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bioRxiv2018

A stress recovery signaling network for enhanced flooding tolerance in Arabidopsis thaliana

Abiotic stresses in plants are often transient and the recovery phase following stress removal is critical. Flooding, a major abiotic stress that negatively impacts plant biodiversity and agriculture, is a sequential stress where tolerance is strongly dependent on viability underwater and during the postflooding period. Here we show that in Arabidopsis thaliana accessions (Bay-0 and Lp2-6), different rates of submergence recovery correlate with submergence tolerance and fecundity. A genome-wide assessment of ribosome-associated transcripts in Bay-0 and Lp2-6 revealed a signaling network regulating recovery processes. Differential recovery between the accessions was related to the activity of three genes: RESPIRATORY BURST OXIDASE HOMOLOG (RBOHD), SENESCENCE-ASSOCIATED GENE113 (SAG113) and ORESARA1 (ORE1/NAC6) which function in a regulatory network involving a reactive oxygen species (ROS) burst upon de-submergence and the hormones abscisic acid and ethylene. This regulatory module controls ROS homeostasis, stomatal aperture and chlorophyll degradation during submergence recovery. This work uncovers a signaling network that regulates recovery processes following flooding to hasten the return to pre-stress homeostasis.\n\nSignificance statementFlooding due to extreme weather events can be highly detrimental to plant development and yield. Speedy recovery following stress removal is an important determinant of tolerance, yet mechanisms regulating this remain largely uncharacterized. We identified a regulatory network in Arabidopsis thaliana that controls water loss and senescence to influence recovery from prolonged submergence. Targeted control of the molecular mechanisms facilitating stress recovery identified here can potentially improve performance of crops in flood-prone areas.

plant biology↗

bioRxiv2018

The IMMUNE-ASSOCIATED NUCLEOTIDE-BINDING 9 protein is a regulator of basal immunity in Arabidopsis thaliana

A robust regulation of plant immune responses requires multitude of positive and negative regulators that act in concert. The immune-associated nucleotide-binding (IAN) gene family members are associated with immunity in different organisms, although no characterization of their function has been carried out to date in plants. In this work, we analyzed the expression patterns of IAN genes and found that IAN9 is repressed upon pathogen infection or treatment with immune elicitors. IAN9 encodes a plasma membrane-localized protein that genetically behaves as a negative regulator of immunity. A novel ian9 mutant generated by CRISPR/Cas9 shows increased resistance to Pseudomonas syringae, while transgenic plants overexpressing IAN9 show a slight increase in susceptibility. In vivo immunoprecipitation of IAN9-GFP followed by mass spectrometry analysis revealed that IAN9 associates with a previously uncharacterized C3HC4-type RING finger domain-containing protein that we named IAP1, for IAN9-associated protein 1, which also acts as a negative regulator of basal immunity. Interestingly, neither ian9 or iap1 mutant plants show any obvious developmental phenotype, suggesting that they display enhanced inducible immunity rather than constitutive immune responses. Since both IAN9 and IAP1 have orthologs in important crop species, they could be suitable targets to generate plants more resistant to diseases caused by bacterial pathogens without yield penalty.

plant biology↗

bioRxiv2018

Efficient CRISPR/Cas9-based genome editing and its application to conditional genetic analysis in Marchantia polymorpha

Marchantia polymorpha is one of the model species of basal land plants. Although CRISPR/Cas9-based genome editing has already been demonstrated for this plant, the efficiency was too low to apply to functional analysis. In this study, we show the establishment of CRISPR/Cas9 genome editing vectors with high efficiency for both construction and genome editing. Codon optimization of Cas9 to Arabidopsis achieved over 70% genome editing efficiency at two loci tested. Systematic assessment revealed that guide sequences of 17 nt or shorter dramatically decreased this efficiency. We also demonstrated that a combinatorial use of this system and a floxed complementation construct enabled conditional analysis of a nearly essential gene. This study reports that simple, rapid, and efficient genome editing is feasible with the series of developed vectors.

plant biology↗

bioRxiv2018

A combinatorial lipid code shapes the electrostatic landscape of plant endomembranes

Membrane surface charge is critical for the transient, yet specific recruitment of proteins with polybasic regions to certain organelles. In all eukaryotes, the plasma membrane (PM) is the most electronegative compartment of the cell, which specifies its identity. As such, membrane electrostatics is a central parameter in signaling, intracellular trafficking and polarity. Here, we explore which are the lipids that control membrane electrostatics using plants as a model. We show that phosphatidic acidic (PA), phosphatidylserine (PS) and phosphatidylinositol-4-phosphate (PI4P) are separately required to generate the electrostatic signature of the plant PM. In addition, we reveal the existence of an electrostatic territory that is organized as a gradient along the endocytic pathway and is controlled by PS/PI4P combination. Altogether, we propose that combinatorial lipid composition of the cytosolic leaflet of cellular organelles not only defines the plant electrostatic territory but also distinguishes different compartments within this territory by specifying their varying surface charges.

plant biology↗

bioRxiv2018

Loss of Peroxisomal Hydroxypyruvate Reductase Inhibits Triose Phosphate Isomerase but Stimulates Cyclic Photosynthetic Electron Flow and the Glc-6P-Phosphate Shunt

The oxygenation of ribulose 1,5-bisphosphate by Rubisco is the first step in photorespiration and reduces the efficiency of photosynthesis in C3 plants. Our recent data indicates that mutants in photorespiration have increased rates of photosynthetic cyclic electron flow around photosystem I. We investigated mutant lines lacking peroxisomal hydroxypyruvate reductase to determine if there are connections between 2-PG accumulation and cyclic electron flow. We found that 2-PG is a competitive inhibitor of triose phosphate isomerase (TPI), an enzyme in the Calvin-Benson cycle that converts glyceraldehyde 3-phosphate to dihydroxyacetone phosphate. This block in metabolism could be overcome if glyceraldehyde 3-phosphate is exported to the cytosol where the cytosolic triose phosphate isomerase could convert it to dihydroxyacetone phosphate. We found evidence that carbon is reimported as Glc-6P-phosphate forming a cytosolic bypass around the block of stromal TPI. However, this also stimulates a Glc-6P-phosphate shunt, which consumes ATP, which can be compensated by higher rates of cyclic electron flow.\n\nOnce Sentence SummaryTriose phosphate isomerase is inhibited in plants lacking hydroxypyruvate reductase 1 and this is overcome by exporting triose phosphate to the cytosol and importing Glc-6P, which stimulates a Glc-6P-phosphate shunt and cyclic electron flow.

plant biology↗

bioRxiv2018

Can Actin Depolymerization Actually Result In Increased Plant Resistance To Pathogens?

Introductory paragraphThe integrity of the actin cytoskeleton is essential for plant immune signalling1. Consequently, it is generally assumed that actin disruption reduces plant resistance to pathogen attack2-4. However, in a previous study, it was shown that actin depolymerisation triggers the salicylic acid (SA) signalling pathway5, which is interesting because increased SA is associated with enhanced plant resistance to pathogen attack6,7. Here, we attempt to resolve this seeming inconsistency by showing that the relationship between actin depolymerization and plant resistance is more complex than currently thought. We investigate the precise nature of this relationship using two completely different plant pathosystems: i) a model plant (Arabidopsis thaliana) and a bacterial pathogen (Pseudomonas syringae), and ii) an important crop (Brassica napus) and a fungal pathogen (Leptosphaeria maculans). We demonstrate that actin depolymerization induces a dramatic increase in SA levels and that the increased SA is biosynthesized by the isochorismate synthase pathway. In both pathosystems, this phenomenon leads to increased plant resistance.

plant biology↗

bioRxiv2018

High affinity Na+ transport by wheat HKT1;5 is blocked by K+

The wheat sodium transporters TmHKT1;5-A and TaHKT1;5-D are encoded by genes underlying major shoot Na+ exclusion loci Nax2 and Kna1 from Triticum monococcum (Tm) and Triticum aestivum (Ta), respectively. In contrast to HKT2 transporters that have been shown to exhibit high affinity K+-dependent Na+ transport, HKT1 proteins have, with one exception, only been shown to catalyse low affinity Na+ transport and no K+ transport. Here, using heterologous expression in Xenopus laevis oocytes we show that both TmHKT1;5-A and TaHKT1;5-D encode dual (high and low) affinity Na+-transporters with the high-affinity component being abolished when external K+ is in excess of external Na+. Based on 3-D structural modelling we propose that tighter binding of K+, compared to that of Na+ in the selectivity filter region by means of additional van der Waals forces, explains the K+ block at the molecular level. The low-affinity component for Na+ transport of TmHKT1;5-A had a lower Km than that of TaHKT1;5-D and was less sensitive to external K+. We propose that these properties underpin the improvements in shoot Na+-exclusion and crop plant salt tolerance following the introgression of TmHKT1;5-A into diverse wheat backgrounds.

plant biology↗

bioRxiv2018

Wheat shovelomics I: A field phenotyping approach for characterising the structure and function of root systems in tillering species

Wheat represents a major crop, yet the current rate of yield improvement is insufficient to meet its projected global food demand. Breeding root systems more efficient for water and nitrogen capture represents a promising avenue for accelerating yield gains. Root crown phenotyping, or shovelomics, relies on excavation of the upper portions of root systems in the field and measuring root properties such as numbers, angles, densities and lengths. We report a new shovelomics method that images the whole wheat root crown, then partitions it into the main shoot and tillers for more intensive phenotyping. Root crowns were phenotyped using the new method from the Rialto x Savannah population consisting of both parents and 94 doubled-haploid lines. For the whole root crown, the main shoot, and tillers, root phenes including nodal root number, growth angle, length, and diameter were measured. Substantial variation and heritability were observed for all phenes. Principal component analysis revealed latent constructs that imply pleiotropic genetic control of several related root phenes. Correlational analysis revealed that nodal root number and growth angle correlate among the whole crown, main shoot, and tillers, indicating shared genetic control among those organs. We conclude that this phenomics approach will be useful for breeding ideotype root systems in tillering species.

plant biology↗

bioRxiv2018

Wheat shovelomics II: Revealing relationships between root crown traits and crop growth

Optimization of root system architecture represents an important goal in wheat breeding. Adopting new field methods for root phenotyping is key to delivering this goal. A novel shovelomics method was applied for phenotyping root crown traits to characterize the Savannah x Rialto doubled-haploid (DH) population in two field experiments under irrigated and rain-fed conditions. Trait validation was carried out through soil coring on a subset of 14 DH lines and the two parents. We observed that drought reduced grain yield per plant by 21.0%. Under rain-fed conditions, nodal root angle and roots shoot-1 were positively associated with root length density (RLD) at 40-60 cm depth; RLD was also positively correlated with grain yield. Nodal root angle and roots shoot-1 were also positively associated with canopy stay green and grain yield under rain-fed conditions. We conclude that shovelomics is a valuable technique for quantifying genetic variation in nodal root traits in wheat, revealing nodal root angle and root number per shoot provide useful selection criteria in breeding programs aimed at improving drought tolerance in wheat.\n\nHighlightNodal root angle and number shoot-1 measured using shovelomics were positively associated with root density at depth and yield under drought in a Savanah x Rialto wheat DH population.

plant biology↗

bioRxiv2018

Topographic effects on dispersal patterns of Phytophthora cinnamomi at a stand scale in a Spanish heathland

Phytophthora cinnamomi is one of the most important plant pathogens in the world, causing root rot in more than a thousand plant species. This observational study was carried out on a P. cinnamomi infected heathland of Erica umbellata used as goat pasture. The patterns and shapes of disease foci and their distribution were described in a spatial and temporal context using an aerial photograph record. A set of topographic traits was selected on the basis of a disease dynamic hypothesis and their effects on observed spatial disease patterns were analyzed. Incipient infections situated in flat terrain expanded as compact circular front patterns with a low growth rate. On slopes, disease patches developed more rapidly down slope, forming parabolic shapes. The axis direction of the parabolas was highly correlated with terrain aspect, while the parabolic amplitude was associated with land curvature and slope. New secondary foci appeared over the years producing an accelerated increase of the affected surface. These new foci were observed in sites where disease density was higher or near sites more frequently visited by animals such as the stable or the forage crop. In contrast, a smaller number of disease foci occur in areas which animals are reluctant to visit, such as where they have a short range of vision. Our results suggest that 1) the growth of existing P. cinnamomi foci is controlled by a combination of root-to-root contact and water flows, 2) the increase in the diseased area arises mainly from the multiplication of patches, 3) the formation of new foci is mediated by long-distance transport due to the movement of animals and humans along certain preferential pathways, and 4) geomorphology and topography traits are associated with the epidemiology of this soil-borne pathogen.

plant biology↗

bioRxiv2018

Unraveling rice tolerance mechanisms against Schizotetranychus oryzae mite infestation

Infestation of Schizotetranychus oryzae (Acari: Tetranychidae) causes great losses in rice productivity. Infestation in Puita INTA-CL cultivar reduced the number of seeds/plant, percentage of full seeds, 1,000 seeds weight, and seed length, whereas infestation in IRGA423 increased 1,000 seeds weight and seed length. Reduction in seed weight/plant caused by infestation was higher in Puita INTA-CL than IRGA423. Thus, Puita INTA-CL was established as susceptible, and IRGA423 as tolerant to S. oryzae infestation. Photosynthetic parameters were less affected by infestation in IRGA423 than in Puita INTA-CL. Infestation also caused accumulation of H2O2, decreased cell membrane integrity and accelerated senescence in leaves of Puita INTA-CL, while leaves of IRGA423 presented higher levels of phenolics compounds. Using proteomic analysis, we identified proteins related to plant defense, such as jasmonate synthesis, and related to other mechanisms of tolerance such as oxidative stress, photosynthesis, and DNA structure maintenance, more abundant in IRGA423 after seven days of infestation. We detected higher levels of silicon (as amorphous silica cells) in leaves of infested IRGA423 plants compared to Puita INTA-CL, an element previously linked to plant defense. Our data shows that IRGA423 presents tolerance to S. oryzae infestation, and that multiple mechanisms might be employed by this cultivar.\n\nHighlightThis is the first report evaluating the defense responses (tolerance and susceptiblity) of two contrasting rice cultivars to Schizotetranychus oryzae mite infestation.

plant biology↗

bioRxiv2018

Evolution analysis and expression divergence of the chitinase gene family against Leptosphaeria maculans and Sclerotinia sclerotiorum infection in Brassica napus

AbstractBlackleg and sclerotinia stem rot caused by Leptosphaeria maculans and Sclerotinia sclerotiorum respectively are two major diseases in rapeseed worldwide, which cause serious yield losses. Chitinases are pathogenesis-related proteins and play important roles in host resistance to various pathogens and abiotic stress responses. However, a systematic investigation of the chitinase gene family and its expression profile against L. maculans and S. sclerotiorum infection in rapeseed remains elusive. The recent release of assembled genome sequence of rapeseed allowed us to perform a genome-wide identification of the chitinase gene family. In this study, 68 chitinase genes were identified in Brassica napus genome. These genes were divided into five different classes and distributed among 15 chromosomes. Evolutionary analysis indicated that the expansion of the chitinase gene family was mainly attributed to segmental and tandem duplication. Moreover, the expression profiling of the chitinase gene family was investigated using RNA sequencing (RNA-Seq) and the results revealed that some chitinase genes were both induced while the other members exhibit distinct expression in response to L. maculans and S. sclerotiorum infection. This study presents a comprehensive survey of the chitinase gene family in B. napus and provides valuable information for further understanding the functions of the chitinase gene family.

plant biology↗

bioRxiv2018

The complex architecture of plant transgene insertions

Over the last 35 years the soil bacterium Agrobacterium tumefaciens has been the workhorse tool for plant genome engineering. Replacement of native tumor-inducing (Ti) plasmid elements with customizable cassettes enabled insertion of a sequence of interest called Transfer DNA (T-DNA) into any plant genome. Although these T-DNA transfer mechanisms are well understood, detailed understanding of structure and epigenomic status of insertion events was limited by current technologies. To fill this gap, we analyzed transgenic Arabidopsis thaliana lines from three widely used collections (SALK, SAIL and WISC) with two single molecule technologies, optical genome mapping and nanopore sequencing. Optical maps for four randomly selected T-DNA lines revealed between one and seven insertions/rearrangements, and for the first time the actual length of individual transgene insertions from 27 to 236 kilobases. De novo nanopore sequencing-based genome assemblies for two segregating lines resolved T-DNA structures up to 36 kb into the insertions and revealed large-scale T-DNA associated translocations and exchange of chromosome arm ends. The multiple internally rearranged nature of T-DNA arrays made full assembly impossible, even with long nanopore reads. For the current TAIR10 reference genome, nanopore contigs corrected 83% of non-centromeric misassemblies. This unprecedented nucleotide-level definition of T-DNA insertions enabled the mapping of epigenome data. We identify variable small RNA transgene targeting and DNA methylation. SALK_059379 T-DNA insertions were enriched for 24nt siRNAs and contained dense cytosine DNA methylation. Transgene silencing via the RNA-directed DNA methylation pathway was confirmed by in planta assays. In contrast, SAIL_232 T-DNA insertions are predominantly targeted by 21/22nt siRNAs, with DNA methylation and silencing limited to a reporter, but not the resistance gene. With the emergence of genome editing technologies that rely on Agrobacterium for gene delivery, this study provides new insights into the structural impact of engineering plant genomes and demonstrates the utility of state-of-the-art long-range sequencing technologies to rapidly identify unanticipated genomic changes.

plant biology↗

bioRxiv2018

Clonal analysis reveals gradual recruitment of lateral root founder cells and a link between root initiation and cambium formation in Arabidopsis thaliana

O_LIThe pericycle gives rise to lateral roots (LRs) and lateral meristems (LMs; cambium and phellogen), however, a thorough clonal analysis of pericycle cell lineage has not been investigated. This study fills in this gap and addresses pericycle impact in LR and LM development.\nC_LIO_LIHeath-shock inducible DS1 transposition in 35S-DS1-H2B:YFP; HS-Ac seedlings results in production of YFP-labelled cell clones. These clones in pericycle cell derivatives were identified with a confocal microscopy and subjected to 3D reconstructions and analysis.\nC_LIO_LIParticipation of pericycle founder cells (FC) in LR formation is more variable than previously considered. LR initiation was found most commonly involved the specification of just one FC in the longitudinal and one or two cells in transverse direction. After LR initiation, FCs continue to be recruited in both directions from pre-existing cells. Anticlinal divisions in the pericycle resulting in LMs start already in the young differentiation zone where only the protoxylem is differentiated.\nC_LIO_LIThe clonal analysis demonstrated that pericycle cell activity related to LR formation is not separated in time and space from that related to LM formation and that LR FC recruitment is a gradual process. The analysis demonstrated that immediate pericycle progeny lack self-renewal capacity.\nC_LI

plant biology↗

bioRxiv2018

Acquisition and loss of secondary metabolite clusters shaped the evolutionary path of three recently emerged phytopathogens of wheat

O_LIWhite grain disorder is a recently emerged wheat disease in Australia, caused by three Botryosphaeriaceae spp.; Eutiarosporella darliae, E. pseudodarliae, and E. tritici-australis. The disease cycle of these pathogens and the molecular basis of their interaction with wheat are poorly understood. To address this, we undertook a comparative genomics approach to identify potential pathogenicity factors.\nC_LIO_LISubsequent genome analysis revealed that each of the white grain disorder species harbour modular polyketide synthase genes. To our knowledge, this is the first report of fungi harbouring such genes. Further comparative analysis using the modular polyketide synthase genes discovered their presence in the closely related Macrophomina phaseolina. Phylogenetic analysis implicates horizontal acquisition of these genes from a bacterial or a protist species.\nC_LIO_LIBoth E. darliae and E. pseudodarliae possess a secondary metabolite cluster with multiple polyketide/non-ribosomal peptide synthase genes (Hybrid-1, -2, and -3). In contrast, only remnant and partial genes homologous to this cluster were identified at a syntenic locus in E. tritici-australis suggesting loss of this cluster. Homologues of Hybrid-2 in other fungi have been proposed to facilitate disease induction in woody plants. Subsequent assays confirmed that E. darliae and E. pseudodarliae were both pathogenic on woody plant hosts, but E. tritici-australis was not, implicating woody plants as potential host reservoirs for the fungi. We hypothesise that loss of the cluster in E. tritici-australis represents a committed lifestyle jump to grasses.\nC_LIO_LICombined, our observations relating to the secondary metabolite potential of the WGD Eutiarosporella spp. have contributed novel data to the field by expanding the range of known fungal secondary metabolite genes, and helped develop our understanding of the lifestyle and potential host-range of a recently emerged pathogen of wheat.\nC_LI

plant biology↗

bioRxiv2018

NRG1 is required for the function of the TIR-NLR immune receptors Roq1 and RPP1 in Nicotiana benthamiana

The plant immune system involves a large family of nucleotide-binding leucine-rich repeat (NLR) intracellular immune receptors. These immune receptors often function to directly or indirectly mediate the perception of specific pathogen effector proteins secreted into the cell. Activation of these immune receptors typically results in activation of the immune system and subsequent suppression of pathogen proliferation. Although many examples of NLR receptors are known, a mechanistic understanding of how receptor activation ultimately leads to an immune response is not well understood. A subset of the NLR proteins contain a TIR domain at their N terminus (TNL). One such TNL, the N gene, was previously shown to depend on a non-TIR NLR protein, N requirement gene 1 (NRG1) for immune function. We tested additional NLR proteins in Nicotiana benthamiana for dependency on NRG1. We found that two additional TIR-NLR proteins, Roq1 and RPP1, also require NRG1 but that two coiled-coil NLR proteins, Bs2 and Rps2, do not. This finding suggests that NRG1 may be a conserved component of TNL signaling pathways.

plant biology↗

bioRxiv2018

Productivity definition of the chilling requirement reveals underestimation of the impact of climate change on winter chill accumulation

Evaluation of chilling requirements (CR) of cultivars of temperate fruit trees provides key information to assess regional suitability, according to winter chill, for both industry expansion and ongoing profitability as climate change continues. Traditional methods for calculating CR use climate controlled chambers and define CR using a fixed budburst percentage, usually close to 50% (CR-50%), without considering the productivity level associated to this percentage. This CR-50% definition may underestimate the real CR of tree crops for optimal productivity. This underestimation is particularly important to consider as winter chill accumulation is declining in many regions due to climate change. In this work we used sweet cherry to analyse the traditional method for calculating CR in many Rosaceae species (CR-50%) and compared the results with more a restrictive, productivity focused method, with CR defined with a 90% bud break level (90%, CRm-90%) close to the optimal budburst which assures productivity. Climate projections of winter chill suitability across Europe using CR-50% and CRm-90% were calculated. Regional suitability landscape was highly dependent on the method used to define CR and differences were found for a wide area of the European geography, both cold and mild winter areas. Our results suggest a need to use an optimal budburst level for the assessment of CR for sweet cherry. The use of traditional methods to determine CR can result in an underestimation of productivity CR with negative consequences for the fruit industry, particularly as climate change advances.

plant biology↗

bioRxiv2018

A genetic link between whole-plant water use efficiency and leaf carbon isotope composition in the C4 grass Setaria

Increasing whole plant water use efficiency (yield per transpiration; WUEplant) through plant breeding can benefit the sustainability of agriculture and improve crop yield under drought. To select for WUEplant, an efficient phenotyping method that reports on the genetic contribution of component traits such as transpiration efficiency (TEi; rate of CO2 assimilation per stomatal conductance) must be developed. Leaf carbon stable isotope composition ({delta}13Cleaf) has been proposed as a high-throughput proxy for TEi, and a negative correlation between {delta}13Cleaf and both WUEplant and TEi has previously been demonstrated in several C4 grass species. Therefore, the aim of the research presented here was to determine if the same loci control {delta}13Cleaf, WUEplant, and TEi under well-watered and water-limited conditions in a recombinant inbred line (RIL) population of closely related C4 grasses Setaria viridis and S. italica. Three quantitative trait loci (QTL) for {delta}13Cleaf were co-localized with transpiration, biomass, and a linear model of WUE. When WUEplant was calculated for allele classes based on the three QTL for {delta}13Cleaf, {delta}13Cleaf was negatively correlated with WUEplant as theory predicts when WUEplant is in part driven by differences in TEi. In any population, multiple traits can influence WUEplant; however, the analysis of {delta}13Cleaf in this RIL population demonstrates that there is genetic control of TEi that significantly contributes to WUEplant. Furthermore, this research suggests that {delta}13Cleaf can be used in marker-assisted breeding to select for TEi and as a tool to better understand the physiology and genetic architecture of TEi and WUEplant in C4 species.\n\nSignificance StatementOverextended water resources and drought are major agricultural problems worldwide. Therefore, selection for increased plant water use efficiency (WUEplant) in food and biofuel crop species is an important trait in plant breeding programs. Leaf carbon isotopic composition ({delta}13Cleaf) has potential as a rapid and effective high throughput phenotyping method for intrinsic transpiration efficiency (TEi), an important leaf-level component trait of WUEplant. Our research shows that {delta}13Cleaf and WUEplant share a common genetic architecture through their shared relationship with TEi. This suggests that {delta}13Cleaf can be used as a screen for TEi in marker-assisted plant breeding programs to improve crop drought resistance and decrease agricultural water consumption.

plant biology↗