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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.

Price surges in staple foods trigger civil unrest and conflict1. The food riots of 2007–2008 and Arab spring uprisings (2010–2012) were, in part, a consequence of price increases due to a tightening supply of staple grains, particularly wheat. Prolonged drought in Australia contributed to the global wheat shortage; Australia accounts for 10% of global wheat exports2. Australian wheat yields have plateaued3 owing to reduced rainfall4,5 and increasing temperatures3 attributed to anthropogenic climate change6. If Australia is to increase wheat production in line with projected global population growth and demand, an increase in yield is required7. Crop simulations reveal that an early sowing system combined with slower-developing wheat genotypes could exploit a longer growing season8. We developed near-isogenic lines and tested this hypothesis in experiments across the grain belt of Australia, and extended the results using whole-farm simulations. Our proposed early sowing system can increase national yields by 0.54 (s.d. = 0.38) t ha−1 representing an additional 7.1 Mt annually under reduced rainfall and increasing temperature regimes. This adaptation could facilitate increasing yields across Australia under climate change with global food security benefits. Crop models suggest that early sowing and slower-developing cultivars could maintain Australian wheat yields despite less-favourable climatic conditions. Field trials now confirm the potential of this adaptation for wheat production across Australia.

Australian wheat (Triticum aestivum) producers have been sowing crops earlier to adapt to reduced autumn rainfall, extreme spring weather and increasing farm size. Analysis of sowing date records indicate a shift of around 1.5 days/year over a 10 year period. The most suitable development patterns to maintain or increase yield at earlier sowing times have not been identified. Field experiments were conducted over two years at a range of sites and times of sowing (TOS), comparing a novel cultivar with fast-winter (FW) development to current elite spring and winter cultivars, and near-isogenic lines that differed only in major development genes. In cooler environments, the FW exhibited a more stable flowering time across a broader range of TOS compared to spring or slower developing winter cultivars. The optimal sowing window was shorter in warmer environments for the FW. Early-sown FW wheat yielded 8% more than fast-developing spring wheat sown later but flowering concurrently. FW wheat yielded 17% more than the elite mid-winter cultivar, and 18% more than elite slower developing spring cultivars when averaged across all TOS. The FW development pattern has potential to extend sowing periods while achieving 10–20% higher yields and flowering time stability. Wheat cultivars with altered development patterns must be developed to ensure crops flower during optimal periods from earlier sowing times.

The impact of different allelic combinations of VERNALIZATION1 (VRN1) and PHOTOPERIOD1 (PPD1) on the development of reproductive structures, spike dry weight (SDW) and its links to the duration of stem elongation (SE) and yield components was assessed using Near Isogenic Lines (NILs) of the cultivar Sunstate for both genes. Emphasis was on observations on the main shoot and tillers, linking to plant and canopy level in Steinfort at al. (2016). Experiments under controlled and irrigated field conditions relevant to the northern wheat growing region of Australia were designed. In vernalised plants, the number of fertile florets in the main shoot spike was positively related to the number of fertile spikelets, the duration of SE, the SDW at anthesis and the sum of the radiation intercepted during SE, while no associations were found under non vernalised and slow vernalising field conditions. Vernalised plants also produced more grains in the average tiller spike, compared to non vernalised plants, particularly under short days. Both main shoot and tiller grain number per spike contributed to the higher grain number per plant observed in vernalised plants in Steinfort et al. (2016), as spike number was similar. In the field, higher yield of lines with spring alleles of VRN1 was underpinned by longer spikes, with fewer spikelets but more fertile florets per spikelet in the main shoot spike, and more grains per tiller. When non vernalised, winter lines had an apex that grew slower, shorter spikes and higher spikelet density compared to the other genotypes. The utilisation of sensitivity to VRN1 or PPD1 to increase the duration of the SE and the number of fertile florets and grains is a more complex process than originally proposed.

Genetic variation in the VERNALIZATION1 (VRN1) and PHOTOPERIOD1 (PPD1) genes, which control the vernalisation and photoperiod response, underpin wheat adaptation to different environments. Near isogenic lines were used to investigate the role of allelic combinations of VRN1 and PPD-D1, including new alleles for VRN1-A1, on the length of developmental phases, dynamics of leaf and tiller appearance and yield components in complementary irrigated field trials relevant to low latitude wheat growing areas and controlled conditions. Allelic differences in VRN1 had a stronger effect on the duration of the vegetative phase, while photoperiod sensitivity at PPD-D1 lengthened the stem elongation phase (SE) by up to 23%. If a phase was lengthened, flowering was delayed. The level of response to daylength during stem elongation (SE) introduced by photoperiod sensitive alleles was dependent on the VRN1 composition and vernalisation status. A longer SE under short days was achieved by PPD1 sensitive genotypes when one VRN1 spring allele was present and plants were vernalised. The duration of SE was weakly related to spike dry weight m−2 at DC65 in the field but did not translate into higher grain number m−2. In the field, lines with two to three VRN1 spring alleles had shortest development phases, including SE, close flowering dates, sampled similar temperature environments at different stages, and achieved high yields. Yield advantage was explained by higher biomass, harvest index, grain number m−2 and thousand kernel weight. Genotypes with three winter VRN1 alleles were comparatively disadvantaged, with a longer vegetative phase placing SE under higher temperatures. Allelic differences in both genes caused large variation in leaf and tiller number generation but also in tiller mortality and individual leaf size, lessening the impact on leaf area. Changes in plant morphology and yield components that did not seem mediated via the influence of development genes on the duration of different stages and their impact on resource capture deserve further investigation.

EARLY FLOWERING3 (ELF3) is a circadian clock gene that contributes to photoperiod-dependent flowering in plants, with loss-of-function mutants in barley (Hordeum vulgare), legumes, and Arabidopsis thaliana flowering early under noninductive short-day (SD) photoperiods. The barley elf3 mutant displays increased expression of FLOWERING LOCUS T1 (FT1); however, it remains unclear whether this is the only factor responsible for the early flowering phenotype. We show that the early flowering and vegetative growth phenotypes of the barley elf3 mutant are strongly dependent on gibberellin (GA) biosynthesis. Expression of the central GA biosynthesis gene, GA20oxidase2, and production of the bioactive GA, GA1, were significantly increased in elf3 leaves under SDs, relative to the wild type. Inhibition of GA biosynthesis suppressed the early flowering of elf3 under SDs independently of FT1 and was associated with altered expression of floral identity genes at the developing apex. GA is also required for normal flowering of spring barley under inductive photoperiods, with chemical and genetic attenuation of the GA biosynthesis and signaling pathways suppressing inflorescence development under long-day conditions. These findings illustrate that GA is an important floral promoting signal in barley and that ELF3 suppresses flowering under noninductive photoperiods by blocking GA production and FT1 expression.

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/3.0/), which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited., Transcriptional activation of the VERNALIZATION1 gene mediates the acceleration of flowering by prolonged cold (vernalization) in temperate cereals. This study examined the earliest stages of the transcriptional response of VRN1 to low temperatures. Time-course analyses, using a sensitive quantitative PCR assay, showed that in sprouting barley seedlings VRN1 transcripts begin to accumulate within 24 hours of the onset of cold. The kinetics of the initial transcriptional response of VRN1 to cold was similar to the cold-induced genes DEHYDRIN5 (DHN5) and COLD REGULATED 14B (COR14B), but occurred at lower levels compared to cold acclimation genes or the response to longer cold treatments. Temperatures between 15 and –2 °C induced expression of VRN1 within 24 hours, with a maximal response observed between 2 and –2 °C. Transcriptional induction was also observed in undifferentiated callus cells. There were significant increases in histone acetylation levels at the VRN1 locus in response to 24-hour cold treatment. Sodium butyrate, a histone deacetylation inhibitor, triggered an increase in histone acetylation at VRN1 chromatin and elevated VRN1 transcript levels. The transcriptional response of VRN1 to short-term cold treatment was examined in near-isogenic lines that have different VRN1 genotypes, showing that an allele of the barley VRN1 gene with an insertion in the first intron and high basal expression levels has a reduced transcriptional response to short term cold treatment. This study suggests that low-temperature induction of VRN1 is a cellular response to cold triggered by the same mechanisms that mediate low-temperature induction of cold acclimation genes., This work was supported by an Office of the Chief Executive (OCE) Postdoctoral fellowship (S.N Oliver).

With 1 figure and 5 tables Abstract Two winter alleles of Vrn-A1 were recently found in wheat cultivars adapted to the Great Plains of the USA. Using a diagnostic marker for a single-nucleotide polymorphism (SNP) in exon 4, which distinguishes the alleles, the allele in the Great Plains cultivar ‘Jagger’ was common in Australian and CIMMYT cultivars, but ‘Veery’ cultivars carried the alternate allele from their Russian ancestor, which was the same as the allele in the Great Plains cultivar ‘Wichita’. The ‘Wichita’ allele was in North American winter cultivars, and chromosome substitution lines with a high level of tolerance to freezing, but not in substitution lines with a lower level of tolerance. The SNP between the alleles alters the predicted Vrn-A1 protein sequence, and this potentially explains differences in freezing tolerance. We suggest that these winter alleles could be coded as Vrn-A1v for the ‘Jagger’ allele and Vrn-A1w for the ‘Wichita’ allele. Cultivars with the spring Vrn-A1a or Vrn-A1b alleles carried the same SNP allele as ‘Jagger’, suggesting that the mutation from winter to spring for these alleles occurred in a Vrn-A1v genotype.

MADS-box transcription factors specify plant meristem identity. In doing so, they determine when floral organs are produced at the shoot apex and control the timing of flowering. The transcriptional activity of key MADS-box genes is controlled by temperature in many plants, and this synchronises flowering with changing seasons. Here we review how seasonal temperature variation influences the developmental programme of plants via transcriptional regulation of MADS-box genes. In particular we examine the role of MADS-box genes in regulating the acceleration of flowering by vernalization (prolonged periods of cold), using FLOWERING LOCUS C of Arabidopsis and VERNALIZATION1 of cereals as examples. A potential role for SHORT VEGETATIVE PHASE-like genes in controlling winter bud dormancy is also examined, as are potential roles for MADS-box genes in regulating developmental responses to elevated growth temperatures. We conclude that understanding how temperature regulates the transcription of MADS-box genes provides insight into how seasonal fluctuations in temperature influence plant development. Plant breeders may be able to use natural variation in temperature-responsive MADS-box genes to breed future crop varieties.

Responses to prolonged low-temperature treatment of imbibed seeds (vernalization) were examined in barley (Hordeum vulgare). These occurred in two phases: the perception of prolonged cold, which occurred gradually at low temperatures, and the acceleration of reproductive development, which occurred after vernalization. Expression of the VERNALIZATION1 gene (HvVRN1) increased gradually in germinating seedlings during vernalization, both at the shoot apex and in the developing leaves. This occurred in darkness, independently of VERNALIZATION2 (HvVRN2), consistent with the hypothesis that expression of HvVRN1 is induced by prolonged cold independently of daylength flowering-response pathways. After vernalization, expression of HvVRN1 was maintained in the shoot apex and leaves. This was associated with accelerated inflorescence initiation and with down-regulation of HvVRN2 in the leaves. The largest determinant of HvVRN1 expression levels in vernalized plants was the length of seed vernalization treatment. Daylength did not influence HvVRN1 expression levels in shoot apices and typically did not affect expression in leaves. In the leaves of plants that had experienced a saturating seed vernalization treatment, expression of HvVRN1 was higher in long days, however. HvFT1 was expressed in the leaves of these plants in long days, which might account for the elevated HvVRN1 expression. Long-day up-regulation of HvVRN1 was not required for inflorescence initiation, but might accelerate subsequent stages of inflorescence development. Similar responses to seed vernalization were also observed in wheat (Triticum aestivum). These data support the hypothesis that VRN1 is induced by cold during winter to promote spring flowering in vernalization-responsive cereals.

Genetic analyses have identified three genes that control the vernalization requirement in wheat and barley; VRN1, VRN2 and FT (VRN3). These genes have now been isolated and shown to regulate not only the vernalization response but also the promotion of flowering by long days. VRN1 is induced by vernalization and accelerates the transition to reproductive development at the shoot apex. FT is induced by long days and further accelerates reproductive apex development. VRN2, a floral repressor, integrates vernalization and day-length responses by repressing FT until plants are vernalized. A comparison of flowering time pathways in cereals and Arabidopsis shows that the vernalization response is controlled by different MADS box genes, but integration of vernalization and long-day responses occurs through similar mechanisms.

Timothy is a perennial forage grass grown commonly in Boreal regions. This study explored the effect of vernalization and photoperiod (PP) on flowering and growth characteristics and how this related to changes in expression of three flowering related genes in accessions from different geographic origin. Large variation was found in accessions in their vernalization and PP responses. In southern accessions vernalization response or requirement was not observed, the heading date remained unchanged, and plants flowered without vernalization. On the contrary, northern types had obligatory requirement for vernalization and long PP, but the tiller elongation did not require vernalization at 16-h PP. Longer vernalization or PP treatments reduced the genotypical differences in flowering. Moreover, the vernalization saturation progressed stepwise from main tiller to lateral tillers, and this process was more synchronized in southern accessions. The expression of PpVRN1 was associated with vernalization while PpVRN3 accumulated at long PP. A crucial role for PpVRN3 in the transition to flowering was supported as in southern accession the transcript accumulated in non-vernalized plants after transfer to 16-h PP, and the apices transformed to generative stage. Differences in vernalization requirements were associated with variation in expression levels of PpVRN1 and PpVRN3, with higher expression levels in southern type. Most divergent transcript accumulation of PpMADS10 was found under different vernalization conditions. These differences between accessions can be translated into agronomic traits, such as the tiller composition of canopy, which affects the forage yield. The southern types, with minimal vernalization response, have fast re-growth ability and rapidly decreasing nutritive value, whereas northern types grow slowly and have better quality. This information can be utilized in breeding for new cultivars for longer growing seasons at high latitudes.

Flowering time of wheat (Triticum aestivum L.) is a critical determinant of grain yield. Frost, drought and heat stresses from either overly early or overly late flowering can inflict significant yield penalties. The ability to predict time of flowering from different sowing dates for diverse cultivars across environments in Australia is important for maintaining yield as autumn rainfall events become less reliable. However, currently there are no models that can accurately do this when new cultivars are released. Two major Photoperiod1 and three Vernalisation1 development genes, with alleles identified by molecular markers, are known to be important in regulating phasic development and therefore time to anthesis, in response to the environmental factors of temperature and photoperiod. Allelic information from molecular markers has been used to parameterise models that could predict flowering time, but it is uncertain how much variation in flowering time can be explained by different alleles of the five major genes. This experiment used 13 elite commercial cultivars of wheat, selected for their variation in phenology and in turn allelic variation at the major development genes, and 13 near-isogenic lines (NILs) with matching multi-locus genotypes for the major development genes, to quantify how much response in time to flowering could be explained by alleles of the major genes. Genotypes were grown in four controlled environments at constant temperature of 22°C with factorial photoperiod (long or short day) and vernalisation (±) treatments applied. NILs were able to explain a large proportion of the variation of thermal time to flowering in elite cultivars in the long-day environment with no vernalisation (97%), a moderate amount in the short-day environment with no vernalisation (62%), and less in the short-day (51%) and long-day (47%) environments with vernalisation. Photoperiod was found to accelerate development, as observed in a reduction in phyllochron, thermal time to heading, thermal time to flowering, and decreased final leaf numbers. Vernalisation response was not as great, and rates of development in most genotypes were not significantly increased. The results indicate that the alleles of the five major development genes alone cannot explain enough variation in flowering time to be used to parameterise gene-based models that will be accurate in simulating flowering time under field conditions. Further understanding of the genetics of wheat development, particularly photoperiod response, is required before a model with genetically based parameter estimates can be deployed to assist growers to make sowing-time decisions for new cultivars.

Fr-B2 is a complex locus on chromosome 5B that affects frost tolerance, days to heading, grain yield and probably other traits of commercial importance in wheat (Triticum aestivum L.). It interacts epistatically with other major genes, especially VRN1. There are two known alleles of Fr-B2: an intact, wild-type allele, and an allele with a large deletion. Published methods for identifying these alleles are slow and expensive, making the development of a high-throughput, co-dominant SNP (single-nucleotide polymorphism) marker highly desirable, especially for commercial wheat breeding. A diverse panel of cultivars and breeding lines was characterised for SNPs and alleles of Fr-B2. Four SNP markers co-segregated as a haplotype block with Fr-B2 across unrelated cultivars and related backcrosses differing for alleles of Fr-B2. A robust KASP (Kompetitive allele-specific PCR) assay was developed for one of the SNPs, KASP_IWB26333, which should facilitate the inclusion of Fr-B2 on genotyping platforms for breeding and research.

Summary Functional genomics is transforming the way biological research is done in the 21st century. Functional genomics brings together high-throughput genetics with multiparallel analyses of gene transcripts, proteins, and metabolites to answer the ultimate question posed by all genome-sequencing projects: what is the biological function of each and every gene? Functional genomics is driving a shift in the research paradigm away from vertical analysis of single genes, proteins, or metabolites, towards horizontal analysis of full suites of genes, proteins, and metabolites. By identifying and measuring many, if not all of the molecular players that participate in a given biological process, functional genomics offers the prospect of obtaining a truly holistic picture of life. This review describes the tools that are currently being used for functional genomics work and considers the impact that this new discipline is likely to have in the future.

Summary Nitrogen-fixing symbioses between legumes and rhizobia are important for sustainable agriculture and contribute significantly to the global nitrogen cycle. The genomes of two rhizobial species, Mesorhizobium loti and Sinorhizobium meliloti have now been completely sequenced. Hundreds of thousands of expressed sequence tags representing tens of thousands of different genes from three major legume species; soybean, Medicago truncatula and Lotus japonicus have also been deposited in the public domain. Lotus japonicus recently became the focus of a genome project that aims to sequence one third of the entire genome over the next 5 yr. With this as a backdrop, the stage is set for a renaissance in symbiosis research, which will provide new insight into the complex molecular interplay that underpins symbiotic nitrogen fixation. This review considers how functional genomics might contribute to this renaissance.

Nitrogen-fixing symbioses between legumes and bacteria of the family Rhizobiaceae involve differentiation of both plant and bacterial cells. Differentiation of plant root cells is required to build an organ, the nodule, which can feed and accommodate a large population of bacteria under conditions conducive to nitrogen fixation. An efficient vascular system is built to connect the nodule to the root, which delivers sugars and other nutrients to the nodule and removes the products of nitrogen fixation for use in the rest of the plant. Cells in the outer cortex differentiate to form a barrier to oxygen diffusion into nodules, which helps to produce the micro-aerobic environment necessary for bacterial nitrogenase activity. Cells of the central, infected zone of nodules undergo multiple rounds of endoreduplication, which may be necessary for colonisation by rhizobia and may enable enlargement and greater metabolic activity of these cells. Infected cells of the nodule contain rhizobia within a unique plant membrane called the peribacteroid or symbiosome membrane, which separates the bacteria from the host cell cytoplasm and mediates nutrient and signal exchanges between the partners. Rhizobia also undergo differentiation during nodule development. Not surprisingly, perhaps, differentiation of each partner is dependent upon interactions with the other. High-throughput methods to assay gene transcripts, proteins, and metabolites are now being used to explore further the different aspects of plant and bacterial differentiation. In this review, we highlight recent advances in our understanding of plant cell differentiation during nodulation that have been made, at least in part, using high-throughput methods.

FROST RESISTANCE 2 (FR2) genes of wheat are C-Repeat Binding Factor (CBF) genes with two major alleles known for both Fr-A2 (‘T’ and ‘S’) and Fr-B2 (‘WT’ and ‘DEL’). VERNALIZATION 1 (VRN1) genes have a regulatory role on CBF genes, with known epistatic interactions between Vrn-A1 and Fr-A2 for tolerance to freezing temperatures during vegetative growth. VRN1 genes were also known to affect days to heading and grain yield. Therefore, FR2 genes might also affect these traits. A wide range of cultivars was characterised for VRN1, Fr-A2 and Fr-B2 genes. A third allele of Fr-A2 was found in cvv Excalibur and Axe. The winter cultivar Norstar, which was known to have a high level of frost tolerance during vegetative growth, had the combination Vrn-A1w + Fr-A2T + Fr-B2WT, as did a spring landrace from Afghanistan that was known to have superior tolerance to frost during reproductive development. No Australian spring cultivar was found with this combination, but it could be selected from crosses between adapted cultivars. This would enable the role of VRN1 and FR2 alleles in reproductive frost tolerance to be evaluated in an adapted background. Using large, existing, plant-breeding datasets, the T allele of Fr-A2 delayed heading relative to the S allele, and the WT allele of Fr-B2 delayed heading relative to the DEL allele, but only in combination with particular alleles of the VRN1 genes. Fr-B2 affected grain yield, with the highest grain yields for spring lines produced by Fr-B2DEL in combination with the spring allele of Vrn-B1.

Our paper describes how the genetic pathways which regulate vernalization and long-day flowering responses are integrated to promote spring flowering in cereals. This process is mediated by the VERNALIZATION1 (VRN1) and VRN2 genes. VRN2 encodes a CONSTANS-like protein that represses FLOWERING LOCUS T (FT1) to block the long-day flowering response until plants are vernalized. When plants are vernalized VRN1, a FRUITFUL-like MADS box transcription factor, is induced. This downregulates VRN2, allowing long-day induction of FT1 to occur post-vernalization. A comparison of the pathways regulating seasonal induction of flowering in cereals with those of Arabidopsis shows the vernalization response pathway has evolved convergently to regulate the activity of a conserved daylength response pathway in these divergent plant lineages.

Photoperiod and vernalisation genes are important for the adaptation of wheat to variable environments. Previously, using diagnostic markers and a large, unbalanced dataset from southern Australia, we estimated the effects on days to heading of frequent alleles of Vrn-A1, Vrn-B1, and Vrn-D1, and also two allelic classes of Ppd-D1. These genes accounted for ~45% of the genotypic variance for that trait. We now extend these analyses to further alleles of Ppd-D1, and four alleles of Ppd-B1 associated with copy number. Variation in copy number of Ppd-B1 occurred in our population, with one to four linked copies present. Additionally, in rare instances, the Ppd-B1 gene was absent (a null allele). The one-copy allele, which we labelled Ppd-B1b, and the three-copy allele, which we labelled Ppd-B1a, occurred through a century of wheat breeding, and are still frequent. With several distinct progenitors, the one-copy allele might not be homogenous. The two-copy allele, which we labelled Ppd-B1d, was generally introduced from WW15 (syn. Anza), and the four-copy allele, which we labelled Ppd-B1c, came from Chinese Spring. In paired comparisons, Ppd-B1a and Ppd-B1c reduced days to heading, but Ppd-B1d increased days to heading. Ppd-D1a, with a promoter deletion, Ppd-D1d, with a deletion in Exon 7, and Ppd-D1b, the intact allele, were frequent in modern Australian germplasm. Differences between Ppd-D1a and Ppd-D1d for days to heading under our field conditions depended on alleles of the vernalisation genes, confirming our previous report of large epistatic interactions between these classes of genes. The Ppd-D1b allele conferred a photoperiod response that might be useful for developing cultivars with closer to optimal heading dates from variable sowing dates. Inclusion of Ppd-B1 genotypes, and more precise resolution of Ppd-D1, increased the proportion of the genotypic variance attributed to these vernalisation and photoperiod genes to ~53%.