embryogenesis. Micropropagation of oil palm through tissue culture is vital for the generation of superior and uniform elite planting materials. Studies were carried out to identify genes to distinguish between leaf explants with the potential to develop into embryogenic or non-embryogenic callus. Oil palm cDNA microarrays were co-hybridized with cDNA probes of reference tissue, separately with embryo forming (media T527) and non-embryo (media T694) forming leaf explants sampled at Day 7, Day 14 and Day 21. Analysis of the normalized datasets has identified 77, 115 and 127 significantly differentially expressed genes at Day 7, Day 14, and Day 21, respectively. An early nodulin 93 protein gene (ENOD93), was highly expressed at Day 7, Day 14, and Day 21 and in callus (media T527), as assessed by RT-qPCR. Validation of EgENOD93 across tissue culture lines of different genetic background and media composition showed the potential of this gene as an embryogenic marker. In situ RNA hybridization and functional characterization in Medicago truncatula provided additional evidence that ENOD93 is essential for somatic embryogenesis. This study supports the suitability of EgENOD93 as a marker to predict the potential of leaf explants to produce embryogenic callus. Crosstalk among stresses, auxin, and Nod-factor like signalling molecules likely induces the expression of EgENOD93 for embryogenic callus formation.

Two decades of functional-structural plant modelling: now addressing fundamental questions in systems biology and predictive ecology.

BACKGROUND: Functional-structural plant models (FSPMs) explore and integrate relationships between a plant's structure and processes that underlie its growth and development. In the last 20 years, scientists interested in functional-structural plant modelling have expanded greatly the range of topics covered and now handle dynamical models of growth and development occurring from the microscopic scale, and involving cell division in plant meristems, to the macroscopic scales of whole plants and plant communities. SCOPE: The FSPM approach occupies a central position in plant science; it is at the crossroads of fundamental questions in systems biology and predictive ecology. This special issue of Annals of Botany features selected papers on critical areas covered by FSPMs and examples of comprehensive models that are used to solve theoretical and applied questions, ranging from developmental biology to plant phenotyping and management of plants for agronomic purposes. Altogether, they offer an opportunity to assess the progress, gaps and bottlenecks along the research path originally foreseen for FSPMs two decades ago. This review also allows discussion of current challenges of FSPMs regarding (1) integration of multidisciplinary knowledge, (2) methods for handling complex models, (3) standards to achieve interoperability and greater genericity and (4) understanding of plant functioning across scales. CONCLUSIONS: This approach has demonstrated considerable progress, but has yet to reach its full potential in terms of integration and heuristic knowledge production. The research agenda of functional-structural plant modellers in the coming years should place a greater emphasis on explaining robust emergent patterns, and on the causes of possible deviation from it. Modelling such patterns could indeed fuel both generic integration across scales and transdisciplinary transfer. In particular, it could be beneficial to emergent fields of research such as model-assisted phenotyping and predictive ecology in managed ecosystems.

Modeling Maize Canopy Morphology in Response to Increased Plant Density.

Increased plant density markedly affects canopy morphophysiological activities and crop productivity. This study aims to model maize canopy final morphology under increased interplant competition by revising a functional-structural plant model, i.e., ADEL-Maize. A 2-year field experiment was conducted at Mengcheng, Anhui Province, China, in 2016 and 2018. A randomized complete block design of five plant densities (PDs), i.e., 4.5, 6, 7.5, 9, and 15 plants m-2, with three replications was applied using a hybrid, i.e., Zhengdan 958. Canopy morphology at different PDs was measured with destructive samplings when maize canopy was fully expanded. The relationship of changes of organ morphology in relation to increased plant density was analyzed based on 2016 data. The ADEL-Maize was first calibrated for the hybrid at 4.5 plants m-2 and then revised by introducing relationships identified from 2016 data, followed by independent validation with 2018 field data. A heatmap visualization was shown to clearly illustrate the effects of increased plant density on final morphology of laminae, sheaths, and internodes. The logarithmic + linear equations were found to fit changes for the organ size versus increased plant density for phytomers excluding ear position or linear equations for the phytomer at ear position based on 2016 field data. The revision was then further tested independently by having achieved satisfactory agreements between the simulations and observations in canopy size under different PDs with 2018 field data. In conclusion, this study has characterized the relationship between canopy morphology and increased interplant competition for use in the ADEL-Maize and realized the simulations of final size of laminae, sheaths, and internodes, as affected by increased plant density, laying a foundation to test an ideotype for maize withstanding high interplant competition.

Heat Stress in Legume Seed Setting: Effects, Causes, and Future Prospects.

Grain legumes provide a rich resource of plant nutrition to human diets and are vital for food security and sustainable cropping. Heat stress during flowering has a detrimental effect on legume seed yield, mainly due to irreversible loss of seed number. To start with, we provide an overview of the developmental and physiological basis of controlling seed setting in response to heat stress. It is shown that every single process of seed setting including male and female gametophyte development, fertilization, and early seed/fruit development is sensitive to heat stress, in particular male reproductive development in legume crops is especially susceptible. A series of physiochemical processes including heat shock proteins, antioxidants, metabolites, and hormones centered with sugar starvation are proposed to play a key role in regulating legume seed setting in response to heat stress. The exploration of the molecular mechanisms underlying reproductive heat tolerance is in its infancy. Medicago truncatula, with a small diploid genome, and well-established transformation system and molecular platforms, has become a valuable model for testing gene function that can be applied to advance the physiological and molecular understanding of legume reproductive heat tolerance.

Optimal nitrogen regimes compensate for the impacts of seedlings subjected to waterlogging stress in summer maize.

A field experiment was performed to explore the compensation effects of different nitrogen (N) regimes on the growth and photosynthetic capacity in different leaf layers of the summer maize hybrid of LuYu9105 under waterlogging at the seedling stage. The results showed that waterlogging significantly decreased the maximum green leaf area (gLA) by 10.0~15.3% and 9.3~22.5%, mainly due to the reduction in the below-ear layer leaves at the silking stage in 2014 and 2015, respectively. Waterlogging also significantly decreased the ear leaf photosynthetic rate (PN), and Fv/Fm, Fv/Fo, ΦPSII and qP at the below-ear layer leaves at the mid- and late-filling stages, which was accompanied by a reduction in the duration of grain-filling (T) by 2.6~5.9%, thus resulting in a loss of grain yield by 7.0~18.5%. Interestingly, a shift in N from basal application to topdressing at the big flare stage was shown to compensate the adverse effects of waterlogging by through increased gLA and leaf photosynthetic capacity at the ear layer and the above-ear layer, as well as a greater grain-filling rate, resulting in an increase in grain yield by 9.9~27.0% and 17.8~25.8% compared to other N treatments. Therefore, this study showed that optimal nitrogen regimes during maize growth are capable of compensating for the impacts caused by waterlogging at the seedling stage.