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Perennial ryegrass (Lolium perenne L.) is one of the most important forage grasses, providing high yields with excellent forage quality. The main limiting factor for increasing the cultivation area of perennial ryegrass in the Nordic-Baltic region is insufficient winter hardiness due to unstable climatic conditions as well as insufficient persistence and drought resistance. Currently, the genetic diversity of perennial ryegrass cultivars is relatively limited; therefore, developing new, highly adaptable germplasm is of high importance in the context of changing climatic conditions. In the framework of the Nordic-Baltic Public-Private Partnership (PPP) project in pre-breeding of perennial ryegrass, 250 tetraploid plants (hereinafter genotypes), created by chromosome doubling using colchicine at the Institute of Agriculture, Lithuanian Research Centre for Agriculture and Forestry, were evaluated in open field conditions at the Research Institute of Agronomy of Latvia University of Life Sciences and Technologies. Detailed phenological scoring of all genotypes was performed over a three-year (2016--2018) period. For data analysis, the plants were grouped according to heading time and growth habit. Significant differences among the groups were found in winterhardiness, regrowth rate, development rate of generative shoots, susceptibility to rust, etc. The seed from plants that survived well and showed some promising properties were harvested in the 2nd ley year -- a total of 199 genotypes or 358 individuals (80% and 48% of all, respectively). Results of genotyping of randomly selected genotypes with the highest and lowest winter hardiness showed that the clones were genetically differentiated from the cultivars developed in Baltic countries -- unique alleles were found in the tetraploid clones that were not present in the analysed cultivars. This suggests that these developed tetraploid clones or genotypes could provide valuable breeding material to improve the suitability of perennial grass cultivars to local environmental conditions in the future. [ABSTRACT FROM AUTHOR]

Genomic selection (GS) is a potential pathway to accelerate genetic gain for perennial ryegrass (Lolium perenne L.). The main objectives of the present study were to investigate the level of genetic gain and accuracy by applying GS in commercial perennial ryegrass breeding programs. Different scenarios were compared to a conventional breeding program. Simulated scenarios differed in the method of selection and structure of the breeding program. Two scenarios (Phen-Y12 and Phen) for phenotypic selection and three scenarios (GS-Y12, GS and GS-SP) were considered for genomic breeding schemes. All breeding schemes were simulated for 25 cycles. The amount of genetic gain achieved was different across scenarios. Compared to phenotypic scenarios, GS scenarios resulted in substantially larger genetic gain for the simulated traits. This was mainly due to more efficient selection of plots and single plants based on genomic estimated breeding values. Also, GS allows for reduction in waiting time for the availability of the superior genetic materials from previous cycles, which led to at least a doubling or a trebling of genetic gain compared to the traditional program. Reduction in additive genetic variance levels were higher with GS scenarios than with phenotypic selection. The results demonstrated that implementation of GS in ryegrass breeding is possible and presents an opportunity to make very significant improvements in genetic gains. [ABSTRACT FROM AUTHOR]

Agricultural systems models are complex and tend to be over-parameterized with respect to observational datasets. Practical identifiability analysis based on local sensitivity analysis has proved effective in investigating identifiable parameter sets in environmental models, but has not been applied to agricultural systems models. Here, we demonstrate that identifiability analysis improves experimental design to ensure independent parameter estimation for yield and quality outputs of a complex grassland model. The Pasture Simulation model (PaSim) was used to demonstrate the effectiveness of practical identifiability analysis in designing experiments and measurement protocols within phenotyping experiments with perennial ryegrass. Virtual experiments were designed combining three factors: frequency of measurements, duration of the experiment. and location of trials. Our results demonstrate that (i) PaSim provides sufficient detail in terms of simulating biomass yield and quality of perennial ryegrass for use in breeding, (ii) typical breeding trials are insufficient to parameterize all influential parameters, (iii) the frequency of measurements is more important than the number of growing seasons to improve the identifiability of PaSim parameters, and (iv) identifiability analysis provides a sound approach for optimizing the design of multi-location trials. Practical identifiability analysis can play an important role in ensuring proper exploitation of phenotypic data and cost-effective multi-location experimental designs. Considering the growing importance of simulation models, this study supports the design of experiments and measurement protocols in the phenotyping networks that have recently been organized. [ABSTRACT FROM AUTHOR]

Abstract: In perennial ryegrass seed production, the establishment of seed yield potential occurs until the point of anthesis. However, utilizing potential seed yield is predominately focused on processes after anthesis, namely seed set (%) and seed filling. In practice, seed yield is the product of the number of harvested seeds remaining after cleaning and average seed weight. For this study, the anthesis patterns and seed set were recorded in a diploid variety grown in seed production fields in three different Danish regions with contrasting weather conditions and investigated in 2013 and 2014. Increases in the total precipitation during anthesis reduced the anthesis synchrony and the seed set, which ranged from 50% to 66%. Under semi‐controlled environmental conditions in which the influence of precipitation was excluded, the seed set was found to be influenced by the floret position in the spikelet and ranged from 73% in the florets in basal positions to 25% in the distal florets. It is suggested that a lower number of florets per spikelet will reduce the anthesis period. These results may provide insights for breeding programmes focused on increasing seed yield. [ABSTRACT FROM AUTHOR]

Abstract: Perennial ryegrass (Lolium perenne) is an important source of protein and energy for dairy cattle. To improve the protein/energy ratio of this forage, focus is now on improving its cell wall digestibility. The in vitro assessment of the digestible fraction of the neutral detergent fibre (NDFD) is a superior method for determining the cell wall digestibility, but requires the use of ruminal fluid, which has a highly variable composition and is often not readily available. As lignin is considered the main cell wall component that impedes NDFD, we investigated whether this “subtrait” could serve as alternative breeding selection criterion to improve NDFD. Therefore, we assessed the accuracy of two lignin quantification methods: van Soest (ADL) and Klason lignin (KL). We also considered KL estimates corrected for the solubilized lignin (total lignin or TL) and non‐solubilized protein (TL'). Although the latter is considered the truest possible lignin content, it was not always the most correlated to NDFD, due to the limited accuracy of protein quantification on lignin residue. TL is most correlated to NDFD and we therefore recommend it for use in conventional breeding if NDFD determination is not a possibility. However, NDFD is still a superior selection criterion, as it combines the effect of several subtraits and not just lignin. For marker–trait association studies, a more accurate estimate of lignin content is more important than a high correlation with NDFD, but also here, TL performs best. [ABSTRACT FROM AUTHOR]

Genomic selection (GS) has been recognized as offering numerous potential benefits for ryegrass (Lolium perenne L.) breeding. While the theoretical benefits of GS in ryegrass breeding are clear, the best way to apply GS in practical breeding programs remains to be determined. The present study aimed to investigate genomic breeding options that best balance genetic gain, breeding costs, and the level of inbreeding using stochastic simulation. Nine GS scenarios were tested, including different numbers of selection candidates (10,000, 5000, and 2000 F1 seedlings) and three reference population sizes for GS composed of plots representing a sward-based trial (500, 200, and 100 plots). Low to moderate prediction accuracy was achieved for productivity traits across cycles (i.e., 0.1–0.45 for yield [h2 = 0.3]). Scenarios with larger reference populations (i.e., 500 plots) achieved higher prediction accuracy but, when considering field trial costs, were more expensive per unit of genetic gain. All nine GS scenarios delivered significantly higher genetic gain (up to fourfold) than the conventional breeding scenario over a 20-yr period. Scenarios with moderate selection intensity on F1 seedlings with fewer plots tested in field gave the most genetic gain per dollar invested (i.e., 2000 or 5000 F1 seedlings and 100 plots). However, all GS scenarios reduced genetic diversity in the breeding population more than phenotypic selection, highlighting the need to mitigate inbreeding when applying GS in perennial ryegrass. [ABSTRACT FROM AUTHOR]

RNA-Seq methodology has been used to generate a comprehensive transcriptome sequence resource for perennial ryegrass, an important temperate pasture grass species. A total of 931 547 255 reads were obtained from libraries corresponding to 19 distinct tissue samples, including both vegetative and reproductive stages of development. Assembly of data generated a final filtered reference set of 48 713 contigs and scaffolds. The transcriptome resource will support whole genome sequence assembly, comparative genomics, implementation of genotyping-by-sequencing (GBS) methods based on transcript sampling, and identification of candidate genes for multiple biological functions. [ABSTRACT FROM AUTHOR]

An assessment of genetic improvement in turf-type perennial ryegrass was performed at a network of six locations. A comparison was made of the turf performances of five natural populations, five forage-type cultivars used for turf seeding until the 1980s and 31 turf-type cultivars released from 1974 to 2004. Populations and cultivars were also compared in two spaced-plant experiments and in two seed-yield trials. Trait regressions on registration year of turf-type cultivars showed that breeding had been highly successful in improving the turf aesthetic merit (from +8·8 to +12·5% per decade according to seasons), wear tolerance (+5·4% per decade) and crown-rust resistance (+8·9% per decade) and in lessening the turf height increase rate (−0·43 mm day−1 per decade). Turf winter greenness had been marginally improved, whereas summer greenness and seed yield had not been significantly changed. A multivariate analysis provided evidence that turf density and fineness played a major role in the visual assessment of turf aesthetic merit and that wear tolerance was closely associated with turf density. Conflicting trait associations may have precluded improvements in turf ground-cover 3 months after sowing, turf winter greenness and turf persistency. [ABSTRACT FROM AUTHOR]

During the last 100 years, in Western Europe and elsewhere, considerable effort has been devoted to improving perennial ryegrass (Lolium perenne L.) for agriculture. The first persistent cultivars to be widely used were more digestible than other common pasture species but were no higher yielding than the better wild populations of perennial ryegrass. Two main approaches (here called mainstream breeding and population improvement) have been used to further improve the species for the UK, but no recent experiments to assess progress have been published. In 2006, two plot trials were established at IBERS to compare the performance of some newer cultivars and candidate varieties with the first persistent cultivars to be widely used in the UK. One trial involved comparing 10 intermediate-heading (6 diploid and 4 tetraploid) cultivars and candidate varieties with the intermediate-heading cv. Talbot, and the other involved comparing 11 late-heading (4 diploid and 7 tetraploid) cultivars and candidate varieties with the late-heading cv. 23. During 2007 to 2009, one silage cut and 6 other cuts were harvested each year, annual dry matter (DM) yields were determined and DM samples analysed for in vitro DM digestibility (DMD), water soluble carbohydrate (WSC) and crude protein (CP) concentrations in the DM. Percentage ground covered by perennial ryegrass in November 2009 was estimated visually. Twenty of the 21 cultivars were significantly (12 to 38%) higher yielding, 15 were significantly (10 to 27 g/kg) higher in mean DMD, 15 were significantly (25 to 58 g/kg) higher in mean WSC and 7 (all diploids) were significantly higher in ground cover in autumn of the third harvest year than their respective control cultivars. There were no significant differences among the varieties in mean CP over all harvests. The newest intermediate heading cultivar (the diploid Abermagic) produced 29% more DM, was 10 g/kg higher in DMD and 51 g/kg higher in WSC, and had significantly better ground cover at the end of the third harvest year than Talbot. The newest late-heading cultivar (the tetrapioid Aberbite) produced 28% more DM than S23 and was 22 g/kg higher in DMD and 58 g/kg higher in WSC, although it was similar to 23 for ground cover at the end of the third harvest year. Both of these new varieties were developed entirely or partly by population improvement at IBERS over 26 years (1980 to 2005). These results suggest that the rates of gain in DM yield under nitrogen-limiting conditions and in herbage WSC concentration from perennial ryegrass breeding can be improved by utilizing new technologies and better breeding strategies. [ABSTRACT FROM AUTHOR]

In all ryegrass breeding programmes it is necessary to select a range of traits within different cultivar types, varying in ploidy and flowering time. The traits selected in ryegrass breeding can be broadly grouped into production traits such as yield, quality and persistency; those seed production traits crucial for delivery of the cultivar, as well as those traits that can benefit the environment, or allow ryegrass to be used for biofuel production. The emphasis placed on each trait will depend on its economic value within the various farming systems where each cultivar will ultimately be used, as well as the potential to make genetic gain in each trait. In all cases multiple trait selection will be required, to develop a cultivar improved for key traits of interest but importantly the cultivar must not have unacceptable performance for any trait. Where the genetic variation is inadequate within perennial ryegrass it may be necessary to enhance ryegrass diversity. In the future this could be achieved through targeted introgression from the closely related Festuca species, or through introduction of genes via genetic modification. Funding of ryegrass breeding internationally will increasingly be subject to the economic success of a few larger seed companies as Government funding for field-based breeding is diminishing and shifting focus to more basic research, often of a molecular nature. Ensuring this expensive basic research and associated molecular technologies are used effectively in ryegrass breeding programmes will remain a challenge when seed companies operating field-based programmes are vulnerable to considerable economic pressure. [ABSTRACT FROM AUTHOR]

Water-soluble carbohydrates (WSC) are an important factor determining the nutritional value of grass forage and development of genetic markers for selection of WSC traits in perennial ryegrass would benefit future breeding programmes. Quantitative trait loci (QTLs) for WSC have been published for an F ryegrass mapping family. Markers showing significant associations with these QTLs were used to design narrow-based populations with homozygosity for target QTLs. Founders were selected from within the mapping family. The divergent populations produced were analysed for WSC content in the glasshouse and the field. There was evidence of complex interactions between WSC content and other factors and traits, including the scale of assessment, time/degree of sward establishment and other forage quality parameters. Differences between the divergent pairs of the various populations were small. However, differences observed between the founder selection groups were maintained and the roles of the QTL regions in regulating forage WSC content were confirmed. In general, the individual divergent populations exploited only a limited extent of the large phenotypic variation available within the mapping family. However, this study sets the scene for exploring the opportunities for marker-assisted breeding strategies for complex traits in obligate out-breeding species, and the challenges of doing this are discussed. [ABSTRACT FROM AUTHOR]

By using data of Belgian trials for Value of Cultivation and Use (1963–2007) the change in agronomic performance of ryegrass varieties was quantified. Data included 144 Lolium perenne and 69 Lolium multiflorum candidate varieties, plus ‘Vigor’ and ‘Lemtal’, two varieties which were always included in the Value of Cultivation and Use (VCU) trials. As the genetics of ‘Vigor’ and ‘Lemtal’ have remained identical to those in 1963, these were used as constant standards to measure improvements of new varieties. Dry matter yield (DMY) of ‘Vigor’ and ‘Lemtal’ varied annually but did not show a progressive change, indicating that cultural changes in VCU trials of ryegrasses were small. By expressing DMY, persistency and rust resistance of the candidate varieties relative to Vigor and ‘Lemtal’, the change in agronomic performance due to breeding was determined. DMY increased with 0.3% of ‘Vigor’ and ‘Lemtal’ annually. Persistency showed an annual increase around 0.5% of the standard varieties. Before 1990, rust resistance of ryegrasses varied around 100% relative to ‘Vigor’ or ‘Lemtal’. After 1990, an annual increase of 3.6% against these standard varieties was found. [ABSTRACT FROM AUTHOR]

Lolium perenne L. (perennial ryegrass) is the principle forage grass species in temperate agriculture. Improving biomass yield still remains one of the most important aims of current forage breeding programmes. A quantitative trait locus (QTL) study investigating biomass yield traits in perennial ryegrass was carried out in greenhouse and field environments. The study is based on an F2 population consisting of 360 individuals derived from two inbred grandparents where the F1 has a large biomass yield phenotype. For both experimental environments co-localized QTL for biomass yield traits including fresh and dry weight and dry matter were identified on linkage groups 2, 3 and 7. A major QTL for fresh and dry weight was identified on LG 3 which explained around 30% of the phenotypic variance in the field experiment. The findings of this study are discussed with regard for their potential in research and breeding. [ABSTRACT FROM AUTHOR]

Contents Summary 1 I. Background 2 II. Objectives, breeding methodologies and progress in grass breeding 4 III. Markers and their suitability for genetic analysis in grasses 5 IV. Associating markers to genes for ‘sustainability’ in the grasses 5 V. The genetic control of flowering time 8 VI. Target traits: resilience against climate change and resistance to abiotic stresses 9 VII. Target traits: carbohydrates and sustainable ruminant animal production 11 VIII. Target traits: grass biomass, climate change and energy sustainability 13 IX. Conclusions 13 References 14 Here, we review the current genetic approaches for grass improvement and their potential for the enhanced breeding of new varieties appropriate for a sustainable agriculture in a changing global climate. These generally out-breeding, perennial, self-incompatible species present unique challenges and opportunities for genetic analysis. We emphasise their distinctiveness from model species and from the in-breeding, annual cereals. We describe the modern genetic approaches appropriate for their analysis, including association mapping. Sustainability traits discussed here include stress resistance (drought, cold and pathogeneses) and favourable agronomic characters (nutrient use efficiency, carbohydrate content, fatty acid content, winter survival, flowering time and biomass yield). Global warming will predictably affect temperature-sensitive traits such as vernalisation, and these traits are under investigation. Grass biomass utilisation for carbon-neutral energy generation may contribute to reduced atmospheric carbon emissions. Because the wider potential outcomes of climate change are unpredictable, breeders must be reactive to events and have a range of well-characterised germplasm available for new applications. [ABSTRACT FROM AUTHOR]

Explores the advances in introgression technologies for precision breeding within the Lolium and Festuca species. Four major species of agricultural importance; Discussion on introgression from diploid and hexaploid Festuca species; Benefits for introgression from using F[sub1] hybrids and their derivatives as male parent in backcross breeding programmes; Progress in the transfer of abiotic stress resistance from Festuca to Lolium.

In order to study the effects of competition on the genotypic composition of chance-hybrids in Lolium multiflorum, eight chance-hybrids were produced according to Burton (1948). inbred lines and cultivars were used as parents. The proportions of parental and hybrid components in the seeds and growing-field stands of the chance-hybrids were determined over three cuts using gel electrophoresis The realized hybrid proportions in the seeds, at a maximum of 38.3%, were considerably lower than the theoretically expected 50%. In the field stands, the hybrid proportions of the chance-hybrids derived from inbred lines increased considerably and continously over cuts, whereas those of the chance-hybrids produced from cultivars increased only slightly. A maximum hybrid tiller proportion of 76.8% was found at the third cut. Correspondingly. the total yield of the chance-hybrids produced from cultivars was only slightly higher than that of the parents The production of chance-hybrids, therefore, proved to be sensible only if inbred lines had been used as parents. [ABSTRACT FROM AUTHOR]

In order to establish an appropriate breeding programme, the reproductive system of a population of Lolisium rigidum Gaud. from Sardinia was analyzed to determine the mechanisms operational in this species. A dual approach was adopted of genotype assay, exploiting the polymorphism at the PGI-2 locus, and an assessment of the effects of inbreeding on agronomic characters. The statistical analysis showed highly significant differences for the number of plants due to selfing among the 35 progenies obtained by controlled crossing of genotypes homozygous for different alleles at the PGI-2 locus. The greatest inbreeding effect was on dry matter yield. Although at a bulk population level the species is outbreeding, highly self-compatible plants can be found within this naturally-derived population. [ABSTRACT FROM AUTHOR]

Perennial ryegrass (Lolium perenne L.), an important forage grass species in temperate regions, is genetically improved by population breeding. Although valued for their broad genetic base, the resulting synthetic varieties only partially exploit heterosis. Hybrid breeding offers opportunities to fix beneficial heterotic patterns more effectively and, hence, to increase the yield potential. A suspected bottleneck in the production of perennial ryegrass hybrids is the genetic intermixture of existing germplasm, impeding the definition of heterotic groups. In this study, selected parental populations of a diploid and tetraploid cytoplasmic male sterility (CMS)-based hybrid breeding program were characterized using genotyping-by-sequencing (GBS). Hybrid populations, derived from 26 parental combinations of the tetraploid breeding program, were tested for yield performance and compared to synthetic varieties at five sites over four growing seasons. The hybrids significantly outperformed the synthetics by 4.15% on average for total dry matter yield. Additionally, GBS revealed the existence of sub-populations within the tetraploid CMS germplasm. This sub-population structure represents the untapped potential that could be exploited for heterosis to further increase biomass yields. Here, we show that CMS hybrids generate substantial yield gains in perennial ryegrass and provide a method to further improve hybrid breeding, using GBS to select for heterotic groups. [ABSTRACT FROM AUTHOR]

Forage plants underpin the livestock industry. Selective breeding, including polyploidization, where genome size is increased by whole genome duplication, changes the productivity and stress tolerance of new varieties. We conducted a growth chamber experiment to investigate the likely responses of Lolium perenne L. to drought, testing four diploid and four tetraploid varieties. We simulated projected spring and summer temperatures for the South-West of England in 2080, applying three projected rainfall scenarios, which varied in drought severity. Drought caused a reduction in productivity, but there was substantial variation between varieties (up to 82%), with the optimal variety changing depending on drought severity. Across three harvests, productivity declined by 43% and 27% (dry biomass) for the severe and likely drought scenarios, respectively. In the final harvest, tetraploids exhibited a greater biomass under severe drought, whereas diploids had a greater biomass under the current rainfall and likely drought scenarios. Longer stomata were observed in tetraploids; however, stomatal conductance was not significantly different between ploidy levels. Trait selection will be important for future drought adaptation. Local climate projections will need to be consulted when selecting L. perenne varieties to tolerate the spatially variable reductions in future rainfall. [ABSTRACT FROM AUTHOR]

Perennial ryegrass (Lolium perenne) is a perennial crop used in temperate regions as forage. In L. perenne breeding programs, persistency is an important trait. Poor persistency results in sward degradation and associated yield and nutritive value losses. Breeders assess persistency of accessions using visual scoring in field plots during the 2nd or 3rd growing season. This evaluation system is easy and cheap but is not free from human bias. In this study, the correlation between the scoring done by different breeders was only 0.243. As an alternative we have developed a methodology to assess persistency of L. perenne breeding materials based on vegetation indices (VIs) derived from Unmanned Aerial Vehicle (UAV) imagery. The VIs Excess green (ExG2), Green Leaf Index and Normalized green intensity (GCC) were found to provide consistent results for flights carried out under different light conditions and were validated by ground reference information. The correlation between the VIs and the percentage of ground cover extracted from on-ground imagery was 0.885. To test the implementation of the method we compared the ExG2 value based approach to selection with a visual score based selection methodology as applied by two breeders. The breeding decisions of Breeder A agreed well with decisions based on ExG2 values (74.6%), but those of Breeder B displayed a lower agreement (54.0%). In contrast, agreement between decisions based on different flights was very high (91.6%). The methodology was validated for general applicability. In summary, the results demonstrate that basing persistency selection in L. perenne breeding programs on ExG2 values from UAV imagery is likely to be more objective in comparison to the currently-used visual scoring method. [ABSTRACT FROM AUTHOR]

New opportunities for plant breeding using androgenesis in Lolium × Festuca hybrids have been identified. Plants derived by anther culture from a Lolium multiflorum × Festuca arundinacea (5x) hybrid were screened for freezing-tolerance, and their post-freezing recovery compared. The androgenic population showed extreme diversity in freezing-tolerance. While the majority of androgenic plants had inferior freezing-tolerance compared with the freezing-sensitive L. multiflorum parent, 6% of the population were more freezing resistant than the freezing-tolerant Festuca parent. Novel Lolium and Festuca gene combinations resulting from rare meiotic events were recovered within the androgenic population. The two most freezing-tolerant androgenic plants carried virtually the entire F. pratensis subgenome of F. arundinacea. F. pratensis is known to carry genes for freezing-tolerance and would be expected to be the primary source of genes governing this trait within the F. arundinacea genome. The most freezing-tolerant androgenic plants were more freezing-tolerant than the hybrid plant from which they were derived. Consequently, androgenesis was also effective in removing factor(s) reducing the expression of freezing-tolerance within the L. multiflorum × F. arundinacea (5x) hybrid. [ABSTRACT FROM AUTHOR]