Classification of Global Environments and Cultivars of Spring Barley Based on Heading Time Interactions

Global heading time data collected by the International Center for Maize and Wheat Improvement (CIMMYT) during the International Barley Yield Trials were used to assess similarity of environments, variation among cultivars, and genotype × environment interactions. Data for 29 spring barley cultivars grown in 89 environments over three years were analyzed by cluster analysis. The deviation m days to heading of the mean of 28 photoperiod sensitive cultivars from the cultivar ‘Mona’ and the mean of ‘Mona’, homozygous recessive for the ea_k gene conferring photoperiod insensitivity and thermal stability, were used as environmental coordinates. In addition, diversity of heading time responses among genotypes was illustrated by differences among overall means and patterns of deviation for days to heading from ‘Mona’ in selected environments. Three main clusters were identified. Mexican environments were similar to warmer Mediterranean, eastern and southern African, West Asian, and Latin American environments. Heading time responses in Syria were similar to those observed m other cool Mediterranean environments. Early heading cultivars exhibited greater variation for heading response, especially in extremely warm-and equaiorial- short daylength environments, com-pared to late heading cultivars, presumably because of larger photothermal × genotype interactions. Photoperiod flux about the winter solstice appeared to be a major environmental cue for heading time in photoperiod sensitive spring barleys.     

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.     

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.     

杂交水稻恢复系桂99的抽穗期基因及其效应分析

以基因型明确的抽穗期主基因近等基因系EG0~EG7、ER~LR、T65系列为测验系（TLs）,在江西南昌（28^o 36’ N）夏季自然高温长日（14 h/d）和人工遮光短日（10 h/d）,以及海南三亚（18^o 14’ N）旱季自然低温短日（11.6 h/d）处理条件下,对籼型杂交水稻恢复系桂99的抽穗期基因及其感温性和基因位点间的互作效应进行了分析。结果表明,桂99在E₁、E₂和E₃位点分别带有感光迟熟等位基因E₁、E₂和E₃,在Se-1位点带有非感光等位基因Se-1^e,在Ef-1位点带有早熟基因Ef-1,由此推断其抽穗期基因型为E₁E₁E₂E₂E₃E₃Se-1^eSe-1^eEf-1Ef-1。迟熟基因E₁、E₃与早熟基因Ef-1同时存在,E₁、E₃与Se-1^e基因位点间的互作使桂99农艺性状表现弱感光性。感光基因Se-1^u（或Se-1ⁿ）的存在能增强E位点感光迟熟基因的感光性,感光基因对“TLs×桂99”F₂植株抽穗的影响,是延长平均抽穗期,增加迟抽穗植株分布频率。分析了感温性对TLs抽穗期的影响,讨论了以桂99为恢复系配置的杂交水稻组合丰产性和广适性的遗传基础。     

Analysis of genes controlling photoperiod sensitivity in common bean using DNA markers

The genetic basis of the photoperiod response in common bean (Phaseolus vulgaris L.) was investigated using DNA markers and recombinant inbred populations. Two loci affecting photoperiod response were resolved, the previously defined primary locus (Ppd), at which the dominant allele confers sensitivity to photoperiod, and a second locus (herein defined as Hr), which influences the degree to which a plant responds to photoperiod. The DNA marker P5₁₆₀₀ cosegregated with the recessive allele, ppd, displaying a recombination frequency with the photoperiod locus of about 3%. A second marker, B303₆₀₀, was linked to the recessive allele at Hr and mapped approximately 13 cM from this locus. The markers demonstrated that in crosses involving Redkloud and several photoperiod sensitive lines, insensitivity to photoperiod is primarily controlled by ppd and that Hr does not significantly affect flowering time in ppd/ppd plants under the environmental conditions used. This revised version was published online in July 2006 with corrections to the Cover Date.     

Nuclear Mutations Affecting Chloroplastic Pigments of Photoperiod-insensitive Barley

Three photoperiod-sensitive spring barley cultivars (Hordeum vulgare L.) and three independently derived, single-gene, nuclear mutants expressing photoperiod insensitivity and extremely early heading time under short daylengths were investigated for chloroplastic pigment variation in three environments using reverse-phase high-performance liquid chromatography (HPLC) to account for differences in laminae colour. In a greenhouse, non-stress environment and in a full sunlight, high-temperature stress environment, no systematic differences were observed among pigments of the mutant-parent pairs. However, under 12 h of daily light (600 μE m ²sec^?1) and stress temperatures (20/10°C, night/day) in a growth chamber, the three mutants appeared similar to each other in chlorotic appearance and laminae pigment contents, but differed from the three non -chlorotic parents, which were similar to each other. The mutants had less chlorophyll a and b, β-carotene, lutein, taraxanthin, violaxanthin, and neoxanthin but more zeaxanthin than their parents. When shaded, the mutants became less chlorotic. How phenotypic differences for heading time and perception of day-length might be related to altered chloroplast contents remains unclear. The single-gene mutants conferring photopenod insensitivity were more sensitive to photothermal stress than their photoperiod-sensitive counterparts and as a result, their zeaxanthin content increased but the level of other pigments decreased.     

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.     

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.     

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.     

Photoperiod insensitivity gene essential to the varieties grown in the northern limit region of paddy rice (Oryza sativa L.) cultivation

Summary The release of extremely early maturing varieties has made it possible to cultivate rice in Hokkaido (NL45-42°) in Japan, the northern limit region of paddy rice cultivation. Until then, rice cultivation in this region has been impracticable due to the climate condition, especially short summer and long-day more than 15 hours during summer. Experiment results confirmed that the success of rice cultivation in this area depends on raising photoperiod insensitivity varieties with short basic vegetative growth period. Moreover, in this study, the genetic factors controlling the photoperiod insensitivity of Hokkaido varieties were analyzed by using 8 kinds of tester lines for three loci, E1, E2, and E3, controlling photoperiod sensitivity. It was found out that all the varieties examined carry el, a photoperiod insensitivity allele of E1 locus, but as for the other loci, E2 and E3, the existence of plural alleles were recognized. We have already clarified that almost all the japonica-type varieties grown in Japan (except Hokkaido) and Taiwan carry E1 bringing about strong photoperiod sensitivity. Accordingly, it can be concluded that el is the gene essential to Hokkaido varieties: the interchange of E1 with e1 enabled rice cultivation under long-day condition.     

Genetic analyses of low photoperiod sensitivity of rice cultivars from the northernmost regions of Japan

The genetic factors controlling the extremely low photoperiod sensitivity (PS) of rice culyibstd from the northernmost regions of Japan were studied using 26 cultivars from the Hokkaido district (42 N 45 N). They were grown under 10-h and 24-h daylengths, and their PS. expressed by the difference between the days to heading under these daylengths, was estimated. The genetic factors responsible for the intervarietal difference in PS were then analysed under 10-h, 14-h and 24-h daylengths using the progenies from crosses between Hokkaido cultivars. These results showed that a recessive gene, se-9(t), drastically reduced PS under a 24-h daylength and that several genes, including se9(t), determined the extremely low PS of the cultivars from the northernmost regions.     

Inheritance of temperature sensitivity of the photoperiod response in common bean (Phaseolus vulgaris L.)

Summary Photoperiod response of flowering in common bean (Phaseolus vulgaris L.) is thought to be controlled by the genes Ppd and Hr. However, cultivars also vary in the degree that cooler temperatures reduces their sensitivity to photoperiod. To examine the inheritance of this temperature sensitivity, crosses of cvs. Gordo x de Celaya and Flor de Mayo × Rojo 70 were evaluated at two sites differing in mean temperature and using 12.5-h natural photoperiod or 18-h artificially extended photoperiod. Under 18-h photoperiod at the warmer site, Palmira, no plants of the parents or of the F₂ populations flowered, confirming that the parents were sensitive to photoperiod. Under 12.5-h photoperiod at the cooler site, Popayan, the parents for each cross flowered at similar dates and no segregation for days to flower was observed. However, under 18-h photoperiod, de Celaya and Rojo 70 and the F₁ populations did not flower within 100 days after planting, while the F₂ and F₃ populations showed segregation that was consistent with single gene inheritance, late flowering being dominant. Late flowering at Popayan under 18-h photoperiod indicates a lack of temperature sensitivity, so temperature insensitivity of the photoperiod response was dominant to sensitivity. The name Tip, for temperature insensitivity of photoperiod response, is proposed for this gene, with the recessive form of this gene conditioning earlier flowering at cooler temperatures with long daylengths. It is recognized that the observed segregation patterns could represent the effect of multiple alleles at the Ppd or Hr loci, and studies are proposed to test this possibility with molecular markers and recombinant inbred lines.     

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.     

Genetic Control of Vernalization, Day-length Response, and Earliness per se by Homoeologous Group-3 Chromosomes in Wheat

To identify homoeologous group-3 chromosomes that carry genes for vernalization, day-length responses, and earliness per se, a series of aneuploid lines (mono-somics and tetrasomics) and chromosome-substitution lines in ‘Chinese Spring’ (CS) were surveyed under different vernalization and day-length regimes in controlled environments. The results indicated that genes on all three chromosomes of group 3 can have striking effects on ear-emergence time. The replacement of CS 3B by its homologues in ‘Lutescens 62’ and ‘Cheyenne’ produced an increased insensitivity to vernalization, while 3B homologues from ‘Ceska Presivka’ gave CS a remarkable sensitivity to vernalization. This provided evidence for multiple allelism at a new Vrn locus on chromosome 3B. A negative association between gene dosage and day-length response was found in CS 3D which was thought to carry a gene for promoting insensitivity to day-length. The behaviour of CS monosomic 3A and CS (Timstein 3A), in reducing numbers of days to heading independently of environmental stimuli, suggested the presence of earliness per se genes on this chromosome.     

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.     

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.     

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.     

Identification of quantitative trait loci for flowering-related traits in the D genome of synthetic hexaploid wheat lines

The gene pool of Aegilops tauschii, the D-genome donor of common wheat (Triticum aestivum L.), can be easily accessed in wheat breeding, but remains largely unexplored. In our previous studies, many synthetic hexaploid wheat lines were produced through interspecific crosses between the tetraploid wheat cultivar Langdon and various A. tauschii accessions. The synthetic hexaploid wheat lines showed wide variation in many characteristics. To elucidate the genetic basis of variation in flowering-related traits, we analyzed quantitative trait loci (QTL) affecting time to heading, flowering and maturity, and the grain-filling period using four different F₂ populations of synthetic hexaploid wheat lines. In total, 10 QTLs located on six D-genome chromosomes (all except 4D) were detected for the analyzed traits. The QTL on 1DL controlling heading time appeared to correspond to a flowering time QTL, previously considered to be an ortholog of Eps-A ^m 1 which is related to the narrow-sense earliness in einkorn wheat. The 5D QTL for heading time might be a novel locus associated with wheat flowering, while the 2DS QTL appears to be an allelic variant of the photoperiod response locus Ppd-D1. Some of the identified QTLs seemed to be novel loci regulating wheat flowering and maturation, including a QTL controlling the grain filling period on chromosome 3D. The exercise demonstrates that synthetic wheat lines can be useful for the identification of new, agriculturally important loci that can be transferred to, and used for the modification of flowering and grain maturation in hexaploid wheat.     

Effects of loci determining photoperiod sensitivity (Ppd-H1) and vernalization response (Sh2) on agronomic traits in the 'Dicktoo'×'Morex' barley mapping population

The objectives of this research were to determine the individual and interaction effects of the Ppd-H1 and Sh2 loci on agronomic traits under short- and long-photoperiod regimes. Nineteen doubled haploid (DH) lines from the ‘Dicktoo’בMorex’ mapping population, which represented the four genotypes at the Ppd-H1 and Sh2 loci, were pheno-typed in controlled environment photoperiods. Both Ppd-H1 and Sh2 had significant effects on several agronomic traits, in addition to their role in determining first node appearance and flowering time. The magnitude of these effects depended on daylight. Under long-day conditions (18 h) Ppd-H1, and under short-day conditions (12 h) Sh2 was a significant determinant of most characters. The interactions between these two loci were significant for several characters, particularly for yield components, under both long- and short-photoperiod regimes. Under the long-day treatment, Ppd-H1 influenced plant height through the determination of node number. There was an epistatic association between the two loci for both 1000-kernel weight and tillering. The combination of photoperiod insensitivity and vernalization requirement caused a significant increase in tillering. This was paralleled by a decrease in 1000-kernel weight. Under the long-day treatment, neither Ppd-H1 nor Sh2 influenced plant yield. Under short-day conditions, the combination of photoperiod insensitivity and vernalization requirement had a pronounced negative effect on plant yield.     

Associations between 23 Quantitative Traits and 10 Genetic Markers in a Barley Cross

Associations of 23 quantitative traits and 10 genetic marker characters were examined in 63 chromosome-doubled lines (DH-lines) derived from the F₁- generation of a cross between an old and a modern spring barley variety. One fourth of the marker × trait combinations showed significant associations. More than two thirds of these associations were to earliness (heading date). Earliness was found to be controlled by two loci: the previously known eak locus and a new locus designated Ea. It is concluded that the associations of quantitative traits with the earliness loci were caused by pleiotropy. Associations were found between two absolutely linked C-bands on chromosome 3 and QTLs (quantitative trait loci) for lodging, straw diameter, and length of top internode of the straw. The three loci on chromosome 5, eak and the two linked powdery mildew loci, Mla9 and Mlk, were associated with a possible QTL for magnesium concentration in grain. Association to the C-band on chromosome 6 suggests QTLs for TGW (thousand grain weight), straw diameter and magnesium concentration in grain. Locus Estl on chromosome 3 was not associated with any of the quantitative traits.