Phenology Genes Research Articles

Impact of Breeding on Free Amino Acids of Wholegrain Flour in Wheat and Role of Phenology Genes

Impact of Breeding on Free Amino Acids of Wholegrain Flour in Wheat and Role of Phenology Genes

Preadapted to adapt: underpinnings of adaptive plasticity revealed by the downy brome genome

Bromus tectorum L. is arguably the most successful invasive weed in the world. It has fundamentally altered arid ecosystems of the western United States, where it now found on an excess of 20 million hectares. Invasion success is related to avoidance of abiotic stress and human management. Early flowering is a heritable trait utilized by B. tectorum, enabling the species to temporally monopolize limited resources and outcompete the native plant community. Thus, understanding the genetic underpinning of flowering time is critical for the design of integrated management strategies. To study flowering time traits in B. tectorum, we assembled a chromosome scale reference genome for B. tectorum. To assess the utility of the assembled genome, 121 diverse B. tectorum accessions are phenotyped and subjected to a genome wide association study (GWAS). Candidate genes, representing homologs of genes that have been previously associated with plant height or flowering phenology traits in related species are located near QTLs we identified. This study uses a high-resolution GWAS to identify reproductive phenology genes in a weedy species and represents a considerable step forward in understanding the mechanisms underlying genetic plasticity in one of the most successful invasive weed species.

Vernalisation and photoperiod responses of diverse wheat genotypes

Context Wheat (Triticum aestivum L.) adaptation is highly dependent on crop lifecycle duration, particularly the time at which flowering occurs in a specific environment. Frost, low solar radiation, heat and drought can significantly reduce yield if a crop flowers too early or late. Wheat genotypes have different lifecycle durations determined by plant responses to temperature (thermal time accumulation and vernalisation) and photoperiod. These responses are largely controlled by five phenology genes (two PPD1 and three VRN1 genes). Advances in crop phenology modelling suggest that flowering time under field conditions could be accurately predicted with parameters derived from photoperiod and vernalisation responses obtained in controlled environments. Aims This study quantified photoperiod and vernalisation responses of 69 Australian wheat genotypes selected for diversity at the PPD1 and VRN1 loci. Methods Spring and winter genotypes were grown in four controlled environments at a constant temperature of 22°C with photoperiod (17 or 8 h) and vernalisation (0 or 8 weeks) treatments as factors. Key results Thermal time from coleoptile emergence to flowering in spring genotypes was typically decreased more by long photoperiod than by vernalisation; the opposite was true for winter genotypes. Spring genotypes that were sensitive to vernalisation contained a sensitive allele at the Vrn-A1 locus. Conclusions There is large diversity in phenological responses of wheat genotypes to photoperiod and vernalisation, including among those with matching multi-locus genotype. Implications Data from this study will be used to parameterise and test a wheat phenology model in a future study.

Allelic Variations in Phenology Genes of Eastern U.S. Soft Winter and Korean Winter Wheat and Their Associations with Heading Date.

Wheat heading time is genetically controlled by phenology genes including vernalization (Vrn), photoperiod (Ppd) and earliness per se (Eps) genes. Characterization of the existing genetic variation in the phenology genes of wheat would provide breeding programs with valuable genetic resources necessary for the development of wheat varieties well-adapted to the local environment and early-maturing traits suitable for double-cropping system. One hundred forty-nine eastern U.S. soft winter (ESW) and 32 Korean winter (KW) wheat genotypes were characterized using molecular markers for Vrn, Ppd, Eps and reduced-height (Rht) genes, and phenotyped for heading date (HD) in the eastern U.S. region. The Ppd-D1 and Rht-D1 genes exhibited the highest genetic diversity in ESW and KW wheat, respectively. The genetic variations for HD of ESW wheat were largely contributed by Ppd-B1, Ppd-D1 and Vrn-D3 genes. The Rht-D1 gene largely contributed to the genetic variation for HD of KW wheat. KW wheat headed on average 14 days earlier than ESW wheat in each crop year, largely due to the presence of the one-copy vrn-A1 allele in the former. The development of early-maturing ESW wheat varieties could be achieved by selecting for the one-copy vrn-A1 and vrn-D3a alleles in combination with Ppd-B1a and Ppd-D1a photoperiod insensitive alleles.

Advancing understanding of oat phenology for crop adaptation.

Oat (Avena sativa) is an annual cereal grown for forage, fodder and grain. Seasonal flowering behaviour, or phenology, is a key contributor to the success of oat as a crop. As a species, oat is a vernalization-responsive long-day plant that flowers after winter as days lengthen in spring. Variation in both vernalization and daylength requirements broadens adaptation of oat and has been used to breed modern cultivars with seasonal flowering behaviours suited to different regions, sowing dates and farming practices. This review examines the importance of variation in oat phenology for crop adaptation. Strategies to advance understanding of the genetic basis of oat phenology are then outlined. These include the potential to transfer knowledge from related temperate cereals, particularly wheat (Triticum aestivum) and barley (Hordeum vulgare), to provide insights into the potential molecular basis of variation in oat phenology. Approaches that use emerging genomic resources to directly investigate the molecular basis of oat phenology are also described, including application of high-resolution genome-wide diversity surveys to map genes linked to variation in flowering behaviour. The need to resolve the contribution of individual phenology genes to crop performance by developing oat genetic resources, such as near-isogenic lines, is emphasised. Finally, ways that deeper knowledge of oat phenology can be applied to breed improved varieties and to inform on-farm decision-making are outlined.

Identification of wild barley derived alleles associated with plant development in an Australian environment

The objective of this study was to explore wild barley as a potential source of alleles controlling plant development that can be used to improve the adaptation of barley to the Australian environment. The HEB-25 population was evaluated in field conditions at Strathalbyn (South Australia) in 2015 and 2016 and phenotyped for five traits related to plant development, in addition to plant height and ear number. GWAS identified QTL for all traits, of which many co-localised with known phenology genes. In the Australian environment, it was found that the Gibberellic acid (GA)-dependant pathway is important in regulating key development stages including shooting, flowering and maturing compared to the photoperiod and vernalisation pathway. Beneficial alleles from wild barley were identified, such as at the QTL tightly linked to the swd1 locus belonging to the GA-dependant pathway, wild alleles from five subfamilies increased EAR by up to 2.23 ears per m2. At the identified QTL, a wide range of effects was observed due to the wild barley alleles thus providing new genetic diversity that can be utilised to fine-tune the adaptation of barley to the Australian environment.

Hybridisation-based target enrichment of phenology genes to dissect the genetic basis of yield and adaptation in barley.

SummaryBarley (Hordeum vulgare L.) is a major cereal grain widely used for livestock feed, brewing malts and human food. Grain yield is the most important breeding target for genetic improvement and largely depends on optimal timing of flowering. Little is known about the allelic diversity of genes that underlie flowering time in domesticated barley, the genetic changes that have occurred during breeding, and their impact on yield and adaptation. Here, we report a comprehensive genomic assessment of a worldwide collection of 895 barley accessions based on the targeted resequencing of phenology genes. A versatile target‐capture method was used to detect genome‐wide polymorphisms in a panel of 174 flowering time‐related genes, chosen based on prior knowledge from barley, rice and Arabidopsis thaliana. Association studies identified novel polymorphisms that accounted for observed phenotypic variation in phenology and grain yield, and explained improvements in adaptation as a result of historical breeding of Australian barley cultivars. We found that 50% of genetic variants associated with grain yield, and 67% of the plant height variation was also associated with phenology. The precise identification of favourable alleles provides a genomic basis to improve barley yield traits and to enhance adaptation for specific production areas.

Developing Wheat for Improved Yield and Adaptation Under a Changing Climate: Optimization of a Few Key Genes

Abstract Wheat grown under rain-fed conditions is often affected by drought worldwide. Future projections from a climate simulation model predict that the combined effects of increasing temperature and changing rainfall patterns will aggravate this drought scenario and may significantly reduce wheat yields unless appropriate varieties are adopted. Wheat is adapted to a wide range of environments due to the diversity in its phenology genes. Wheat phenology offers the opportunity to fight against drought by modifying crop developmental phases according to water availability in target environments. This review summarizes recent advances in wheat phenology research, including vernalization (Vrn), photoperiod (Ppd), and also dwarfing (Rht) genes. The alleles, haplotypes, and copy number variation identified for Vrn and Ppd genes respond differently in different climatic conditions, and thus could alter not only the development phases but also the yield. Compared with the model plant Arabidopsis, more phenology genes have not yet been identified in wheat; quantifying their effects in target environments would benefit the breeding of wheat for improved drought tolerance. Hence, there is scope to maximize yields in water-limited environments by deploying appropriate phenology gene combinations along with Rht genes and other important physiological traits that are associated with drought resistance.

Quantitative trait loci for yield and grain plumpness relative to maturity in three populations of barley (Hordeum vulgare L.) grown in a low rain-fall environment.

Identifying yield and grain plumpness QTL that are independent of developmental variation or phenology is of paramount importance for developing widely adapted and stable varieties through the application of marker assisted selection. The current study was designed to dissect the genetic basis of yield performance and grain plumpness in southern Australia using three doubled haploid (DH) populations developed from crosses between adapted parents that are similar in maturity and overall plant development. Three interconnected genetic populations, Commander x Fleet (CF), Commander x WI4304 (CW), and Fleet x WI4304 (FW) developed from crossing of Australian elite barley genotypes, were used to map QTL controlling yield and grain plumpness. QTL for grain plumpness and yield were analysed using genetic linkage maps made of genotyping-by-sequencing markers and major phenology genes, and field trials at three drought prone environments for two growing seasons. Seventeen QTL were detected for grain plumpness. Eighteen yield QTL explaining from 1.2% to 25.0% of the phenotypic variation were found across populations and environments. Significant QTL x environment interaction was observed for all grain plumpness and yield QTL, except QPlum.FW-4H.1 and QYld.FW-2H.1. Unlike previous yield QTL studies in barley, none of the major developmental genes, including Ppd-H1, Vrn-H1, Vrn-H2 and Vrn-H3, that drive barley adaption significantly affected grain plumpness and yield here. Twenty-two QTL controlled yield or grain plumpness independently of known maturity QTL or genes. Adjustment for maturity effects through co-variance analysis had no major effect on these yield QTL indicating that they control yield per se.

High resolution mapping of traits related to whole-plant transpiration under increasing evaporative demand in wheat.

Atmospheric vapor pressure deficit (VPD) is a key component of drought and has a strong influence on yields. Whole-plant transpiration rate (TR) response to increasing VPD has been linked to drought tolerance in wheat, but because of its challenging phenotyping, its genetic basis remains unexplored. Further, the genetic control of other key traits linked to daytime TR such as leaf area, stomata densities and - more recently - nocturnal transpiration remains unknown. Considering the presence of wheat phenology genes that can interfere with drought tolerance, the aim of this investigation was to identify at an enhanced resolution the genetic basis of the above traits while investigating the effects of phenology genes Ppd-D1 and Ppd-B1 Virtually all traits were highly heritable (heritabilities from 0.61 to 0.91) and a total of mostly trait-specific 68 QTL were detected. Six QTL were identified for TR response to VPD, with one QTL (QSLP.ucl-5A) individually explaining 25.4% of the genetic variance. This QTL harbored several genes previously reported to be involved in ABA signaling, interaction with DREB2A and root hydraulics. Surprisingly, nocturnal TR and stomata densities on both leaf sides were characterized by highly specific and robust QTL. In addition, negative correlations were found between TR and leaf area suggesting trade-offs between these traits. Further, Ppd-D1 had strong but opposite effects on these traits, suggesting an involvement in this trade-off. Overall, these findings revealed novel genetic resources while suggesting a more direct role of phenology genes in enhancing wheat drought tolerance.

Genetic analysis of developmental and adaptive traits in three doubled haploid populations of barley (Hordeum vulgare L.).

Study of three interconnected populations identified 13 maturity QTL of which eight collocate with phenology genes, and 18 QTL for traits associated with adaptation to drought-prone environments. QTL for maturity and other adaptive traits affecting barley adaptation were mapped in a drought-prone environment. Three interconnected doubled haploid (DH) populations were developed from inter-crossing three Australian elite genotypes (Commander, Fleet and WI4304). High-density genetic maps were constructed using genotyping by sequencing and single nucleotide polymorphisms (SNP) for major phenology genes controlling photoperiod response and vernalization requirement. Field trials were conducted on the three DH populations in six environments at three sites in southern Australia and over two cropping seasons. Phenotypic evaluations were done for maturity, early vigour, normalized difference vegetation index (NDVI) and leaf chlorophyll content (SPAD), leaf waxiness and leaf rolling. Thirteen maturity QTL were identified, all with significant QTL×environment interaction with one exception. Eighteen QTL were detected for other adaptive traits across the three populations, including three QTL for leaf rolling, six for leaf waxiness, three for early vigour, four for NDVI, and two QTL for SPAD. The three interlinked populations with high-density linkage maps described in this study are a significant resource for examining the genetic basis for barley adaptation in low-to-medium rainfall Mediterranean type environments.

Bridging the gap between ideotype and genotype: Challenges and prospects for modelling as exemplified by the case of adapting wheat (Triticum aestivum L.) phenology to climate change in France

Simulations using crop models can assist designing ideotypes for current and future agricultural conditions. This approach consists in running simulations for different “in silico genotypes” obtained by varying the most sensitive genotypic parameters of these models, and analyzing results obtained for different environments, so as to identify the best genotypes for a target population of environments. However, this approach has rarely been used to guide commercial breeding programs so far. In this paper, we attempt to address some of the gaps yet to be filled before this kind of approach can be implemented, and identify some remaining issues that should be addressed in future research. Our focus is on optimizing wheat phenology, integrating simulations from a modified version of the ARCWHEAT model of wheat growth stages with available knowledge on the genetic control of wheat phenology obtained via molecular markers. Based on simulations, stem extension could be advanced by 10 days in 2025–2049 without increasing frost risks, thus opening up opportunities for lengthening the rapid growth period. Analysis of the current genetic variability for major phenology genes in French elite varieties, showed that the insensitive PpdD1—spring Vrn3 allele combination appears undesirable and current genotypes with early stem extensions are unstable (i.e. show a strong response to temperature and can start the stem extension very early in case of mild winter temperatures). We finally use a case study on gene-based modelling of wheat phenology in France to illustrate how it can be used to dissect the genetic basis of the quantitative nature of the three components of earliness, beyond the effects of major genes. We identify the need to link the variability for optimized model parameters and the allelic variations at the gene level as a critical step of this type of approach.

Genetic control of grain protein, dough rheology traits and loaf traits in a bread wheat population grown in three environments

The processing and baking quality of bread wheat can be affected by both genetic and environmental factors and by interactions between them. Here we report on the genetic control of grain protein content, flour water absorption, dough and bread making quality in wheat grown in three experiments under differing environmental conditions. The research used a recombinant inbred line mapping population derived from a cross between two Australian cultivars, Drysdale and Gladius. Three field experiments were conducted, including one that was sown late in order to expose the lines to high temperatures during grain filling and one in a year in which conditions were unusually cool and wet. Genomic regions containing photoperiod sensitivity loci affected grain protein content while the Ha (puroindoline) locus on chromosome 5D was associated with loaf quality traits. Other QTLs (on chromosomes 2B, 3B and 5A) were novel and not associated with any known quality or phenology genes.

From Ideotypes to Genotypes: Approaches to Adapt Wheat Phenology to Climate Change

IntroductionSimulations using crop models can assist in designing ideotypes for current and future agricultural conditions. This approach has been often in recent years to identify avenues for adapting wheat to climate change. However, this approach has rarely been used to guide commercial breeding programs. We hypothesize that the lack of link between models and the available tools for breeding, i.e. available genetic variability and selection methods. Materials and methods- We use a modified ARCWHEAT2 phenology model and future climate data from the ARPEGE global circulation model to identify targets for future phenologies-We genotyped over 400 French cultivars for known phenology genes and confronted the genetic make-up of these varieties to their success in France over the past 25 years- We developed a methodology to link model parameters to underlying marker data. We tested the performance of the methodology against circa 60 varieties ResultsEarlier phenology may be an avenue for stress avoidance in the future.Current photoperiod sensitivity of early cultivars already poses problems in terms of adaptation, as exemplified by the interaction between Ppd-D1 and Vrn-A3We show that a gene-based model can be used to predict wheat phenology without a significant loss in predictive performance. DiscussionAnalyzing current phenology genes of existing cultivars and their adaptation allowed us to identify a limit to past breeding efforts in obtaining early cultivars. This requires that a more knowledge based approach be taken. Gene-based modelling of phenology is possible on a collection of elite, adapted varieties and provides the tools for constructing genotypes with specific allelic combinations leading to more appropriate constructions of earliness.

Evidence for selection on a CONSTANS-like gene between two red oak species.

Hybridizing species such as oaks may provide a model to study the role of selection in speciation with gene flow. Discrete species' identities and different adaptations are maintained among closely related oak species despite recurrent gene flow. This is probably due to ecologically mediated selection at a few key genes or genomic regions. Neutrality tests can be applied to identify so-called outlier loci, which demonstrate locus-specific signatures of divergent selection and are candidate genes for further study. Thirty-six genic microsatellite markers, some with putative functions in flowering time and drought tolerance, and eight non-genic microsatellite markers were screened in two population pairs (n = 160) of the interfertile species Quercus rubra and Q. ellipsoidalis, which are characterized by contrasting adaptations to drought. Putative outliers were then tested in additional population pairs from two different geographic regions (n = 159) to support further their potential role in adaptive divergence. A marker located in the coding sequence of a putative CONSTANS-like (COL) gene was repeatedly identified as under strong divergent selection across all three geographically disjunct population pairs. COL genes are involved in the photoperiodic control of growth and development and are implicated in the regulation of flowering time. The location of the polymorphism in the Quercus COL gene and given the potential role of COL genes in adaptive divergence and reproductive isolation makes this a promising candidate speciation gene. Further investigation of the phenological characteristics of both species and flowering time pathway genes is suggested in order to elucidate the importance of phenology genes for the maintenance of species integrity. Next-generation sequencing in multiple population pairs in combination with high-density genetic linkage maps could reveal the genome-wide distribution of outlier genes and their potential role in reproductive isolation between these species.

Ppd1, Vrn1, ALMT1 and Rht genes and their effects on grain yield in lower rainfall environments in southern Australia

Allele-specific markers for important genes can improve the efficiency of plant breeding. Their value can be enhanced if effects of the alleles for important traits can be estimated in identifiable types of environment. Provided potential bias can be minimised, large, unbalanced, datasets from previous plant-breeding and agronomic research can be used. Reliable, allele-specific markers are now available for the phenology genes Ppd-D1, Vrn-A1, Vrn-B1 and Vrn-D1, the aluminium-tolerance gene TaALMT1, and the plant-stature genes Rht-B1 and Rht-D1. We used a set of 208 experiments with growing-season rainfall of <347 mm from southern Australia to estimate the effects of seven frequent combinations of the phenology genes, an intolerant and a tolerant allele of TaALMT1, and two semi-dwarf combinations Rht-B1b + Rht-D1a (Rht-ba) and Rht-B1a + Rht-D1b (Rht-ab) on grain yield in lower rainfall, Mediterranean-type environments in southern Australia. There were 775 lines in our analyses and a relationship matrix was used to minimise bias. Differences among the phenology genes were small, but the spring allele Vrn-B1a might be desirable. The tolerant allele, TaALMT1-V, was advantageous in locations with alkaline soils, possibly because of toxic levels of aluminium ions in subsoils. The advantage of TaALMT1-V is likely to be highest when mean maximum temperatures in spring are high. Rht-ab (Rht2 semi-dwarf) was also advantageous in environments with high mean maximum temperatures in spring, suggesting that for these stress environments, the combination of Vrn-B1a plus TaALMT1-V plus Rht-ab should be desirable. Many successful cultivars carry this combination.