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.     

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.     

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.     

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.     

A reciprocal F1 monosomic analysis of the genetic control of time of ear emergence,number of leaves and number of spikelets in wheat (Triticum aestivum L.)

Summary A reciprocal F₁ monosomic analysis of chromosomal differences between Spica and Bersée was carried out under controlled environment conditions. Chromosomes associated with differences in days to ear emergence, number of leaves and number of spikelets were identified. The results indicated that chromosome 2B of Spica carries a photoperiod insensitivity allele at the Ppd ₂ locus. Both Spica and Bersée appear to have a vernalization insensitity allele at the Vrn ₂ locus on chromosome 5B. On chromosome 3A, 4B, 4D and 6B factors were found with major effects on earliness per se, diffeences in ear emergence and number of spikelets which were independent of photoperiod and vernalization. The possibility that these factors influence growth rate is discussed.     

Genetic control of flowering time and its component processes in subterranean clover (Trifolium subterraneum L.)

Summary The inheritance of flowering time and its component processes, vernalization and photoperiod response, were studied in two crosses of subterranean clover (Trifolium subterraneum L.) using a field sowing and four controlled environment sowings with different combinations of vernalization and photoperiod. Time to flowering was under polygenic control and was highly heritable. For both vernalization and photoperiod response, there was dominance for a low response, or earliness. A simple genetic control was indicated for photoperiod response. The results for vernalization response were not clear cut, although the character appeared to be under polygenic control. An interaction between vernalization and photoperiod response was evident in three of the four cultivars studied. This made it impossible to separate the effect of these two component processes and complicated the study of their inheritance. Node of first flower on the main stem was closely related to flowering time and its use led to similar conclusions in the inheritance studies.     

The influence of flowering time genes on environmental adaptability in European wheats

Summary In order to obtain high levels of environmental adaptability in wheat varieties it is essential they flower at times appropriate to particular environmental conditions. The influence of three distinct genetic systems that together determine time of flowering is reviewed here.Vernalization genes are seen to be particularly important to winter wheats for their direct or indirect effects on winter hardiness. Vernalization genes play a minor role in determining flowering time in autumn sown winter wheats but insensitivity is essential if spring sown wheats are to flower.Day length sensitive photoperiod genes play a major role in determining flowering time and adaptability of autumn sown wheats. Insensitivity can promote yield advantages of over 35% in Southern European environments. 15% in Central Europe and offers benefits even in the UK. At present only a single allele of Ppd1 appears to have been introduced into commercial European wheat varieties. The merits of alternative Ppd1 alleles or different loci are discussed.The influence of earliness per se genes that determine flowering time independently of environmental stimuli is less well documented than the effect of photoperiod and vernalization genes. It is likely that genes on chromosomes belonging to groups 2, 3, 4, 6 and 7 may act to modify flowering time independently of environmental stimuli probably by determining numbers of vegetative and floral primordia being initiated or the rate of initiation of the primordia. Earliness per se genes appear to be widespread in European wheats and play a significant role in determining the exact time plants flower.     

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.     

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.     

Characterization of QEet.ocs-5A.1, a quantitative trait locus for ear emergence time on wheat chromosome 5AL

QEet.ocs‐5A.1, a quantitative trait locus controlling ear emergence time, has been detected on wheat chromosome 5AL using single chromosome recombinant lines (SCRs) developed from a cross between ‘Chinese Spring’ (CS) (‘Cappelle‐Desprez’ 5A) and CS (Triticum spelta 5A). This locus has little influence on grain yield and its components, and thus has breeding potential for changing ear emergence time without yield reduction. To characterize the phenotypic expression of QEet.ocs.1 and to test its interaction with the Vrn‐A1 gene for vernalization response, six near‐isogenic SCRs differing for these two gene regions were grown together with the parental controls under different vernalization and photoperiod regimes. The T. spelta allele of QEet.ocs.1 accelerated heading time when vernalization and photoperiod were satisfied, demonstrating that the function of this QTL is earliness per se. There was no interaction between Vrn‐A1 and QEet.ocs.1.     

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。     

Validation of the two-gene epistatic model for vernalization response in a winter × spring barley cross

A two gene epistatic model in which a dominant “winter growth habit” allele at Vrn-H2 encodes a repressor with a corresponding binding site in a recessive vrn-H1 allele explains the vernalization response phenotypes in an array of barley germplasm. In order to validate the model genetically, we developed an F ₂ population (and F ₂-derived F ₃ families) from the cross of Hardy (winter) × Jubilant (spring). Using gene-specific primers, we determined the Vrn-H1 and Vrn-H2 allele architecture of each F ₂ plant and we measured the growth habit phenotype of each F ₂ plant via phenotyping of its F ₃ progeny under controlled environment conditions. We used a set of treatments involving plus/minus vernalization under long photoperiod and vernalization under short photoperiod. Alleles at the two loci showed expected patterns of segregation and independent assortment. Under long day conditions, the two Vrn genes were the primary determinants of heading date, regardless of the vernalization treatment. Under short photoperiod, the effects of these loci were not significant. There was incomplete dominance at Vrn-H1: heterozygotes were significantly later to head than Vrn-H1Vrn-H1 genotypes. Vrn-H2 genotypes were also significantly later to head, even when plants were vernalized. These results validate the two-gene epistatic model for vernalization response under long-day conditions. The results under short photoperiod, and the variance in flowering with vernalization, confirm that while the two Vrn genes are the primary determinants of vernalization response, they are part of a larger interactome that determines the timing of the vegetative to reproductive transition.     

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.     

Genetic control of photoperiodic sensivity and maturity in spring wheat within narrow limits of adaptation

Summary Photoperiodic respose, as assessed by a regression technique, exhibited complete dominance averaged over the crosses of an eight parent diallel in the vernalized condition. Photoperiodic response as final leaf number for the vernalized 8-hour photoperiod diallel was closely related to photoeriodic response of the regression method. However, the diallel analyses of both sets of data showed little agreement in terms of respectieve array positions.The inheritance of photoperiodic response in diallels using regression values showed little agreement between the vernalized and unvernalized conditions. This difference was postulated to be due to interaction of vernalization and photoperiodic response in the unvernalized situation. In the unvernalized condition photoperiodic response exhibited non-allelic interaction, attributable mainly to the cultivar Pinnacle in general behaviour in its crosses. Its removal gave a situation of high average dominance for photoperiodic response with a clear indication that high photoperiodic sensitivity was dominant to comparative insensitivity.Days to ear emergence (vernalized and 18-hour photoperiod) exhibited non-allelic interaction in its expression, due mainly to the general behaviour of the cultivar Pinnacle in its crosses. Removal of its array gave a situation of a moderately strong degree of overdominance in the expression of days to ear emergence. Maturity differences amongst parents and F₁'s, vernalized and under 18-hour photoperiod, are postulated to be due to a factor other than vernalization or photoperiodic response beheved to be growth temperature in differentially in fluencing growth and/or developmental rates between genotypes.     

Detection of an earliness per se quantitative trait locus in the proximal region of wheat chromosome 5AL

A homoeologous quantitative trait locus to that of eps5L on barley chromosome 5H was identified in a syntenic region of wheat chromosome 5A. Wheat single chromosome recombinant lines (SCRs) were developed from a cross between ‘Chinese Spring’(‘Cappelle-Desprez’ 5A) and ‘Chinese Spring’(Triticum spelta 5A), these were grown together with the parental controls under different vernalization and photoperiod regimes. The variation for ear emergence time accelerated heading induced by the T. spelta segment indicated an effect associated with the Xcdo412-Xbcd9 interval. Since no differences between the SCRs and controls in responses to vernalization and photoperiod treatments were detected, this effect was identified as an earliness per se gene, Q Eetocs-5 A.2, which may be homoeologous to the eps5L quantitative trait locus of barley. Xbcd926 has been found to be closely linked to the rice flowering time quantitative trait loci, QHd9a or FLTQ2, on chromosome 9, suggesting possible relationships among the quantitative trait loci across wheat, barley and rice genomes.     

Temperature as a component of the expression of developmental responses in wheat

Summary Studies were made of days to ear emergence under the constant temperatures of 9, 14, 19 and 25°C and 16 h photoperiod in three sets of wheat lines each possessing genotypes differing for developmental responses.Days to ear emergence in three near-isogenic lines of the wheat cultivar Triple Dirk, which differed for vernalization response, increased as the strength of the response increased. At the four temperatures Triple Dirk D (Vrn 1 vrn 2) was not significantly different from normal Triple Dirk (Vrn 1 Vrn 2) but Triple Dirk B (vrn 1 Vrn 2) was significantly (P=0.01) later than normal Triple Dirk at each temperature. This indicates that the vrn 1 allele confers stronger vernalization response than vrn 2 over the range of temperatures (9–24°C). However, Triple Dirk C (vrn 1 vrn 2) failed to head after 120 days at each temperature indicating strong interaction between vrn 1 and vrn 2 with each other (and possibly the Triple Dirk back-ground) to give a much stronger vernalization response than predictions from additivity of their individual effects.The second set comprised the four Chinese Spring/Thatcher chromosome substitution lines CS/T 3B, 6B, 7B and 5D, plus Chinese Spring and Thatcher, and were grown in the unvernalized condition. CS/T 5D was similar in days to ear emergence as Chinese Spring at all four temperatures but the other three lines were earlier to ear emergence, particularly as the temperature increased. Days to ear emergence was fastest at 14°C in all lines, except CS/T 3B, in which it decreased progressively from 9 to 24°C.The third set of Chinese Spring and Thatcher and the homoeologous group 2 chromosomes of Thatcher substituted in Chinese Spring, the group which is considered to be involved in the control of photoperiod sensitivity. The three substitution lines responded differently to temperature compared with Chinese Spring and with each other, with chromosome 2D being the least, and chromosome 2B the most, responsive to temperature.     

Effect of alien 5R(5A) chromosome substitution on ear-emergence time and winter hardiness in wheat-rye substitution lines

Ear emergence time and response to vernalization were investigated in 12 alien substitution lines in which a pair of chromosomes 5A of recipient spring wheat cultivars was replaced by a pair of chromosomes 5R of Siberian spring rye ‘Onokhoiskaya’. The recipients were 12 spring cultivars of common wheat, each carrying different Vrn genes. Spring rye ‘Onokhoiskaya’ had the Sp1 (now called Vrn-R1) gene for spring growth habit located on chromosome 5R, but its expression was weaker. The Vrn-R1 gene had no effect on growth habit, ear emergence time and response to vernalization in wheat-rye substitution lines. Ears emerged significantly later in the 5R(5A) alien substitution lines than in the recipient wheat cultivars with the Vrn-A1/Vrn-B1/vrn-D1 or Vrn-A1/vrn-B1/Vrn-D1 genotypes. No difference in ear emergence time was found between most of the 5R(5A) alien substitution lines and the cultivars carrying the recessive vrn-A1 gene. The presence of the Vrn2a and Vrn2b alleles at the Vrn2 (now called Vrn-B1) locus located on wheat chromosome 5B was confirmed.The replacement of chromosome 5A by chromosome 5R in wheat cultivars ‘Rang’ and ‘Mironovskaya Krupnozernaya’, which carries the single dominant gene Vrn-A1, converted them to winter growth habit. In field studies near Novosibirsk the winter hardiness of 5R(5A) wheat–rye substitution lines of ‘Rang’ and ‘Mironovskaya Krupnozernaya’ was increased by 20–47% and 27–34%, respectively, over the recurrent parents.     

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.     

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.     

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.