A practical protocol to accelerate the breeding process of rice in semitropical and tropical regions

Breeding of excellent rice varieties is essential for modern rice production. Typical breeding procedures to introduce and maintain valuable agricultural traits require at least 8 generations from crossing to stabilization, always taking more than 4–5 years of work. This long and tedious process is the rate-limiting step in the development of new varieties, and therefore fast culturing methods are in urgent need. Taking advantage of early flowering characteristics of light-sensitive rice under short-day conditions, we have developed a practical protocol to accelerate the breeding cycle of rice, which we have termed the “1 + 2”, “2 + 2”, “1 + 3”, and “0 + 5” methods according to the different rice varieties and different breeding purposes. We have also incorporated several techniques, including glume cutting, seed desiccation at 50°C in a drier seed dormancy breakage with low concentration of HNO₃, and direct seeding. Using the above strategy, we have shortened the life cycle of light-sensitive rice varieties to about 70 days, making it possible for several rice cultivars to proliferate 4–5 generations in a single calendar year. This protocol greatly accelerates the process of new variety breeding, and can be used in rice research for shortening the process of genetic analysis and the construction of mapping populations.     

DNA marker for resistance to Puccinia horiana in chrysanthemum (Chrysanthemum morifolium Ramat.) “Southern Pegasus”

White rust caused by Puccinia horiana Henn. adversely affects chrysanthemum (Chrysanthemum morifolium Ramat.) production. The breeding of resistant varieties is effective in controlling the disease. Here we aimed to develop DNA markers for the strong resistance to P. horiana. We conducted a linkage analysis based on the genome-wide association study (GWAS) method. We employed a biparental population for the GWAS, wherein the single nucleotide polymorphism (SNP) allele frequency could be predicted. The population was derived from crosses between a strong resistant “Southern Pegasus” and a susceptible line. The GWAS used simplex and double-simplex SNP markers selected out of SNP candidates mined from ddRAD-Seq data of an F₁ biparental population. These F₁ individuals segregated in a 1:1 ratio of resistant to susceptible. Twenty-one simplex SNPs were significantly associated with P. horiana resistance in “Southern Pegasus” and generated one linkage group. These results show the presence of a single resistance gene in “Southern Pegasus”. We identified the nearest SNP marker located 2.2 cM from P. horiana resistance locus and demonstrated this SNP marker-resistance link using an independent population. This is the first report of an effective DNA marker linked to a gene for P. horiana resistance in chrysanthemum.     

Distribution of two groups of melon landraces and inter-group hybridization enhanced genetic diversity in Vietnam

To understand the genetic diversity and differentiation of Vietnamese melon (Cucumis melo L.), we collected 64 landraces from the central and southern parts of the country and assessed molecular polymorphism using simple sequence repeat and random amplified polymorphic DNA markers. The Vietnamese melon was divided into seven cultivar groups, namely “Dua le”, “Dua vang”, “Dua bo”, “Dua gang-andromonoecious”, “Dua gang-monoecious”, “Dua thom”, “Montok”, and the weedy-type melon “Dua dai”. Among these, Dua le, Dua vang, Dua bo, and Dua gang-andromonoecious are cultivated on plains and they formed cluster II along with the reference accessions of Conomon and Makuwa. Based on genetic distance, Dua le and Dua vang were regarded as Makuwa and Dua bo and Dua gang-andromonoecious as Conomon. In contrast, Dua thom and Montok are cultivated in highlands, and they formed cluster III along with landraces from the southern and eastern foot of the Himalayas. Dua gang-monoecious which is commonly cultivated in the southern parts of Vietnam, exhibited the greatest genetic diversity, as explained by its possible origin through the hybridization between Dua gang-andromonoecious and Montok. Genetic differences in melon landraces between plains and highlands and hybridization between these two geographical groups have contributed to the enhancement of genetic diversity in Vietnamese melon.     

Evaluation of soybean germplasm conserved in NIAS genebank and development of mini core collections

Genetic variation and population structure among 1603 soybean accessions, consisted of 832 Japanese landraces, 109 old and 57 recent Japanese varieties, 341 landrace from 16 Asian countries and 264 wild soybean accessions, were characterized using 191 SNP markers. Although gene diversity of Japanese soybean germplasm was slight lower than that of exotic soybean germplasm, population differentiation and clustering analyses indicated clear genetic differentiation among Japanese cultivated soybeans, exotic cultivated soybeans and wild soybeans. Nine hundred ninety eight Japanese accessions were separated to a certain extent into groups corresponding to their agro-morphologic characteristics such as photosensitivity and seed characteristics rather than their geographical origin. Based on the assessment of the SNP markers and several agro-morphologic traits, accessions that retain gene diversity of the whole collection were selected to develop several soybean sets of different sizes using an heuristic approach; a minimum of 12 accessions can represent the observed gene diversity; a mini-core collection of 96 accession can represent a major proportion of both geographic origin and agro-morphologic trait variation. These selected sets of germplasm will provide an effective platform for enhancing soybean diversity studies and assist in finding novel traits for crop improvement.     

Molecular and genealogical analysis of grain dormancy in Japanese wheat varieties,with specific focus on MOTHER OF FT AND TFL1 on chromosome 3A

In the wheat (Triticum aestivum L.) cultivar ‘Zenkoujikomugi’, a single nucleotide polymorphism (SNP) in the promoter of MOTHER OF FT AND TFL1 on chromosome 3A (MFT-3A) causes an increase in the level of gene expression, resulting in strong grain dormancy. We used a DNA marker to detect the ‘Zenkoujikomugi’-type (Zen-type) SNP and examined the genotype of MFT-3A in Japanese wheat varieties, and we found that 169 of 324 varieties carry the Zen-type SNP. In Japanese commercial varieties, the frequency of the Zen-type SNP was remarkably high in the southern part of Japan, but low in the northern part. To examine the relationship between MFT-3A genotype and grain dormancy, we performed a germination assay in three wheat-growing seasons. On average, the varieties carrying the Zen-type SNP showed stronger grain dormancy than the varieties carrying the non-Zen-type SNP. Among commercial cultivars, ‘Iwainodaichi’ (Kyushu), ‘Junreikomugi’ (Kinki-Chugoku-Shikoku), ‘Kinuhime’ (Kanto-Tokai), ‘Nebarigoshi’ (Tohoku-Hokuriku), and ‘Kitamoe’ (Hokkaido) showed the strongest grain dormancy in each geographical group, and all these varieties, except for ‘Kitamoe’, were found to carry the Zen-type SNP. In recent years, the number of varieties carrying the Zen-type SNP has increased in the Tohoku-Hokuriku region, but not in the Hokkaido region.     

蚂蚱麦和小白麦衍生系的遗传多样性分析

陕西关中蚂蚱麦和山西平遥小白麦是我国北方小麦品种的原始骨干亲本,解析蚂蚱麦和小白麦及其衍生系的遗传多样性对于小麦品种改良具有重要的参考意义。本研究利用小麦660KSNP芯片对蚂蚱麦、小白麦及其衍生品种(系)进行全基因组扫描,分析其遗传多样性。结果表明,小麦3个基因组的多态性SNP标记数为BAD,第4同源群的多态性标记数最少, 149份供试材料基因多样性(H)范围为0.095～0.500,平均值为0.336;核苷酸多样性指数(π)范围为0.272～0.435,平均值为0.340;而遗传相似系数(GS)变幅为0.335～0.997,平均值达0.619,表明蚂蚱麦和小白麦衍生系的遗传多样性较低。聚类分析表明蚂蚱麦和小白麦紧密地聚在亚群I,其衍生品种(系)分为5个亚群,其中2000年以前以蚂蚱麦或小白麦的单一衍生系为主,分在亚群I、II、III, 2000年以后多数品种同时拥有蚂蚱麦和小白麦血缘,分在亚群IV、V,遗传多样性较高,且与大面积推广品种聚为一类。因此,应加强优异基因资源导入,拓宽小麦品种的遗传基础,最终提高育种水平。     

新疆冬小麦地方品种与育成品种基于SNP芯片的遗传多样性分析

新疆是我国西北重要的小麦优势产区和消费区。解析新疆冬小麦地方品种与育成品种之间的遗传多样性和亲缘关系,对新疆冬小麦育种中杂交组合的合理选配以及后代选择具有重要的指导意义。本研究利用小麦55K SNP(single nucleotide polymorphism)芯片对134份新疆冬小麦地方品种及54份育成品种进行全基因组扫描,估算其品种间的遗传距离,揭示其遗传多样性。结果表明,所有SNP位点的多态性比率达到95.62%(50,743/53,063)。每条染色体分布1068～2616个多态性位点,多态性标记在基因组间分布呈现ABD。188个品种间的两两遗传距离在0.002～0.723之间,平均为0.378。其中134个地方品种两两之间的遗传距离在0.002～0.400之间,平均为0.070; 54个育成品种两两之间的遗传距离在0.004～0.337之间,平均为0.114; 134个地方品种与54个育成品种之间的遗传距离在0.605～0.723之间,平均为0.699。聚类结果显示可将所有材料分为10个不同类群。综合SNP和系谱分析,育成品种与地方品种之间的遗传差异最大,其次是育成品种之间,而地方品种之间遗传差异最小。鉴于育成与地方品种之间较大的遗传差异,新疆冬小麦品种可以利用地方种来丰富其育种的种质基础,拓宽遗传背景,进而提高当地小麦育种水平。本研究为新疆冬小麦品种选育和改良提供了重要的理论指导。     

Spanish spelt: a separate gene pool within the spelt germplasm

The renewed interest in spelt (Triticum spelta L.) for wheat improvement programmes requires the study of the available genetic diversity. The purpose of this study was to assess genetic diversity within a Spanish spelt collection. Sixty‐six Spanish spelt accessions, 19 accessions of T. spelta and T. macha from different origins, three bread wheat cultivars (T. aestivum) and one accession of T. dicoccum were screened using simple sequence repeats (SSRs). The diversity observed within the Spanish group was comparable with that observed in the other wheat varieties, despite their broader geographical diversity. Indeed, the highest polymorphic information content value calculated with SSRs for Spanish material (0.90) is similar to that observed for the other wheat varieties (0.98). Principal component analysis explained 46.5% of the cumulative variation and confirmed the Spanish accessions as a separate group. This study showed the Spanish spelt collection to be a variable and unique genetic resource for wheat and spelt breeding programmes.     

“十五”以来我国杂交小麦审定品种分析

对“十五”以来我国杂交小麦审定品种的产量构成和品质特性等进行了汇总和分析。结果表明,“十五”以来,我国共审定杂交小麦品种22个,“十二五”以来审定速度明显加快,北方冬麦区是我国杂交小麦品种审定主要区域;审定品种产量水平总体呈上升趋势,北方冬麦区杂交小麦品种生产试验的产量平均每年增长76.0kg/hm²,增幅为1.25%,审定品种综合品质亦不断提升;杂交小麦审定品种产量与库容量呈正相关关系,穗粒数和千粒重潜力的挖掘与提升对产量增长的贡献最为明显,但是有效穗数并未明显提升,因此进一步提高成穗数,实现库容量扩充,对提高杂交小麦产量潜力和加快杂交小麦品种审定具有重要意义。     

小麦茎秆断裂强度相关性状QTL的连锁和关联分析

小麦茎秆断裂强度与倒伏特性关系密切,并对产量有很大影响。本研究旨在解析茎秆断裂强度的遗传机制,开发与该性状紧密连锁/关联的分子标记。利用山农01-35′藁城9411重组自交系(RIL)群体(含173个F8:9株系)和由205个品种(系)构成的自然群体,借助90 k小麦SNP基因芯片、DArT芯片及传统分子标记技术,在2个环境中对两群体的茎秆断裂强度相关性状进行连锁分析和全基因组关联分析。利用已构建的高密度连锁图谱,在4B染色体的TDURUM_CONTIG63670_287–IACX557和EX_C101685–RAC875_C27536等区段上,检测到9个控制小麦茎秆断裂强度、株高、茎秆第2节间充实度、茎秆第2节壁厚相关性状的加性QTL,可解释表型变异9.40%~36.30%。同时,利用包含24 355个SNP位点的复合遗传图谱,在自然群体中检测到37个与茎秆断裂强度相关性状(P0.0001)的标记,分别位于1A、1B、2B、2D、3A、3B、4A、4B、5A、5B、5D、6B、7A、7B和7D染色体,可解释表型变异7.76%~36.36%。在4B染色体上,以连锁分析检测到控制茎秆断裂强度的RAC875_C27536与关联分析检测到的Tdurum_contig4974_355标记,在复合遗传图谱上的距离为6.7 cM,说明该区段存在控制小麦茎秆断裂强度的重要基因。     

Genetic diversity and population structure of ‘Khao Kai Noi’, a Lao rice (Oryza sativa L.) landrace,revealed by microsatellite DNA markers

Rice (Oryza sativa L.) is the main food for people in Laos, where it has been grown and eaten since prehistory. Diverse landraces are grown in Laos. ‘Khao Kai Noi’, a landrace favored for its eating quality, is held in the nationwide collection of traditional landraces in the Lao national genebank. Genetic diversity is crucial for sustainable use of genetic resources and conservation. To investigate the genetic diversity of ‘Khao Kai Noi’ for conservation, we genotyped 70 accessions by using 23 polymorphic simple sequence repeat markers. The markers generated 2 to 17 alleles (132 in total), with an average of 5.7 per locus. The total expected heterozygosity over all ‘Khao Kai Noi’ accessions was 0.271. Genetic variation was largest among accessions and smallest within accessions. Khao Kai Noi accessions were classified into three different genetic backgrounds, but there was unclear association between the three inferred population and name subgroups and geographical distribution. Most of the accessions were clustered with temperate japonica and showed genetic relatedness to rice from neighboring provinces of Vietnam, suggesting a Vietnamese origin. The results of this study will contribute to the conservation, core collection and future breeding of the Khao Kai Noi population.     

利用SSR标记分析野生小豆及其近缘野生植物的遗传多样性

利用28对SSR引物对96份野生小豆资源、小豆近缘野生植物及栽培小豆品种进行遗传多样性分析,共检测到255个等位变异,平均每对SSR引物9.10个,多态信息含量的变异范围从0.374到0.865,平均为0.722。野生小豆材料和近缘植物Vigna minima遗传变异丰富。来自不同地域的野生小豆材料具有大量特异等位变异,基于非加权成组配对算术平均法的聚类分析可将不同地理来源的野生小豆单独分群,且与主坐标分析的结果相一致。4份栽培小豆材料与日本野生小豆遗传距离较近,表明目前国内小豆育种中较多使用了含有日本血缘的小豆材料,以及国内野生小豆资源的搜集和利用工作落后于日本。本研究对国内野生小豆资源的搜集和保存具有指导意义,并可以为这些资源的评价、利用和优异基因的发掘提供参考。     

油橄榄(Olea europaea L.)核心SNP位点筛选与评价

本研究基于前期对57份油橄榄种质资源的全基因组GBS-SNP分型结果开展核心SNP位点的筛选。统计分析73482个GPS-SNP位点发现,有68030个(92.58%)SNP位点的检出率达到100%,33979个(46.2%)位点的最小等位基因频率≥0.2,29647个(40.35%)位点的杂合率为0,其中同时符合上述3个条件(检出率=100%,最小等位基因频率≥0.2,杂合率=0)的位点有14125个(19.2%),其位置覆盖全基因组,既分布于基因区也位于基因间区,与73482个SNP位点的分布高度一致(R=0.997),多态信息量平均为0.43(0.33~0.67)。进一步以14125个SNP位点信息为依据,计算57个油橄榄品种间的遗传距离,并与基于全部位点(73482个SNPs)信息获得的对应品种间的遗传距离作比较,结果显示两者呈极显著的相关性(R=0.9),表明这些SNP位点具有多态性好、代表性广、可靠性高的特点,可作为油橄榄的核心SNP位点,适用于油橄榄品种鉴定、种质评价、基因定位和分子辅助育种。本研究对核心SNP位点在油橄榄品种鉴定中的具体应用进行了探讨,并指出至少需要11个核心SNP位点组合才能实现对57个油橄榄品种的完全区分。     

Advances in Japanese pear breeding in Japan

The Japanese pear (Pyrus pyrifolia Nakai) is one of the most widely grown fruit trees in Japan, and it has been used throughout Japan’s history. The commercial production of pears increased rapidly with the successive discoveries of the chance seedling cultivars ‘Chojuro’ and ‘Nijisseiki’ around 1890, and the development of new cultivars has continued since 1915. The late-maturing, leading cultivars ‘Niitaka’ and ‘Shinko’ were released during the initial breeding stage. Furthermore, systematic breeding by the Horticultural Research Station (currently, NARO Institute of Fruit Tree Science, National Agriculture and Food Research Organization (NIFTS)) began in 1935, which mainly aimed to improve fruit quality by focusing on flesh texture and black spot disease resistance. To date, 22 cultivars have been released, including ‘Kosui’, ‘Hosui’, and ‘Akizuki’, which are current leading cultivars from the breeding program. Four induced mutant cultivars induced by gamma irradiation, which exhibit some resistance to black spot disease, were released from the Institute of Radiation Breeding. Among these cultivars, ‘Gold Nijisseiki’ has become a leading cultivar. Moreover, ‘Nansui’ from the Nagano prefectural institute breeding program was released, and it has also become a leading cultivar. Current breeding objectives at NIFTS mainly combine superior fruit quality with traits related to labor and cost reduction, multiple disease resistance, or self-compatibility. Regarding future breeding, marker-assisted selection for each trait, QTL analyses, genome-wide association studies, and genomic selection analyses are currently in progress.     

河南省近年审定小麦品种基于系谱和SNP标记的遗传多样性分析

为了解河南省近年小麦品种的遗传基础,利用Illumina 90k i Select SNP标记技术对豫麦34及该省2000—2013年审定的小麦品种共96个进行全基因组扫描,分析了其遗传多样性和遗传基础。结果表明,在所有SNP位点中,多态性比率为47.39%(38 661/81 587),多态性标记在基因组间分布呈现BAD。96个品种亲缘关系较近,两两遗传相似系数的平均值为0.719,变幅为0.552~0.998,且94.3%的品种间遗传相似系数在0.652~0.812之间;按UPGMA法将96个品种划分为7个类群。综合SNP和系谱分析,近10年河南省审定的96个小麦品种遗传多样性不够丰富,多数品种亲缘关系较近,在育种中迫切需要引入新的种质资源,拓宽遗传背景。     

Distribution of photoperiod-insensitive alleles Ppd-B1a and Ppd-D1a and their effect on heading time in Japanese wheat cultivars

The genotypes of photoperiod response genes Ppd-B1 and Ppd-D1 in Japanese wheat cultivars were determined by a PCR-based method, and heading times were compared among genotypes. Most of the Japanese wheat cultivars, except those from the Hokkaido region, carried the photoperiod-insensitive allele Ppd-D1a, and heading was accelerated 10.3 days compared with the Ppd-D1b genotype. Early cultivars with Ppd-D1a may have been selected to avoid damage from preharvest rain. In the Hokkaido region, Ppd-D1a frequency was lower and heading date was late regardless of Ppd-D1 genotype, suggesting another genetic mechanism for late heading in Hokkaido cultivars. In this study, only 11 cultivars proved to carry Ppd-B1a, and all of them carried another photoperiod-insensitive allele, Ppd-D1a. The Ppd-B1a/Ppd-D1a genotype headed 6.7 days earlier than the Ppd-B1b/Ppd-D1a genotype, indicating a significant effect of Ppd-B1a in the genetic background with Ppd-D1a. Early-maturity breeding in Japan is believed to be accelerated by the introduction of the Ppd-B1a allele into medium-heading cultivars carrying Ppd-D1a. Pedigree analysis showed that Ppd-B1a in three extra-early commercial cultivars was inherited from ‘Shiroboro 21’ by early-heading Chugoku lines bred at the Chugoku Agriculture Experimental Station.     

Penetrance of creeping-rootedness in clonal progenies of lucerne and observations on underground morphology of plants differing for this character

Elaeis oleifera or ‘caiaué’, a close relative of oil palm (E. guineensis), has some agronomic traits of great interest for the oil palm genetic breeding such as slow growth, oil quality (mostly unsaturated) and disease resistance. An analysis of a Brazilian oil palm germplasm collection was carried out using RAPD (Random Amplified Polymorphic DNA) markers with the objective of understanding the genetic variation of ‘caiaué’ accessions collected in the Amazon Forest in the last two decades. A sample of 175 accessions obtained along the Amazon River Basin was analyzed and compared to 17 accessions of oil palm from Africa. Ninety-six RAPD markers were used in the analysis, of which fourteen were shown to be specific to oil palm, while twelve were specific to ‘caiaué’. Results showed that the Brazilian ‘caiaué’ accessions studied have moderate levels of genetic diversity as compared to oil palm accessions. The data allowed the establishment of similarity groups for ‘caiaué’ accessions, which is useful for selecting parental plants for population breeding. Cluster analysis showed that, in general, genetic similarities are not correlated to geographical distances, but consistent with geographical dispersal along the Amazon River network. AMOVA showed that most of the genetic variation is found within populations, as expected for anallogamous and long-lived perennial species. The study provides important information to define strategies for future collection expeditions, for germplasm conservation and for the use of E. oleifera in breeding programs. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genome-wide association mapping of genetic factors controlling Septoria tritici blotch resistance and their associations with plant height and heading date in wheat

Septoria tritici blotch (STB), caused by the ascomycete fungus Zymoseptoria tritici (also known as Mycosphaerella graminicola), is one of the most devastating foliar wheat diseases worldwide. Host resistance is the most effective strategy for management of the disease. A factor that complicates the determination of resistance is its reported interaction with heading date (Hd) and plant height (Ph). In this study, we report findings from a genome-wide association study of resistance to STB in a world-wide collection of 96 wheat accessions. The collection was evaluated under conditions of artificial infection for seedling and adult plant STB resistance, Hd and Ph in field trials. Marker-trait associations (MTAs) were detected using a mixed linear model. STB disease severities showed significant phenotypic variation. In total, 73 MTAs involving STB resistance were detected. The chromosomal locations of some of them were similar to known Stb genes or quantitative trait loci; whereas others were detected in new genomic regions. The field experiment showed evidence of genetic association between STB resistance and Hd, but only for a few genotypes. This was corroborated at the molecular level, where a total of eight genomic regions associated with STB resistance were located in similar positions to MTAs for Hd. New genomic regions associated with STB resistance found here could be useful in wheat breeding aimed at controlling STB after validation in relevant genetic backgrounds     

Variations for Fusarium head blight resistance associated with genomic diversity in different sources of the resistant wheat cultivar ‘Sumai 3’

Fusarium head blight (FHB), caused by Fusarium graminearum, is a serious disease of wheat (Triticum aestivum L.) associated with contamination by the mycotoxin deoxynivalenol (DON). The FHB-resistant wheat cultivar ‘Sumai 3’ has been used extensively around the world. The existence of variation in FHB resistance among ‘Sumai 3’ accessions has been discussed. In this study, genetic variation among ‘Sumai 3’ accessions collected from six countries were identified using SSR markers; our results demonstrate unique chromosome regions in Sumai 3-AUT and Sumai 3-JPN (‘Sumai 3’ accessions from Austria and Japan, respectively). Field evaluation indicated strong resistance to FHB in Sumai 3-AUT. The polymorphic rate (number of polymorphic markers/number of available markers × 100) based on a DArT array was 12.5% between the two ‘Sumai 3’ accessions. Genotyping for DNA markers flanking FHB-resistant quantitative trait loci (QTLs) revealed genetic variations for the QTL regions on 5AS and 2DS; however, no variation was observed for the QTL regions on 3BS and 6B. Thus, the variation in FHB resistance among ‘Sumai 3’ accessions in the field is due to genetic diversity.     

Differential contribution of two Ppd-1 homoeoalleles to early-flowering phenotype in Nepalese and Japanese varieties of common wheat

Wheat landraces carry abundant genetic variation in heading and flowering times. Here, we studied flowering-related traits of two Nepalese varieties, KU-4770 and KU-180 and a Japanese wheat cultivar, Shiroganekomugi (SGK). These three wheat varieties showed similar flowering time in a common garden experiment. In total, five significant quantitative trait loci (QTLs) for three examined traits, the heading, flowering and maturation times, were detected using an F₂ population of SGK/KU-4770. The QTLs were found at the Ppd-1 loci on chromosomes 2B and 2D and the 2B QTL was also confirmed in another F₂ population of SGK/KU-180. The Ppd-D1 allele from SGK and the Ppd-B1 alleles from the two Nepalese varieties might be causal for early-flowering phenotype. The SGK Ppd-D1 allele contained a 2-kb deletion in the 5′ upstream region, indicating a photoperiod-insensitive Ppd-D1a allele. Real-time PCR analysis estimating the Ppd-B1 copy number revealed that the two Nepalese varieties included two intact Ppd-B1 copies, putatively resulting in photoperiod insensitivity and an early-flowering phenotype. The two photoperiod-insensitive Ppd-1 homoeoalleles could independently contribute to segregation of early-flowering individuals in the two F₂ populations. Therefore, wheat landraces are genetic resources for discovery of alleles useful for improving wheat heading or flowering times.