Photoperiod-insensitive Allele Research Articles

GIGANTEA accelerates wheat heading time through gene interactions converging on FLOWERING LOCUS T1.

Precise regulation of flowering time is critical for cereal crops to synchronize reproductive development with optimum environmental conditions, thereby maximizing grain yield. The plant-specific gene GIGANTEA (GI) plays an important role in the control of flowering time, with additional functions on the circadian clock and plant stress responses. In this study, we show that GI loss-of-function mutants in a photoperiod-sensitive tetraploid wheat background exhibit significant delays in heading time under both long-day (LD) and short-day photoperiods, with stronger effects under LD. However, this interaction between GI and photoperiod is no longer observed in isogenic lines carrying either a photoperiod-insensitive allele in the PHOTOPERIOD1 (PPD1) gene or a loss-of-function allele in EARLY FLOWERING 3 (ELF3), a known repressor of PPD1. These results suggest that the normal circadian regulation of PPD1 is required for the differential effect of GI on heading time in different photoperiods. Using crosses between mutant or transgenic plants of GI and those of critical genes in the flowering regulation pathway, we show that GI accelerates wheat heading time by promoting FLOWERING LOCUS T1 (FT1) expression via interactions with ELF3, VERNALIZATION 2 (VRN2), CONSTANS (CO), and the age-dependent microRNA172-APETALA2 (AP2) pathway, at both transcriptional and protein levels. Our study reveals conserved GI mechanisms between wheat and Arabidopsis but also identifies specific interactions of GI with the distinctive photoperiod and vernalization pathways of the temperate grasses. These results provide valuable knowledge for modulating wheat heading time and engineering new varieties better adapted to a changing environment.

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.

Distribution of Photoperiod-Insensitive Alleles Ppd-D1a, Ppd-B1a, and Ppd-B1c in Winter Common Wheat Cultivars (Triticum aestivum L.) of Various Origin

Distribution of Photoperiod-Insensitive Alleles Ppd-D1a, Ppd-B1a, and Ppd-B1c in Winter Common Wheat Cultivars (Triticum aestivum L.) of Various Origin

Understanding the Effects of Growing Seasons, Genotypes, and Their Interactions on the Anthesis Date of Wheat Sown in North China.

Simple SummaryWheat anthesis date is an important turning point for wheat from vegetative growth stage to reproductive growth stage, which is crucial for wheat to adapt to environment and increase grain yield. In this study, a panel of adaptive wheat varieties including historical varieties from the 1940s and current varieties was used to understand the contribution of growing season, genotypes and their interaction effects to anthesis date. Based on our results, we can conclude that growing seasons contributed tremendously to the anthesis date of wheat. In future wheat breeding, more consideration should be given to growing seasons, and the gene combination with the strongest adaptability to the growing seasons should be selected.Quantitative studies on the effects of growing season, genotype (including photoperiod genes and vernalization genes), and their interaction (GGI) on the anthesis date of winter wheat (Triticum aestivum L.) are helpful to provide a scientific reference for selecting or developing adaptive varieties in target environments. In this study, we collected 100 winter wheat varieties with ecological adaptability in North China and identified the anthesis date under field conditions for three consecutive years from 2016 to 2019 with mapped photoperiod and vernalization alleles. Our results showed that the number of the photoperiod-insensitive Ppd-D1a allele increased with variety replacement, while the haplotype Ppd-A1b + Ppd-D1b + vrn-D1 (A4B2) decreased from the 1940s to 2000s. The anthesis date of A4B2 was significantly delayed due to the photoperiod-insensitive alleles Ppd-A1b and Ppd-D1b. The additive main effect and multiplicative interaction (AMMI) model and GGI biplot analysis were used for data analysis. A large portion of the total variation was explained by growing seasons (66.3%), while genotypes and GGIs explained 21.9% and 10.1% of the anthesis dates, respectively. The varieties from the 1940s and before had a great influence on the anthesis date, suggesting these germplasms tend to avoid premature anthesis and could facilitate the development of phenological resilient varieties.

Interactive Effects of Vernalization and Photoperiod Loci on Phenological Traits and Grain Yield and Differentiation of Iranian Wheat Landraces and Cultivars

Phenological stages and allelic variation of vernalization and photoperiod response genes play an important role in wheat environmental adaptation and grain yield. In the present study, phenological stages of 260 Iranian wheat landraces and cultivars were studied during two cropping years. In addition, allelic variation of vernalization and photoperiod markers were also investigated to identify the genetic basis of different phenological stages. Ppd-D1 and then Ppd-B1 were highly variable in cultivars and landraces, while Ppd-A1 showed lower variation. Distribution patterns of allele frequency in Vrn-B1b, Ppd-D1b-D001, and Ppd-D1 markers were remarkably different in cultivars and landraces. About 98% and 96% of landraces showed photoperiod-sensitive allele of Ppd-D1b-D001 and Ppd-D1 markers, respectively, while most of cultivars had photoperiod-insensitive alleles. The frequency of Vrn-B1a was 82.2% in cultivars and 100% in landraces. However, Vrn-A1a-E4 was 43% and 79% in cultivar and landraces, respectively. We have found that Ppd-D1, Ppd-B1, and Vrn-A1 have a decisive effect on phenological stages and the combination of Ppd-B1a, Ppd-D1b, and Vrn-A1b is the most abundant allelic compound. The combination of aaa, bbb, and aab alleles for these three loci led to increased yield, thousand kernel weight (TKW), and grain filling period and shortening of phenological stages. Cluster analysis based on phenological stages and growing degree-days (GDD) of phenological stages clearly separated cultivars from landraces. This clustering pattern was consistent with marker data. The findings of this study provide a comprehensive insight into the basis of genetic control of the phenological stages of Iranian wheat landraces and cultivars. Therefore, this information can be used to select desirable genotypes in future breeding programs.

Stepwise increases in FT1 expression regulate seasonal progression of flowering in wheat (Triticum aestivum).

Flowering is regulated by genes that respond to changing daylengths and temperature, which have been well studied using controlled conditions; however, the molecular processes underpinning flowering in nature remain poorly understood. Here, we investigate the genetic pathways that coordinate flowering and inflorescence development of wheat (Triticum aestivum) as daylengths extend naturally in the field, using lines that contain variant alleles for the key photoperiod gene, Photoperiod-1 (Ppd-1). We found flowering involves a stepwise increase in the expression of FLOWERING LOCUS T1 (FT1), which initiates under day-neutral conditions of early spring. The incremental rise in FT1 expression is overridden in plants that contain a photoperiod-insensitive allele of Ppd-1, which hastens the completion of spikelet development and accelerates flowering time. The accelerated inflorescence development of photoperiod-insensitive lines is promoted by advanced seasonal expression of floral meristem identity genes. The completion of spikelet formation is promoted by FLOWERING LOCUS T2, which regulates spikelet number and is activated by Ppd-1. In wheat, flowering under natural photoperiods is regulated by stepwise increases in the expression of FT1, which responds dynamically to extending daylengths to promote early inflorescence development. This research provides a strong foundation to improve yield potential by fine-tuning the photoperiod-dependent control of inflorescence development.

Phenology and Dwarfing Gene Interaction Effects on the Adaptation of Selected Wheat (Triticum aestivum L.) Advanced Lines across Diverse Water-Limited Environments of Western Australia

Photoperiod, vernalization, and plant height controlling genes are major developmental genes in wheat that govern environmental adaptation and hence, knowledge on the interaction effects among different alleles of these genes is crucial in breeding cultivars for target environments. The interaction effects among these genes were studied in nineteen Australian advanced lines from diverse germplasm pools and four commercial checks. Diagnostic markers for the Vrn-A1 locus revealed the presence of the spring allele Vrn-A1a in 10 lines and Vrn-A1c in one line. The dominant alleles of Vrn-B1a and Vrn-D1a were identified in 19 and 8 lines, respectively. The most common photoperiod-insensitive allele of Ppd-D1a was identified in 19 lines and three and four copy photoperiod-insensitive alleles (Ppd-B1a and Ppd-B1c) were present in five and one lines, respectively. All the lines were photoperiod-sensitive for the Ppd-A1 locus. All lines were semi-dwarf, having either of the two dwarfing alleles; 14 lines had the Rht-B1b (Rht-1) and the remaining had the Rht-D1b (Rht-2) dwarfing allele. The presence of the photoperiod-insensitive allele Ppd-D1a along with one or two spring alleles at the Vrn1 loci resulted in an earlier heading and better yield. Dwarfing genes were found to modify the heading time—the Rht-D1b allele advanced heading by three days and also showed superior effects on yield-contributing traits, indicating its beneficial role in yield under rain-fed conditions along with an appropriate combination of photoperiod and vernalization alleles. This study also identified the adaptability value of these allelic combinations for higher grain yield and protein content across the different the water-limited environments.

Agronomic performance of durum wheat landraces and modern cultivars and its association with genotypic variation in vernalization response (Vrn-1) and photoperiod sensitivity (Ppd-1) genes

This study analyzed the relationship between important agronomic traits and major genes regulating flowering time in a panel of 151 Mediterranean durum wheat landraces and 20 modern cultivars. Field experiments were conducted under rainfed conditions during six crop seasons in northeastern Spain. Multivariate analysis of agronomic traits and genotypic data allowed the modern cultivars to be differentiated from the landraces and germplasm pools to be identified within the landraces associated with their geographic origin. The high frequency of the Vrn-A1c allele and the photoperiod insensitive alleles GS105 and GS100 at Ppd-A1 reduced time to anthesis and enlarged the grain filling period of the modern cultivars compared with the landraces. Ancient durums collected close to the domestication area of wheat showed a high frequency of the winter allele vrn-B1 and the photoperiod sensitivity allele Ppd-B1b. None of the allele variants or allelic combinations accounted significantly for variations in any agronomic trait of modern cultivars. Vernalization and photoperiod genes acted additively in explaining the genotypic variance for the agronomic traits of the landraces. Vrn-A1 alleles and Vrn-A1+Vrn-B1 allelic combinations significantly affected the number of grains per spike (NGS), thousand kernel weight (TKW) and grain filling rate (R), accounting for 9%?12% of the genotypic variance for these traits. Ppd-1 accounted for 6%?21% of the genotypic variance for R, grain filling duration (GFD), plant height (PH), biomass at anthesis (CDW) and harvest index (HI). Vrn-1+Ppd-1 allelic combinations accounted for 21%?26% of the genotypic variance for these traits. Except for NGS, the effect of vernalization and photoperiod genes on the agronomic traits was linked to their effect on anthesis time. The three-day delay in anthesis time caused by the allele Vrn-A1d irrespective of the allele Vrn-A1c resulted in increases of 10 % in R and 7% in TKW. The eight-day delay in anthesis time caused by the allele Ppd-A1(DelCD) compared with Ppd-A1(GS105) increased R by 19 % and PH by 28 %, but reduced GFD and HI by 10 %. None of the allele variants or allelic combinations at the Vrn-1 or Ppd-1 genes accounted significantly for variations in yield or number of spikes m?2 (NSm2).

Molecular Processes Underpinning Seasonal Regulation of Flowering in Bread Wheat ( <i>Triticum Aestivum L.</i>)

Plants integrate seasonal progression in daylength and temperature to determine the optimal time to flower and produce seed. In cereals, genes that regulate responses to changing daylengths and temperature have been well-studied using controlled conditions; however, the molecular processes underpinning flowering in nature are poorly understood. Here, we investigate the molecular pathways that coordinate flowering and inflorescence development in wheat as daylengths extend naturally in the field. We find that discrete photoperiods contribute to the initiation of flowering and control of yield-related traits as days becomes longer from winter into spring. This process involves step-wise increases in the expression of FLOWERING LOCUS T1 (FT1). Photoperiod-1 (Ppd-1) partially regulates the seasonal induction of FT1, which responds dynamically to changes in daylength to influence the rate of inflorescence development. The incremental rise in FT1 expression is overridden in plants that contain a photoperiod-insensitive allele of Ppd-1, which hastens the completion of spikelet development and accelerates flowering time. The completion of spikelet formation is promoted by FLOWERING LOCUS T2, which influences spikelet number and is activated by Ppd-1. This research advances our understanding of the seasonal regulation of flowering in wheat and provides a strong foundation to improve yield potential by fine-tuning the photoperiod-dependent control of inflorescence development.

Allelic Differentiation at the E1/Ghd7 Locus Has Allowed Expansion of Rice Cultivation Area.

The photoperiod-insensitivity allele e1 is known to be essential for the extremely low photoperiod sensitivity of rice, and thereby enabled rice cultivation in high latitudes (42–53° north (N)). The E1 locus regulating photoperiod-sensitivity was identified on chromosome 7 using a cross between T65 and its near-isogenic line T65w. Sequence analyses confirmed that the E1 and the Ghd7 are the same locus, and haplotype analysis showed that the e1/ghd7-0a is a pioneer allele that enabled rice production in Hokkaido (42–45° N). Further, we detected two novel alleles, e1-ret/ghd7-0ret and E1-r/Ghd7-r, each harboring mutations in the promoter region. These mutant alleles alter the respective expression profiles, leading to marked alteration of flowering time. Moreover, e1-ret/ghd7-0ret, as well as e1/ghd7-0a, was found to have contributed to the establishment of Hokkaido varieties through the marked reduction effect on photoperiod sensitivity, whereas E1-r/Ghd7-r showed a higher expression than the E1/Ghd7 due to the nucleotide substitutions in the cis elements. The haplotype analysis showed that two photoperiod-insensitivity alleles e1/ghd7-0a and e1-ret/ghd7-0ret, originated independently from two sources. These results indicate that naturally occurring allelic variation at the E1/Ghd7 locus allowed expansion of the rice cultivation area through diversification and fine-tuning of flowering time.

The Photoperiod-Insensitive Allele Ppd-D1a Promotes Earlier Flowering in Rht12 Dwarf Plants of Bread Wheat.

The gibberellin-responsive dwarfing gene Rht12 can significantly reduce plant height without changing seedling vigor and substantially increase ear fertility in bread wheat (Triticum aestivum. L). However, Rht12 delays heading date and anthesis date, hindering the use of Rht12 in wheat improvement. To promote early flowering of the Rht12 dwarf plants, the photoperiod-insensitive allele Ppd-D1a was introduced through a cross between Jinmai47 (Ppd-D1a) and Karcagi (Rht12). The results showed that Ppd-D1a can rescue the delaying effect of Rht12 on flowering time and promote earlier flowering by 9.0 days (163.2°Cd) in the Rht12 dwarf plants by shortening the late reproduction phase. Plant height was reduced by Rht12 (43.2%) and Ppd-D1a (10.9%), achieving dwarf plants with higher lodging resistance. Ear fertility, like the grain number per spike, was significantly increased by Rht12 (21.3%), while it was reduced by Ppd-D1a (6.5%). However, thousand kernel weight was significantly reduced by Rht12 (12.9%) but significantly increased by Ppd-D1a (16.9%). Finally, plant yield was increased by 16.4 and 8.2%, and harvest index was increased by 24.9 and 15.4% in the Rht12 dwarf lines and tall lines with Ppd-D1a, respectively. Clearly, there was an additive interaction between Rht12 and Ppd-D1 and the introduction of Ppd-D1a advanced the flowering time and improved the yield traits of Rht12 dwarf plants, suggesting that the combination of Rht12 and Ppd-D1a would be conducive to the successful use of Rht12 in wheat breeding programs.

Allelic variation of vernalization and photoperiod response genes in a diverse set of North American high latitude winter wheat genotypes.

The major physiological determinants of wheat (Triticum aestivum L.) phenology in a given area are a response to vernalization temperature and day length, which are at least in part, regulated by the allelic variation at the vernalization (VRN) and photoperiod (PPD) loci, respectively. Characterization of the existing genetic variation for plant phenology in winter wheat can assist breeding programs improve adaptation to local environments and to optimize wheat phenology for the changing climate. The objectives of this research were to characterize the allelic variation at the major VRN and PPD loci in a diverse panel of high latitude winter wheat genotypes (n = 203) and to associate the allelic variation with phenologic, agronomic and adaptation traits. The panel was genotyped using allele-specific markers at vernalization (VRN-A1, VRN-B1, VRN-D1 and VRN-B3) and photoperiod (PPD-A1, PPD-B1, and PPD-D1) loci and phenotyped for agronomically-important traits. Though photoperiod sensitivity was more prevalent, most of the variation in the phenology of the winter wheat panel was explained by allelic variation at PPD-D1, PPD-A1, and the interaction between these loci. While a typical high latitude winter wheat genotype is one that carries winter alleles at all major VRN loci and photoperiod sensitive alleles at the major PPD loci, in lower latitudes where winters are milder, the presence of one or two photoperiod insensitive alleles seems to contribute to higher yield and wider adaptation.

Dynamics of floret initiation/death determining spike fertility in wheat as affected by Ppd genes under field conditions.

As wheat yield is linearly related to grain number, understanding the physiological determinants of the number of fertile florets based on floret development dynamics due to the role of the particular genes is relevant. The effects of photoperiod genes on dynamics of floret development are largely ignored. Field experiments were carried out to (i) characterize the dynamics of floret primordia initiation and degeneration and (ii) to determine which are the most critical traits of such dynamics in establishing genotypic differences in the number of fertile florets at anthesis in near isogenic lines (NILs) carrying photoperiod-insensitive alleles. Results varied in magnitude between the two growing seasons, but in general introgression of Ppd-1a alleles reduced the number of fertile florets. The actual effect was affected not only by the genome and the doses but also by the source of the alleles. Differences in the number of fertile florets were mainly explained by differences in the floret generation/degeneration dynamics, and in most cases associated with floret survival. Manipulating photoperiod insensitivity, unquestionably useful for changing flowering time, may reduce spike fertility but much less than proportionally to the change in duration of development, as the insensitivity alleles did increase the rate of floret development.

Physiological, biochemical and agronomic traits associated with drought tolerance in a synthetic-derived wheat diversity panel

Synthetic hexaploid wheat and their advanced derivatives (SYN-DERs) are promising sources for introducing novel genetic diversity to develop climate-resilient cultivars. In a series of field and laboratory experiments, we measured biochemical, physiological and agronomic traits in a diversity panel of SYN-DERs evaluated under well-watered (WW) and water-limited (WL) conditions. Analysis of variance revealed significant differences among genotypes, treatments and their interaction for all agronomic and physiological traits. Grain yield (GY) was reduced by 62.75% under WL, with a reduction of 38.10% in grains per spike (GS) and 19.42% in 1000-grain weight (TGW). In a Pearson’s coefficient correlation, GY was significantly correlated with GS, number of tillers per plant and TGW in both conditions. Path coefficient analysis showed that TGW and GS made the highest contribution to GY in WW and WL conditions, respectively. The traits examined in this experiment explained 59.6% and 63.01% of the variation in GY under WL and WW conditions, respectively; TGW, canopy temperature at spike and superoxide dismutase were major determinants of GY under WL conditions. The major flowering-time determinant gene Ppd-D1 was fixed in the diversity panel, with presence of the photoperiod-insensitive allele (Ppd-D1a) in 99% accessions. Wild-type alleles at Rht-B1 and Rht-D1, and presence of the rye translocation (1B.1R), favoured GY under WL conditions. Continuous variation for the important traits indicated the potential use of genome-wide association studies to identify favourable alleles for drought adaptation in the SYN-DERs. This study showed sufficient genetic variation in the SYN-DERs diversity panel to improve yields during droughts because of better adaptability than bread wheat.

Loss-of-Function Mutations in Three Homoeologous PHYTOCLOCK 1 Genes in Common Wheat Are Associated with the Extra-Early Flowering Phenotype.

Triticum aestivum L. cv ‘Chogokuwase’ is an extra-early flowering common wheat cultivar that is insensitive to photoperiod conferred by the photoperiod insensitive alleles at the Photoperiod-B1 (Ppd-B1) and Ppd-D1loci, and does not require vernalization for flowering. This reduced vernalization requirement is likely due to the spring habitat allele Vrn-D1 at the VERNALIZATION-D1 locus. Genotypes of the Ppd-1 loci that determine photoperiod sensitivity do not fully explain the insensitivity to photoperiod seen in ‘Chogokuwase’. We detected altered expression patterns of clock and clock-output genes including Ppd-1 in ‘Chogokuwase’ that were similar to those in an einkorn wheat mutant that lacks the clock-gene homologue, wheat PHYTOCLOCK 1 (WPCL1). Presumptive loss-of-function mutations in all WPCL1 homoeologous genes were found in ‘Chogokuwase’ and ‘Geurumil’, one of the parental cultivars. Segregation analysis of the two intervarietal F2 populations revealed that all the examined F2 plants that headed as early as ‘Chogokuwase’ had the loss-of-function wpcl1 alleles at all three homoeoloci. Some F2 plants carrying the wpcl1 alleles at three homoeoloci headed later than ‘Chogokuwase’, suggesting the presence of other loci influencing heading date. Flowering repressor Vrn-2 was up-regulated in ‘Chogokuwase’ and ‘Geurumil’ that had the triple recessive wpcl1 alleles. An elevated transcript abundance of Vrn-2 could explain the observation that ‘Geurumil’ and some F2 plants carrying the three recessive wpcl1 homeoealleles headed later than ‘Chogokuwase’. In spite of the up-regulation of Vrn-2, ‘Chogokuwase’ may have headed earlier due to unidentified earliness genes. Our observations indicated that loss-of-function mutations in the clock gene wpcl1 are necessary but are not sufficient to explain the extra-early heading of ‘Chogokuwase’.

Agronomic assessment of the wheat semi-dwarfing gene Rht8 in contrasting nitrogen treatments and water regimes

Reduced height 8 (Rht8) is the main alternative to the GA-insensitive Rht alleles in hot and dry environments where it reduces plant height without yield penalty. The potential of Rht8 in northern-European wheat breeding remains unclear, since the close linkage with the photoperiod-insensitive allele Ppd-D1a is unfavourable in the relatively cool summers. In the present study, two near-isogenic lines (NILs) contrasting for the Rht8/tall allele from Mara in a UK-adapted and photoperiod-sensitive wheat variety were evaluated in trials with varying nitrogen fertiliser (N) treatments and water regimes across sites in the UK and Spain.The Rht8 introgression was associated with a robust height reduction of 11% regardless of N treatment and water regime and the Rht8 NIL was more resistant to root-lodging at agronomically-relevant N levels than the tall NIL. In the UK with reduced solar radiation over the growing season than the site in Spain, the Rht8 NIL showed a 10% yield penalty at standard agronomic N levels due to concomitant reduction in grain number and spike number whereas grain weight and harvest index were not significantly different to the tall NIL. The yield penalty associated with the Rht8 introgression was overcome at low N and in irrigated conditions in the UK, and in the high-temperature site in Spain. Decreased spike length and constant spikelet number in the Rht8 NIL resulted in spike compaction of 15%, independent of N and water regime. The genetic interval of Rht8 overlaps with the compactum gene on 2DS, raising the possibility of the same causative gene. Further genetic dissection of these loci is required.

Genetic variation for flowering time and height reducing genes and important traits in western Canadian spring wheat

Genetic variation is prerequisite for wheat improvement. High grain yield and protein content and early maturity are some of the major objectives in global as well as Canadian wheat breeding programs. We investigated genetic diversity for earliness related and plant height reducing (Rht) genes in 82 spring wheat cultivars, registered in western Canada, through eight diagnostic DNA markers. Allelic variation was observed at the Vrn-A1, Vrn-B1, Vrn-D1 and Ppd-D1 loci but not at Ppd-A1 and Ppd-B1 loci in the 82 cultivars. Spring type allele of Vrn-A1 was present in 94 % cultivars, whereas only two cultivars carried spring allele of Vrn-D1. Among the four earliness related genes, the most frequent combination was Vrn-A1a, Vrn-B1, vrn-D1 and Ppd-D1b, which was found in 32 cultivars. As for the Rht genes, eight cultivars had Rht-B1b and 13 cultivars had Rht-D1b. All cultivars carrying dominant allele of Vrn-B1, photoperiod-insensitive allele of Ppd-D1 and height reducing allele of Rht-1 had shorter plants and higher grain yield but lower grain protein content. Days to heading and maturity showed positive genetic (rg = 0.65) and phenotypic (rp = 0.44) correlation, and were also positively correlated with grain yield and kernel weight but negatively correlated with test weight and protein content. Plant height was positively correlated with grain protein content (rg = 0.53; rp = 0.42), but negatively correlated with grain yield (rg = −0.47; rp = −0.14). Grain yield and protein content showed negative genetic correlation (rg = −0.57). Among the sixty cultivars from Canada Western Red Spring Class released over 100 years, the newest cultivar yielded 23 % more grain and had 15 % higher grain protein than the oldest cultivar ‘Red Fife’. Breeders in western Canada have incorporated vernalization and photoperiod insensitive and Rht genes in modern cultivars to promote early maturity, to make use of off-season nurseries abroad, and to improve lodging tolerance.

Discovery, evaluation and distribution of haplotypes and new alleles of the Photoperiod-A1 gene in wheat

Photoperiod response in wheat is determined to a large extent by the homoeologous series of Photoperiod 1 (Ppd1) genes. In this study, Ppd-A1 genomic sequences from the 5' UTR and promoter region were analysed in 104 accessions of six tetraploid wheat species (Triticum dicoccoides, T. dicoccum, T. turgidum, T. polonicum, T. carthlicum, T. durum) and 102 accessions of six hexaploid wheat species (T. aestivum, T. compactum, T. sphaerococcum, T. spelta, T. macha, T. vavilovii). This data was supplemented with in silico analysis of publicly available sequences from 46 to 193 accessions of diploid and tetraploid wheat, respectively. Analysis of a region of the Ppd-A1 promoter identified thirteen haplotypes, which were divided in two haplogroups. Distribution of the Ppd-A1 haplogroups and haplotypes in wheat species, and their geographical distributions were analysed. Polymerase chain reaction combined with a heteroduplex mobility assay was subsequently used to efficiently discriminate between Ppd-A1 alleles, allowing identification of the Ppd-A1b haplotypes and haplogroups. The causes of anomalous migration of Ppd-A1 heteroduplexes in gels were found to be the localization of mismatches relative to the center of fragment, the cumulative effect of neighbouring polymorphic sites, and the location of mismatches within A/T-tracts. Analysis of the Ppd-A1 5' UTR in hexaploid wheat revealed a novel mutation within the "photoperiod critical" region in a subset of T. compactum accessions. This putative photoperiod insensitive allele (designated Ppd-A1a.4) includes a 684bp deletion which spans region in common with deletions previously identified in other photoperiod insensitive Ppd1 alleles.

Allelic Variation in Loci for Adaptive Response and Its Effect on Agronomical Traits in Chinese Wheat (Triticum aestivum L.)

Heading date was an important trait that decided the adaptation of wheat to environments. It was modified by genes involved in vernalization response, photoperiod response and development rate. In this study, four loci Xgwm261, Xgwm219, Xbarc23 and Ppd-D1 which were previously reported related to heading time were analyzed based on three groups of wheat including landraces (L), varieties bred before 1983 (B82) and after 1983 (A83) collected from Chinese wheat growing areas. Generally, heading date of landrace was longer than that of varieties. Significant differences in the heading time existed within the groups, which implied that diversification selection was much helpful for adaptation in each wheat zone. Photoperiod insensitive allele Ppd-D1a was the first choice for both landrace and modern varieties, which promoted the heading date about four days earlier than that of sensitive allele Ppd-D1b. The three SSR loci had different characters in the three groups. Predominant allele combination for each zone was predicted for wheat group L and A83, which made great contribution to advantageous traits. Xgwm219 was found to be significantly associated with heading date in Yellow and Huai River Winter Wheat Zone (Zone II) and spike length in Middle and lower Yangtze Valley Winter Wheat Zone (Zone III), which implied functional diversification for adaption. Variation for earliness genes provided here will be helpful for whet breeding in future climatic change.

Allele frequencies in the VRN-A1, VRN-B1 and VRN-D1 vernalization response and PPD-B1 and PPD-D1 photoperiod sensitivity genes, and their effects on heading in a diverse set of wheat cultivars (Triticum aestivum L.).

Heading of cereals is determined by complex genetic and environmental factors in which genes responsible for vernalization and photoperiod sensitivity play a decisive role. Our aim was to use diagnostic molecular markers to determine the main allele types in VRN-A1, VRN-B1, VRN-D1, PPD-B1 and PPD-D1 in a worldwide wheat collection of 683 genotypes and to investigate the effect of these alleles on heading in the field. The dominant VRN-A1, VRN-B1 and VRN-D1 alleles were present at a low frequency. The PPD-D1a photoperiod-insensitive allele was carried by 57 % of the cultivars and was most frequent in Asian and European cultivars. The PPD-B1 photoperiod-insensitive allele was carried by 22 % of the genotypes from Asia, America and Europe. Nine versions of the PPD-B1-insensitive allele were identified based on gene copy number and intercopy structure. The allele compositions in PPD-D1, PPD-B1 and VRN-D1 significantly influenced heading and together explained 37.5 % of the phenotypic variance. The role of gene model increased to 39.1 % when PPD-B1 intercopy structure was taken into account instead of overall PPD-B1 type (sensitive vs. insensitive). As a single component, PPD-D1 had the most important role (28.0 % of the phenotypic variance), followed by PPD-B1 (12.3 % for PPD-B1_overall, and 15.1 % for PPD-B1_intercopy) and VRN-D1 (2.2 %). Significant gene interactions were identified between the marker alleles within PPD-B1 and between VRN-D1 and the two PPD1 genes. The earliest heading genotypes were those with the photoperiod-insensitive allele in PPD-D1 and PPD-B1, and with the spring allele for VRN-D1 and the winter alleles for VRN-A1 and VRN-B1. This combination could only be detected in genotypes from Southern Europe and Asia. Late-heading genotypes had the sensitivity alleles for both PPD1 genes, regardless of the allelic composition of the VRN1 genes. There was a 10-day difference in heading between the earliest and latest groups under field conditions.Electronic supplementary materialThe online version of this article (doi:10.1007/s11032-014-0034-2) contains supplementary material, which is available to authorized users.