Developmental responses to vernalization in wheat deletion lines for chromosomes 5A and 5D

The aim of this investigation was to test the developmental patterns of deletion lines, generated for chromosome arms 5AL and 5DL in the variety ‘Chinese Spring’ (CS) under vernalized and non‐vernalized treatments. Plants were grown in controlled conditions under saturating daylength. Time to heading and the duration of particular phases before flowering were recorded, and leaf and spikelet production rates and numbers were analysed. The lines lacking Vrn‐A1 and Vrn‐D1 were delayed in time to heading under non‐vernalized conditions, because of the lengthening of the emergence to floral initiation phase (EM‐FI) and the terminal spikelet to heading phase (TS‐H). Differences in final leaf numbers corresponded to longer durations of the EM‐FI phase. The absence of Vrn‐A1 and Vrn‐D1 apparently decreased the number of spikelets by a lower primordium production rate, even though the duration of the FI‐TS phase was longer or equal to CS. The sensitivity to vernalization in lines where the Vrn genes were deleted was much higher.     

The influence of a spring habit gene, Vrn-D1, on heading time in wheat

The adaptability of wheat cultivars to environmental conditions is known to be associated with a vernalization requirement, that is, spring/winter habit. To clarify the genetic effect of the spring habit gene, Vrn‐D1, on heading time in the field, recombinant inbred lines (RILs) with or without the Vrn‐D1 gene were produced from F₂ plants of the cross between ‘Nanbukomugi’ and ‘Nishikazekomugi’, non‐carrier and carrier cultivars of this gene, respectively. Using growth chambers with a controlled temperature and photoperiod, three components of heading time, i.e. vernalization requirement, photoperiodic sensitivity and narrow‐sense earliness (earliness per se), were evaluated in each RIL. RILs with the Vrn‐D1 gene (E lines) showed greatly reduced vernalization requirements and slightly shorter narrow‐sense earliness than RILs without Vrn‐D1 (L lines), although no difference in photoperiodic sensitivity was observed between the two groups. RILs were planted at four different sites in Japan and examined for their heading time in the field. E lines headed significantly earlier than L lines at all locations, indicating that the earliness of E lines is stable in various environmental conditions. These results indicated that spring habit caused by Vrn‐D1 gene, as well as narrow‐sense earliness, was responsible for heading time in the field.     

The effect of VRN1 genes on important agronomic traits in high‐yielding Canadian soft white spring wheat

For reproductive success, flowering time must synchronize with favourable environmental conditions. Vernalization genes play a major role in accelerating or delaying the time to flowering. We studied how different vernalization (VRN1) gene combinations alter days to flowering and maturity and consequently the effect on grain yield and other agronomic traits. The study focussed on the effect of the VRN1 gene series (Vrn‐A1, Vrn‐B1 and Vrn‐D1) and their combinations. The Vrn gene group Vrn‐A1a, Vrn‐B1, vrn‐D1 was the earliest to flower and mature, while Vrn‐A1b, Vrn‐B1, vrn‐D1 was the latest to flower. Spring wheat lines with vrn‐A1, Vrn‐B1, Vrn‐D1 were the highest yielding and matured at a similar time as those having vernalization genes Vrn‐A1a, Vrn‐B1 and Vrn‐D1. The findings of this study suggest that the presence of Vrn‐D1 has a direct or indirect role in producing higher grain yield. We therefore suggest the introduction of Vrn‐D1 allele into higher‐yielding classes within Canadian spring wheat germplasm.     

Genetic systems regulating flowering response in wheat

Genetic systems regulating bread wheat ontogenesis have been studied at Ukraine's Plant Breeding and Genetics Institute, for more than two decades. The influence of Vrn genes is the most obvious; dominant alleles of Vrn genes inhibit the vernalisation requirement. The Vrn genotypes of more than 1000 cultivars were determined and the peculiarities of gene geography were shown. Dominant Vrn1 or Vrn2 seemed to be replaced by Vrn3 in regions closer to the equator. In the developed sets of near-isogenic (congenic) lines, the value of different genes was characterised for certain environments (favourable – phytotron, natural – early or late drought) based on their effects. Methods of Vrn gene manipulation were elaborated, including methods for winter genotype selection from spring x spring crosses. The possibility of alien homoeologous Vrn loci introgression was shown. In the introgressed lines, the new genes were identified and found to be nonallelic to known Vrn genes in wheat. In studying congenic lines for three Ppd genes, differences were observed in duration and intensity of photoperiodic response, vernalisation requirement and effects on agronomic traits. For typical winter wheats, two loci were identified that influenced the duration of the vernalisation requirement. One system, controlling intrinsic earliness, might be responsible for the differences in reaction to light intensity, as selection of earlier genotypes is supposed to be more effective at lower light intensity. This revised version was published online in August 2006 with corrections to the Cover Date.     

Alien introgression of spring habit dominant genes into bread wheat

Summary Alien dominant genes of spring habit were introgressed into bread wheat. The introgression was undertaken by simple crossing of winter bread wheat to related spring species or genera, followed by backcrossing to winter bread wheat, and did not involve the use of the ph mutants or embryo culture. The introgressed genes were located mostly on chromosomes of homoeologous group 5, and were allelic to the known Vrn genes in bread wheat. Nevertheless three groups of lines were discovered with the genes possibly located on other chromosomes. These genes were non-allelic to each other and to known Vrn genes and were designated Vrn6 ^Sc , Vrn7 ^Sc (introgressed from Secale cereale) and Vrn8 ^Ts (from Triticum sphaerococcum).     

Waterlogging Affects Leaf and Tillering Dynamics in Wheat and Barley

The aim of this study was to analyse (i) the crop attributes that determine flowering time (i.e. final leaf number, FLN; and phyllochron, Phy), (ii) the dynamics of tiller appearance and (iii) the synchrony between leaf and tiller appearance in wheat and barley plants exposed to waterlogging. Two experiments were carried out in pots, in which wheat and barley cultivars were exposed to five waterlogging treatments, during different periods throughout the crop cycle, from emergence to maturity. The appearance of leaves and tillers on the main stem was measured twice a week in labelled plants. Waterlogging from emergence to flag leaf appearance significantly delayed time to flowering. The delay was greater when waterlogging occurred at the beginning of tillering, lengthening the period from emergence to flowering 24 % (13–15 days) in barley and 10–15 % (6–10 days) in wheat, as compared to control. Phy was the main attribute explaining the delay in flowering, as FLN was not altered. Waterlogging during the early stages of development reduced tiller appearance rate (TAR) in both species, but this effect was partially counterbalanced by a lengthening of the tillering phase, so the effect on final tiller number at maturity was limited. In conclusion, the exposure of wheat and barley to waterlogging during early stages of development delayed time to flowering and reduce TAR in both species. Waterlogging during more advanced crop stages produced slight effects on tillering dynamics, which would indicate that waterlogging affected structure generation more than mortality.     

Differences between spring wheat cultivars in early growth

Summary Twelve spring wheat cultivars were grown as isolated plants in the field and their pre-anthesis growth was interpreted in terms of plant growth analysis. Relative growth rate (RGR) decreased steadily with time due to a decline of leaf area per unit plant weight (LAR), which could be explained by the reduction of the portion of leaf weight in total plant weight (LWR). Growth per unit leaf area (NAR) and leaf area per unit leaf weight (SLA) changed only little with time.Differences between cultivars for NAR and LAR were of similar magnitude: both 8% when measured by the genetic coefficient of variation. Because both quantities were negatively correlated, the genetic variation of RGR was only 5%. Genetic variation for LWR and SLA were also of similar size, both about 4%. Estimates of genetic variances and covariances based on cultivar means appeared to be biased strongly when the error variation of the means was neglected.Special attention is paid to the methodology of plant growth analysis.     

Geographic distribution of Vrn-genes in landraces and improved varieties of spring bread wheat

Summary A study of the global Vrn-genes distribution supports the hypothesis that these loci have different breeding values. Evident zonal differences were discovered in percentage of dominant Vrn-genes and genotypes between the sets of varieties from various locations. For the same zone analogous differences were not so manifested between the sets of local and improved varieties. It showed the similar environmental fitness of those sets as a result of natural or artificial selection. Obvious changes in distribution of Vrn-genes were discovered in the history of breeding. Within this century the dominant Vrn3 was introgressed into many modern varieties of countries close to the equator. The main direction of such introgression ran from the Japanese landrace Akakomugi to Italian varieties. Later it came from Italy through Mentana to Mexican semidwarf varieties, and the last ones (especially Sonora 64 and Lerma Rojo 64) promoted wide distribution of that gene all over the world.     

Growth habit in common wheat (Triticum aestivum L. em. Thell.)

Summary A study of the Vrn genotypes of 642 spring wheats supports the theory that only Vrn1, Vrn2 and Vrn3 exist in Tricticum aestivum. In none of the varieties investigated Vrn4 was present. Seven varieties, which according to literature carry Vrn4, showed to carry Vrn1, Vrn2 and/or Vrn3. Some varieties were mixtures of Vrn-genotypes, which could mislead geneticists in pooled data analysis. Other causes for misinterpretation of the data could be hybrid necrosis, hybrid dwarfness or a wrong determination of plants with a winter habitus. Only Hope was dominant on another Vrn locus. Its haploid Vrn-genotype is Vrn1 vrn2vrn3 Vrn5.     

Location of a gene for frost resistance on chromosome 5A of wheat

Summary A gene for frost resistance on chromosome 5A of wheat was located using single chromosome recombinant lines from the cross between the substitution line Hobbit (Triticum spelta 5A) and Hobbit. In this sample of recombinant lines the locus for frost resistance, designated Fr1, is completely linked to the locus Vrn1 controlling vernalisation requirement. The results can be explained by a pleiotropic action of the Vrn1 locus or close genetic linkage between Vrn1 and Fr1. Further detailed study is necessary to resolve these alternative hypotheses.     

Genetic resources of wheat related species: The Vrn genes controlling growth habit (spring vs. winter)

Variations in growth habit and its genetic control in landraces of hexa-, tetra- and diploid wheats were studied. Studies demonstrated simple (one, two, or three genes) genetic control of growth habit (spring vs. winter). Triticum aestivum originated from only a part of the gene pools of the diploid species. Results show the feasibility of transferring new Vrn genes from the Sitopsis section of species of the Aegilops genus and the weak allele at the Vrn1 gene from T. dicoccum into cultivated species. The possibilities of introgressing new Vrn genes from diploid related species into cultivated wheats are considered. This revised version was published online in August 2006 with corrections to the Cover Date.     

Partial resistance to wheat leaf rust in 18 spring wheat cultivars

Summary Eighteen spring wheat cultivars were tested in microfields and race nurseries for their partial resistance PR to wheat leaf rust under low and high disease pressure respectively. Large differences existed between the 18 cultivars, Skalavatis 56 being the most susceptible and Ponta Grossa 1 being the most resistant cultivar. Of the three epidemic parameters, disease severity (DS) at the time that the susceptible check was severely diseased and area under the transformed disease severity curve (AUTC) and the logistic growth rate (r), AUTC and DS were highly correlated. Both seemed to be reliable estimators of PR but DS should be preferred for economical reasons. The logistic growth rate seemed to be unsuitable as an estimator of partial resistance.High and low disease pressure gave similar cultivar ranking. PR can be screened and selected equally well in race nurseries with low space, low time and low cost input as in microfields with high space, time and cost input.Cultivar differences in development rate had a large impact on the cultivar differences for amount of disease and can therefore greatly bias the estimation of cultivar resistance. The resistance of early cultivars tended to be underestimated whereas the resistance of late cultivars tended to be overestimated. The effect of differences in developmental rate was most pronounced in the flag leaf. It is advisable to avoid the assessment of disease levels on the flag leaf only and to incorporate in the tests several susceptible and resistant checks that cover the range of development rates in the material to be selected, because otherwise selection for resistance will tend to select also for lateness.Regression of the epidemiological parameters on three components of partial resistance revealed that latency period (LP) is an important factor in determining the resistance observed in the field explaining on average 67% of the observed variation. Adding infection frequency (IF) and urediosorus size (US) to the linear model increased the proportion of the observed variation in the field explained by the components to 80%. This result supports the idea that the components of PR inherit independently, at least, in part.     

Evaluation of cold hardiness in two sets of near-isogenic lines of wheat (Triticum aestivum) with polymorphic vernalization alleles

In wheat, variation at the orthologus Vrn‐1 loci, located on each of the three genomes, A, B and D, is responsible for vernalization response. A dominant Vrn‐1a allele on any of the three wheat genomes results in spring habit and the presence of recessive Vrn‐1b alleles on all three genomes results in winter habit. Two sets of near‐isogenic lines (NILs) were evaluated for DNA polymorphisms at their Vrn‐A1, B1 and D1 loci and for cold hardiness. Two winter wheat cultivars, ‘Daws’ and ‘Wanser’ were used as recurrent parents and ‘Triple Dirk’ NILs were used as donor parents for orthologous Vrn‐1 alleles. The NILs were analysed using molecular markers specific for each allele. Only 26 of 32 ‘Daws’ NILs and 23 of 32 ‘Wanser’ NILs had a plant growth habit that corresponded to the marker genotype for the markers used. Freezing tests were conducted in growth chambers programmed to cool to ?21.5°C. Relative area under the death progress curve (AUDPC), with a maximum value of 100 was used as a measure of death due to freezing. The average relative AUDPC of the spring habit ‘Daws’Vrn‐A1a NILs was 86.15; significantly greater than the corresponding winter habit ‘Daws’Vrn‐A1b NILs (42.98). In contrast, all the ‘Daws’Vrn‐A1bVrn‐B1aVrn‐D1b and Vrn‐A1bVrn‐B1bVrn‐D1a NILs (spring habit) had relative AUDPC values equal to those of their ‘Daws’ sister genotypes with Vrn‐A1bVrn‐B1bVrn‐D1b NILs (winter habit). The average AUDPC of spring and winter habit ‘Wanser’ NILs differed at all three Vrn‐A1, Vrn‐B1 and Vrn‐D1 locus comparisons. We conclude that ‘Daws’ and ‘Wanser’ have different background genetic interactions with the Vrn‐1 loci influencing cold hardiness. The marker for Vrn‐A1 is diagnostic for growth habit and cold hardiness but there is no relationship between the Vrn‐B1 and Vrn‐D1 markers and the cold tolerance of the NILs used in this study.     

The extent of similarity between near-isogenic lines of the Australian spring wheat variety Triple Dirk with their recurrent parent

Summary The near-isogenic lines of the wheat variety Triple Dirk supposed to differ only for Vrn genes have been investigated for their similarity to Triple Dirk and for purity. The NILs were found to differ from Triple Dirk. It was also discovered that Triple Dirk and the NILs each exist of more than one genotype. It is recommended that each NIL is tested for its Vrn-genotype and purified. New sets of Vrn NILs can only be made with these checked NILs and with one-plant progeny of the newly used recurrent parents.     

Genetic and Molecular Characterization of Vernalization Genes Vrn‐A1, Vrn‐B1, and Vrn‐D1 in Spring Wheat Germplasm from the Pacific Northwest Region of the U.S.A.

The objective of this study was to determine the Vrn‐1 allelic composition of spring wheat germplasm from the Pacific Northwest region of the USA. Individual plants from 56 spring wheat lines were crossed to near‐isogenic tester lines carrying the dominant allele Vrn‐A1, Vrn‐B1 or Vrn‐D1. F₂ progeny were evaluated for growth habit in the field and Vrn‐1 allelic composition was determined through chi‐square analysis. Lines also were analysed with DNA sequence‐based Vrn‐1 allele‐specific markers. A majority of the germplasm carried the dominant allele Vrn‐A1a alone or in combination with Vrn‐B1, Vrn‐D1 or Vrn‐B3 alleles. Vrn‐B1 and Vrn‐D1 were almost always associated with other dominant Vrn‐1 allele(s). Based on DNA sequence analysis, a novel Vrn‐B1 allele referred to as Vrn‐B1b, which carried a single nucleotide polymorphism (SNP) and a 36 bp deletion, was identified in cultivar ‘Alpowa’. These results will be useful to wheat breeders for choosing parents with different Vrn‐1 alleles for crossing to maximize diversity at the Vrn‐1 loci with an expectation of identifying superior Vrn‐1 allelic combinations for cultivar improvement.     

Genotypic variation for competitive ability in spring wheat

Herbicides are the primary method of weed control for crop production in developed countries. For economic and environmental reasons alternative control strategies are being devised. One of these strategies is the development of competitive crop cultivars. The objectives of this research were to establish whether spring wheat (Triticum aestivum L.) genotypes differed in competitive ability and if those differences were related to specific growth characteristics. Sixteen genotypes of spring wheat were grown under simulated weed competition conditions at Saskatoon, Canada over a 3–year period. Four high and four low tillering genotypes from each of two crosses (Neepawa/M1417 and Ingal/M1417) were studied. Weeds consisted of cultivated oat (Avena saliva cv. ‘Waldern’) and oriental mustard (Brassicajuncea cv. ‘Cutlass’) sown at two densities (48 and 96 seeds/m² per weed species). Seedling establishment, ground cover, and seed yield for the three species were determined, as was wheat tiller number, spike number, maximum height, leaf area index, leaf orientation, and flag leaf length and size. Significant (P = 0.001) weed rate by genotype interactions involving changes in genotype rank were detected for wheat grain yield, indicating that the 16 wheat genotypes differed in competitive ability. Wheat grain yield reductions averaged over the two weed densities ranged from 45% to 59%. The highest-yielding genotypes under weed-free conditions were not necessarily the highest yielding under weedy conditions. Genotypes which suffered smaller yield reductions were more effective in suppressing weed growth. Although competitive genotypes were generally taller than non-competitive genotypes, other traits such as large seedling ground cover and flag leaf length were associated with wheat yield under competitive conditions.     

Adaptation and ecological differentiation in wheat with special reference to geographical variation of growth habit and Vrn genotype

Geographical variation of growth habit was studied for 749 landraces from various parts of the world, with special reference to their adaptation and ecogeographical differentiation. The total frequency of spring‐type landraces was 49.9%, and varied between localities. Spring‐type landraces were frequent in two distinct areas where the average January temperature was either below ‐7°C or above 4°C, with winter‐type landraces in areas from ‐7°C to 4°C. These results indicated that geographical variation of growth habit is closely related to the degree of winter coldness. An analysis of the Vrn genotype for 216 spring‐type landraces demonstrated the uneven distribution of four Vrn genes, with Vrn4 being the least frequent. The adaptive Vrn genotype was different between localities. Genotypes carrying Vrn‐A1 and additional Vrn gene(s) were frequent in two distinct areas where the average January temperature was either below ‐7°C or over 10°C, while genotypes with any of three Vrn genes, except Vrn‐A1, adapted to areas with temperatures from 4°C to 10°C. Therefore, it was concluded that the adaptability of wheat landraces differed depending on their growth habit and Vrn genotype, and that ecotypes with different Vrn genotypes were allopatrically distributed as a result of adaptation to different winter temperature. However, the differential distribution of Vrn‐B1, Vrn‐D1 and Vrn4 could not be explained by their adaptability, and might reflect the polyphyletic origin of common wheat.     

The inheritance of morphological and agronomic characters in spring wheat

Summary Inheritance of yield and its component (number of ears per plant, number of spikelets per ear, number of kernels per ear, 1000-kernel weight), together with the inheritance of morphological characters (flag leaf length, flag leaf breadth, peduncle length, extrusion length and leaf sheath length) and anthesis date was studied in a 5-parent diallel cross of spring wheat varieties. The diallel analysis of gene actions indicated that a large part of the total genetic variation observed for all the characters studied was in the form of additive genetic effects. The degree of dominance was also determined for each character. Number of ears per plant showed most overdominance, followed by leaf sheath length. Full or nearly full dominance was found for anthesis date, flag leaf length and yield per plant. Other characters showed partial dominance. Correlation coefficients showed that yield per plant, as well as being correlated with the yield components, was also assiciated with flag leaf breadth and leaf sheath length. There was no evidence of genetic barriers to the combining of ideal characters among the varieties of different origin.     

Responses to salt stress controlled by the homoeologous group 5 chromosomes of hexaploid wheat

The responses to salt stress in NFT (nutrient film) hydroponics of ‘Chinese Spring’ wheat and a number of its aneuploids involving the chromosomes of homoeologous group 5 were studied. This showed that the absence of chromosome 5D allowed plants to survive better than in the euploid condition. Much of this response could be related to the effects of Vrn3, which conditions the spring habit of ‘Chinese Spring’. The ability to survive relatively high levels of stress was promoted by the group 5 homoeologue from Thinopyrum bessarabicum.