Origin and genetic structure of feral rye in the western United States

Feral rye (Secale cereale) is a serious, introduced weed of dry land agricultural regions of the western United States. It closely resembles cultivated cereal rye (Secale cereale cereale L.) with the exception of having a shattering seed head. Feral rye may have originated from hybridization of cultivated rye with mountain rye, Secale strictum, as past studies of northern Californian populations suggest, or directly from volunteer cultivated rye. We characterized the genetic structure of feral rye populations across a broad geographical range and reexamined evidence for hybrid origin versus direct evolution from domesticated cultivars. Eighteen feral populations were examined from three climatically distinct regions in the western United States. Seven cultivars, four mountain rye accessions, and one wild annual relative (Secale cereale ancestrale) were included in our analysis as possible progenitors of feral rye. Individual plants were scored for 14 allozyme and three microsatellite loci. Estimates of genetic diversity in feral populations were relatively high compared to those of the possible progenitors, suggesting that the weed had not undergone a genetic bottleneck. Weed populations had no geographical structure at either a broad or a local scale, suggesting idiosyncratic colonization and gene-flow histories at each site. Feral rye populations were no more closely related to mountain rye than cultivars were. They were, however, weakly clustered as a distinct lineage relative to cultivars. Our results do not support an interspecific hybrid origin for feral rye, but do suggest that the sampled populations of feral rye share a common ancestry that may explain its weedy nature.     

Evolutionary trends of microsatellites during the speciation process and phylogenetic relationships within the genus Secale

Eleven weedy or wild species or subspecies of the genus Secale L. were compared with a set of cultivated rye accessions, based on inter-simple sequence repeat (ISSR) markers to analyze their phylogenetic relationships. A total of 846 bands were amplified from reactions using 12 screening primers, including 79 loci with a mean of 10.1 alleles per locus. The number of amplified bands for each primer ranged from 12 to 134, with a mean of 70.5 amplified bands per primer. The presence and distribution of amplified bands in different accessions demonstrate that a rapid evolutionary trend of microsatellite repeats occurred during the speciation process from the perennial wild form to annual cultivated rye. In addition, variation, amplification, and deletion of microsatellites in genomes revealed phylogenetic relationships in the genus Secale. Analysis of the presence, number, and distribution of amplified bands in genomes, as well as the comparison with genetic similarity (GS) indices based on ISSR, indicate that Secale strictum subsp. africanum (Stapf) Hammer, Secale strictum anatolicum (Boiss.) Hammer, Secale sylvestre Host, and Secale strictum subsp. strictum (C. Presl) Hammer emerged in succession from a common ancestor of Secale following geographic separation and genetic differentiation. The annual weedy rye evolved from S. strictum subsp. strictum, which was domesticated as present-day cultivated rye. Data from ISSR analyses separated all investigated accessions of the genus Secale into three distinct groups. These results support the division of the genus Secale into three species: the annual wild species S. sylvestre; the perennial wild species S. strictum, including several differential subspecies forms such as strictum, africanum, and anatolicum; and S. cereale, including cultivated and weedy rye as subspecies forms.     

非洲黑麦染色体特异性标记的建立与应用

以非洲黑麦、小麦-非洲黑麦双二倍体、安岳排灯麦等为材料筛选100条ISSR引物。其中, 引物UBC815可在非洲黑麦中扩增出1条长561 bp的特异性片段(命名为pSaUBC815₅₆₁), 而小麦对照均未扩出该片段。引物UBC815同样能在黑麦属的瓦维洛夫黑麦(Secale vavilovii Grossh.)、森林黑麦(Secale sylvestre Host.)等5个种扩增出pSaUBC815₅₆₁。根据pSaUBC815₅₆₁设计特异PCR引物U815-F、U815-R, 对小麦族多物种进行扩增, 表明pSaUBC815₅₆₁为黑麦属特有。进而利用一套中国春-Imperial黑麦二体附加系及小麦-黑麦异源材料进行扩增, 结果显示, pSaUBC815₅₆₁分布在黑麦整套染色体上, 并且所有后代材料都能扩增出pSaUBC815₅₆₁, 表明pSaUBC815₅₆₁可作为特异性标记用来检测小麦背景中的黑麦染色质。     

A taxonomic revision of Secale (Triticeae, Poaceae)

Several taxa have previously been recognized within Secale , but most of them are difficult or even impossible to distinguish morphologically. We recognize only three species: S. sylvestre, S. strictum , and S. cereale. Secale strictum has priority over S. montanum and has two subspecies, ssp. strictum and ssp. africanum , and two varieties within ssp. strictum , van strictum and var. ciliatoglume comb. nov. Secale cereale is also treated as having two subspecies. The cultivated taxa, marked by their tough rachises, are placed in ssp. cereale and the wild or weedy taxa that have more or less fragile rachis, in ssp. ancestrale. A complete synonymy is given for S. cereale , but typification has been omitted because, in many instances, type material does not exist or has been impossible to trace.     

Isoenzymes of aspartate aminotransferase in perennial and annual rye and their hybrids.

Aspartate aminotransferase patterns were screened in a collection of rye genotypes that included 24 accessions of wild perennial rye (Secale montanum Guss.), 6 accessions of cultivated perennial Derzhavin and Tsitsin rye (Secale cereale x S. montanum), 15 accessions of winter and spring rye cultivars (S. cereale L.), and 9 accessions of perennial and annual rye genotypes bred from S. montanum ssp. kuprijanovii, Derzhavin rye, and winter rye for their resistance to fungal diseases. Aspartate aminotransferase is coded for by four loci. The data fit the model where AAT 1/4 is coded by Aat 1 and Aat 4, two duplicate loci, with null and two active alleles for each locus, alleles 1 and 3 for locus Aat 1 and alleles 2 and 4 for locus Aat 4; dimeric AAT 1/4 enzyme molecules are the products of both intralocus and interloci complementation. Allele 1 of Aat 1 was the most prominent in the isoenzyme patterns of the rye species. Alleles null and 2 of Aat 4 were twice as frequent in the perennial rye accessions, including Derzhavin and Tsitsin rye, than in winter and spring rye. In contrast, allele 4 of Aat 4 was characteristic of S. cereale. Within the screened collection, locus Aat 2 was monomorphic. Among three alleles of Aat 3, allele 2 dominated isoenzyme profiles of both rye species, whereas the other two alleles were species-specific: allele 1 was characteristic of S. montanum and allele 3 was found only in S. cereale. Key words : rye, Secale cereale, Secale derzhavinii, Secale montanum, aspartate aminotransferase, isoenzymes, perennial habit, polymorphism.     

Crop seed spillage along roads: a factor of uncertainty in the containment of GMO

Feral populations of crop species along roadsides contribute to the uncertainty regarding the containment of genetically modified (GM) crops, as the feral populations could promote the persistence of transgenes outside of cultivated fields. Roadside populations of several common crop species are known to occur far from arable fields, and the dispersal pathways that promote their recruitment in road verges are unclear. Human-aided dispersal, in particular adhesive dispersal by vehicles, has been suggested as a possible vector, but this has not yet been proven experimentally. We sampled the seed rain from vehicles inside two motorway tunnels in an urban environment to reveal the contribution of crop species to seeds unintentionally dispersed by traffic beyond agricultural production areas. Three species of arable crops, wheat Triticum aestivum , rye Secale cereale and oilseed rape Brassica napus , were among the most frequent species deposited by vehicles inside the motorway tunnels. Each of the three species was clearly more predominant in one direction of traffic. While seeds of Triticum aestivum and Secale cereale were primarily transported into the city, Brassica napus was significantly more abundant in samples from lanes leading out of the city. Seed sources in the local surroundings of the tunnels were virtually nonexistent, and the high magnitude of seed deposition combined with high seed weights suggests a dispersal mechanism different from other species in the sample, at least for Triticum aestivum and Secale cereale . This provides evidence that spillage during transport is a major driver for long-distance dispersal of crops. Our results suggest that seed dispersal by vehicles is the major driver in the recruitment of roadside populations of arable crops, providing a possible escape route for GM crops. Risk management should thus aim at curbing transport losses of GM crops.     

Features of crossability, haploidy and polyembryony in hybrid combinations between common barley Hordeum vulgare L. (2n = 14) and wheat-rye substitution lines Triticum aestivum L., cultivar Saratovskaya 29/Secale cereale L., cultivar Onokhoiskaya

The role of individual chromosomes of rye in the manifestation of crossability and seedling development in hybrid combinations between common barley Hordeum vulgare L., cultivar Nepolegayushchii (2n = 14) and five wheat-rye substitution lines Triticum aestivum L., cultivar Saratovskaya 29/Secale cereale L., cultivar Onokhoiskaya (2n = 40 wheat + 2 rye chromosomes). Crossability, which was measured by two parameters--frequency of set grains and frequency of grains with embryos--was shown to be significantly affected by each of the five rye chromosomes examined: 1R, 2R, 3R, 5R, and 6R; the development of barley haploids was affected by rye chromosomes 1 R, 3R, and 5R. We were the first to demonstrate that polyembryony could be induced by mutual effects of barley cytoplasm and rye chromosome 1R. Possible mechanisms controlling the development of haploids and twins in hybrid combinations H. vulgare x T. aestivum/S. cereale are discussed. The conclusion is drawn that hybrid combinations between common barley and wheat-rye substitution lines can serve as new models for studying incompatibility mechanisms in distant crosses and genetic control of parthenogenesis.     

Discrimination of Repetitive Sequences Polymorphism in Secale cereale by Genomic In Situ Hybridization-Banding

Genomic in situ hybridization banding (GISH-banding), a technique slightly modified from conventional GISH, was used to probe the Chinese native rye (Secale cereale L.) DNA, and enabled us to visualize the Individual rye chromosomes and create a universal reference karyotype of the S. cereale chromosome 1R to 7R. The GISH-banding approach used in the present study was able to discriminate S. cereale chromosomes or segments in the wheat (Triticum aeativum L.) background, including the Triticale, wheat-rye addition and translocation lines. Moreover, the GISH-banding pattern of S.cereale subsp. Afghanicum chromosomes was consistent with that of Chinese native rye cv. Jingzhou rye; whereas the GISH-banding pattem of Secale vavilovli was different from that of S. cereale, indicating that GISH-banding can be used to study evolutionary polymorphism in species or subspecies of Secale. In addition, the production and application of GISH-banding to the study of adenine-thymine-riched heterochromatin is discussed.     

Resistance genes for rye stem rust (SrR) and barley powdery mildew (Mla) are located in syntenic regions on short arm of chromosome.

Genetic stocks were developed for the localization and eventual cloning of the stem rust resistance gene SrR that occurs in wheat lines carrying the 1RS translocation from Secale cereale 'Imperial' rye. We have used a mutation-based approach for molecular analysis of the SrR region in rye. Forty-one independent mutants resulting in loss of SrR resistance were isolated: many of these were deletions of various sizes that were used to locate SrR with respect to chromosome group 1S markers. The analysis of the mutants showed that markers about 1 Mb apart flanking the barley Mla locus also flank SrR. Additionally, three of the approximately 20 closely related sequences of Mla in rye are deleted in each of six interstitial deletion mutants of SrR. The results indicate that the SrR region in rye is syntenic to the Mla region in barley or that SrR is possibly orthologous to the Mla locus.     

RFLP maps of rye chromosomes 6R and 7R including terminal C-bands.

A F2 mapping population was created from a cross between 'UC-90' and E-line ryes (Secale cereale L.), two lines that showed polymorphism for eight C-band loci. Clones from rye, as well as other grasses, were used as probes. RFLP maps of rye chromosomes 6R and 7R were generated that include the 6RS and 6RL terminal C-bands and the 7RS terminal C-band. The 6R map spans 230 cM and includes 9 loci. The 7R map covers 225 cM and includes 21 loci. Segregation distortion was detected for several chromosomal regions. Heterochromatic C-bands did not appear to be responsible for the distortion.     

Cold pretreatment to enhance green plant regeneration from rye anther culture

Cold pretreatment of detached tillers of rye, Secale cereale, was tested under two light regimes. The ryes included two spring and two winter cultivars. Significant increases in green plant regeneration were recorded in each experiment when cold pretreatments of two to four weeks were applied. Dim light during the stress period improved green plant regeneration for two of the four cultivars tested. The highest regeneration rate, 30.6 green plants per 100 plated anthers, was reached following three weeks at +4 °C under dim light, for spring rye Jo02. Starvation stress applied to plated anthers in mannitol medium suppressed anther response. This revised version was published online in June 2006 with corrections to the Cover Date.     

RFLP mapping of a Hordeum bulbosum gene highly expressed in pistils and its relationship to homoeologous loci in other Gramineae species

A cDNA sequence (Hbc8-2) isolated from pistils of the self-incompatible species Hordeum bulbosum was analysed for expression pattern and genetic map location. Hbc8-2 was expressed just prior to anthesis in mature pistils, and expression was maintained at a high level throughout anthesis. The same expression pattern was found in self-incompatible rye ( Secale cereale), but no expression was detected in the self-compatible cereals wheat ( Triticum aestivum) or barley ( Hordeum vulgare) at comparable stages of development. However, three wheat expressed sequence tags from a pre-anthesis library had high homology to Hbc8-2. Southern blot analyses using Hbc8-2 as a probe detected hybridising bands in the genomes of various Gramineae species including rye, barley, bread wheat, wild wheat relatives ( Aegilops tauschii and Ae. speltoides), oats ( Avena fatua and A. strigosa), rice ( Oryza sativa) and maize ( Zea mays). This suggests that Hbc8-2-like sequences are present in many species but that high levels of expression may be associated with self-incompatibility. Hbc8-2 was mapped on the long arms of chromosome 2H(b) of H. bulbosum, 2R of rye, and 2B and 2D of wheat and was assigned to chromosome 2H of barley using wheat/barley addition lines. On a H. bulbosum genetic map, Xhbc8-2 was located between Xbcd266 and Xpsr87, while in rye and wheat it was located in a 13.2-cM interval between Xpsr331 and Xpsr932, consistent with previous comparative mapping studies of these species. Mapping in rye suggested that Hbc8-2 is probably proximal to the Z self-incompatibility locus which was previously shown to be tightly linked to Xbcd266.     

Origin of B chromosomes in cultivated rye.

Cultivated rye (Secale cereale) and its weedy relative (S. segetale) carry B chromosomes. The B chromosomes are known to be morphologically alike at somatic metaphase and they are of the standard type in natural populations. To clarify the cytogenetic relationship between the standard B chromosomes of S. cereale and those of S. segetale, we made four crosses between Afghan S. segetale with two standard B chromosomes as a pistillate parent and Turkish, Iranian, Korean, and Japanese S. cereale, all with two standard B chromosomes as pollen parents. We observed the pairing of B chromosomes at diakinesis in pollen mother cells in all F1 hybrids with four standard B chromosomes, two from each of the pistillate and the pollen parents. The degree of pairing of B chromosomes in all F1 hybrids with four standard B chromosomes was similar to or somewhat lower than, that in parental strains with four standard B chromosomes. These results showed that the standard B chromosomes in S. segetale from Afghanistan are homologous with those in S. cereale from Turkey, Iran, Korea, and Japan. We therefore propose monophyletic origin of the standard B chromosomes in S. segetale and S. cereale.     

黑麦1R染色体特异性PCR引物的分子证据

Based on the differences of rRNA intergenic sequences between wheat ( Triticum aestivum L. ) and rye ( Secale cereale L. ), rye specific primer set NOR-R1 was synthesized according to Koebner' design. PCR analyses were carried out on different DNA substrates of common wheat and its relatives such as Agropyron elongataum (Host) Beauv., Haynaldia villosa Shur. and Hordeum vulgare L. The results confirmed that NOR-R1 primer set is specific to rye. It was found that PCR using DNAs from wheat materials containing 1R chromosome resulted in the specific amplification products of rye, whereas no amplification product was detected in PCR when using DNAs with other rye chromosomes. FISH (Fluorescent in situ Hybridization) further revealed that the binding sites for the primer set NOR-R1 were only on nucleolar organizing region of chromosome 1R. These results indicated that the primer set NOR-R1 provides a useful means for molecular tagging of rye chromosomes 1 R in wheat genetic background.     

Respiration and Involvement of an Alternative Pathway as Related to Age and Phenological Strategy of the Leaf

Russian Journal of Plant Physiology - Plants of spring wheat (Triticum aestivum L.) and winter rye (Secale cereale L.) pursuing different phenological strategies were studied. Respiratory activity,...     

The history of rye cultivation in Europe

During recent years finds from several prehistoric and medieval periods have thrown new light on the history of the spread of rye. It is now proven that wild rye is indigenous to Anatolia and was already domesticated there by the early Neolithic at the beginning of agriculture. Secale migrated to Central Europe as a weed among other cereals, and single grains of weed rye have been recorded there since the early Neolithic. The number of finds increased during the Bronze Age and Iron Age, and the status of rye changed from weed to crop plant, probably in the course of the early Iron Age. This acculturation of Secale cereale in central and eastern Europe was obviously independent of the earlier one in Anatolia. The first stages towards deliberate cultivation happened unintentionally through harvesting close to the ground, so that the rye was permanently represented in the seed corn. From this point rye was able to take advantage of its competitive strength on poor soils and in areas with unfavourable climate. The start of rye as a crop in its own right during the pre-Roman Iron Age and Roman period presumably took place independently in different areas. The expansion of intensive rye cultivation occurred in the Middle Ages. However, new finds from north-west Germany, which are presented here, show that in this area rye has been cultivated as a main crop on poor soils since the Roman period. In two maps all rye finds up to 1000 A.D. are shown, which after critical consideration can be regarded as cultivated rye.     

Reproduction and feeding behavior of Myzus persicae on four cereals

Green peach aphid, Myzus persicae (Sulzer), does not overwinter outdoors in Minnesota; it arrives each spring on low-level jet streams from the south. After arrival, anholocylic reproduction occurs on numerous herbaceous species, including many common weeds, before movement to potato, Solanum tuberosum L. In investigating aphid feeding behavior on barrier crops, we observed winter wheat, Triticum aestivum L., colonized by green peach aphid. The Northern Great Plains grows 94,000 ha of potatoes and 20.5 million ha of small grain cereals each year, the latter potentially providing an early emerging and widely distributed green peach aphid host to influence early season potato colonization. Life tables statistics indicated green peach aphid had its highest reproductive potential among cereals on winter wheat, with rye (Secale cereale L.) > barley (Hordeum vulgare L.) > oats (Avena sativa L.). Green peach aphid was found to colonize barley, rye, and winter wheat, but not oats. Mean generation time, net reproductive rate, doubling time, and finite rate of increase were significantly different between host plants. Electrical penetration graph technique indicated mean nonpenetration duration by green peach aphid was significantly different among plant species, and significantly longer on winter wheat than on the other cereals. Mean xylem phase duration was not significantly different among plant species but sieve element salivation was of longest duration on potato. Phloem sap ingestion (E2) was also significantly different among plant species with longest E2 duration on winter wheat. This study demonstrates that this aphid can effectively use key cereals at the vegetative stage.     

A phylogenetic analysis based on nucleotide sequence of a marker linked to the brittle rachis locus indicates a diphyletic origin of barley

BACKGROUND AND AIMS: Barley (Hordeum vulgare ssp. vulgare) cultivation started between 9500 and 8400 years ago, and was a major part of ancient agriculture in the Near East. The brittle rachis is a critical trait in the domestication process. METHODS: A DNA sequence closely linked to the brittle rachis complex was amplified and resequenced in a collection of cultivated barleys, wild barleys (H. vulgare ssp. spontaneum) and weedy brittle rachis varieties (H. vulgare ssp. vulgare var. agriocrithon). The sequence was used to construct a phylogenetic tree. KEY RESULTS: The phylogeny separated the W- (btr1-carrying) from the E- (btr2-carrying) cultivars. The wild barleys had a high sequence diversity and were distributed throughout the W- and E-clades. Some of the Tibetan var. agriocrithon lines were closely related to the E-type and others to the W-type cultivated barleys, but an Israeli var. agriocrithon line has a complex origin. CONCLUSIONS: The results are consistent with a diphyletic origin of barley. The W- and E-type cultivars are assumed to have evolved from previously diverged wild barley via independent mutations at Btr1 and Btr2.     

中国春及其ph2b突变体与秦岭黑麦杂种胚胎发育的比较研究

对普通小麦中国春(CS)及其突变体(CSph2b)与秦岭黑麦远缘杂交的受精过程及其杂种早期胚胎发育进行了研究.结果显示,大部分秦岭黑麦花粉能在CS及CSph2b小麦柱头上萌发,花粉管可顺利伸入花柱和胚囊;CS和CSph2b已授粉子房中,分别有80.12%和84.80%完成双受精过程而形成正常胚及胚乳,也有小部分仅有卵核受精或极核受精而只形成胚或胚乳,两者总受精率分别为86.74%和88.89%,成胚率分别为83.73%和87.14%.表明中国春及其突变体CSph2b与秦岭黑麦的可杂交性都很高,并且CSph2b略高于CS.