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 influence of photoperiod on the Vrn-H2 locus (4H) which is a major determinant of plant development and reproductive fitness traits in a facultative × winter barley (Hordeum vulgare L.) mapping population

To determine the effect of Vrn‐H2 locus on plant developmental and agronomic traits, detailed controlled environment tests involving a factorial set of vernalization and photoperiod treatments were carried out using doubled haploid lines developed from a facultative (Vrn‐H2^?) × winter (Vrn‐H2⁺) barley cross. The allele phase at the Vrn‐H2 locus influenced heading date as well as the developmental and agronomic traits. The performance of Vrn‐H2⁺ lines was significantly influenced by vernalization: reproductive fitness traits showed significant decreases without vernalization. However, the effects of alleles at the Vrn‐H2 locus extended beyond simple satisfaction of the vernalization requirement. Vrn‐H2⁺ lines showed increased reproductive fitness compared with Vrn‐H2^? lines when vernalization was followed by a long photoperiod. The responses of the two Vrn‐H2 allele classes to photoperiod duration were quite different in terms of heading date, developmental and agronomic traits. These results suggest that alleles at the Vrn‐H2 locus – and/or tightly linked gene(s) – respond primarily to the exogenous signal of vernalization (temperature), but when the vernalization requirement has been fulfilled, they also respond to photoperiod duration.     

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.     

Effect of added barley chromosomes on the flowering time of new wheat/winter barley addition lines in various environments

The heading characters and morphological traits of two partial sets of wheat–barley disomic addition lines, namely Mv9kr1/Igri and Asakaze/Manas, were evaluated under controlled environmental conditions in a phytotron under long-day, short-day and non-vernalised conditions and in field-sown experiments. The winter barley chromosome additions significantly influenced the flowering time of wheat both in the controlled environment test and under field-sown conditions. Of all the barley addition lines, the effect of the 4H and 7H additions was the most characteristic. The 7H addition lines were the earliest in both cultivar combinations in each treatment. In the Mv9kr1/Igri combination the 4H addition was the latest under all the environmental conditions. In the Asakaze/Manas combination 4H addition was the latest under short-day and long-day illumination in the phytotron but the 6H addition was the latest without vernalisation and in the field in 2012. There was 12 and 11 days difference between the flowering times of the 7H and 4H Mv9kr1/Igri and Asakaze/Manas addition lines in the field in 2012, which increased to 52 and 44 days under short-day illumination in the phytotron. In the winter wheat background, the addition of 2H carrying the photoperiod sensitivity gene Ppd-H1 decreased the flowering time under the short photoperiod regime, but had a very strong delaying effect under field-grown conditions. Considering the yield components under field conditions, 4H was the most fertile of the addition lines, while 7H showed the highest tillering capacity, and Igri 3H had good tillering capacity and the highest number of seeds per plant.     

Effects of Temperature and Photoperiod on Days to Flowering, Yield and Yield Components of Lupinus albus (L.) under Field Conditions

A better understanding of the agronomic importance of planting date and the influence of cold temperatures and photoperiod during germination and plant growth may lead to better management strategies for cultivation of the sweet white lupin (Lupinus albus). The effects of planting date (temperature and photoperiod) were determined on the number of days to flowering, yield and yield components of four early to medium and one late sweet white lupin genotype in a field trial at Potchefstroom, South Africa, planted during February 1996 to January 1997. Moisture stress was avoided through regular irrigation. Duration of the developmental phases planting date to emergence, emergence to floral initiation, initiation to first flower, duration of flower and days to physiological and harvest maturity was related to field measurements of temperature and photoperiod. Differences in the main determinants of yield, i.e. seeds per pod, pods per plant, single seed mass (SSM), plant and pod height and yield, were measured. Results showed that both temperature and photoperiod influence the growth and development of the Lupinus albus genotypes ‘Esta’, ‘Hantie’, ‘Tifwhite’, ‘Kiev’ and ‘LAL 186’. Temperature influences include the effect of vernalization at seedling emergence. Minimum grass temperatures under 5 °C at emergence are effective for vernalization. However, after grass temperatures at emergence increased again from June to December, to gether with an increase in the photoperiod length, ‘Tifwhite’ as well as the other genotypes still flowered earlier, confirming that these cultivars are long‐day plants, which is in accordance with controlled‐environment data. Cool vernalizing temperatures thus not only influence obligate vernalization requiring genotypes such as ‘Tifwhite’, but also influence the non‐obligate genotypes studied. Plan‐ting date had a significant influence on pods per plant, single seed mass (SSM) and seed yield. In all trials laterplanting, from June to November, decreased SSM and seed yield. The highest seed yield of 1.5 t ha^?1 was obtained for the 10 June planting date and the lowest average yield of 0.450 t ha^?1 for the 5 November planting date.     

Effect of Temperature and Photoperiod on the Development of Lupinus albus L. in a Controlled Environment

An experiment to determine the effect of temperature and photoperiod on Lupinus albus under controlled environmental conditions was carried out, using the three Lupinus albus genotypes 'Tifwhite', 'Esta' and 'Kiev', and three temperature (10/20, 18/28 and 20 °C continuously) and two photoperiod (8 and 16 h daylength) regimes, in all combinations. Half of the seeds were vernalized for 21 days at 4 °C to alleviate the obligate vernalization requirement of Tifwhite. Although Esta and Kiev do not have obligate vernalization requirements, they were influenced by this vernalization period. Observations included the duration of the period from planting to seedling emergence, the duration of the period from planting to the beginning of flowering and the duration of flowering. The vernalization treatment accelerated plant development in all genotypes. The period from planting to emergence was shorter under the higher temperature regime. For all genotypes, the period from planting to flowering was shorter under the longer photoperiod, the same trend as would be expected for long-day plants. Duration of flowering periods were, in contrast to pre-flowering periods, shorter for all genotypes at cooler temperatures. The results of this study confirm that photoperiod does contribute to the growth period from planting to flowering in L. albus and that this species does behave as a long-day plant.     

不同光温条件谷子光温互作模式研究及SiCCT基因表达分析

光周期和温度是影响作物生长发育、生态适应性和产量的2个重要环境因素,揭示光温互作对作物生长发育的效应及其分子机制对育种实践和理论研究具有重要意义。本研究设置长日照高温、长日照低温、短日照高温、短日照低温4个光温处理,调查‘黄毛谷’抽穗期、株高、叶片数和穗长。结果表明,光周期对谷子的发育起关键作用,温度的改变不影响长日照比短日照延迟谷子生殖生长的效应,温度的作用随光周期的不同而异,短日照条件下,高温缩短谷子营养生长期而低温延长营养生长期,长日照条件下则相反;对谷子生殖生长的促迚作用是短日照高温短日照低温长日照低温长日照高温。利用RT-PCR技术从‘黄毛谷’叶片兊隆了一个CCT结构域基因(SiCCT),该基因编码286个氨基酸,属于CMF亚家族成员,基于CCT域基因氨基酸序列的系统迚化分析,谷子与高粱、玉米亲缘关系较近。实时荧光定量PCR分析发现, SiCCT基因在‘黄毛谷’叶片中高表达,其次为幼穗和叶鞘;长日照、短日照处理SiCCT基因均表现24h昼夜节律性特点,短日照七叶期表达水平最高,八叶期(抽穗)及穗后表达迅速降低,长日照七叶至十叶期‘黄毛谷’处于营养生长期,SiCCT基因维持较高表达水平;无论高温低温,长日照条件下SiCCT基因在各叶期表达量整体高于短日照处理,长日照条件下低温处理SiCCT基因的相对表达量明显低于高温处理, SiCCT基因的总体表达量与‘黄毛谷’营养生长期存在正相关。总之SiCCT基因受光周期调控,同时也受温度调控,因而推测SiCCT基因参与了光周期途径和感温性途径,并通过二者互作调控谷子营养生长和生殖生长的全过程。     

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.     

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.     

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.     

绵麦系列小麦品种（系）的农艺性状和品质性状分析

为利用好小麦资源材料,对19份绵麦系列小麦品种（系）的农艺性状和品质性状等15个性状进行分析。结果表明,农艺性状与品质性状间有着复杂的相关关系,有13对性状表现为显著或极显著相关,有23对性状表现为显著或极显著偏相关。产量与8个农艺性状的关联系数的序次为：基本苗>千粒重>穗粒数>容重>株高>最高苗>生育期>有效穗;产量与6个品质性状的关联系数的序次为：稳定时间>最大阻力>籽粒蛋白>面积>吸水率>湿面筋;蛋白质与9个农艺性状的关联系数的序次为：最高苗>有效穗>基本苗>穗粒数>产量>容重>千粒重>株高>生育期;稳定时间与9个农艺性状的关联系数的序次为：容重>最高苗>产量>生育期>有效穗>千粒重>株高>基本苗>穗粒数;面积与9个农艺性状的关联系数的序次为：基本苗>穗粒数>千粒重>产量>株高>最高苗>有效穗>生育期>容重。供试材料从农艺性状和品质性状可被聚为三类,第Ⅰ_2、Ⅰ_3、Ⅰ₄类育种时可以作为高产材料加以应用,同时Ⅰ₃和Ⅰ₄类群育种时可以作矮杆材料加以运用;Ⅰ₁和Ⅰ₄类群育种时可以作为优质品质材料加以运用。第Ⅱ类育种时可从粒重方面加以应用;第Ⅲ类从优质矮杆材料加以应用。     

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.     

Development of photoperiod-sensitive cytoplasmic male sterile (PCMS) wheat lines showing high male sterility under long-day conditions and high seed fertility under short-day conditions

A “two-line system” using photoperiod-sensitive cytoplasmic male sterility (PCMS) caused by Aegilops crassa cytoplasm under long-day photoperiods (≧15 h) has been proposed as a means of producing hybrid varieties in common wheat (Triticum aestivum). The PCMS line is maintained by self-pollination under short-day conditions, and hybrid seeds can be produced through outcrossing of the PCMS line with a pollinator line under long-day conditions. Our previous studies revealed that PCMS lines showing complete male sterility under long-day conditions are necessary for practical hybrid wheat breeding, especially to obtain high hybrid purity in F₁ seeds. Furthermore, practical PCMS lines should have high seed fertility under short-day conditions, which is associated with female fertility. Wheat cv. Norin 26 with Ae. crassa cytoplasm exhibits high seed fertility under short-day conditions, and cv. Fujimikomugi with Ae. crassa cytoplasm shows high male sterility under long-day conditions. Here we developed practical PCMS lines derived from the F₁ generation of Norin 26 and Fujimikomugi (with Ae. crassa cytoplasm) that were then backcrossed to elite wheat lines.     

Independent selection for seed free tryptophan content and vernalization response in chickpea domestication

Chickpea shows a distinct domestication trajectory vis‐a‐vis pod dehiscence and growth cycle mediated by vernalization insensitivity compared with its companion Near Eastern legumes. Our objectives were: (i) to map the quantitative trait loci (QTLs) associated with vernalization response and seed free tryptophan in domesticated × wild chickpea progeny and (ii) estimate the genetic correlation between vernalization response and free tryptophan content. A domesticated × wild chickpea cross was used to document phenotypic segregation in both traits and to construct a skeletal genetic map for QTL detection. A number of vernalization response and seed free tryptophan content QTLs were documented in both F₂ and F₃ generations. No significant genetic correlation between these two traits was observed. Epistatic relationship between two free tryptophan loci was documented. It is evident that selection for high seed tryptophan is easier to accomplish relative to selection for vernalization insensitivity. This suggests that the two traits were selected independently in antiquity, thereby corroborating earlier claims for conscious selection processes associated with chickpea domestication.     

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.     

Pleiotropic effects of the ea7 photoperiod response gene on the morphology and agronomic traits in barley

The photoperiod‐insensitive barley mutant ‘Atsel’, carrying the recessive gene ea₇, was studied together with the donor variety ‘Atlas’ (wild‐type, Ea₇) under different daylengths with the aim of analysing pleiotropic effects. Grown under long and short photoperiods ‘Atsel’ flowered about 10 days and 34 days, respectively, earlier than ‘Atlas’. The significantly shorter life‐cycle of the photoperiod‐insensitive mutant resulted in several changes of plant morphology. Tillering, plant height, number of leaves and number of internodes were reduced. A lower number of florets per main spike was observed for ‘Atsel’, but only in the long photoperiod experiment. Finally, photoperiod insensitivity combined with a lower grain yield per plant was most pronounced under long‐day treatment. The data are comparable with results obtained from single chromosome recombinant lines of wheat that have differences in their photoperiod response caused by the genes Ppd1 or Ppd2.     

玉米赤霉烯酮与冬小麦的生长与发育

实验采用酶联免疫法分析了冬小麦（燕大1817）（Triticum aestivum L.Yan Da 1817）在不同条件（低温、短日或长日）下内源玉米赤霉烯酮（Zearalenone, 以下简称ZEN）的动态变化,并通过在不同时期外施ZEN探讨外源ZEN对冬小麦生长发育的影响。结果表明,ZEN可能是冬小麦生长与发育的重要调节物质之一。短日诱导可以代替低温春     

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.     

小麦主要农艺性状的相关性及聚类分析

为了解小麦产量与农艺性状间的关系。采用变异性分析、相关性分析和聚类分析,对18份小麦新品系的产量及各农艺性状间的关系进行统计分析。结果表明,不同小麦品系的产量与其农艺性状间存在显著差异,株高和千粒重变异系数较大,产量与茎蘖数呈极显著正相关,有效穗数与产量、茎蘖数呈显著正相关,穗粒数与千粒重、株高呈负相关。聚类分析结果显示,18份小麦品系可划分为4个类群,依据各类群的特点提出了不同的改良目标。在小麦育种选配亲本时,应注意亲本间农艺性状的相关性和协调性。     

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.