Estimation of Crown Temperature of Winter Wheat and the Effect on Simulation of Frost Tolerance

Accurate estimation of winter wheat frost kill in cold‐temperate agricultural regions is limited by lack of data on soil temperature at wheat crown depth, which determines winter survival. We compared the ability of four models of differing complexity to predict observed soil temperature at 2 cm depth during two winter seasons (2013‐14 and 2014‐15) at Ultuna, Sweden, and at 1 cm depth at Ilseng and Ås, Norway. Predicted and observed soil temperature at 2 cm depth was then used in FROSTOL model simulations of the frost tolerance of winter wheat at Ultuna. Compared with the observed soil temperature at 2 cm depth, soil temperature was better predicted by detailed models than simpler models for both seasons at Ultuna. The LT50 (temperature at which 50 % of plants die) predictions from FROSTOL model simulations using input from the most detailed soil temperature model agreed better with LT50 FROSTOL outputs from observed soil temperature than what LT50 FROSTOL predictions using temperature from simpler models did. These results highlight the need for simpler temperature prediction tools to be further improved when used to evaluate winter wheat frost kill.     

Evidence of a major genetic factor conditioning freezing sensitivity in winter wheat

The inheritance of freezing tolerance in F₂‐derived F₄ populations from all possible crosses of winter wheat cultivars ‘Kestrel’, ‘Eltan’, ‘Tiber’, ‘Froid’ and germplasm line Oregon Feed Wheat #5 (ORFW) was investigated. When frozen to a temperature equal to the LT₅₀ of the least freezing tolerant parent (ORFW), survival frequency distributions were skewed to greater survival in six of the 10 crosses, however, very few of the progeny from the four crosses to ORFW survived. The inheritance of this freezing sensitivity was investigated with freezing of F_2:4 populations from the crosses of ORFW to ‘Eltan’ or ‘Tiber’ to the LT₅₀ of the hardier parent. Very few of the F_2:4 populations survived as well as ‘Eltan’ or ‘Tiber’, indicating a small number of strongly dominant genetic factors in ORFW that conditioned freezing sensitivity. Molecular analysis indicated these factors were not spring‐type vernalization alleles.     

Heritable improvement of frost tolerance in winter wheat by in vitro-selection of hydroxyproline-resistant proline overproducing mutants

In a previous study (Dörffling et al., J. Plant Physiol. 142, 222–225, 1993) in vitro-selection and regeneration of hydroxyproline (Hyp)-resistant lines of winter wheat (Triticum aestivum cv. Jo 3063) with increased frost tolerance and increased proline contents was reported. In this study the heritability of these traits was investigated. The F ₁progenies derived from the pollination of regenerated plants with pollen from wild type plants developed higher mean frost tolerance (lower LT ₅₀values) and higher proline levels compared with in vitro-controls and wild type plants. In the F ₂generation, which was obtained by self-pollination, segregation of the Hyp-lines in a 3:1 phenotypic ratio with regard to the traits frost tolerance and proline content was observed. Subsequent selection of extremes in the F ₂yielded homozygous plants in the F ₃generation. Lower LT ₅₀values and increased proline contents in those F ₃plants were significantly correlated. Furthermore, improved frost tolerance of one of the selected mutants was also observed in studies with whole plants in the F ₄generation. These results, which are similar to results of parallel studies on winter barley, provide strong evidence for the heritability of the traits 'increased frost tolerance' and 'increased proline content' obtained by this in vitro-selection procedure. The mutation seems to be due to a single incompletely dominant gene. A comparison of frost tolerance values from F ₃Hyp-progenies with those of seven standard varieties differing in frost tolerance indicates a considerable improvement of frost tolerance by the described in vitro-selection method.     

Use of chlorophyll fluorescence to evaluate the cold acclimation and freezing tolerance of winter and spring oats

The efficiency of the excitation capture by open Photosystem II (PSII) reaction centres was measured by the F_v/F_m ratios in a collection of winter and spring oats in order to assess the effects of hardening and freezing on the functionality of PSII and also the suitability of a chlorophyll fluorescence‐based method to screen oat cultivars for frost tolerance. A significant reversible decrease in F_v/F_m was found in all genotypes during acclimation to low, non‐freezing temperatures. F_v/F_m analysis appears to be an attractive test for the evaluation of frost tolerance in oats, being rapid, non‐invasive and capable of monitoring a trait related to a crucial stage in the acquisition of frost tolerance. It is more sensitive and precise than other standard methods and highly correlated with field‐evaluated frost damage. The measurements made during recovery 1 or 2 days after stress when the visual symptoms are not yet expressed, were especially advantageous because of the large variability in genotype response. The r‐values (close to 0.8) were reduced due to the non‐standard behaviour of the winter cultivar ‘Aintree’. The cold acclimation response of this genotype has been analysed in detail and the limits of artificial freezing tests are discussed.     

Combining frost tolerance,high grain yield and good pasta quality in durum wheat

Growing in Central Europe winter instead of spring durum wheat would substantially increase yield potential but is currently hampered by the lack of knowledge of frost tolerance present in elite material. The objectives of our survey were to (i) study the genetic variability and heritability of frost tolerance and its association with other important agronomic and quality traits in durum wheat, (ii) examine the potential to combine frost tolerance with high quality and high grain yield and (iii) investigate the consequences of the heritabilities and associations among traits on the optimum design of a multistage selection programme for winter durum wheat. We investigated 101 elite winter durum wheat lines and four commercial checks in field trials at four locations. Four agronomic as well as nine quality traits were recorded. In addition, frost tolerance was evaluated using a semi‐controlled test resulting in high‐quality phenotypic data. Genotypic variances (σ²_G) were significantly larger than zero for all traits, and heritabilities were moderate to high. Several elite durum wheat lines exhibited a frost tolerance comparable to that of two frost‐tolerant Triticum aestivum varieties. Frost tolerance was not negatively associated with other important agronomic and quality traits. The high quality of the phenotypic data for frost tolerance evaluated in a semi‐controlled test suggests that this is a cost‐efficient approach to consider frost tolerance at early stages of a multistage durum wheat breeding programme.     

Genetic control of frost tolerance in wheat (Triticum aestivum L.)

Summary The frost tolerance of winter wheat is one component of winter hardiness. If seedlings are frost resistant, it means that they can survive the frost effect without any considerable damage. To study the genetic control of frost tolerance, an artificial freezing test was used. Frost tolerance is controlled by an additive-dominance system. The results of diallel analyses indicate the importance of both additive and non-additive gene action in the inheritance of this character. The dominant genes act in the direction of lower frost tolerance and the recessive genes in the direction of a higher level of frost tolerance. The results of monosomic and substitution analyses show that at least 10 of the 21 pairs of chromosomes are involved in the control of frost tolerance and winter hardiness. Chromosomes 5A and 5D have been implicated most frequently. The geneFr1 (Frost 1) was located on the long arm of chromosome 5A. Crosses between cultivars, chromosome manipulation and the induction of somaclonal variation may be suitable methods for broadening the gene pool for frost tolerance.     

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.     

Studies for assessing the influence of hardening on cold tolerance of barley genotypes

Summary Frost tolerance of 30 barley (Hordeum vulgare L.) cultivars have been field evaluated in North Italy during the 1990/1991 winter season that was characterized by exceptionally low temperatures without snow cover. The results showed a significant correlation between cold injury and grain yield loss (r=0.61^**). Five cultivars chosen for their varying degree of frost tolerance were further evaluated using laboratory tests. Measurements of survival rate and membrane damage were used to assess the influence of hardening on frost resistance. The reliability of the tests is shown by the high correlation to the field data. For both the laboratory temperature regimes and field conditions, the tested cultivars showed the same order of classification. The effect of a rise in temperature at the end of the hardening treatment on frost tolerance is also reported. The laboratory tests here proposed can be integrated in a breeding programme for improving frost tolerance in barley.     

Drought priming effects on alleviating the photosynthetic limitations of wheat cultivars (Triticum aestivum L.) with contrasting tolerance to abiotic stresses

Abiotic stress tolerance in plants is said to be induced by pre-stress events (priming) during the vegetative phase. We aimed to test whether drought priming could improve the heat and drought tolerance in wheat cultivars. Two wheat cultivars “Gladius” and “Paragon” were grown in a fully controlled gravimetric platform and subjected to either no stress or two drought cycles during the tillering stage. At anthesis, both batches were either subjected to high temperature stress, drought stress or kept as control. No alleviation of grain yield reduction due to priming was observed. Higher CO₂ assimilation rates were achieved due to priming under drought stress. Yield results showed that priming was not damage cumulative to wheat. Priming was responsible to alleviated biochemical photosynthetic limitations under drought stress and sustained photochemical utilization under heat stress in “Paragon.” Priming as a strategy in abiotic stress alleviation was better evidenced in the stress susceptible cultivar “Paragon” than tolerant cultivar “Gladius”; therefore, the type of response to priming appears to be cultivar dependable, and thus phenotypical variation should be expected when studying the effects of abiotic priming.     

Genetic variation for shoot elongation before winter and its correlation with vernalization requirement in winter oilseed rape (<Emphasis Type="Italic">Brassica napus</Emphasis> L.)

Changing climatic conditions in north-western Europe are accompanied by occasional extreme weather conditions. This requires breeding of winter oilseed rape cultivars which are resilient to diverse abiotic stress factors, e.g. frost, drought and heat. The degree of vernalization requirement of winter oilseed rape has been found to be related to frost tolerance and winter hardiness. Shoot elongation before winter in particular has been identified as one decisive factor for frost tolerance in winter oilseed rape. However, the relationship between vernalization requirement and shoot elongation before winter is not known. In the present study the genetic variation for shoot elongation before winter and vernalization requirement of 19 genetic diverse breeding lines and cultivars were analyzed. Autumn and spring sown field experiments in multiple environments were performed to determine shoot elongation before winter and vernalization requirement, respectively. In spring sown field experiments, genotypes with a low vernalization requirement were characterized by the occurrence of long bolting plants with flower buds. Large and significant genotypic variation was found for shoot length in the autumn sown and spring sown environment. Broad sense heritability was quite high for shoot length in the spring sown environment (h²?=?97%), whereas it was only of medium size for shoot length before winter (h²?=?62%). Although the correlation between shoot length before winter and shoot length in the spring sown environment was positive (Spearman’s rank r_S?=?0.48*), a number of genotypes with reduced shoot elongation before winter and low vernalization requirement were identified. Results indicate that genotypes with a reduced shoot elongation before winter independent of their vernalization requirement can be selected in breeding programs.     

Improved frost tolerance and winter hardiness in proline overaccumulating winter wheat mutants obtained by in vitro-selection is associated with increased carbohydrate,soluble protein and abscisic acid (ABA) levels

In previous studies in vitro-selection of proline overaccumulating lines of winter wheat (Triticum sativum L. cv. Jo 3063) with increased frost tolerance was reported. These traits were found to be genetically stable. In the present study the improvement of frost tolerance (winter hardiness) under field conditions is confirmed for F₇ progenies of the mutants. Moreover, the mutants accumulated higher levels of glucose and fructose, soluble protein and abscisic acid (ABA) in addition to proline than the wild type under cold hardening conditions in a growth chamber as well as under cold hardening field conditions. ABA and proline levels peaked when the temperature decreased, whereas carbohydrate levels increased more slowly at decreasing temperature. Soluble protein levels also increased during cold hardening, but in addition showed sharp declines during frost periods. Increased carbohydrate levels of the mutants were associated with lower osmotic potential values. The differences in carbohydrate, protein and ABA levels between the mutants and the wild type are probably due to pleiotropic effects of the mutation.     

Improved frost tolerance and winter survival in winter barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.) by <Emphasis Type="Italic">in vitro</Emphasis> selection of proline overaccumulating lines

Embryogenic calli derived from anther cultures of the two-rowed winter barley cultivar Igri were plated on solid L3 medium containing the proline analogue hydroxyproline (Hyp), 10–20 mmol l^–1. Exposure to Hyp caused severe degeneration of most of the calli. Hyp resistant calli, distinguishable by their lighter colour and higher growth rate, and control calli not exposed to Hyp were plated on L3 regeneration medium. From 22,500 anthers exposed to Hyp 46 Hyp resistant regenerates were obtained, which were transferred to soil. After cultivation for 5–10 weeks at normal growth conditions they were cold hardened at 2 C under short day conditions together with control regenerates. Frost tolerance assays with segments of fully grown leaves of unhardened and cold hardened plants revealed that Hyp resistant regenerants were significantly more frost tolerant than the control regenerants. Improved frost tolerance was found also in the progenies R₁ to R₉, and genotypic segregation in the R₁ generation in a 1:2:1 ratio was indicated. Increased proline content was observed in the R₂ generation and in subsequent generations and was significantly (P 0.001) correlated with increased frost tolerance in the Hyp lines. Comparative studies of R₉ progenies from homozygous R₂ plants with the wild type Igri under field conditions in winter at three locations in Europe as well as crossing experiments confirmed the heritable improvement of frost tolerance and winter survival, respectively, in the Hyp lines. The results support the hypothesis that proline accumulation in cold acclimated winter barley plants is causally related to the acquisition of frost tolerance. Moreover, the described biotechnological procedure may be applicable in breeding programs for improved winter hardiness and possibly also for other stress tolerances.     

Adaptability and phenotypic stability of Lolium perenne L. cultivars of diverse origin grown at the margin of the species distribution

In northern countries, Lolium perenne L. generally survives poorly when grown inland and north of 60°N because of extensive winter damage. With the projected future climate change, it could become a promising option for improving production efficiency of the agricultural sector in these regions. Here, we compare the biomass production potential of cultivars of diverse origin across five locations stretching from Estonia to Iceland over a period of three harvest years, and their freezing tolerance under artificial conditions. The aim was to relate the observed pattern of adaptation to the geographic origin of the cultivars and their response to prevailing agroclimatic conditions. Significant interactions were observed between cultivars and test environments (locations × years), and significant interactions between cultivars and years were detected at four of the five locations. Models of joint regression, additive main effects and multiplicative interaction (AMMI) and factorial regression using several agroclimatic indices showed that cultivars developed in northern countries showed greater yield potential across the test environments and were, thus, generally better adapted than cultivars from Central Europe. Diploid cultivars were more frost tolerant than tetraploid cultivars giving them an advantage in locations which were characterized by low temperatures during the hardening period in autumn and mild and rainy winters, such as at the Icelandic location. Only a few cultivars showed general adaptability to the environmental conditions at the test sites, the most stable cultivar being an admixture of diploids and tetraploids. In future breeding, the best strategy would be to hybridize cultivars developed in northern countries with more exotic materials that combine high yield potential, adequate winter survival and superior disease resistance under northern conditions.     

Overwintering of winter cereals in Hungary in the case of global warming

Summary Under phytotronic conditions investigations were made on the effect of important environmental factors, such as temperature, water and an increasing concentration of atmospheric CO₂, on the hardening of young cereal plants.In all the varieties derived from the major wheat growing regions of the world the hardening process was favourably influenced by a doubling of atmospheric CO₂ content, so that a significantly larger number of plants survived the frost test than for plants of the same variety raised under normal conditions.A reduction in freezing temperature and an increase in soil moisture content caused a slight reduction in survival % for varieties with excellent frost resistance and a great reduction for those with medium or poor frost resistance.Predictions suggest that in Central Europe, as the result of global climatic changes, there will be a reduction in the quantity of winter precipitation, a considerable rise in winter temperatures and an increase in atmospheric CO₂ concentration. Judging by the experimental results, these changes could improve the overwintering of winter cereals; at the same time, however, a number of factors (mainly the reduction of precipitation) leading to yield losses must be expected during the vegetation period.     

Breeding winter hardy grasses

Summary Perennial grasses are vital for Norwegian agricultural production. The nature and extent of winter damage on grasslands is highly dependent on climatic conditions, and determines both persistency and yield. Physical stresses such as frost and ice encasement predominate in coastal regions with an unstable winter climate, while biotic stresses such as low temperature fungi are more common in the inland regions. Development of hardening depends on plant adaptation and climatic conditions during autumn and winter. New winter-hardy cultivars should be bred for wide adaptation to winter stresses. The genetic background for the most important character, freezing tolerance, seems to be of polygenic nature with mainly additive gene action. Selection for increased freezing tolerance has been effective over generations in grasses, and in most grass species ample variation still exists to be exploited by breeding. However, in some species like perennial ryegrass, modern biotechnological methods should be used to improve freezing tolerance and winter hardiness.     

Association between frost tolerance and the alleles of high molecular weight glutenin subunits present in Polish winter wheats

Subunits of high molecular weight glutenins strongly influence wheat bread making quality and can be associated with important agronomic traits. Polish winter wheats show a significant quantitative dominance of the null allele over the coding alleles of the Glu-A1 locus. To identify the causes of such skewed distribution, 116 F₅ lines obtained from six cross combinations were analyzed for their HMW glutenin subunits and 11 agronomic characteristics, such as plant height and uniformity, leaf blotch and leaf rust resistance, grain yield per plot, number of grains per ear, grain yield per ear, 1000 kernel weight, frost tolerance, total protein content and the SDS-sedimentation value. The SDS-sedimentation value, resistance to leaf blotch and frost tolerance showed statistically significant associations with the status of the Glu-A1 locus. It appears that chromosome 1A with the null allele at Glu-A1 carries a closely linked locus responsible for frost tolerance. With early strong selection for winter hardiness, the null allele of Glu-A1 becomes fixed in advanced breeding materials despite its strong negative impact on the end use quality.     

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.     

Genetic improvement trends in agronomic performances and end-use quality characteristics among hard red winter wheat cultivars in Nebraska

Evaluation of wheat cultivars from different eras allows breeders to determine changes in agronomic and end-use quality characteristics associated with grain yield and end-use quality improvement over time. The objective of this research was to examine the trends in agronomic and end-use quality characteristics of hard red winter wheat cultivars grown in Nebraska. Thirty historically important and popular hard red winter wheat cultivars introduced or released between 1874 and 2000 were evaluated at Lincoln, Mead and North Platte, Nebraska in 2002 and 2003. An alpha lattice design with 15 incomplete blocks of two plots and three replications was used at all locations. Agronomic (days to flowering, plant height, spike length, culm length, grain yield and yield components, and grain volume weight) and end-use quality (flour yield, SDS-sedimentation value, flour protein content, and mixograph time and tolerance) traits were measured in each environment. Highly significant differences were observed among environments, genotypes and their interactions for most agronomic and end-use quality characteristics. Unlike modern cultivars, older cultivars were low yielding, and less responsive to favorable environments for grain yield and yield components. Semidwarf cultivars were more stable for plant height than traditional medium to tall cultivars. All cultivars had high grain volume weight since it is part of the grading system and highly selected for in cultivar release. Modern cultivars were less stable than older cultivars for SDS-sedimentation and mixing tolerance. However, the stability of older cultivars was attributed to their having weak mixing tolerance and reduced SDS-sedimentation values. The reduced protein content of modern cultivars was offset by increased functionality, as measured by mixograph and SDS sedimentation. In conclusion, breeders have tailored agronomic and end-use quality traits essential for hard red winter wheat production and marketing in Nebraska.     

不同类型冬小麦品种拔节后幼穗低温敏感期的鉴定

摘要：以百农矮抗58、豫麦49－198、豫农949和偃展4110这4个小麦品种为材料,利用人工气候室处理,研究了冬小麦拔节期幼穗抗霜冻力随幼穗分化进程的变化规律。在药隔期前后对4个品种进行了抗霜冻力鉴定,进而分析了不同品种、冬春性以及幼穗发育进程等对冬小麦抗霜冻力的影响。结果表明：4个品种中百农矮抗58抗冻能力较强,其他几个品种抗冻能力相对较弱。随幼穗发育进程的推进,幼穗抗霜冻力总体呈下降趋势, 从雌雄蕊末期发育进入药隔期,幼穗的抗霜冻力骤然下降,雌雄蕊分化期之前以及药隔期之后,抗霜力的下降趋势比较平缓,表明雌雄蕊至药隔期这个阶段是冬小麦幼穗分化的低温敏感期。品种的抗霜性研究适宜在该时期展开。     

System analysis of frost resistance in winter wheat and its use in breeding

Morphophysiological and physicochemical parameters of sets of winter wheat genotypes were measured. High correlations were found between survival percentage of the winter wheat check variety under freezing and electroconductivity tests, stem electrical resistance after electric shock, and intensity of delayed leaf fluorescence at low temperatures. These correlations decreased when the number of varieties was increased to 35-100. Complex attribute of frost resistance was divided into large components using factor analysis. Biological interpretation was achieved for the first six components. Quantitative values for each component of frost resistance in the various genotypes were determined. The determination coefficient of multiple regression equations, where function was a level of frost resistance and arguments were the component's value for each genotype, varied from 0.80-0.94 over 10 years. It was proposed that the received equations might serve as a model for frost resistance and indicate which component of frost resistance determines the survival of plants in particular ecological conditions. The component structure concept of frost resistance is suggested and possible applications of this concept are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.