Temperature Before Cold Acclimation Affects Cold Tolerance and Photoacclimation in Timothy (Phleum pratense L.), Perennial Ryegrass (Lolium perenne L.) and Red Clover (Trifolium pratense L.)

The expected temperature rise in late summer/early autumn can change the conditions for acclimation and affect the winter survival of perennial crops. This study examined the effect of the temperature just before the onset of cold acclimation (pre‐acclimation) on freezing tolerance of timothy (Phleum pratense L.), perennial ryegrass (Lolium perenne L.) and red clover (Trifolium pratense L.) populations (both cultivars and breeding populations) adapted to either northern or southern parts of Norway. Using phytotron experiments, we studied whether increasing pre‐acclimation temperature delays growth cessation, affects photoacclimation and reduces freezing tolerance. Furthermore, we assessed whether these effects were related to the latitudinal adaptation of the plant material. The results showed that a rise in pre‐acclimation temperature decreased both cold acclimation capacity and photoacclimation in these species. This affected the freezing tolerance, which was reduced significantly more in northern‐adapted population of timothy and perennial ryegrass compared with southern‐adapted populations. Red clover was less affected by temperature changes than the grasses.     

Management of Italian and Perennial Ryegrasses for Seed and Forage Production in Crop Rotations

Italian ryegrass (Lolium multiflorum Lam.) and perennial ryegrass (L. perenne L.) can be grown for seed and forage in cold winter regions provided the stand persists well over winter. Seed yield and plant characteristics during primary growth, and forage yield during regrowth, were determined for two Italian and one perennial ryegrass cultivars in Atlantic Canada. Establishment methods and dates included sowing ryegrass in cultivated soil alone or with barley in mid‐May and, after harvesting the barley crop, by sowing ryegrass following conventional or reduced cultivation and by no‐till drilling into barley stubble in mid‐August and early September. Despite some winterkill, particularly in Italian ryegrass, seed and forage yields were adequate in post‐establishment growing seasons. Seed yield for Italian ryegrass was greatest (1270 kg ha^?1) when it was sown into cultivated soil in mid‐August and least (890 kg ha^?1) when sown alone in May. Italian ryegrass yielded 15–17 % more seed when plots were established in mid‐August rather than in mid‐May or early September. Italian ryegrass cv. Lemtal had a greater density of fertile tillers (1030 m^?2) in the sward than cv. Ajax (860 m^?2) and its tiller density was greater when seeded into cultivated soil in September than in mid‐August. There were fewer spikelets per seed head for sowing Italian ryegrass with barley in May than for the other methods of establishment. Forage yield in regrowth was greater for Italian ryegrass cv. Ajax (2770 kg ha^?1) than for cv. Lemtal (2480 kg ha^?1). Seed yield of perennial ryegrass was greater when seeded in mid‐May than in mid‐August or early September. The seed yield of perennial ryegrass was greater when it was sown with barley in May and harvested for grain, than when it was sown alone or with barley harvested at late milk stage. The establishment methods for mid‐August and early September sowing had little effect on seed yield. However, the no‐till and reduced tillage methods resulted in a greater tiller density than sowing into the cultivated seedbed. Fertile tillers tended to be denser under reduced cultivation for sowing in August. Forage yield of perennial ryegrass regrowth was not influenced by the sowing method and timing. In conclusion, Italian and perennial ryegrasses produce adequate seed and forage regrowth under different establishment methods and timing. However, the poor persistence of Italian ryegrass may limit commercial production after the establishment year in Atlantic Canada.     

Use of a micro-sward technique for determining bite mass of four grass species in short-term tests

Four grass species typically found in semi-natural pastures in the UK were sown individually in boxes (85 × 44 × 14 cm) to create micro-swards. Yorkshire fog (Holcus lanatus L.); creeping bent (Agrostis stolonifera L. cv Kromi); rough meadow-grass (Poa trivialis L.); and perennial ryegrass (Lolium perenne L. cv Glen) were offered to yearling Simmental × Holstein steers in short-term tests to characterize bite mass in relation to morphological traits of the grass species. Boxes were weighed (±0.1 g) before and after a 5 min period during which the steers were allowed to take ~50 bites. Perennial ryegrass had the greatest sward surface height (17.7 cm vs. mean 12.7 cm for the other species). Extended tiller length was greatest for ryegrass (17.3 cm) and least for rough meadow-grass (13.5 cm). Leaf length was greatest for ryegrass (6.5 cm) and Yorkshire fog (7.0 cm) and least for creeping bent (4.0 cm). Sward DM% was lowest for ryegrass (19.7%) and highest for creeping bent and rough meadow-grass (mean 23.2%). Number of bites taken from the boxes did not differ between grass species (mean 32.7 bites). Mean dry matter bite mass was greatest for perennial ryegrass (0.50 vs. mean 0.37 g DM bite⁻¹ for the other species). Regression analyses indicated that the best sward variables for predicting mean DM bite mass were sward DM% (F prob. = 0.066) and sward surface height (F prob. = 0.086). While morphological characteristics affected bite mass, more is yet to be learned about other factors that may affect intake characteristics of forage species to improve economic and environmental sustainability.     

Phenotypic and QTL analyses of herbage production-related traits in perennial ryegrass (<Emphasis Type="Italic">Lolium perenne</Emphasis> L.)

Phenotypic and genetic evaluation of morphological traits associated with herbage biomass production was undertaken in a perennial ryegrass (Lolium perenne L.) biparental F₁ mapping population (n = 200) with parent plants from cultivars ‘Grasslands Impact’ and ‘Grasslands Samson’. Morphological traits measured on three clonal replicates of the parental genotypes and 200 F₁ progeny in a glasshouse in two separate trials (autumn and spring) included: dry weight (DW), leaf elongation rate (LER), initial tiller number (TNs), final tiller number (TNe), site filling (Fs), tiller weight (TW), leaf lamina length, leaf tip and ligule appearance rates (ALf, ALg) and leaf elongation duration (LED). Principal component analysis of patterns of trait association identified negative correlation between TNs or TNe, and TW as the primary basis for morphological difference and indicated that either high LER or long LED could reduce TN. Plants with higher LER tended to have increased DW. Quantitative trait loci (QTL) were detected on all seven linkage groups (LG) of a perennial ryegrass linkage map for all but three traits. A total of 61 QTL were identified, many of which clustered at 15 shared genome locations. Significant genotype by environment effects were encountered, evidenced both by variation between experiments in genotype rankings and by a general lack of commonality for QTL for the same traits in the different experiments. Only five QTL, for ALf, ALg and TN, were conserved between autumn and spring trials. A QTL for TN and DW on LG6 is a strong candidate for application of MAS in future plant improvement work and was found to be co-linear with QTL for equivalent traits reported on chromosome 2 in rice.     

The influence of sward-age and management on the type of timothy and cocksfoot

Summary Cocksfoot and timothy were collected from a number of Dutch grasslands. The populations obtained in this way were investigated to see whether there might be a correlation between age and management of the fields, and heading time and growthhabit type of the plants in them. It appeared that old grasslands contain a type of timothy quite different from that of young grasslands, the plants of which closely resembled the hay types from commercial strains. Such extreme hay types were not encountered in old grasslands. One old meadow developed intermediate types which collectively might be considered to represent an indigenous hay form.The difference between old and young grasslands was less clear in the case of cocksfoot; the populations from young pastures had rather more plants with slightly more erect growth-habit.An evident influence of management on the plant type, as was found in perennial ryegrass from Dutch grasslands by Jonker (8), could not be established in the case of cocksfoot and timothy.Choke (Epichloë typhina) appears to be a fairly extensively distributed disease in timothy, especially along the borders of the large rivers of the Netherlands..

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Variation in forage grasses as hosts of the root-knot nematode Meloidogyne naasi (Franklin) and selection for resistance in ryegrass

Summary Reactions of 13 grasses to Meloidogyne naasi varied with species; ryegrasses, fescues and their hybrids were generally susceptible and cocksfoot and timothy resistant. Marked variation in host resistance levels occurred between genotypes within cultivars.Selection of single plants, followed by tests on replicate tillers, identified resistant and susceptible genotypes in both Italian and perennial ryegrass cultivars. Resistant plants had few nematode-induced galls and fewer females and eggs than susceptibles. There was more or less continuous variation, with many genotypes intermediate between extremes of resistance and susceptibility. Selected resistant and susceptible genotypes are of use in assessing variation in nematode populations and as controls for breeding and selection programmes.     

Impact of waterlogging and temperature on autumn growth,hardening and freezing tolerance of timothy (Phleum pratense)

Precipitation has generally increased in Norway during the last century, and climate projections indicate a further increase. The growing season has also become longer with higher temperatures, particularly in autumn. Previous studies have shown negative effects of high temperatures and, depending upon temperature conditions, contrasting effects of waterlogging on hardening capacity of timothy. We studied effects of waterlogging on seedlings of timothy (Phleum pratense, cv. Noreng) under three pre-acclimation temperatures: 3°C, 7°C, 12°C, and in autumn natural light in a phytotron at Holt, Tromsø (69°N). After temperature treatments, all plants were cold acclimated at 2°C for three weeks under continued waterlogging treatments. Freezing tolerance was determined by intact plants being frozen in pots at incremental temperature decreases in a programmable freezer. Waterlogging resulted in a higher probability of death after freezing, and a significantly reduced regrowth after three weeks at 18°C, 24 hrs light in a greenhouse. Increasing pre-acclimation temperatures also had a clear negative effect on freezing tolerance, but there was no interaction between temperatures and waterlogging. The results indicate that waterlogging may have negative implications for hardening of timothy and may contribute to reduced winter survival under the projected increase in autumn temperatures and precipitation.     

Variation in resistance to different winter stress factors within a full-sib family of perennial ryegrass

Wintering ability in the field and resistance to different winter-stress factors under controlled environmental conditions were studied in a full-sib family of perennial ryegrass (Lolium perenne L.). Significant variation in tolerance to freezing and ice encasement, resistance to pink snow mould (Microdochium nivale) and also in winter survival and spring growth were found between the different genotypes. No strong correlations were found between the resistances to the different stress factors. These results indicate that resistance to different winter-stress factors is controlled by separate genes in perennial ryegrass. A low but significant positive correlation was found between spring growth of plants in the field after the first winter and both freezing tolerance and M. nivale resistance measured in controlled environments. Cold hardening seemed to influence freezing tolerance and M. nivale resistance differently in the different genotypes, since no distinct correlation in tolerance to freezing or resistance to M. nivale was found between unhardened and hardened plants. Tolerance or resistance to most of the winter stress factors measured was positively correlated with plant size. This revised version was published online in July 2006 with corrections to the Cover Date.     

The Relationship Between Progeny Seed Yield in Drilled Plots and Maternal Spaced-Plant Traits in Perennial Ryegrass (Lolium perenne L.)

Seed production in perennial ryegrass, Lolium perenne L., is low. Early selection for high seed yield carried out in spaced plants would be valuable for breeders, but only if this potential is also expressed in derived progenies sown later in drilled plots. Therefore, spaced-plant traits assessed on clonal ramets of 31 genotypes in each of two perennial ryegrass varieties in four environments were related to seed production characteristics in drilled plots of open-pollinated progenies of these plants. Significant differences were found among these half-sib families for seed yield, thousand-grain weight, spring performance and earliness. The number of spikelets per ear in the maternal plants was negatively correlated with the seed yield of the progenies, explaining 17 % of the variation for seed yield in the cv. ‘Barenza’ and 14 % in the cv. ‘Wendy’. Multiple regression analyses revealed that no combinations of plant traits consistently explained a major portion of the variation for seed yield of the progenies. The results indicate that in these varieties, spaced-plant data are of limited value in predicting seed production. Direct selection for seed yield in drills of progenies in later stages of the breeding programme remains the best method for obtaining varieties with sufficient seed production.     

多年生黑麦草四个品种抗寒性研究

本研究以四个多年生黑麦草品种为试材．对其进行低温胁迫处理,分别测定其抗寒性的几项生理指标,结果表明在抗寒性方面：高原>太阳岛>赛车>绅士。该研究为多年生黑麦草各品种的推广和应用提供重要理论依据。     

Mitochondrial DNA analysis reveals cytoplasmic variation within a single cultivar of perennial ryegrass (Lolium perenne L.)

Summary The extent of mitochondrial DNA restriction fragment length polymorphisms among individual plant samples of perennial ryegrass was determined. A total of 72 plants from three cultivars Yorktown II, S23 and Riikka were surveyed using three restriction enzymes (BamHI,EcoRI andHindIII) and three mitochondrial gene probes (coxI, coxIII andnad9). Polymorphisms were noted within each of the two cultivars Yorktown II and S23, whereas in Riikka no variation was detected. It seems most likely that the mitochondrial genome diversity within the same cultivar has resulted from non-homogeneous ancestor cytoplasms. The hybridization-based assay employed is simple to perform, gives unambiguous results, and may thus be used in mass screening of perennial ryegrass populations for breeding purpose.     

Improved frost tolerance and winter hardiness in proline overaccumulating winter wheat mutants obtained by in vitro-selection is associated with increased carbohydrate,soluble protein and abscisic acid (ABA) levels

In previous studies in vitro-selection of proline overaccumulating lines of winter wheat (Triticum sativum L. cv. Jo 3063) with increased frost tolerance was reported. These traits were found to be genetically stable. In the present study the improvement of frost tolerance (winter hardiness) under field conditions is confirmed for F₇ progenies of the mutants. Moreover, the mutants accumulated higher levels of glucose and fructose, soluble protein and abscisic acid (ABA) in addition to proline than the wild type under cold hardening conditions in a growth chamber as well as under cold hardening field conditions. ABA and proline levels peaked when the temperature decreased, whereas carbohydrate levels increased more slowly at decreasing temperature. Soluble protein levels also increased during cold hardening, but in addition showed sharp declines during frost periods. Increased carbohydrate levels of the mutants were associated with lower osmotic potential values. The differences in carbohydrate, protein and ABA levels between the mutants and the wild type are probably due to pleiotropic effects of the mutation.     

Reed Canarygrass Forage Yield and Nutrient Uptake on a Year-round Wastewater Application Site

Reed canarygrass (Phalaris arundinacea L.) is often planted at wastewater treatment sites to provide ground cover and remove nutrients. Our overall objective was to determine the forage yield and nutrient uptake under year-round potato wastewater application in northern latitudes. Specifically, we determined the effect of N fertilization rate on forage dry matter yield and N and P uptake by reed canarygrass, and compared the forage yield, persistence and nutrient uptake of reed canarygrass relative to those of orchardgrass (Dactylis glomerata L.), smooth bromegrass (Bromis inermis Leyss), timothy (Phleum pratense L.) and quackgrass [Elytrigia repens (L.) Nevski]. With only wastewater application, reed canarygrass had a forage yield of 5.8 Mg ha⁻¹, with N and P uptake of 113 and 30 kg ha⁻¹, respectively. Forage dry matter yield, N uptake and P uptake increased to 14.5 Mg ha⁻¹, 383 kg ha⁻¹ and 64 kg ha⁻¹, respectively, with an N fertilization rate of 224 kg ha⁻¹. Forage yield and N uptake of reed canarygrass, orchardgrass, timothy and smooth bromegrass were similar and exceeded those of quackgrass. Reed canarygrass P uptake exceeded that of the other grasses. Reed canarygrass was less persistent than quackgrass or smooth bromegrass.     

Assessing inbreeding in perennial ryegrass (Lolium perenne) as a step towards F1 hybrid breeding

Perennial ryegrass is an important turf and forage grass in temperate regions worldwide. Limited genetic gains have been made with current breeding strategies compared with other grass crops, such as rice and maize, which benefit from Filial 1 (F1) hybrid breeding. One of the largest constraints on hybrid breeding in ryegrass is self-incompatibility preventing inbreeding, as homozygous parental lines are required to develop hybrids with maximal hybrid vigour. Obligate outcrossing in ryegrass has resulted in cultivars with high levels of heterozygosity, lacking trait uniformity across the population. A naturally occurring self-fertile (SF) locus that overcomes the self-incompatibility system has been identified in a European perennial ryegrass population. This study crossed the SF locus into an elite cultivar, producing a self-compatible population that was inbred for several generations. Genotyping-by-sequencing (GBS) was used to assess the population structure and degree of inbreeding in the self-compatible population. Phenotypic analysis indicated that increased homozygosity did not necessarily affect growth and performance. This study concludes that self-compatible ryegrass is a promising tool for hybrid breeding and agronomic improvement of perennial ryegrass.     

Late heading of perennial ryegrass caused by introducing an Arabidopsis homeobox gene

Perennial ryegrass (Lolium perenne L.) is the most important temperate forage grass species. Unfortunately, the nutritional value of perennial ryegrass declines as maturity progresses, mainly because of a high concentration of poorly digestible compounds in inflorescences. Therefore, the development of forage‐type ryegrass varieties with extended vegetative growth is of interest for agriculture. To delay floral transition in perennial ryegrass the Arabidopsis ATH1 gene driven by the maize ubiquitin promoter, the rice actin promoter or the rice OSH1 promoter, respectively was introduced. In ATH1‐expressing plants heading was delayed, and in a number of cases the plants never flowered at all. Such non‐ or late‐heading was accompanied by the outgrowth of normally quiescent lateral meristems into extra leaves, resulting in a leafy growth habit. When eventually heading, these plants generally produced a reduced number of inflorescences. These observations suggest that ATH1‐mediated delay of heading may be useful to improve fodder quality of perennial ryegrass.     

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

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

Breeding winter hardy grasses

Summary Perennial grasses are vital for Norwegian agricultural production. The nature and extent of winter damage on grasslands is highly dependent on climatic conditions, and determines both persistency and yield. Physical stresses such as frost and ice encasement predominate in coastal regions with an unstable winter climate, while biotic stresses such as low temperature fungi are more common in the inland regions. Development of hardening depends on plant adaptation and climatic conditions during autumn and winter. New winter-hardy cultivars should be bred for wide adaptation to winter stresses. The genetic background for the most important character, freezing tolerance, seems to be of polygenic nature with mainly additive gene action. Selection for increased freezing tolerance has been effective over generations in grasses, and in most grass species ample variation still exists to be exploited by breeding. However, in some species like perennial ryegrass, modern biotechnological methods should be used to improve freezing tolerance and winter hardiness.     

Assessment of perennial ryegrass (Lolium perenne L.) for breeding. II. Components of winter hardiness

Summary When breeding for good winter hardiness a broad range of adaptations must be considered. Such a multitrait approach requires some understanding of physiological and genetical relationships between relevant traits.Fourteen winter related characters were measured on field grown plants from 86 perennial ryegrass accessions originating from a wide range of climates. Genetic components of variance and covariance were extracted from the data and a principal components analysis carried out. The first three independent components accounted for 80% of the variance represented by the genetic correlations and were broadly identified as winter growth, winter hardiness and early spring activity factors. None of the accessions exhibited a combination of extreme winter growth and winter hardiness although several with excellent winter hardiness showed better than average winter growth. Amounts of water soluble carbohydrate maintained through the winter appeared important for good survival.It was concluded that a selection index approach may prove useful in future breeding programmes as further information concerning physiological relationships becomes available.     

Molecular characterization of genetic diversity in European germplasm of perennial ryegrass

Summary Perennial ryegrass (Lolium perenne L.) is the most important grass species for temperate grassland agriculture. The level and distribution of genetic variation in gene bank ecotype collections is still largely unknown but of great interest for the planning of breeding programs. The objectives of this study were to (i) assess the molecular diversity of Polish ecotypes of perennial ryegrass, and (ii) compare the relationship between this group and German ecotypes and European cultivars investigated previously. A total number of 166 polymorphic marker bands were detected among the 171 individual plants of the 9 Polish ecotypes. In a joint analysis with 9 Polish and 22 German ecotypes, and 22 European cultivars 172 polymorphic RAPD markers could be found. Genetic distance among the Polish ecotypes ranged from 0.31 to 0.51, while for all 53 populations a broader range was detected (0.25–0.67). An analysis of molecular variance (AMOVA) revealed a much larger variation within populations (71%) than among them (29%). The Polish ecotypes contained the highest within population variation (74%). The largest among group difference (15%) was found between the Polish ecotypes versus all other accessions. We conclude that the Polish ecotypes represent a valuable genetic resource for enlarging the genetic variation in the West European germplasm pool of perennial ryegrass.     

Inheritance of resistance to systemic symptom expression of Melon necrotic spot virus (MNSV) in Cucumis melo L. `Doublon'

In melon (Cucumis melo L.) a resistance mechanism to Melon necrotic spot virus (MNSV) controlled by a single recessive gene called nsv has been reported. We found that following mechanical inoculation of the virus C. melo cv. ‘Doublon’ shows only local reaction, but it does not develop any systemic symptom and the virus only can be recovered from the local lesions. This reaction could be considered as a new source of resistance to MNSV. In this work,the genetic control of this behaviour in the progenies from the cross ‘Doublon’ × ‘ANC-42’ (a susceptible line) is studied. We report two complete dominant genes, which we propose to call Melon necrotic resistance 1 (Mnr1) andMelon necrotic resistance 2 (Mnr2), controlling this character. The genetic linkage between one of these genes and nsv in the populations derived from the cross between ‘Doublon’ and ‘PMR-5’ (a resistant line) is also reported. The map distance between these two loci was estimated to be 19 cm. This revised version was published online in July 2006 with corrections to the Cover Date.