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The growing world population and global increases in the standard of living both result in an increasing demand for food, feed and other plant-derived products. In the coming years, plant-based research will be among the major drivers ensuring food security and the expansion of the bio-based economy. Crop productivity is determined by several factors, including the available physical and agricultural resources, crop management, and the resource use efficiency, quality and intrinsic yield potential of the chosen crop. This review focuses on intrinsic yield potential, since understanding its determinants and their biological basis will allow to maximize the plant's potential in food and energy production. Yield potential is determined by a variety of complex traits that integrate strictly regulated processes and their underlying gene regulatory networks. Due to this inherent complexity, numerous potential targets have been identified that could be exploited to increase crop yield. These encompass diverse metabolic and physical processes at the cellular, organ and canopy level. We present an overview of some of the distinct biological processes considered to be crucial for yield determination that could further be exploited to improve future crop productivity.

Crop models can support the design of smart crop management practices. The Farquhar-von Caemmerer-Berry (FvCB) model is increasingly being used in these models for quantifying leaf photosynthesis. Nitrogen (N) is required for many functional machineries of photosynthesis, thus relationships between FvCB-model parameters and leaf N content (LNC) should be established. We conducted combined gas exchange and chlorophyll fluorescence measurements on fully expanded leaves of two C3 crops, rice (Oryza sativa) and wheat (Triticum aestivum), and two C4 crops, maize (Zea mays) and sorghum (Sorghum bicolor), grown under three N levels. Photosynthetic parameters were estimated and linear relationships between these parameters and LNC were quantified in both C3 and C4 crop types. The efficiency of converting incident light into linear electron transport for C3 crops or into ATP production for C4 crops showed a weak increase with LNC. The maximum electron transport rate (Jmax) for C3 crops or the maximum ATP production rate (Jmax,atp) for C4 crops significantly increased with LNC. The increase in Rubisco carboxylation capacity (Vcmax) with LNC was significantly higher in C3 than in C4 crops. Triose phosphate utilization for C3 crops and PEP carboxylation capacity (Vpmax) for C4 crops increased significantly with LNC as well. Except for Jmax at 21% O2 and Vcmax of C3 crops, there was no significant difference among crops in the relationship between estimated photosynthetic parameters and LNC. The tight associations of photosynthesis parameters with LNC were discussed in view of decision making on N management in the context of smart farming.

Thousands of scientific papers have described how plants responded to different levels of a given environmental factor, for a wide variety of physiological processes and morphological, anatomical or chemical characteristics. There is a clear need to summarize this information in a structured and comparable way through meta-analysis. This paper describes how to use relative trait responses from many independent experiments to create generalized dose-response curves. By applying the same methodology to a wide range of plant traits, varying from the molecular to the whole plant level, we can achieve an unprecedented view on the many ways that plants are affected by and acclimate to their environment. We illustrate this approach, which we refer to as ‘MetaPhenomics’, with a variety of previously published and unpublished dose-response curves of the effect of light intensity on 25 plant traits. Furthermore, we discuss the need and difficulties to expand this approach to the transcriptomics and metabolomics level, and show how the generalized dose-response curves can be used to improve simulation models as well as the communication between modelers and experimental plant biologists.

Drought poses a serious threat to crop production worldwide, and is expected particularly to affect rice production and hence food security. Given that wheat is known to tolerate drought better than rice, we compare rice and wheat (cv. Weebill) to understand the species level differences in drought adaptive mechanisms. We also compare two contrasting rice genotypes (IR64, drought susceptible, and Apo, drought tolerant) for such mechanisms under well-watered (100% field capacity, 100%FC) and water-limited (60%FC) conditions. The reduction in biomass of wheat under water limitation was smaller due to a higher rate of photosynthesis associated with maintenance of tissue turgor compared to rice genotypes. Drought caused greater inhibition of Photosystem II quantum efficiency, carboxylation efficiency, and photosynthetic capacity parameters in IR64 than in Apo. Transcript levels of photosynthesis-related genes were also significantly more repressed by water limitation in IR64, whilst the wheat genotype showed smaller reduction than Apo. Despite higher non-photochemical quenching (NPQ), a smaller increase in scavenging enzymes in IR64 resulted in more accumulation of reactive oxygen species (ROS) in 60%FC than in 100%FC compared to Apo. As a photoprotection mechanism, increased levels of NPQ resulted in lower ROS accumulation in wheat despite the similar increase in scavenging enzyme transcript levels as in Apo, signifying the importance of preventing oxidative burst for enhanced drought tolerance. In Apo, upregulation of the 9-cis-epoxycarotenoid dioxygenase 2 gene implies the use of xanthophyll pool for the ABA biosynthesis. Our data suggest that regulating photosynthesis and oxidative protection in the wheat genotype enhanced drought tolerance. Improving these traits for rice is crucial to develop drought-tolerant rice genotypes.

CONTEXT: To avoid excessive chemical-fertilizer application and improve agricultural productivity in rice, a fertilizer recommendation system called Nutrient Expert® (NE) was designed. However, the ability of NE to balance yield, profitability and environmental sustainability in rice production needs to be further evaluated, as it is still difficult for farmers to assess the proper nitrogen (N) application rate. OBJECTIVE: The objective of this study was to demonstrate the advantages of the NE system in balancing yield, profitability and N loss in rice production, and recommend proper N application rates for different cropping seasons of rice. METHODS: This study describes results from field experiments conducted in five main rice cropping provinces from 2017 to 2020 in China, to investigate any advantages of NE compared with local farmers' practice (FP), and to determine the proper N application rates for different cropping seasons of rice. RESULTS AND CONCLUSIONS: Compared with FP, NE had 12.1% lower N-fertilizer application, but increased rice grain yield by 4.3% and net profit by 7.4%, and decreased yield-scaled N loss by 20.7%. We showed how yield, profitability and N loss were affected by N balance (i.e., N applied to the field minus N removed from the field by the harvested crop biomass), and quantified relationships between N balance and N application rate, and between N output and input. Based on relationships between N balance and N application rate, we recommend N application rates in a range from 122 to 214 kg ha−1, depending on cropping seasons of rice. We demonstrated that NE can simultaneously improve yield, profitability and environmental sustainability. SIGNIFICANCE: Our study provided quantitative support for NE-based recommendations on the N application rate for smallholders farming in different rice cropping systems, and these recommendations can serve as a reference for avoiding excessive N application rate in paddy fields in other regions with similar eco-environment.

C4 crops of agricultural importance all belong to the NADP-malic enzyme (ME) subtype, and this subtype has been the template for C4 introductions into C3 crops, like rice, to improve their productivity. However, the ATP cost for the C4 cycle in both NADP-ME and NAD-ME subtypes accounts for > 40% of the total ATP requirement for CO2 assimilation. These high ATP costs, and the associated need for intense cyclic electron transport and low intrinsic quantum yield Φ CO 2 , are major constraints in realizing strong improvements of canopy photosynthesis and crop productivity. Based on mathematical modelling, we propose a C4 ideotype that utilizes low chloroplastic ATP requirements present in the nondomesticated phosphoenolpyruvate carboxykinase (PEP-CK) subtype. The ideotype is a mixed form of NAD(P)-ME and PEP-CK types, requires no cyclic electron transport under low irradiances, and its theoretical Φ CO 2 is c. 25% higher than that of a C4 crop type. Its cell-type-specific ATP and NADPH requirements can be fulfilled by local energy production. The ideotype is projected to have c. 10% yield advantage over NADP-ME-type crops and > 50% advantage over C3 counterparts. The ideotype provides a unique (theoretical) case where Φ CO 2 could be improved, thereby paving a new avenue for improving photosynthesis in both C3 and C4 crops.

Rice yield reduction due to water limitation depends on its severity and duration and on the phenological stage of its occurrence. We exposed three contrasting rice genotypes, IR64, UPLRi7 and Apo (adapted to lowland, upland and aerobic conditions, respectively), to three water regimes (puddle, 100% and 60% field capacity) in pots during the vegetative (GSI), flowering (GSII) and grain filling (GSIII) stages. Stress at all the three stages significantly reduced yield especially in lowland genotype IR64. Effect of water limitation was more severe at GSII than at the other two stages. Stress at GSI stage reduced both source activity (leaf area and photosynthetic rate) and sink capacity (tiller number or panicle number per pot). When stress was imposed at GSII, spikelet fertility was most affected in all the three genotypes. In both GSII and GSIII, although leaf area was constant in all the three water regimes, estimated relative whole-plant photosynthesis was strongly associated with yield reduction. Reduced photosynthesis due to stress at any given stage was found to have direct impact on yield. Compared to the other genotypes, Apo had deeper roots and maintained a better water relation, thus, higher carbon gain and spikelet viability, and ultimately, higher biomass and productivity under water-limited conditions. Therefore, screening for these stage-dependent adaptive mechanisms is crucial in breeding for sustained rice production under water limitation.

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.

Crops show considerable capacity to adjust their photosynthetic characteristics to seasonal changes in temperature. However, how photosynthesis acclimates to changes in seasonal temperature under future climate conditions has not been revealed. We measured leaf photosynthesis (An ) of wheat (Triticum aestivum L.) and rice (Oryza sativa L.) grown under four combinations of two levels of CO2 (ambient and enriched up to 500 µmol/mol) and two levels of canopy temperature (ambient and increased by 1.5-2.0°C) in temperature by free-air CO2 enrichment (T-FACE) systems. Parameters of a biochemical C3 -photosynthesis model and of a stomatal conductance (gs ) model were estimated for the four conditions and for several crop stages. Some biochemical parameters related to electron transport and most gs parameters showed acclimation to seasonal growth temperature in both crops. The acclimation response did not differ much between wheat and rice, nor among the four treatments of the T-FACE systems, when the difference in the seasonal growth temperature was accounted for. The relationships between biochemical parameters and leaf nitrogen content were consistent across leaf ranks, developmental stages, and treatment conditions. The acclimation had a strong impact on gs model parameters: when parameter values of a particular stage were used, the model failed to correctly estimate gs values of other stages. Further analysis using the coupled gs -biochemical photosynthesis model showed that ignoring the acclimation effect did not result in critical errors in estimating leaf photosynthesis under future climate, as long as parameter values were measured or derived from data obtained before flowering.

Global dimming, a decadal decrease in incident global radiation, is often accompanied with an increase in the diffuse radiation fraction, and, therefore, the impact of global dimming on crop production is hard to predict. A popular approach to quantify this impact is the statistical analysis of historical climate and crop data, or use of dynamic crop simulation modelling approach. Here, we show that statistical analysis of historical data did not provide plausible values for the effect of diffuse radiation versus direct radiation on rice or wheat yield. In contrast, our field experimental study of 3 years demonstrated a fertilization effect of increased diffuse radiation fraction, which partly offset yield losses caused by decreased global radiation, in both crops. The fertilization effect was not attributed to any improved canopy light interception but mainly to the increased radiation use efficiency (RUE). The increased RUE was explained not only by the saturating shape of photosynthetic light response curves but also by plant acclimation to dimming that gradually increased leaf nitrogen concentration. Crop harvest index slightly decreased under dimming, thereby discounting the fertilization effect on crop yields. These results challenge existing modelling paradigms, which assume that the fertilization effect on crop yields is mainly attributed to an improved light interception. Further studies on the physiological mechanism of plant acclimation are required to better quantify the global dimming impact on agroecosystem productivity under future climate change.

Atmospheric CO2 concentration [CO2] has increased from 260 to 280 μmol mol−1 (level during crop domestication up to the industrial revolution) to currently 400 and will reach 550 μmol mol−1 by 2050. C3 crops are expected to benefit from elevated [CO2] (e-CO2) thanks to photosynthesis responsiveness to [CO2] but this may require greater sink capacity. We review recent literature on crop e-CO2 responses, related source-sink interactions, how abiotic stresses potentially interact, and prospects to improve e-CO2 response via breeding or genetic engineering. Several lines of evidence suggest that e-CO2 responsiveness is related either to sink intrinsic capacity or adaptive plasticity, for example, involving enhanced branching. Wild relatives and old cultivars mostly showed lower photosynthetic rates, less downward acclimation of photosynthesis to e-CO2 and responded strongly to e-CO2 due to greater phenotypic plasticity. While reverting to such archaic traits would be an inappropriate strategy for breeding, we argue that substantial enhancement of vegetative sink vigor, inflorescence size and/or number and root sinks will be necessary to fully benefit from e-CO2. Potential ideotype features based on enhanced sinks are discussed. The generic 'feast-famine' sugar signaling pathway may be suited to engineer sink strength tissue-specifically and stage-specifically and help validate ideotype concepts. Finally, we argue that models better accounting for acclimation to e-CO2 are needed to predict which trait combinations should be targeted by breeders for a CO2-rich world.

We assessed how the temperature response of leaf day respiration (Rd) in wheat responded to contrasting water regimes and growth temperatures. In Experiment 1, well-watered and drought-stressed conditions were imposed on two genotypes; in Experiment 2, the two water regimes combined with high (HT), medium (MT) and low (LT) growth temperatures were imposed on one of the genotypes. Rd was estimated from simultaneous gas exchange and chlorophyll fluorescence measurements at six leaf temperatures (Tleaf) for each treatment, using the Yin method for non-photorespiratory conditions and the non-rectangular hyperbolic fitting method for photorespiratory conditions. The two genotypes responded similarly to growth and measurement conditions. Estimates of Rd for non-photorespiratory conditions were generally higher than those for photorespiratory conditions but their responses to Tleaf were similar. Under well-watered conditions, Rd and its sensitivity to Tleaf slightly acclimated to LT but did not acclimate to HT. Temperature sensitivities of Rd were considerably suppressed by drought, and the suppression varied among growth temperatures. Thus, it is necessary to quantify interactions between drought and growth temperature for reliably modelling Rd under climate change. Our study also demonstrated that the Kok method, a currently popular method for estimating Rd, underestimated Rd significantly and should be abandoned.HighlightLeaf day respiration (Rd) acclimated little to growth temperature, but significantly to drought by reducing its thermal sensitivity. The Kok method underestimates Rd and should no longer be used.

Increasing fresh water scarcity is a major constraint for rice (Oryza sativa) productivity under future climatic conditions. It is important to understand why rice cannot perform like other dryland cereals, when grown under drier soil conditions. The responses of plant growth, plant water status, leaf and stem anatomical attributes to soil water deficit were examined at two developmental stages of three rice and two wheat (Triticum aestivum) cultivars grown under three water regimes in a pot experiment. Yield and yield components were determined at plant maturity. The stress sensitivity of plant growth was slightly more prominent at vegetative than at flowering stages. Water deficit limited root growth but had little impact on above-ground growth in rice plants. The xylem system of the leaves was more developed in rice than in wheat, but the opposite was true for the root xylem system. Compared with stem xylem attributes, leaf xylem attributes played a larger role in modulating leaf water potential under water deficit. In addition, the variation in yield response to water deficit between various types of rice, and between rice and wheat was closely associated with the response of leaf water potential to water deficit. To grow rice in drier conditions, its xylem system should be balanced by increasing root growth and modifying leaf anatomy.

The Nutrient Expert system (NE) has been proposed to improve yield and nitrogen (N) use efficiency (NUE) in the double rice cropping systems in China. However, the advantage of the NE system has yet to be quantified experimentally. A four-year field experiment was conducted in a double rice cropping system in China, to evaluate the ability of NE in improving yield and NUE. The experimental treatments consisted of NE based fertilization, current farmers’ practices (FP) and soil test based fertilizer recommendation (ST), and a series of nitrogen (N) rate treatments. The NUE decreased with increasing N application, while the yield did not increase significantly beyond N application rates of about 140 kg ha−1 (corresponding to the amount proposed by NE) in both early and late rice. NE increased grain yield (by 10.3% and 6.3%) and N uptake (by 5.7% and 4.0%) compared with FP and ST, respectively. NE significantly increased NUE compared with FP, and decreased the N surplus in comparison to FP and ST. The N dilution curve was Nc = 34.50 W−0.55 for early rice and Nc = 37.71 W−0.59 for late rice (where Nc is the N concentration in g kg−1, and W is the dry matter accumulation in t ha−1). The relationship between relative yield and the nitrogen nutrition index derived from the dilution curves confirmed that NE offered an optimum N application rate (approximately 140 kg ha−1) for both early and late rice. Carbon (C) and N translocation from vegetative organs to grains was enhanced with increasing N rate, while NE significantly increased C and N translocation compared with FP. Overall, the NE system ensured a high rice yield, increased N uptake and NUE. Therefore, the NE, as a user-friendly tool, is a sustainable fertilizer recommendation approach suitable for double rice cropping system, especially when soil testing is not available or timely for smallholders.

Plant responses to abiotic stresses are complex and dynamic, and involve changes in different traits, either as the direct consequence of the stress, or as an active acclimatory response. Abiotic stresses frequently occur simultaneously or in succession, rather than in isolation. Despite this, most studies have focused on a single stress and single or few plant traits. To address this gap, our study comprehensively and categorically quantified the individual and combined effects of three major abiotic stresses associated with climate change (flooding, progressive drought and high temperature) on 12 phenotypic traits related to morphology, development, growth and fitness, at different developmental stages in four Arabidopsis thaliana accessions. Combined sublethal stresses were applied either simultaneously (high temperature and drought) or sequentially (flooding followed by drought). In total, we analysed the phenotypic responses of 1782 individuals across these stresses and different developmental stages. Overall, abiotic stresses and their combinations resulted in distinct patterns of effects across the traits analysed, with both quantitative and qualitative differences across accessions. Stress combinations had additive effects on some traits, whereas clear positive and negative interactions were observed for other traits: 9 out of 12 traits for high temperature and drought, 6 out of 12 traits for post-submergence and drought showed significant interactions. In many cases where the stresses interacted, the strength of interactions varied across accessions. Hence, our results indicated a general pattern of response in most phenotypic traits to the different stresses and stress combinations, but it also indicated a natural genetic variation in the strength of these responses. This includes novel results regarding the lack of a response to drought after submergence and a decoupling between leaf number and flowering time after submergence. Overall, our study provides a rich characterization of trait responses of Arabidopsis plants to sublethal abiotic stresses at the phenotypic level and can serve as starting point for further in-depth physiological research and plant modelling efforts.

Breeding for improved leaf photosynthesis is considered as a viable approach to increase crop yield. Whether it should be improved in combination with other traits has not been assessed critically. Based on the quantitative crop model GECROS that interconnects various traits to crop productivity, we review natural variation in relevant traits, from biochemical aspects of leaf photosynthesis to morpho-physiological crop characteristics. While large phenotypic variations (sometimes >2-fold) for leaf photosynthesis and its underlying biochemical parameters were reported, few quantitative trait loci (QTL) were identified, accounting for a small percentage of phenotypic variation. More QTL were reported for sink size (that feeds back on photosynthesis) or morpho-physiological traits (that affect canopy productivity and duration), together explaining a much greater percentage of their phenotypic variation. Traits for both photosynthetic rate and sustaining it during grain filling were strongly related to nitrogen-related traits. Much of the molecular basis of known photosynthesis QTL thus resides in genes controlling photosynthesis indirectly. Simulation using GECROS demonstrated the overwhelming importance of electron transport parameters, compared with the maximum Rubisco activity that largely determines the commonly studied light-saturated photosynthetic rate. Exploiting photosynthetic natural variation might significantly improve crop yield if nitrogen uptake, sink capacity, and other morpho-physiological traits are co-selected synergistically.

On the occasion of the 40th anniversary of the publication of the landmark model by Farquhar, von Caemmerer & Berry on steady‐state C3 photosynthesis (known as the “FvCB model”), we review three major further developments of the model. These include: (1) limitation by triose phosphate utilization, (2) alternative electron transport pathways, and (3) photorespiration‐associated nitrogen and C1 metabolisms. We discussed the relation of the third extension with the two other extensions, and some equivalent extensions to model C4 photosynthesis. In addition, the FvCB model has been coupled with CO2‐diffusion models. We review how these extensions and integration have broadened the use of the FvCB model in understanding photosynthesis, especially with regard to bioenergetic stoichiometries associated with photosynthetic quantum yields. Based on the new insights, we present caveats in applying the FvCB model. Further research needs are highlighted., The year 2020 marks the 40th anniversary of the publication of the landmark model by Farquhar, von Caemmerer & Berry on photosynthesis. Here, we take a historical view on the original model, go into details of how this model has been extended since then, and review how these extensions have broadened the use of the model.

Global dimming reduces incident global radiation but increases the fraction of diffuse radiation, and thus affects crop yields; however, the underlying mechanisms of such an effect have not been revealed. We hypothesized that crop source–sink imbalance of either carbon (C) or nitrogen (N) during grain filling is a key factor underlying the effect of global dimming on yields. We presented a practical framework to assess both C and N source–sink relationships, using data of biomass and N accumulation from periodical sampling conducted in field experiments for wheat and rice from 2013 to 2016. We found a fertilization effect of the increased diffuse radiation fraction under global dimming, which alleviated the negative impact of decreased global radiation on source supply and sink growth, but the source supply and sink growth were still decreased by dimming, for both C and N. In wheat, the C source supply decreased more than the C sink demand, and as a result, crops remobilized more pre-heading C reserves, in response to dimming. However, these responses were converse in rice, which presumably stemmed from the more increment in radiation use efficiency and the more limited sink size in rice than wheat. The global dimming affected source supply and sink growth of C more significantly than that of N. Therefore, yields in both crops were dependent more on the source–sink imbalance of C than that of N during grain filling. Our revealed source–sink relationships, and their differences and similarities between wheat and rice, provide a basis for designing strategies to alleviate the impact of global dimming on crop productivity.

The role of bundle sheath conductance (gbs) in sustaining sugarcane photosynthesis under nitrogen deficiency was investigated. Sugarcane was grown under different levels of nitrogen supply and gbs was estimated using simultaneous measurements of leaf gas exchange and chlorophyll fluorescence at 21% or 2% [O2] and varying air [CO2] and light intensity. Maximum rates of PEPC carboxylation, Rubisco carboxylation, and ATP production increased with an increase in leaf nitrogen concentration (LNC) from 1 to 3 g m−2. Low nitrogen supply reduced Rubisco and PEPC abundancies, the quantum efficiency of CO2 assimilation and gbs. Because of reduced gbs, low photosynthetic rates were not associated with increased leakiness under nitrogen deficiency. In fact, low nitrogen supply increased bundle sheath cell wall thickness, probably accounting for low gbs and increased estimates of [CO2] at Rubisco sites. Effects of nitrogen on expression of ShPIP2;1 and ShPIP1;2 aquaporins did not explain changes in gbs. Our data revealed that reduced Rubisco carboxylation was the main factor causing low sugarcane photosynthesis at low nitrogen supply, in contrast to the previous report on the importance of an impaired CO2 concentration mechanism under N deficiency. Our findings suggest higher investment of nitrogen into Rubisco protein would favour photosynthesis and plant performance under low nitrogen availability.

Major crop models were developed before the 1990s and many of their algorithms are (semi-)empirical. In the recent two decades, the number of models has grown rapidly, but the increase in their quality does not match the growth in number. Often, an ensemble of multiple models is required to make a useful prediction. This ensemble approach does not facilitate the improvement of models as a tool to understand crop physiological mechanisms. On the other hand, following a bottom-up approach that tries to assemble all elements along different scales of biological organisation may result in numerically clumsy models that are not necessarily more robust than existing models in simulating phenotypes at the crop scale. We argue that to model complex crop phenotypes in a simple yet accurate manner, crop modellers should be inspired by experiences in some fundamental sciences. For example, physicists used sound theories and solid mathematics in thought experiments, and came up with seemingly simple equations to explain the behaviour of very diverse systems, from sub-atomic particles to the largest clusters of galaxies. We review examples, where biological insights and mathematics are combined to derive simple equations that apply to different processes of crop growth. The essence of this modelling approach is a combination of simplicity, elegance and robustness. Models integrating those equations can predict crop phenotypes as well as generate hypotheses or emerging properties to assist in knowing the unknowns. Thus, crop models should not only provide practical predictions, but should especially be considered as a research tool for data interpretation, system design, and heuristic understanding.

Drought significantly decreases crop productivity, especially in high water consuming crops like rice. Grain filling is one of the important critical growth phases in rice and drought during this phase leads to significant reduction in yield. In this study, a comparison was made between IR64 (drought susceptible) and Apo (drought tolerant) rice genotypes to capture the response to water limitation (50% field capacity (FC)) compared with the control (100%FC) during grain filling. Plants were grown in a high-throughput phenomics facility for precise imposition of moisture stress during grain filling. Apo performed better in water limited conditions with lower reduction of photosynthetic rate and maintenance of lower leaf temperature than IR64. Days from sowing to maturity, spikelet fertility and seed weight were more impeded by water limitation in IR64 than in Apo. Unlike Apo, IR64 did not show any decrease in transpiration rate at 50%FC compared with 100%FC. Metabolomic profiling of spikelets at grain filling showed distinct effects of water limitation on accumulation of metabolites, especially in Apo. Secondary metabolism, mainly the phenylpropanoid pathway involved in scavenging mechanisms, was upregulated in Apo. Accumulation of most amino acids was significantly higher in IR64 than in Apo. Due to higher rates of photosynthesis under stress, most carbohydrates accumulated more in Apo than in IR64 at 50%FC. Sucrose transporters were significantly upregulated in water limited conditions especially in Apo. Overall, thanks to higher source capacity, more source to sink transport and better scavenging, Apo showed a lower reduction in yield than IR64.

Summary The Kok effect refers to the abrupt decrease around the light compensation point in the slope of net photosynthetic rate vs irradiance. Arguably, this switch arises from light inhibition of respiration, allowing the Kok method to estimate day respiration (R d). Recent analysis suggests that increasing proportions of photorespiration (quantified as Γ*/C c, the ratio of CO2 compensation point Γ* to chloroplast CO2 concentration, C c) with irradiance explain much of the Kok effect. Also, the Kok method has been modified to account for the decrease in PSII photochemical efficiency (Φ2) with irradiance.Using a model that illustrates how varying R d, Γ*/C c, Φ2 and proportions of alternative electron transport could engender the Kok effect, we quantified the contribution of these parameters to the Kok effect measured in sunflower across various O2 and CO2 concentrations and various temperatures.Overall, the decreasing Φ2 with irradiance explained c. 12%, and the varying Γ*/C c explained c. 25%, of the Kok effect. Maximum real light inhibition of R d was much lower than the inhibition derived from the Kok method, but still increased with photorespiration.Photorespiration had a dual contribution to the Kok effect, one via the varying Γ*/C c and the other via its participation in light inhibition of R d.

Aims To face the challenge of decreasing freshwater availability for agriculture, it is important to explore avenues for developing rice genotypes that can be grown like dryland cereals. Roots play a key role in plant adaptation to dry environments. Methods We examined anatomical and histochemical root traits that affect water acquisition in rice (Oryza sativa) and wheat (Triticum aestivum). These traits and root growth were measured at two developmental stages for three rice and two wheat cultivars that were grown in pots under three water regimes. Results Wheat roots had larger xylem sizes than rice roots, which potentially led to a higher axial conductance, especially under water-deficit conditions. Suberization, lignification and thickening of the endodermis in rice roots increased with increasing water deficit, resulting in stronger radial barriers for water flow in rice than in wheat, especially near the root apex. In addition, water deficit strongly impeded root growth and lateral root proliferation in rice, but only slightly in wheat, and cultivars within a species differed little in these responses. The stress sensitivity of rice attributes was slightly more prominent at vegetative than at flowering stages. Conclusions Rice root characteristics, which are essential for growth under inundated conditions, are not conducive to growth under water deficit. Although rice roots show considerable plasticity under different watering regimes, improving root xylem size and reducing the radial barriers would be required if rice is to grow like dryland cereals.

Drought tolerance is governed by constitutive and acquired traits. Combining them has relevance for sustaining crop productivity under drought. Mild levels of stress induce specific mechanisms that protect metabolism when stress becomes severe. Here, we report a comparative assessment of “acquired drought tolerance (ADT)” traits in two rice cultivars, IR64 (drought susceptible) and Apo (tolerant), and a drought-tolerant wheat cultivar, Weebill. Young seedlings were exposed to progressive concentrations of methyl viologen (MV), a stress inducer, before transferring to a severe concentration. “Induced” seedlings showed higher tolerance and recovery growth than seedlings exposed directly to severe stress. A novel phenomic platform with an automated irrigation system was used for precisely imposing soil moisture stress to capture ADT traits during the vegetative stage. Gradual progression of drought was achieved through a software-controlled automated irrigation facility. This facility allowed the maintenance of the same level of soil moisture irrespective of differences in transpiration, and hence, this platform provided the most appropriate method to assess ADT traits. Total biomass decreased more in IR64 than in Apo. The wheat cultivar showed lower levels of damage and higher recovery growth even compared to Apo. Expression of ROS-scavenging enzymes and drought-responsive genes was significantly higher in Apo than in IR64, but differences were only marginal between Apo and Weebill. The wheat cultivar showed significantly higher stomatal conductance, carbon gain, and biomass than the rice cultivars, under drought. These differences in ADT traits between cultivars as well as between species can be utilised for improving drought tolerance in crop plants.

Classical approaches to estimate mesophyll conductance ignore differences in resistance components for CO2from intercellular air spaces (IAS) and CO2from photorespiration (F) and respiration (Rd). Consequently, mesophyll conductance apparently becomes sensitive to (photo)respiration relative to net photosynthesis, (F + Rd)/A. This sensitivity depends on several hard-to-measure anatomical properties of mesophyll cells. We developed a method to estimate the parameterm(0 ≤ m ≤ 1) that lumps these anatomical properties, using gas exchange and chlorophyll fluorescence measurements where (F + Rd)/Aratios vary. This method was applied to tomato and rice leaves measured at five O2levels. The estimatedmwas 0.3 for tomato but 0.0 for rice, suggesting that classical approaches implyingm = 0 work well for rice. The mesophyll conductance taking themfactor into account still responded to irradiance, CO2, and O2levels, similar to response patterns of stomatal conductance to these variables. Largely due to differentmvalues, the fraction of (photo)respired CO2being refixed within mesophyll cells was lower in tomato than in rice. But that was compensated for by the higher fraction via IAS, making the total re-fixation similar for both species. These results, agreeing with CO2compensation point estimates, support our method of effectively analysing mesophyll resistance.

International audience; This study aimed to understand the response of photosynthesis and growth to e-CO2 conditions (800 vs. 400 mu mol mol(-1)) of rice genotypes differing in source-sink relationships. A proxy trait called local C source-sink ratio was defined as the ratio of flag leaf area to the number of spikelets on the corresponding panicle, and five genotypes differing in this ratio were grown in a controlled greenhouse. Differential CO2 resources were applied either during the 2 weeks following heading (EXP1) or during the whole growth cycle (EXP2). Under e-CO2, low source-sink ratio cultivars (LSS) had greater gains in photosynthesis, and they accumulated less nonstructural carbohydrate in the flag leaf than high source-sink ratio cultivars (HSS). In EXP2, grain yield and biomass gain was also greater in LSS probably caused by their strong sink. Photosynthetic capacity response to e-CO2 was negatively correlated across genotypes with local C source-sink ratio, a trait highly conserved across environments. HSS were sink-limited under e-CO2, probably associated with low triose phosphate utilization (TPU) capacity. We suggest that the local C source-sink ratio is a potential target for selecting more CO2-responsive cultivars, pending validation for a broader genotypic spectrum and for field conditions.

High grain nitrogen (N) concentration in crops may require to translocate more N from the vegetative tissues, cause faster plant senescence, alter sink-source balance during grain filling, and ultimately lower grain yield. Previous studies have shown that indica hybrid rice (IH) have shorter grain-filling period, and lower biomass and grain yield, compared with japonica/indica hybrid rice (JIH) and japonica conventional inbred rice (JC) in China. We hypothesise that these are caused by early senescence and poor sink-source balance after heading in IH as a result of its high grain N concentration. In this study, two JIH, two JC, and two IH varieties were grown in 2013 and 2014 at Ningbo, and in 2015 and 2016 at Yangzhou, China, and the observed dynamics of grain and crop biomass were fitted to mathematical equations to quantify sink-source relationships during grain filling. Compared with JC and IH varieties, JIH varieties showed higher sink growth and source capacity and 7.5–21.6% higher grain yield across years and sites. In JIH and JC genotypes, total source during grain filling was higher than the total sink value, whereas opposite was observed for IH genotypes. Source-sink difference, source/sink ratio, and grain-filling duration all negatively (P < 0.01) correlated with grain N concentration at maturity. Compared with JIH and JC, IH accumulated more N in grains and translocated more N from leaf during the period from heading to maturity. We conclude that early senescence and poor sink-source balance after heading of IH was linked to its high grain N concentration, and sink-source relationships during rice grain filling were associated with grain N concentration. Further breeding and cultivation programmes should aim at improving sink-related traits in JIH and JC genotypes, whereas for IH varieties, whether yield was set by sink size or by any limited availability of pre-heading reserves remains to be further examined.

We present a new simulation model of the reactions in the photosynthetic electron transport chain of C3 species. We show that including recent insights about the regulation of the thylakoid proton motive force, ATP/NADPH balancing mechanisms (cyclic and non-cyclic alternative electron transport), and regulation of Rubisco activity, leads to emergent behaviors that may affect the operation and regulation of photosynthesis under different dynamic environmental conditions. The model was parameterized with experimental results in the literature, with a focus on Arabidopsis thaliana. A dataset was constructed from multiple sources, including measurements of steady-state and dynamic gas exchange, chlorophyll fluorescence and absorbance spectroscopy under different light intensities and CO2. This dataset was used to test predictions of the model under different experimental conditions. Simulations suggested that there are strong interactions between cyclic and non-cyclic alternative electron transport and that an excess capacity for alternative electron transport is required to ensure adequate redox state and lumen pH. Furthermore, the model predicted that, under specific conditions, reduction of ferredoxin by plastoquinol was possible, especially after a rapid increase in light intensity. Further analysis also revealed that the relationship between ATP synthesis and proton motive force was highly regulated by the concentrations of ATP and ADP, and this facilitated an increase in non-photochemical quenching under conditions where metabolism was limiting, such as low CO2 or high light intensity. The model may be used as an in silico platform for future research on the regulation of photosynthetic electron transport.

Estimates of mesophyll conductance (gm), when calculated from chlorophyll fluorescence, are uncertain, especially when the photosystem II (PSII) operating efficiency is measured from the traditional single saturation pulse methodology. The multiphase flash method has recently been recommended to replace the single saturation pulse method, allowing a more reliable estimation of gm. Also, many researchers still directly use the PSII operating efficiency to derive linear electron transport rate J (that is required to estimate gm), without appropriate calibration using measurements under non-photorespiratory conditions. Here we demonstrate for tomato and rice that (i) using the multiphase flash method did not yield realistic estimates of gm if no calibration was conducted; and (ii) using the single saturation pulse method still gave reasonable estimates of gm when calibration based on the non-photorespiratory measurements was properly conducted. Therefore, conducting calibration based on data under non-photorespiratory conditions was indispensable for a reliable estimation of gm, regardless whether the multiphase flash or the single saturation pulse method was used for measuring the PSII operating efficiency. Other issues related to the procedure of using the chlorophyll fluorescence method to estimate gm were discussed.

Leaf photosynthesis of crops acclimates to elevated CO2 and temperature, but studies quantifying responses of leaf photosynthetic parameters to combined CO2 and temperature increases under field conditions are scarce. We measured leaf photosynthesis of rice cultivars Changyou 5 and Nanjing 9108 grown in two free-air CO2 enrichment (FACE) systems, respectively, installed in paddy fields. Each FACE system had four combinations of two levels of CO2 (ambient and enriched) and two levels of canopy temperature (no warming and warmed by 1.0-2.0°C). Parameters of the C3 photosynthesis model of Farquhar, von Caemmerer and Berry (the FvCB model), and of a stomatal conductance (gs ) model were estimated for the four conditions. Most photosynthetic parameters acclimated to elevated CO2 , elevated temperature, and their combination. The combination of elevated CO2 and temperature changed the functional relationships between biochemical parameters and leaf nitrogen content for Changyou 5. The gs model significantly underestimated gs under the combination of elevated CO2 and temperature by 19% for Changyou 5 and by 10% for Nanjing 9108 if no acclimation was assumed. However, our further analysis applying the coupled gs -FvCB model to an independent, previously published FACE experiment showed that including such an acclimation response of gs hardly improved prediction of leaf photosynthesis under the four combinations of CO2 and temperature. Therefore, the typical procedure that crop models using the FvCB and gs models are parameterized from plants grown under current ambient conditions may not result in critical errors in projecting productivity of paddy rice under future global change.

In the wake of declining water availability for agricultural use that will severely disturb the cultivation of irrigated lowland rice (Oryza sativa), aerobic genotypes have been developed allowing rice to grow under drier conditions. However, canopy physiological and morphological characteristics associated with adaptation of aerobic rice to water-limited conditions have so far been understudied. Canopy architectural characters, light and nitrogen distribution parameters (i.e. the extinction coefficient for light, KL; and for nitrogen, KN; and their ratio KN:KL), plant growth and yield were examined in irrigated-lowland (cv. IR64) vs aerobic (cv. Apo) rice genotypes grown under three water regimes. Compared with cv. IR64, cv. Apo allocated less nitrogen to strongly shaded leaves, allowing the vertical nitrogen distribution to better acclimate to light distribution within the canopy. The KN:KL ratio increased with increasing water deficit, and cv. Apo (up to 49 % increase) had a higher increase than cv. IR64 (up to 24 % increase). This was attributed to the maintenance of erect leaves at the upper part of the canopy, allocation of less leaf area and less nitrogen to the shaded leaves under water deficit in cv. Apo. The observed differential response in the KN:KL ratio partly explained the difference in biomass gain and yield in response to water deficit between the two rice cultivars. We concluded that favourable above-ground architecture and canopy nitrogen profile relative to the light profile contributed greatly to the better performance of cv. Apo than cv. IR64 under dry conditions.

Highlight We model the impact of improving photosynthesis on rice productivity. We consider all major photosynthesis-enhancing approaches under one umbrella and scale them from leaf photosynthesis to crop production., Various genetic engineering routes to enhance C3 leaf photosynthesis have been proposed to improve crop productivity. However, their potential contribution to crop productivity needs to be assessed under realistic field conditions. Using 31 year weather data, we ran the crop model GECROS for rice in tropical, subtropical, and temperate environments, to evaluate the following routes: (1) improving mesophyll conductance (gm); (2) improving Rubisco specificity (Sc/o); (3) improving both gm and Sc/o; (4) introducing C4 biochemistry; (5) introducing C4 Kranz anatomy that effectively minimizes CO2 leakage; (6) engineering the complete C4 mechanism; (7) engineering cyanobacterial bicarbonate transporters; (8) engineering a more elaborate cyanobacterial CO2-concentrating mechanism (CCM) with the carboxysome in the chloroplast; and (9) a mechanism that combines the low ATP cost of the cyanobacterial CCM and the high photosynthetic capacity per unit leaf nitrogen. All routes improved crop mass production, but benefits from Routes 1, 2, and 7 were ≤10%. Benefits were higher in the presence than in the absence of drought, and under the present climate than for the climate predicted for 2050. Simulated crop mass differences resulted not only from the increased canopy photosynthesis competence but also from changes in traits such as light interception and crop senescence. The route combinations gave larger effects than the sum of the effects of the single routes, but only Route 9 could bring an advantage of ≥50% under any environmental conditions. To supercharge crop productivity, exploring a combination of routes in improving the CCM, photosynthetic capacity, and quantum efficiency is required.

Harvesting hemp (Cannabis sativa L.) for both stems and seeds is now a common practice in Europe while crop management strategies for dual-purpose hemp cultivation have not been properly addressed so far. In the present study, the effects of planting density and nitrogen fertilization on hemp stem and seed yields were tested with the cultivars Futura 75 and/or Bialobrzeskie in eight contrasting environments (Italy in 2013; Italy and Latvia in 2014; Italy (two sites), Latvia, the Czech Republic, and France in 2015). Stem yield ranged between 1.3 and 22.3 Mg ha−1. The effects of planting density and nitrogen fertilization on stem yield did not interact significantly with each other, or with cultivar and harvest time. Increasing planting density from 30 to 120 plants m−2 and increasing nitrogen fertilization rate from 0 to 60 kg N ha−1 increased stem yield by 29% and 32%, respectively. Further increase in planting density and nitrogen fertilization did not result in a significant increase in stem yield. Seed yield ranged from 0.3 to 2.1 Mg ha−1. The seed yield was not affected significantly by planting density between 30 and 240 plants m−2. Although the seed yield showed an increasing trend with increasing nitrogen fertilization, the effects of nitrogen fertilization on seed yield were not statistically significant. To grow hemp as a dual-purpose crop it is recommended to plant 90–150 plants m−2 across all tested environments. Nitrogen fertilization rate at 60 kg N ha−1 was generally sufficient in the tested environments whereas further optimization of nitrogen fertilization requires accurate assessment of plant nitrogen status. To facilitate assessing plant nutritional status, a critical nitrogen dilution curve was determined for hemp and a practical method to determine nitrogen nutritional status was discussed.

Elucidating the genetic control of rooting behavior under water-deficit stress is essential to breed climate-robust rice (Oryza sativa) cultivars. Using a diverse panel of 274 indica genotypes grown under control and water-deficit conditions during vegetative growth, we phenotyped 35 traits, mostly related to root morphology and anatomy, involving 45,000 root-scanning images and nearly 25,000 cross sections from the root-shoot junction. The phenotypic plasticity of these traits was quantified as the relative change in trait value under water-deficit compared with control conditions. We then carried out a genome-wide association analysis on these traits and their plasticity, using 45,608 high-quality single-nucleotide polymorphisms. One hundred four significant loci were detected for these traits under control conditions, 106 were detected under water-deficit stress, and 76 were detected for trait plasticity. We predicted 296 (control), 284 (water-deficit stress), and 233 (plasticity) a priori candidate genes within linkage disequilibrium blocks for these loci. We identified key a priori candidate genes regulating root growth and development and relevant alleles that, upon validation, can help improve rice adaptation to water-deficit stress.

High day and night temperatures affect the rate and duration of grain filling and starch packaging differently in a developing rice grain., Rice grain yield and quality are predicted to be highly vulnerable to global warming. Five genotypes including heat-tolerant and susceptible checks, a heat-tolerant near-isogenic line and two hybrids were exposed to control (31 °C/23 °C, day/night), high night-time temperature (HNT; 31 °C/30 °C), high day-time temperature (HDT; 38 °C/23 °C) and high day- and night-time temperature (HNDT; 38 °C/30 °C) treatments for 20 consecutive days during the grain-filling stage. Grain-filling dynamics, starch metabolism enzymes, temporal starch accumulation patterns and the process of chalk formation were quantified. Compensation between the rate and duration of grain filling minimized the impact of HNT, but irreversible impacts on seed-set, grain filling and ultimately grain weight were recorded with HDT and HNDT. Scanning electron microscopy demonstrated irregular and smaller starch granule formation affecting amyloplast build-up with HDT and HNDT, while a quicker but normal amylopast build-up was recorded with HNT. Our findings revealed temporal variation in the starch metabolism enzymes in all three stress treatments. Changes in the enzymatic activity did not derail starch accumulation under HNT when assimilates were sufficiently available, while both sucrose supply and the conversion of sucrose into starch were affected by HDT and HNDT. The findings indicate differential mechanisms leading to high day and high night temperature stress-induced loss in yield and quality. Additional genetic improvement is needed to sustain rice productivity and quality under future climates.

For most biophysical domains, differences in model structures are seldom quantified. Here, we used a taxonomy-based approach to characterise thirteen rice models. Classification keys and binary attributes for each key were identified, and models were categorised into five clusters using a binary similarity measure and the unweighted pair-group method with arithmetic mean. Principal component analysis was performed on model outputs at four sites. Results indicated that (i) differences in structure often resulted in similar predictions and (ii) similar structures can lead to large differences in model outputs. User subjectivity during calibration may have hidden expected relationships between model structure and behaviour. This explanation, if confirmed, highlights the need for shared protocols to reduce the degrees of freedom during calibration, and to limit, in turn, the risk that user subjectivity influences model performance. A taxonomy-based approach was used to classify AgMIP rice simulation models.Different model structures often resulted in similar outputs.Similar structures often led to large differences in outputs.User subjectivity likely hides relationships between model structure and behaviour.Shared protocols are still needed to limit the risks during calibration.

Methods using gas exchange measurements to estimate respiration in the light (day respiration Rd ) make implicit assumptions about reassimilation of (photo)respired CO2 ; however, this reassimilation depends on the positions of mitochondria. We used a reaction-diffusion model without making these assumptions to analyse datasets on gas exchange, chlorophyll fluorescence and anatomy for tomato leaves. We investigated how Rd values obtained by the Kok and the Yin methods are affected by these assumptions and how those by the Laisk method are affected by the positions of mitochondria. The Kok method always underestimated Rd . Estimates of Rd by the Yin method and by the reaction-diffusion model agreed only for nonphotorespiratory conditions. Both the Yin and Kok methods ignore reassimilation of (photo)respired CO2 , and thus underestimated Rd for photorespiratory conditions, but this was less so in the Yin than in the Kok method. Estimates by the Laisk method were affected by assumed positions of mitochondria. It did not work if mitochondria were in the cytosol between the plasmamembrane and the chloroplast envelope. However, mitochondria were found to be most likely between the tonoplast and chloroplasts. Our reaction-diffusion model effectively estimates Rd , enlightens the dependence of Rd estimates on reassimilation and clarifies (dis)advantages of existing methods. ispartof: NEW PHYTOLOGIST vol:223 issue:2 pages:619-631 ispartof: location:England status: published

International audience; This study aimed to understand the physiological basis of rice photosynthetic response to C source-sink imbalances, focusing on the dynamics of the photosynthetic parameter triose phosphate utilization (TPU). Here, rice (Oriza sativa L.) indica cultivar IR64 were grown in controlled environment chambers under current ambient CO2 concentration until heading, and thereafter two CO2 treatments (400 and 800 mu mol mol(-1)) were compared in the presence and absence of a panicle-pruning treatment modifying the C sink. At 2 weeks after heading, photosynthetic parameters derived from CO2 response curves, and non-structural carbohydrate content of flag leaf and internodes were measured three to four times of day. Spikelet number per panicle and flag leaf area on the main culm were recorded. Net C assimilation and TPU decreased progressively after midday in panicle-pruned plants, especially under 800 mu mol mol(-1) CO2. This TPU reduction was explained by sucrose accumulation in the flag leaf resulting from the sink limitation. Taking together, our findings suggest that TPU is involved in the regulation of photosynthesis in rice under elevated CO2 conditions, and that sink limitation effects should be considered in crop models.

An analytical solution to the optimum partitioning of leaf nitrogen among photosynthetic proteins is developed, providing a new simple avenue of modelling leaf photosynthetic acclimation to environmental conditions during growth., The distribution of leaf nitrogen among photosynthetic proteins (i.e. chlorophyll, the electron transport system, Rubisco, and other soluble proteins) responds to environmental changes. We hypothesize that this response may underlie the biochemical aspect of leaf acclimation to the growth environment, and describe an analytical method to solve optimum nitrogen partitioning for maximized photosynthesis in C3 leaves. The method predicts a high investment of nitrogen in Rubisco under conditions leading to excessive energy supply relative to metabolic demand (e.g. low temperature, high light, low nitrogen, or low CO2). Conversely, more nitrogen is invested in chlorophyll when the energy supply is limiting. Overall, our optimization results are qualitatively consistent with literature reports. Commonly reported changes in photosynthetic parameters with growth temperature were emergent properties of the optimum nitrogen partitioning. The method was used to simulate dynamic acclimation under varying environmental conditions, using first-order kinetics. Simulated diurnal patterns of leaf photosynthetic rates as a result of acclimation differed greatly from those without acclimation (Awithout). However, differences in predicted photosynthesis integrated over a day or over the growing season from Awithout depended on the value of the kinetic time constant (τ), suggesting that τ is a critical parameter determining the overall impact of nitrogen distribution on acclimated photosynthesis.

Quantitative differences in tuber bulking of 100 genotypes in a segregating F1 population, their parents (SH, RH) and five contrasting cultivars of potato (Solanum tuberosum) grown in six environments were analysed using a piece-wise expolinear function. Tuber bulking was characterised by three parameters: cm, ED and wmax, where cm and ED were growth rate and effective duration, respectively, of the linear phase of tuber bulking, and wmax was the final tuber dry weight at the end of the linear phase (tE). We also analysed radiation (RUET) and nitrogen use efficiency (NUET), and their relationships with the model parameters. Values of cm and RUET were highest for early-maturing genotypes. Late-maturing genotypes had largest ED and NUET. As a result, wmax was higher in late genotypes than in early genotypes. Most traits exhibited high heritability and high genetic correlations with wmax. Path analysis showed that RUET, cm and a previously quantified parameter for total canopy cover Asum, had a major influence on wmax. Sixteen QTLs were detected for all traits explaining the phenotypic variance by up to 66%. One particular QTL on paternal linkage group V was detected for all traits with a major additive effect and maximum total phenotypic variance. Additional QTLs mostly associated with RH (cm, tE and ED) or both SH-RH linkage groups (NUET, wmax). Our study demonstrates that there are opportunities for improving tuber dry matter yield by selecting an optimal combination of important physiological traits.

Interest in hemp as a multi-purpose crop is growing worldwide and for the first time in 2015 it was cultivated in Europe on more than 20.000 ha as a dual-purpose crop, for the seeds and for the fibre. In the present study, fibre and seed productivity of 14 commercial cultivars were tested in four contrasting European environments (Latvia, the Czech Republic, France, Italy). At full flowering, the stem yield ranged from 3.7 Mg ha −1 to 22.7 Mg ha −1 , the bast fibre content ranged from 21% to 43%, and the bast fibre yield ranged from 1.3 Mg ha −1 to 7.4 Mg ha −1 . When harvesting was postponed from full flowering until seed maturity, the stem yield of monoecious cultivars significantly increased but in dioecious cultivars it decreased at all tested sites, except for Italy. Only the early cultivars Fedora 17 and Markant produced seed in the most northern location Latvia. The seed yield ranged from 0.3 Mg ha −1 to 2.4 Mg ha −1 in Italy, France and the Czech Republic. The cultivar effect on stem and seed yield was mainly determined by the genetic variation in time of flowering. Stem yield at full flowering was strictly related to the duration of the vegetative phase while seed yield was lowest in the late flowering cultivar. The late cultivar CS is suitable for stem and fibre production as it had the highest stem yield at full flowering in all locations. Both Fedora 17 and Futura 75 are candidate cultivars for dual-purpose production in Italy, France and the Czech Republic, with Fedora 17 being more suitable for seed production and Futura 75 for fibre production. The application of modelling to design production strategies for dual-purpose hemp is promising. However, accurate parameterisation is needed based on large data sets and diverse genetic background.

Elevated CO 2 and temperature strongly affect crop production, but understanding of the crop response to combined CO 2 and temperature increases under field conditions is still limited while data are scarce. We grew wheat (Triticum aestivum L.) and rice (Oryza sativa L.) under two levels of CO 2 (ambient and enriched up to 500 μmol mol -1 ) and two levels of canopy temperature (ambient and increased by 1.5-2.0 °C) in free-air CO 2 enrichment (FACE) systems and carried out a detailed growth and yield component analysis during two growing seasons for both crops. An increase in CO 2 resulted in higher grain yield, whereas an increase in temperature reduced grain yield, in both crops. An increase in CO 2 was unable to compensate for the negative impact of an increase in temperature on biomass and yield of wheat and rice. Yields of wheat and rice were decreased by 10-12% and 17-35%, respectively, under the combination of elevated CO 2 and temperature. The number of filled grains per unit area was the most important yield component accounting for the effects of elevated CO 2 and temperature in wheat and rice. Our data showed complex treatment effects on the interplay between preheading duration, nitrogen uptake, tillering, leaf area index, and radiation-use efficiency, and thus on yield components and yield. Nitrogen uptake before heading was crucial in minimizing yield loss due to climate change in both crops. For rice, however, a breeding strategy to increase grain number per m 2 and % filled grains (or to reduce spikelet sterility) at high temperature is also required to prevent yield reduction under conditions of global change.

High temperature has a pronounced effect on grain yield and quality in rice. Climate change has increased night temperature more than day temperature in many parts of the world. How rice responds to high night-time temperature (HNT) is largely unknown. This study presents the first effort to assess the response of tropical hybrid rice to HNT. Six commercial tropical hybrid rice cultivars together with a tolerant (N22-aus) and a susceptible (Gharib-indica) genotype were evaluated under control temperature (23 °C) and HNT (29 °C) starting from panicle initiation until maturity under field conditions at the International Rice Research Institute during the dry (DS) and wet (WS) seasons of 2013. Overall, HNT significantly decreased grain yield of Gharib and all tested hybrids across both seasons, with less average reduction in the DS (13.4%) than in the WS (18.6%). Among the yield components, spikelets m−2 most significantly contributed to yield variation under control and/or HNT during both DS and WS followed by grain weight, while the contribution of seed-set was low and season-specific. Grain quality in most hybrids was also strongly affected by HNT, with decreased head rice yield, increased chalkiness and reduced grain width. Given this vulnerability to HNT, there is an urgent need to explore options for improving the adaptation of rice hybrids to increasingly warmer nights.

Interest in hemp (Cannabis sativa L.) as a crop for the biobased economy is growing worldwide because hemp produces a high and valuable biomass while requiring low inputs. To understand the physiological basis of hemp's resource-use efficiency, canopy gas exchange was assessed using a chamber technique on canopies exposed to a range of nitrogen (N) and water levels. Since canopy transpiration and carbon assimilation were very sensitive to variations in microclimate among canopy chambers, observations were adjusted for microclimatic differences using a physiological canopy model, with leaf-level parameters estimated for hemp from our previous study. Canopy photosynthetic water-use efficiency (PWUEc), defined as the ratio of gross canopy photosynthesis to canopy transpiration, ranged from 4.0 mmol CO2 (mol H2O)-1 to 7.5 mmol CO2 (mol H2O)-1. Canopy photosynthetic nitrogen-use efficiency (PNUEc), the ratio of the gross canopy photosynthesis to canopy leaf-N content, ranged from 0.3 mol CO2 d-1 (g N)-1 to 0.7 mol CO2 d-1 (g N)-1. The effect of N-input levels on PWUEc and PNUEc was largely determined by the N effect on canopy size or leaf area index (LAI), whereas the effect of water-input levels differed between short- and long-term stresses. The effect of short-term water stress was reflected by stomatal regulation. The long-term stress increased leaf senescence, decreased LAI but retained total canopy N content; however, the increased average leaf-N could not compensate for the lost LAI, leading to a decreased PNUEc. Although hemp is known as a resource-use efficient crop, its final biomass yield and nitrogen use efficiency may be restricted by water limitation during growth. Our results also suggest that crop models should take stress-induced senescence into account in addition to stomatal effects if crops experience a prolonged water stress during growth.

Leaves are a nexus for the exchange of water, carbon, and energy between terrestrial plants and the atmosphere. Research in recent decades has highlighted the critical importance of the underlying biophysical and anatomical determinants of CO2 and H2O transport, but a quantitative understanding of how detailed 3D leaf anatomy mediates within-leaf transport has been hindered by the lack of a consensus framework for analyzing or simulating transport and its spatial and temporal dynamics realistically, and by the difficulty of measuring within-leaf transport at the appropriate scales. We discuss how recent technological advancements now make a spatially explicit 3D leaf analysis possible, through new imaging and modeling tools that will allow us to address long-standing questions related to plant carbon–water exchange.

Genetic analysis of grain yield and its components in a rice diversity panel identified genetic loci that regulate water deficit tolerance during the reproductive stage., A diversity panel comprising of 296 indica rice genotypes was phenotyped under non-stress and water-deficit stress conditions during the reproductive stage in the 2013 and 2014 dry seasons (DSs) at IRRI, Philippines. We investigated the genotypic variability for grain yield, yield components, and related traits, and conducted genome-wide association studies (GWAS) using high-density 45K single nucleotide polymorphisms. We detected 38 loci in 2013 and 64 loci in 2014 for non-stress conditions and 69 loci in 2013 and 55 loci in 2014 for water-deficit stress. Desynchronized flowering time confounded grain yield and its components under water-deficit stress in the 2013 experiment. Statistically corrected grain yield and yield component values using days to flowering helped to detect 31 additional genetic loci for grain yield, its components, and the harvest index in 2013. There were few overlaps in the detected loci between years and treatments, and when compared with previous studies using the same panel, indicating the complexity of yield formation under stress. Nevertheless, our analyses provided important insights into the potential links between grain yield with seed set and assimilate partitioning. Our findings demonstrate the complex genetic architecture of yield formation and we propose exploring the genetic basis of less complex component traits as an alternative route for further yield enhancement.

Extra ATP required in C4 photosynthesis for the CO2‐concentrating mechanism probably comes from cyclic electron transport (CET). As metabolic ATP : NADPH requirements in mesophyll (M) and bundle‐sheath (BS) cells differ among C4 subtypes, the subtypes may differ in the extent to which CET operates in these cells. We present an analytical model for cell‐type‐specific CET and linear electron transport. Modelled NADPH and ATP production were compared with requirements. For malic‐enzyme (ME) subtypes, c. 50% of electron flux is CET, occurring predominantly in BS cells for standard NADP‐ME species, but in a ratio of c. 6 : 4 in BS : M cells for NAD‐ME species. Some C4 acids follow a secondary decarboxylation route, which is obligatory, in the form of ‘aspartate‐malate’, for the NADP‐ME subtype, but facultative, in the form of phosphoenolpyruvate‐carboxykinase (PEP‐CK), for the NAD‐ME subtype. The percentage for secondary decarboxylation is c. 25% and that for 3‐phosphoglycerate reduction in BS cells is c. 40%; but these values vary with species. The ‘pure’ PEP‐CK type is unrealistic because its is impossible to fulfil ATP : NADPH requirements in BS cells. The standard PEP‐CK subtype requires negligible CET, and thus has the highest intrinsic quantum yields and deserves further studies in the context of improving canopy productivity.

A dynamic model of leaf CO2 assimilation was developed as an extension of the canonical steady-state model, by adding the effects of energy-dependent non-photochemical quenching (qE), chloroplast movement, photoinhibition, regulation of enzyme activity in the Calvin cycle, metabolite concentrations, and dynamic CO2 diffusion. The model was calibrated and tested successfully using published measurements of gas exchange and chlorophyll fluorescence on Arabidopsis thaliana ecotype Col-0 and several photosynthetic mutants and transformants affecting the regulation of Rubisco activity (rca-2 and rwt43), non-photochemical quenching (npq4-1 and npq1-2), and sucrose synthesis (spsa1). The potential improvements on CO2 assimilation under fluctuating irradiance that can be achieved by removing the kinetic limitations on the regulation of enzyme activities, electron transport, and stomatal conductance were calculated in silico for different scenarios. The model predicted that the rates of activation of enzymes in the Calvin cycle and stomatal opening were the most limiting (up to 17% improvement) and that effects varied with the frequency of fluctuations. On the other hand, relaxation of qE and chloroplast movement had a strong effect on average low-irradiance CO2 assimilation (up to 10% improvement). Strong synergies among processes were found, such that removing all kinetic limitations simultaneously resulted in improvements of up to 32%.

High-temperature during flowering in rice causes spikelet sterility and is a major threat to rice productivity in tropical and subtropical regions, where hybrid rice development is increasingly contributing to sustain food security. However, the sensitivity of hybrids to increasing temperature and physiological responses in terms of dynamic fertilization processes is unknown. To address these questions, several promising hybrids and inbreds were exposed to control temperature and high day-time temperature (HDT) in Experiment 1, and hybrids having contrasting heat tolerance were selected for Experiment 2 for further physiological investigation under HDT and high-night-time-temperature treatments. The day-time temperature played a dominant role in determining spikelet fertility compared with the night-time temperature. HDT significantly induced spikelet sterility in tested hybrids, and hybrids had higher heat susceptibility than the high-yielding inbred varieties. Poor pollen germination was strongly associated with sterility under high-temperature. Our novel observations capturing the series of dynamic fertilization processes demonstrated that pollen tubes not reaching the viable embryo sac was the major cause for spikelet sterility under heat exposure. Our findings highlight the urgent need to improve heat tolerance in hybrids and incorporating early-morning flowering as a promising trait for mitigating HDT stress impact at flowering.