Variability in the Duration of Stem Elongation in Wheat Genotypes and Sensitivity to Photoperiod and Vernalization

The stem elongation phase in wheat [Triticum aestivum (L.)] is considered critical for yield determination. A longer duration of this phase could hypothetically increase grain set and therefore yield. Genetic variation in the relative duration of the stem elongation phase having been reported, the aim was to pinpoint whether this variability was associated with sensitivity to photoperiod, vernalizing temperatures or to differences in intrinsic earliness. Pairs of cultivars identified as having different duration of the stem elongation phase (from the appearance of the first visible node to anthesis) were grown under natural (short) or extended photoperiod, with or without vernalization. Variability in the duration of this phase, in the cultivars analysed, was related to different sensitivity to photoperiod, while differences in the previous phases were related to sensitivity to both photoperiod (though different to the sensitivity of the following phase) and vernalization.     

Relationship between cold resistance,heading traits and ear primordia development of wheat cultivars

Summary For breeding early heading wheat cultivars with resistance to frost damage which are well adapted to dry areas of West Asia and North Africa, the relationships between winter hardiness, ear primordia development and heading traits, i.e. veernalization requirement, photoperiodic response and narrow-sense earliness, were assessed using a total of 30 genotypes of wheat (Triticum aestivum L.) grown in an experiment in Syria. The results of artificial freezing tests indicated that cultivars with good winter hardiness were to be found only in the winter wheat cultivars which required 50 or more days of vernalization treatment. These winter wheat cultivars did not initiate internode elongation without vernalization even at 95 days after planting. Thus their ear primordia were still underground and were protected from frost injury at this stage. Photoperiodic response and narrow-sense earliness were not associated with winter hardiness and earliness of internode elongation, but were related to the number of days to heading after planting. This indicated the possibility for breeding early heading cultivars with winter hardiness and tiller frost avoidance by combining high vernalization requirement, short narrow-sense earliness and neutral response to photoperiod.     

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.     

The physiology of variation in the time of ear emergence among wheat varieties from different regions of the world

Summary Differences in response to photoperiod and vernalization and genetic variation independent of photoperiod and vernalization (earliness per se), affecting time of ear emergence of wheat, were identified in controlled environment experiments with 33 varieties of diverse geographical origin. The results were compared with an analysis of time of ear emergence of 10409 T. aestivum accessions from the USDA Small Grain Collection grown from autumn sowings in Pendleton, Oregon, and spring sowings in Fargo, North Dakota. The effect of differences in photoperiod and vernalization sensitivity on time of ear emergence was similar to the effect of earliness per se, both under controlled environment conditions and in the field. Most of the accessions from low latitude regions reached ear emergence rapidly owing to their insensitivity to photoperiod and vernalization and earliness per se factors accelerating ear emergence. Lateness was common among accessions from Northern Europe, Afghanistan and Turkey, which was due to sensitivity to photoperiod and vernalization, and to earliness per se factors delaying ear emergence. The physiological basis of earliness per se is discussed.     

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.     

Potential for nitrogen fertilization and Hessian fly-resistance to improve Morocco's dryland wheat yields

Hessian fly (Mayetiola destructor, Say) is a perennial scourge of cereal production in the Mediterranean region, particularly in North Africa. In Morocco, it accounts for considerable yield losses of wheat (Triticum spp.), especially in the semi-arid southwestern coastal provinces. Breeding for resistance is the only feasible approach to abate its effects. Nine major Hessian fly-resistance genes have been identified in bread wheat. Two bread wheat varieties have these characteristics; the first variety with complete resistance was ‘Saada’, released to farmers in 1989, whereas a tolerant variety ‘Massira’ was released in 1994. Another widespread limiting factor for all cereals in Morocco is lack of adequate nitrogen (N). With favorable farmer acceptance, Saada became the focus of on-farm N fertilizer trials throughout the low rainfall (250-450 mm year⁻¹) zone, where it consistently out-yielded the susceptible common bread wheat, Nesma, except when no Hessian fly infestation occurred. In most cases, 40 kg N ha⁻¹ was adequate for maximum yield. Substituting resistant cultivars for Hessian fly-susceptible cultivars and increased N use could have an immediate and positive effect on wheat production in Morocco, especially in areas where the insect is endemic. The future impact will be greater when Hessian fly resistance is also transferred to other bread wheat cultivars and to durum (T. durum) wheat, the major staple food in the Mediterranean region.     

The impact of vernalization requirement,photoperiod sensitivity and earliness per se on grain protein content of bread wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

In wheat, a shorter pre-anthesis phase is often associated with increased grain protein content (GPC) but decreased grain yield. Cultivar differences in pre-anthesis development are mainly determined by vernalization requirement, photoperiod sensitivity and earliness per se. This research examines whether cultivar differences in these traits affect GPC, especially whether the three traits can partially explain genotype × environment interactions for GPC. Twenty-four winter wheat and five spring wheat cultivars selected from International Winter Wheat Performance Nursery (IWWPN) trials and 12 winter wheats tested over 2 years in Germany were characterized using the CSM-Cropsim-CERES-Wheat model. The model parameter P1V specifies the cultivar vernalization requirement, P1D the photoperiod response, and P₁₂₃ earliness per se. Covariance analyses of the IWWPN dataset indicated that about 7% of variation in GPC was explained by cultivar, with another 7% attributable to interactions of cultivar with region, site and year. P1V, P1D and P₁₂₃ all influenced GPC, but their effects varied with region, site and year. For example, for two regions, the effect of P1V on GPC decreased with latitude. Path analyses using the data from Germany confirmed that GPC increased with earlier anthesis, which was influenced by P1D and P₁₂₃. Lack of an effect of P1V at this location presumably was due to all cultivars being completely vernalized. The results indicate that efforts to improve GPC could target the three traits to specific populations of environments, which should reduce the large influence of environment on GPC.     

Daylength,Temperature and Solar Radiation Effects on the Phenology and Yield Formation of Spring Durum Wheat

Future food security will depend on crop adaptation to changing environments. We studied the limitations imposed by daylength, temperature and solar radiation on wheat yield in eight field experiments conducted at contrasting northern latitudes and involving 42 adapted spring durum wheat genotypes of divergent phenology, and reduced or without photoperiod sensitivity. Air temperatures averaged from sowing to anthesis (SA) increased from northern to southern sites, while daylength and minimum temperatures from anthesis to maturity (grain filling, GF) followed the opposite trend, due to differences in the latitude of sites. The site effect explained 96 % of the variation in the number of days SA, which was much smaller in southern sites. Average minimum daily temperatures above 6.9 °C before anthesis and below 10.8 °C during GF accompanied by photoperiods during GF of less than 14.2 h resulted in less than 14 000 kernels m^?2, which was the threshold below which kernel number limited yield. Radiation during GF lower than 1.8 kJ kernel^?1 day^?1 limited kernel weight, which was then a constraint to the achievement of yield potential.     

Photoperiod and vernalization response of wheat under controlled environment and field conditions

Characterization of large numbers of breeding lines for vernalization and photoperiod response in wheat is needed to enhance adaptation. A total of 20 wheat lines were evaluated for response to vernalization and photoperiod under two controlled environments and high ambient air temperatures under field conditions. Vernalized and non-vernalized seedlings were transplanted into pots and placed in three photoperiod (8, 12 and 16 h of light) cabinets, in the greenhouse or in growth chambers. Days to anthesis decreased with increasing photoperiod. Vernalized plants flowered earlier than non-vernalized. There was a significant correlation between days to anthesis in the greenhouse and the growth chamber (r = 0.88, P < 0.001). Basal vegetative period, effect of vernalization and photoperiod from the two screening techniques were positively correlated with each other. Growth habit, vernalization requirement and heading date in the field were highly correlated with the main effect of vernalization in the two controlled environments. The results indicate that selection for vernalization response in a large number of genotypes can be achieved under high ambient air temperatures in the field and the selected material can subsequently be screened for photoperiod response under greenhouse conditions.     

A temporal limit to the association between flag leaf life extension by fungicides and wheat yields

Three field experiments, each repeated over two or three seasons, on winter wheat investigated a possible limit to the association between grain yield and flag leaf life, as extended by fungicide application. The experiments involved up to six cultivars and different application rates, timings and frequencies of the strobilurin azoxystrobin and the triazole epoxiconazole. In the 2000/01 and 2001/02 seasons, the relationships between the thermal time to 37% green flag leaf area (m) and yield deviated from linearity. ‘Broken stick’ models were fitted to cultivar × experiment combinations within each season and the limit to the benefit to yield associated with extending flag leaf life was 700 °C days (S.E. = 20.7) and 725 °C days (S.E. = 9.33) after anthesis in 2000/01 and 2001/02, respectively. In 2002/03, the relationship between yield and m did not deviate significantly (P > 0.05) from linearity, but in this latter year the fungicide application failed to increase m past 700 °C days.     

Genetic analysis of flowering and maturity time in high latitude spring wheat

Due to the short growing season in the high northern latitudes, the development of early maturing spring wheat (Triticum aestivum L.) cultivars is important to avoid frost damage which can lower production and quality. We investigated earliness of flowering and maturity, and some associated agronomic traits, using a set of randomly selected high northern latitude adapted spring wheat cultivars (differing in maturity) and their F₁ and F₂ crosses made in a one-way diallel mating design. The parents, and their F₁ and F₂ crosses were evaluated under field conditions over 2 years. Anthesis and maturity times were controlled by both vernalization response and earliness per se genes, mainly acting additively. Non-additive genetic effects were more important in controlling grain fill duration, grain yield and plant height. Additive × additive epistatic effects were detected for all traits studied except time to anthesis. Segregation analyses of the F₂ populations for time to anthesis indicated the presence of different vernalization response genes. Molecular genetic analyses revealed the presence of Vrn-A1 and Vrn-B1 genes in the parental cultivars. Narrow-sense heritability was medium to high (60–86%) for anthesis and maturity times but low to medium (13–55%) for grain fill duration, plant height and grain yield. Selection for early flowering/maturity in early segregating generations would be expected to result in genetic improvement towards earliness in high latitude spring wheats. Incorporation of the vernalization responsive gene Vrn-B1 in combination with vernalization non-responsive gene Vrn-A1 into spring wheats would aid in the development of early maturing cultivars with high grain yield potential for the high latitude wheat growing regions of the northern hemisphere.     

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.     

Multivariate analysis of traits determining adaptation in cultivated barley

Thirty‐nine barley varieties of different origin, representing different growth types, were included in a series of experiments aimed at analysing the variability in vernalization response, photoperiod sensitivity and earliness per se and establishing the types of ecoclimatic adaptability using multivariate analysis. In the case of spring barley varieties there was no correlation between any of the three traits. For winter barleys, a negative correlation was found between photoperiod sensitivity and vernalization response and between photoperiod sensitivity and earliness per se. Vernalization response and earliness per se showed a positive correlation. Among the winter barley varieties large variations were apparent in photoperiod sensitivity, vernalization response and earliness per se, which resulted in a tremendous variation in flowering patterns and frost tolerance. Between the spring barley varieties only wider variations in photoperiod sensitivity were detected. Based on the cluster analysis, the 39 varieties could be separated into seven groups. The spring barley varieties were placed in two groups, and the winter barleys in five groups representing different adaptational types. Among these five groups two represented the two opposing extreme combinations of photoperiod sensitivity and vernalization response. The combination of large photoperiod sensitivity and no vernalization response resulted in better frost tolerance than did the combination of photoperiod insensitivity and large vernalization response.     

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.     

Effect of vernalization and photoperiod on flax flowering time

Vernalization and photoperiodism are two important physiological processes related to yield of many cool-season annual crops. The flowering response of 20 flax (Linum usitatissimum L.) genotypes to two vernalization regimes (vernalized and unvernalized) and two photoperiod treatments (10 and 14 h) was evaluated in a growth chamber study in 2010 and 2011. The results suggest that photoperiod, vernalization, and genotype all had an effect on earliness as measured by days to anthesis. Unlike flax grown in the Upper US Midwest and Canada, Texas flax is grown in the fall due to high spring and summer temperatures. Genotype interaction was observed with both vernalization and photoperiod. Specifically, flax genotypes from Texas (winter type) were sensitive to both vernalization and photoperiods for flowering. Texas genotypes delayed anthesis for 7 days or more in unvernalized seedlings, whereas flowering time of most other spring grown flax genotypes was unaffected by the vernalization treatments. Texas genotypes also delayed anthesis for 12 days or more under vernalized and short day conditions, whereas most other genotypes were not influenced by photoperiodism in vernalized seedlings. The selection for vernalization and photoperiodic sensitivity in Texas genotypes and introgression of these traits into recently adapted spring grown genotypes is needed for development of high yielding flax genotypes for southern Great Plains production areas.     

Productivity and water use efficiency of sweet sorghum as affected by soil water deficit occurring at different vegetative growth stages

The growth and production of sweet sorghum [Sorghum bicolor (L.) Moench] crops under semi-arid conditions in the Mediterranean environment of southern Italy are constrained by water stress. The effects of temporary water stress on growth and productivity of sweet sorghum were studied during three seasons at Rutigliano (Bari, Italy). The aim of this research was to evaluate the sensitivity of phenological stages subjected to the same water deficit. In a preliminary study it was observed that stomata closed when pre-dawn leaf water potential (Ψ_b) became lower than −0.4 MPa. This criterion was used in monitoring plant water status in three different plots: one never stressed and two stressed at different phenological stages (‘leaf’ and ‘stem’) when mainly leaves or stems were growing, respectively. An evaluation of the sensitivity of phenological stages subjected to identical water stress was obtained by comparing the above-ground biomass and WUE of drought crops with those of the well-irrigated crop (up to 32.5 t ha⁻¹ of dry matter and 5.7 g kg⁻¹). The sensitivity was greatest at the early stage (‘leaf’), when a temporary soil water stress reduced the biomass production by up to 30% with respect to the control and WUE was 4.8 g kg⁻¹ (average of three seasons). These results help quantify the effects of water constraints on sweet sorghum productivity. An irrigation strategy based on phenological stage sensitivity is suggested.     

Photoperiod and vernalisation response of Mediterranean wheats, and implications for adaptation

Hexaploid wheat has the largest cultivated area among crop plants due to its adaptability to different agroclimatic regions. A large part of this adaptability depends upon the variation in vernalisation and photoperiod requirements. A better understanding of the genetic control of flowering in wheat, as expressed by vernalisation requirements and photoperiod response, will guide breeders in targeting crosses of different types and will also improve our understanding of regional adaptation requirements. Characterisation of large numbers of breeding lines for photoperiod and vernalisation response in wheat is needed to assign the lines to geographic areas of most probable adaptation. Simple screening methods to quantify the effects of these two factors and their interaction are needed to assist breeding progress. Twenty wheat lines were evaluated for response to photoperiod and vernalisation under two controlled environments and under high ambient air temperatures in field conditions. Vernalised and non vernalised seedlings were transplanted into pots and placed in three photoperiod (8, 12 and 16 h light) cabinets, in the greenhouse or in growth chambers. Days to anthesis decreased with increasing length of photoperiod. Vernalised plants flowered earlier than non vernalised plants. There was a significant correlation between days to anthesis in the greenhouse and the growth chamber (r=0.88, P<0.001). Length of basal vegetative period, effects of vernalisation, and photoperiod from the two screening techniques were positively correlated with each other. Growth habit score, vernalisation requirement and heading date in the field were highly correlated with the main effect of vernalisation in the two controlled environments. The results indicated that selection for vernalisation response in a large number of genotypes can be achieved under high ambient air temperatures in the field. The selected material can subsequently be screened for photoperiod response under greenhouse conditions. Using these techniques, 49 local and improved cultivars from the Mediterranean region in west Asia and north Africa (WANA), showing differences in response to photoperiod, vernalisation, and earliness independent of vernalisation and photoperiod, affecting time to anthesis, were identified. Most old local cultivars were sensitive to both photoperiod and vernalisation. All the improved genotypes were insensitive to photoperiod. Responses to vernalisation were generally small under short photoperiods, but were more pronounced in long photoperiod, particularly in winter and facultative types from northern latitudes. These results should help to explain the adaptability of cultivars based on photoperiod and vernalisation requirements and their interaction. This revised version was published online in August 2006 with corrections to the Cover Date.     

温光互作对不同生态型小麦品种发育效应的研究 Ⅰ.品种最长、最短苗穗期及温光敏感性分析

试验在温光互作设计下,通过品种苗穗期(出苗—抽穗)对温光反应的变异分析看出,不同类型品种在不同温光组合下,不仅存在着最短与最长的苗穗期,而且不同类型品种,对温光反应类型不同,春性类型品种对春化基本无反应而对光长反应敏感;强冬性类型品种对春化反应敏感,而对光长必须在一定春化量基础上才表现敏感,并且具有短日代替春化效应。一般冬性类型品种(主要指黄淮麦区的半冬性过渡类型)多为温光兼敏性,无短日春化特性。     

Molecular examination and genotype diversity of vernalization sensitivity and photoperiod response in old and modern bread wheat cultivars grown in Iran

Understanding the genetic factors governing developmental patterns and flowering time in breeding materials is required for the development of new wheat varieties for a specific environment. Iran is among the largest wheat producers in the arid and semi-arid regions of the Middle East and North Africa. The wheat germplasm grown in Iran is either developed nationally or is introduced from the CIMMYT global wheat program. For decades, the wheat breeding program in Iran focused on generating new varieties better able to grow in the predominant Iranian climatic conditions such as humidity at the reproductive stage, high temperature during reproductive stages (terminal heat stress), moderate temperature during the cropping season, and high probability of frost damage during early stages of growth. There have also been sub-programs aimed at developing drought and salinity-tolerant wheat cultivars in Iran. Knowledge of cultivars’ growth habits in Iran is currently limited to flowering in spring-sown nurseries. We identified allelic diversity in loci involved in vernalization response (Vrn) and photoperiod sensitivity (Ppd) in 60 bread wheat cultivars developed in Iran, CIMMYT, or ICARDA. This study revealed that the spring growth habit observed in most of the cultivars is conferred by a combination of recessive vrn-A1 and dominant Vrn-D1, Vrn-B1, and/or Vrn-B3 loci. This implies that most of the cultivars have minimal vernalization requirements for overwintering. Perhaps cold winters, even in the southern regions of Iran, provide sufficient vernalization conditions for cultivars possessing the recessive vrn-A1 allele. The germplasm investigated in this study revealed no evidence indicating selection for or against any specific Vrn and Ppd allele in our wheat breeding program.     

Modeling the effect of cultivar resistance on yield losses of winter wheat in natural multiple disease conditions

The use of winter wheat (Triticum aestivum L.) cultivars resistant to diseases may make it possible to reduce yield losses without the need to use fungicides, which are expensive and may damage the environment. The cultivar resistances favored depend on the region considered and the nature of the most widespread diseases in that region. We have constructed a statistical linear model for the estimation of relative yield loss due to diseases, making it possible to assess the effects of winter wheat cultivars in various disease conditions. We considered the interactions “potential disease intensity × winter wheat cultivar” for four main fungal diseases: septoria tritici blotch (Mycosphaerella graminicola), brown rust (Puccinia triticina), yellow rust (Puccinia striiformis) and powdery mildew (Erysiphe graminis). This model can therefore be readily adapted to different regions with diverse combinations of these diseases. The potential intensity of each disease in each trial was calculated based on the symptoms observed on susceptible cultivars not treated with fungicide. The cultivar effect was characterized by the ratings of cultivar susceptibility to each disease and by cultivar earliness. The parameters of the model were estimated from 276 wheat cultivar trials carried out over 12 years (1991–2002) in the major wheat-growing areas of France. This model can help to choose from the cultivars resistant to several diseases (but very rarely resistant to all the diseases), those cultivars best adapted to a given environment, according to the hierarchy of potential disease intensities encountered.