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.     

Simplified culture method of detached ears and its application to vernalization in wheat

Summary The effects of the addition of sulfurous acid into culture solution and of cold treatment of the solution were examined to simplify the culture of detached wheat ears. In the simplified method, detached ears could be cultured at room temperature on the liquid medium containing 100 g/l sucrose and 0.075% sulfurous acid without any sterilization. The immature seeds in detached ears cultured by this method were treated with low temperature or with chemicals known to have vernalizing effect. The chemical treatment did not affect the chilling requirement of immature embryos, although photoperiodic response and narrow-sense earliness were reduced by kinetin and trypsin. The low temperature treatment drastically affected the chilling requirement, and fully vernalized mature seeds having normal germinability were obtained by treating the detached ears in culture with low temperature from 10 days after anthesis.     

Genetic effects of Vrn genes on heading date and agronomic traits in bread wheat

Summary The Vrn1, Vrn2 and Vrn3 genes have different values of effects on heading date and related yield components. The genetic background and environment do not affect the ranking of Vrn genotypes according to earliness within near-isogenic line sets; however, they do influence the level of differences between heading dates of particular genotypes and between effect values, respectively. The frequencies of defined Vrn genotypes in the global set of spring bread wheat cultivars are associated with grain weight per plant predicted on the basis of Vrn gene effects averaged over backgrounds and over environments. Peculiarities of backgrounds and environments alter the grain yield ranges of Vrn genotypes. For early photoperiod-insensitive wheats, planted in stress conditions at grain filling, the highest yield was predicted for double dominant Vrn genotypes with Vrn3. This gene is rarely used by the breeders in middle latitudes and its wider adoption is encouraged.     

Vernalization response and earliness per se in cultivars representing different eras of wheat breeding in Argentina

Argentine wheat cultivars are assumed to be essentially vernalization insensitive or very slightly sensitive. However, there are only speculations on this lack of vernalization requirement and a greater unawareness on the variation in earliness per se. The aims of this research have been to determine the extent of variability in vernalization requirement and earliness per se, and how the variability in both traits was produced by breeding programs, through the release of wheat cultivars from the 1930's to the 1990's in Argentina. Sixty-eight cultivars, selected among those of highest performance in each era, were evaluated under field and glasshouse conditions for their vernalization response and earliness per se. Forty per cent of the cultivars showed some vernalization response. There was a decrease in this requirement along the first decades of the analysed breeding period, likely in response to the considerable introgression of CIMMYT germplasm. This initial trend to release earlier cultivars was also evidenced in a clear decrease in earliness per se. As this tendency in both characteristics was reverted during the last two decades, it may denote that certain vernalization response and not extreme earliness per se, may contribute to achieve higher yield cultivars in our area. This revised version was published online in July 2006 with corrections to the Cover Date.     

Factors influencing the development of bread wheat plant types to be grown in the ‘transitional zone’ in northern Syria

Summary This study sought to identify factors that influence wheat development in the transitional wheat growing zone of northern Syria. Three development factors were studied, intrinsic earliness, and responses to vernalization and to photoperiod. Two sets of wheat were studied, each composed of lines with differing combinations of development factors. Set 1 comprised 20 parental and breeding lines utilized by the CIMMYT/ICARDA facultative and winter wheat breeding program based at Tel Hadya. Set 2 comprised 19 parental and breeding lines utilized by an Australian winter wheat breeding program based at Temora. Field development was recorded in greatest detail at one site. Tel Hadya, using the state of differentiation of the apex of the main tiller of sampled plants. To extend findings, development was also recorded as the time from sowing to ear emergence for later sowings of wheat at Tel Hadya, and in sowings at four other regional sites.The significance of each development factor was tested in multiple regressions that predicted either stage of apical development at Tel Hadya, or time to ear emergence in all trials. It was found that intrinsic earliness was the major factor associated with development, in both sets of wheat. Response to photoperiod had a much smaller and less consistent effect. Response to vernalization had least effect on development, possibly because low temperature in winter delayed development for a longer period than was required to fully vernalize winter wheats. Our results suggested it may not be directly relevant whether spring or winter wheats are grown in the transitional zone of northern Syria. The desired phenotype for the region, of slow development prior to double ridge, then fast development to ear emergence, cannot be simply achieved from combinations of the three development factors. Selection for improved adaptation to the region must continue to rely on direct field observations.     

Measurement of response to vernalization in Australian wheats with winter habit

Summary Development in wheat is strongly controlled by sensitivity to vernalization and photoperiod, and to a lesser degree by non-vernalizing temperature and intrinsic earliness. A method to measure effect of vernalization in wheats with winter habit is described. Twenty seven wheats with winter habit and eight with spring/facultative habit were studied, comprising breeding lines and cultivars with maturities suited to south-central New South Wales. Effect of vernalization on the development of these wheats was quantified by integrating responses to vernalizing treatments of differing duration. Intrinsic earliness was measured as time for vernalized seedlings to grow to ear emergece in an 18h photoperiod with day/night temperature of 21/16°C, and response to photoperiod as the difference in time to ear emergence between 9 and 18h daylengths. Integrated response to vernalization is sensitive to both cumulative and thresh-hold responses and is applicable to wheats of all habit type. Integrated response to vernalization and intrinsic earliness were positively associated within wheats with winter habit. Wheats were largely of restricted origin, so that there were few allelic differences at Vrn loci to disrupt this association, which suggests intrinsic earliness may modify response to vernalization. Though integrated response to vernalization was measured with artificial treatments, it was strongly associated with ear emergence for wheats with winter habit when grown at a site in New South Wales.     

Response of four winter wheat cultivars to triallate

Summary Genetic variation among four winter wheat cultivars for response to treatment with triallate was examined. The cultivars showed differential tolerance based on stand count, however, differential tolerance was not seen for any of the other traits examined including visual score, and the mature plant traits, tiller count and yield. Significant effects due to increasing the rate of chemical were observed. An increase in yield was seen at the 0.5 and 1.0 kg/ha rates of triallate. This yield increase was paralleled by trends toward increasing seed number/head and tiller number/unit area. The results indicate that the effect of triallate on wheat has two components. The first component, toxicity, is seen on all cultivars, but the extent of damage varies significantly among cultivars. The second component is positive and leads to an increase in yield without interacting significantly with genotype.     

Dough stickiness in rye-derived wheat cultivars

Summary Rye-derived wheat cultivars are being used in many breeding programmes throughout the world in order to achieve improvements in yield and disease resistance. However, the serious quality defect of intense dough stickiness associated with many of these wheat cultivars is limiting the usefulness of their flour in large mechanised bread bakeries. A dough preparation procedure was developed which enabled the dough surface properties of a range of rye-derived wheat cultivars to be assessed on doughs mixed quantitatively to their optimum mixing time. Intense dough stickiness was found in samples of 1AL/1RS and 1DL/1RS translocation lines tested and in all of the 1BL/1RS wheat cultivars examined except the West German cultivar, Disponent. Most of the 1BL/1RS wheat cultivars were derived from the Russian cultivars, Kavkaz, Aurora and Skorospelka 35 and included the CIMMYT-bred cultivars such as the Veery lines (Glennson, Ures, Genaro and Seri) and the Nebraskan cultivar, Siouxland. Based on the results of studying selected 1BL/1RS wheat cultivars in detail, this intense dough stickiness appeared to be independent of growing season, trial location, protein content, mixing tolerance, milling process and extraction rate. In addition pilot bakery trials confirmed that our laboratory testing procedures can be used to detect this intense dough stickiness.     

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.     

Intrinsic earliness and basic development rate assessed for their response to temperature in wheat

The related concepts of basic vegetative period, intrinsic earliness and basic development rate in wheat are examined. These concepts have the common assumption that, if plants are vernalised fully and then grown at long daylength in order to remove any responses to vernalisation and photoperiod, the calendar or thermal time then taken to anthesis will be a characteristic of a genotype that will be heritable. Thus, regardless of temperature, early genotypes will always be earlier than late genotypes (providing there are no vernalisation and photoperiod responses).Using four genotypes, exposed to 50 days of vernalisation, and then grown at 18 h photoperiod under six temperature regimes ranging between 10 and 25°C, it is shown that; (1) no genotype had an absolute basic period as, depending on temperature, durations to anthesis for any one genotype varied by more than 50 days; (2) no genotype had an absolute value for intrinsic earliness (to anthesis), ranging for any genotype by more than 300°Cd depending on temperature; (3) basic development rate was not a single value for a genotype but varied with stage of development; (4) some genotypes changed their ranking for earliness depending on the temperature regime; and (5) genotypes were differentially sensitive to temperature for the subphases prior to anthesis.     

On defining a winter wheat

Summary An international survey of wheat breeders and scientists was conducted to determine whether there was a common understanding of the term winter wheat. Response to photoperiod was mentioned by only seven of the 30 survey respondents. It was consequently suggested that it be excluded from any definition. Response to vernalization was mentioned by 28 of the 30 respondents. However, there was considerable variation in the method of quantifying this response and the degree of response necessary for wheat to be called winter. It was concluded that the only objective and internationally consistent definition of winter wheat was a genetic definition based on the complement of Vrn genes a wheat possesses. Varieties not possessing any of the dominant Vrn alleles appear to take a distinctively longer and more variable time to head when grown under non-vernalizing, long day conditions. These are genetically winter wheats.     

Distribution of photoperiod-insensitive allele Ppd-A1a and its effect on heading time in Japanese wheat cultivars

The Ppd-A1 genotype of 240 Japanese wheat cultivars and 40 foreign cultivars was determined using a PCR-based method. Among Japanese cultivars, only 12 cultivars, all of which were Hokkaido winter wheat, carried the Ppd-A1a allele, while this allele was not found in Hokkaido spring wheat cultivars or Tohoku-Kyushu cultivars. Cultivars with a photoperiod-insensitive allele headed 6.9–9.8 days earlier in Kanto and 2.5 days earlier in Hokkaido than photoperiod-sensitive cultivars. The lower effect of photoperiod-insensitive alleles observed in Hokkaido could be due to the longer day-length at the spike formation stage compared with that in Kanto. Pedigree analysis showed that ‘Purple Straw’ and ‘Tohoku 118’ were donors of Ppd-A1a and Ppd-D1a in Hokkaido wheat cultivars, respectively. Wheat cultivars recently developed in Hokkaido carry photoperiod-insensitive alleles at a high frequency. For efficient utilization of Ppd-1 alleles in the Hokkaido wheat-breeding program, the effect of Ppd-1 on growth pattern and grain yield should be investigated. Ppd-A1a may be useful as a unique gene source for fine tuning the heading time in the Tohoku-Kyushu region since the effect of Ppd-A1a on photoperiod insensitivity appears to differ from the effect of Ppd-B1a and Ppd-D1a.     

Multiline cultivars — how their resistance influence leaf rust diseases in wheat

Summary To understand how multiline cultivars of wheat develop better protection against leaf rust, seven experimental multilines with 0, 28, 40, 50, 58, 60 and 70% susceptibility were subjected to leaf rust epiphytotics in the field along with their pure line components. A mixture comprising 12 leaf rust races, 10, 11, 12, 17, 20, 63, 77, 106, 107, 108, 162 and 162 A was used.Both the initial inoculum (Xo) and rate of increase (r) of leaf rust were substantially reduced in the multiline cultivars. Xo was reduced by 45–75% and the over-all infection rate (r) by as much as 16% over the average of components.As a result of reduced Xo and r, the intensity of leaf rust in the multilines was also significantly affected at all stages of rust development. It was reduced from 32,10 to 89.54% over the average of components differing from one multiline to another and also from time to time. The susceptible recurrent parent, Kalyansona at the peak period of rust infection exhibited 86.75% severity while in the multilines it ranged from 5.80 to 35%.The rate of increase in the multilines was found to be proportional to the logarithm of the proportion of susceptible plants in the host mixture.Further, it was found that even if as many as 50% susceptible plants are present in a multiline they would not suffer much from leaf rust damage.     

Comparison of phenotypic and molecular marker-based classifications of hard red winter wheat cultivars

Genetic diversity is the basis for successful crop improvement and can be estimated by different methods. The objectives of this study were to estimate the genetic diversity of 30 ancestral to modern hard red winter wheat (Triticum aestivum L.) cultivars adapted to the Northern Great Plains using pedigree information, morphological traits (agronomic measurements from six environments), end-use quality traits (micro-quality assays on 50 g grain or milled flour samples for the six environments), and molecular markers (seed storage proteins separated using SDS-PAGE, 51 SSRs, and 23 SRAP DNA markers), and to determine the relationships of genetic distance estimates obtained from these methods. Relationships among diversity estimates were determined using simple (Pearson) and rank (Spearman) correlation coefficients between distance estimates and by clustering cultivars using genetic-distances for different traits. All methods found a wide range in genetic diversity. The genetic distance estimates based on pedigree had the highest values due to possible over-estimation arising from model assumptions. The genetic diversity estimates based on seed storage protein were lowest because they were the major determinants of end-use quality, which is a highly selected trait. In general, the diversity estimates from each of the methods were positively correlated at a low level with the exceptions of SRAP diversity estimates being independent of morphologic traits (simple correlation), SDS-PAGE, and SSR diversity estimates (rank correlation). However, SSR markers, thought to be among the most efficient markers for estimating genetic diversity, were most highly correlated with seed storage proteins. The procedures used to accurately estimate genetic diversity will depend largely upon the tools available to the researcher and their application to the breeding scheme.     

Effect of VRN‐1 and PPD‐D1 genes on heading time in European bread wheat cultivars

The variation of the vernalization (VRN‐1) and photoperiod (PPD‐1) genes offers opportunities to adjust heading time and to maximize yield in crop species. The effect of these genes on heading time was studied based on a set of 245 predominantly spring cultivars of bread wheat from the main eco‐geographical regions of Europe. The genotypes were screened using previously published diagnostic molecular markers for detecting the dominant or recessive alleles of the major VRN‐1 loci such as: VRN‐A1, VRN‐B1, VRN‐D1 as well as PPD‐D1. We found that 91% of spring wheat cultivars contain the photoperiod sensitive PPD‐D1b allele. Photoperiod insensitive PPD‐D1a allele has been found mainly in southern region of Europe. For this region the monogenic control of vernalization by VRN‐B1 or VRN‐D1 dominant alleles is common, whereas in the remaining part of Europe, the combination of photoperiod sensitive PPD‐D1b allele with dominant VRN‐A1, VRN‐B1 and recessive vrn‐D1 alleles represents the most frequent genotype. Also, we revealed a significantly later (5–8 days) heading of the monogenically dominant genotypes at VRN‐B1 as compared to the digenic VRN‐A1 VRN‐B1 genotypes.     

Evaluation of seedling and adult plant resistance to leaf rust in European wheat cultivars

Summary A set of 105 European wheat cultivars, comprising 68 cultivars with known seedling resistance genes and 37 cultivars that had not been tested previously, was tested for resistance to selected Australian pathotypes of P. triticina in seedling greenhouse tests and adult plant field tests. Only 4% of the cultivars were susceptible at all growth stages. Twelve cultivars lacked detectable seedling resistance to leaf rust, and among the remaining cultivars, 10 designated genes were present either singly or in combination. Lr13 was the most frequently detected gene, present in 67 cultivars, followed by the rye-derived gene Lr26, present in 19 cultivars. Other genes present were Lr1, Lr3a, Lr3ka, Lr10, Lr14a, Lr17b, Lr20 and Lr37. There was evidence for unidentified seedling resistance in addition to known resistance genes in 11 cultivars. Field tests with known pathotypes of P. triticina demonstrated that 57% of the cultivars carried adult plant resistance (APR) to P. triticina. The genetic identity of the APR is largely unknown. Genetic studies on selected cultivars with unidentified seedling resistances as well as all of those identified to carry APR are required to determine the number and inheritance of the genes involved, to determine their relationships with previously designated rust resistance genes, and to assess their potential value in breeding for resistance to leaf rust.     

Relationship between relative water content during reproductive development and winter wheat grain yield

Summary Water is often the most limiting factor to winter wheat (Triticum aestivum L.) production in the southern Great Plains of the U.S.A., yet the lack of reliable screening criteria has precluded direct selection for drought resistance in breeding programs. Previous work showed that leaf relative water content (RWC) was highly heritable when measured under field-drought conditions, but its adoption as a screening tool for yield improvement requires further investigation of the genetic relationship between grain yield and RWC. Plants representing high and low yield potential under drought stress, and a random group of plants, were selected from an F₂ population having the pedigree, TAM W-101/Sturdy. Two sets of entries, each comprised of the two parents and 24 F₂-derived lines, were evaluated under a rainshelter in the F₃ (1986) and F₄ (1987) generations to determine differences in leaf RWC during reproductive development. One set of entries did not receive any water after the jointing stage, and the other set was grown under well-watered conditions. A positive relationship was observed between grain yield and RWC measured during anthesis and mid-grain fill, as the high-yield selections maintained a significantly higher RWC than the low-yield selections. Grain yield and RWC were also positively associated among random selections segregating for both traits. Subsequent adjustment of genotype means for differences in reproductive development at time of sampling underscored the need to consider differences in maturity when RWC is the selection criterion.     

Shortening vernalization of winter wheat with kinetin

Summary The addition of 5 to 100 ppm kinetin during vernalization is a substitute for a part of the cold treatment in the winter wheat cultivar Sava.A cold treatment that is not long enough in itself makes vernalization possible even if only 5 ppm kinetin is added. The effect can be gradually improved by increasing the dose from 20 to 100 ppm, but 200 ppm kinetin is harmful. The smallest quantity of kinetin added to the otherwise optimal 40-day cold treatment results in total oververnalization.A heading percentage comparable to that of the 40-day vernalization period without kinetin can be obtained in 20-day with 100 ppm kinetin. This method can reduce the vernalization time by half and accelerate winter wheat generations.     

Relationship between common wheat (Triticum aestivum L.) gluten proteins and dough rheological properties

This paper reports the correlation between the rheological properties of bread wheat dough and the types and quantities of endosperm proteins in 28 common wheat cultivars. Different methods were used to analyse the allelic composition of these cultivars and the relative quantities of the different proteins contributing to the gluten structure. Neither dough strength (W) nor tenacity/extensibility (P/L) correlated with allelic composition. Different wheats with the same allelic composition (i.e., with respect to glutenins) showed different rheological properties. The glutenins were the most influential components of W and P/L, especially the high molecular weight (HMW) glutenin subunits and in particular the type x form. These proteins seem to increase W and are the main constituents of the gluten network. The gliadins and low molecular weight (LMW) glutenin subunits appear to act as a “solvent”, and thus modify the rheological properties of the dough by either interfering with the polymerisation of the HMW glutenin subunits, or by altering the relative amounts of the different types of glutenin available. Thus, the protein subunits coded for by the alleles Glu-B1x7 and Glu-D1x5 stabilised the gluten network, whereas those coded for by Glu-B1x17 and Glu-D1x2 had the opposite effect. Dough properties therefore appear to depend on the glutenin/gliadins balance, and on the ratio of the type x and type y HMW proteins. The influence of external factors seems to depend on the allelic composition of each cultivar.     

Evaluation of winter wheat cultivars for drought resistance

Summary Cultivars of winter wheat (Triticum aestivum L. em. Thell.), considered by wheat breeders to be drought sensitive or drought resistant, were grown under two irrigation regimes (daily or weekly waterings) to determine physiological responses to drought and to evaluate methods to use in screening for drought resistance. Leaf water potential, stomatal resistance, plant resistance to water flow, and soil water potential were measured for three weeks on vernalized plants in a growth chamber. When water was lacking, drought-sensitive plants had a lower leaf water potential than did drought-resistant plants. With both daily and weekly waterings, stomatal resistance was higher in drought-resistant plants than in drought-sensitive plants. Plant resistance to water flow, calculated as the difference between the soil water potential and leaf water potential divided by the amount of water used by the plant, was usually higher in drought-resistant plants than in the drought-sensitive plants. The results showed that, when screening for drought resistance, stomatal resistance was a better method to use than determinations of leaf water potential or plant resistance to water flow.Journal article 3578 of the Agricultural Experiment Station, Oklahoma State University. Work was partially supported by a Grant-in-Aid of Research from Sigma Xi, The Scientific Research Society of North America, to the senior author.