Floral inhibition in red fescue (Festuca rubra L.) through expression of a heterologous flowering repressor from Lolium

Extension of the vegetative growth phase through delay of flowering is an important goal in today's breeding programs of both forage and turf grasses. In forage grasses, the stem and inflorescence production comprise a significant reduction in the digestibility, nutritional value and productivity of the crop, and in turf grasses the stems that start to emerge during the growth season suppress the formation of new shoots and affect the quality, density and persistence of the sward. We have tested the potential of the strong floral repressor LpTFL1 from perennial ryegrass (Lolium perenne L.) to manipulate the transition to flowering in red fescue (Festuca rubra L.), a cool-season turf grass. Expression of LpTFL1 from the constitutive maize ubiquitin promoter represses flowering in red fescue, and the flowering repression phenotype correlates well with the level of LpTFL expression. Transgenic lines showing low to intermediate expression of LpTFL1 flowered approximately two weeks later than the controls, and transgenic lines showing very high LpTFL1 expression levels still remained non-flowering after exposure to natural vernalization conditions (Danish winter) in two successive years. There were no other phenotypic effects associated with the LpTFL transgene expression during vegetative growth. However, there was a tendency towards an LpTFL1-mediated reduction in stem length among the flowering lines. Expression of a truncated LpTFL, caused by transgene rearrangements during the transformation, lead to increased flowering and stem production and a decrease in panicle size. This is to our knowledge the first report on full inhibition of floral development in a commercially important grass species.     

Tiller demography in seashore populations of Agrostis stolonifera,Festuca rubra and Poa irrigata

Abstract. The dynamics of tillers in natural populations of three cohabiting perennial grass species, Agrostis stolonifera, Festuca rubra and Poa irrigata (= Poa pratensis ssp. irrigata) were studied for five years in a Baltic seashore meadow. The process of tiller population maintenance was very dynamic. Both birth and death rates of tillers were high, particularly in A stolonifera, and the turnover rate of the populations was high. Recruitment was mainly by vegetative tillers, produced continuously throughout the growing season. The proportion of flowering tillers was low, but varied between years. Considerable year-to-year variation was also found in birth and death rates. Despite this between-year variation and the differences found between species in flowering frequency, pattern of survivorship and tiller longevity, population sizes of the species remained relatively constant.     

Effects of plant age on performance of the tropical perennial fodder grass,Cenchrus ciliaris L. subjected to elevated ultraviolet‐B radiation

• Abiotic stress, notably high ultraviolet‐B (eUV‐B), limit growth and productivity of many crop plants, but information on response of forage grasses to eUV‐B radiation is rather limited.
• The present study was therefore conducted to increase our understanding of differential age‐related responses on growth, metabolism and fodder quality of Cenchrus ciliaris‐3108 (Buffel grass) to elevated UV‐B (eUV‐B: 7.2 kJ·m^?2·day^?1).
• Plant growth at both growth stages was notably reduced in response to eUV‐B, except for the number of nodes and tillers at vegetative and reproductive stages. At anthesis, tillering increased due to the perennial habit of this plant, but leaf senescence reduced the number of leaves per tiller. Unlike ambient UV‐B, eUV‐B at the vegetative stage resulted in diversion of photosynthate for the formation of secondary metabolites (tannins and phenolics), providing dual protection from photooxidative damage and from herbivory.
• The forage biomass as well as quality showed a marked decline under eUV‐B and relative nutritive value was reduced at both growth stages.

     

Morphological Constraints of Shoot Demography of a Clonal Plant: Extra- and Intravaginal Tillers of Festuca rubra

Abstract Festuca rubra forms tillers in two different ways: extravaginally and intravaginally. Demography of these two tiller types was observed in seventeen selected tussocks of Festuca rubra s.s. over four growing seasons. Extravaginal tillers were bigger at birth and on the average produced twice as many daughter tillers per tiller. In general, the natality and mortality of extravaginal tillers were less regular than that of intravaginal tillers. Overall tillering rate per tiller was correlated with the density of the surrounding vegetation; mortality, natality and tiller life span were not. High density of the surrounding vegetation did not result in increased formation of extravaginal tillers. The proportion of the extravaginal tillers was not correlated with the density of the F. rubra tussocks. There is no evidence for foraging by extravaginal tillers, but they do act as founders of small clusters of tillers.     

Effects of photoperiod in the glasshouse on the growth of leaves and tillers in three perennial grasses

Leaf and tiller development in vegetative plants of S.₃₇ cocksfoot (Dactylis glomerata L.), S.₂₁₅ meadow fescue (Festuca pratensis Huds.) and S.₂₄ perennial rye-grass (Lolium perenne L.) were measured in photoperiods varied by extending a basic period of natural light with low-intensity artificial illumination so that all plants received approximately the same amount of light each day. Increasing the photoperiod usually increased leaf length and sometimes leaf width, but decreased rate of production of leaves. However, the rate of production of new leaf surface increased with photoperiod. Increasing the photoperiod generally decreased the rate of tillering; thus, plants grown in long days had few tillers with large leaves and sheaths, in short days, plants developed more but smaller tillers.     

Epidermal Cell Division and the Coordination of Leaf and Tiller Development

Initiation and development of grass leaves and tillers are oftendescribed individually with little attention to possible interrelationshipsamong organs. In order to better understand these interrelationships,this research examined epidermal cell division during developmentaltransitions at the apical meristem of tall fescue (Festuca arundinaceaSchreb.). Ten seedlings were harvested each day for a 9-d period,and lengths of main shoot leaves and primary tillers were measured.In addition, numbers and lengths of epidermal cells were determinedfor 0·5 mm segments along the basal 3 mm of each leafand tiller. Primordia development and onset of rapid leaf elongationwere characterized by an increase in the number of cells perepidermal file with mean cell length remaining near 20 µmper cell. After the leaf had lengthened to 1-1·5 mm,cells near the leaf tip ceased dividing and increased in length,at which time leaf elongation rate increased rapidly. Liguleformation, marking the boundary between blade and sheath cells,occurred prior to leaf tip emergence above the whorl of oldersheaths, while the earliest differentiation between blade andsheath cells probably began when leaves were < 1 mm long.Major transitions in leaf and tiller development appeared tobe synchronized among at least three adjacent nodes. At theoldest node, cessation of cell division in the leaf sheath wasaccompanied by initiation of cell division and elongation inthe associated tiller bud. At the next younger node the ligulewas being initiated, while at the youngest node cell divisioncommenced in the leaf primordium, as elongation of a new leafblade began. This synchronization of events suggests a key rolefor the cell division process in regulating leaf and tillerdevelopment.Copyright 1994, 1999 Academic Press Festuca arundinacea Schreb., tall fescue, cell division, leaf initiation, tillering, ligule development     

Estimation of Potential Tiller Production and Site Usage During Tall Fescue Canopy Development

Tiller appearance in tall fescue (Festuca arundinacea Schreb.)occurs in an orderly, predictable manner with the potentialfor a high degree of synchronization among tillers on a givenplant. Estimates of potential cumulative tiller production (Tmax)are made for synchronous (Tmax_lx = 2^Lx+1 – 1, where Lxis the axil number of the youngest leaf on the main stem whichbears an emerged tiller) and non-synchronous (Tmax₁ = 2^L–NLAT–1,where L is the number of leaves on the main stem and NLAT thenumber of leaves above the youngest primary tiller at its appearance)conditions. A method for determining the degree of synchronizationand an equation for estimating site usage are also presented.Early in seedling development, site usage of a tall fescue populationwith high tillering capacity was near 90%, and tillering wasregulated largely by rate of tiller site formation. As the canopydeveloped the phyllochron (time between successive leaf appearances)and NLAT increased, slowing the rate of tiller production intemporal terms and in relation to leaf appearance, respectively.Beyond 45 d after planting, site usage decreased rapidly, furtherreducing tiller production. High tiller production appearedto be associated with synchronized tiller appearance, with alack of synchrony being associated with decreased site usage.Tillers formed in prophyll axils were less likely to be in synchronywith other tillers and frequently failed to appear. In contrastwith branching in dicotyledons, apical dominance appears toplay a minor role in regulating tillering in tall fescue Tillerproduction initially appears to proceed at near maximum ratesthen is down-regulated during later development by longer phyllochrons,slower rate of tiller elongation and reduced site usage. Festuca arundinaceaSchreb., tall fescue, tiller production, leaf appearance, site usage, leaf elongation rate, synchronization, phyllochron, canopy development     

松嫩平原栽培条件下羊草无性系构件的结构

羊草是长根茎型禾草,是典型的无性系植物,在松嫩平原的生长季末期,栽培条件下羊草无性系分株由分蘖株和分蘖苗组成,在具有充分生长空间而又没有种间竞争的风沙土上,羊草分株的分蘖节在一个生长季内可以繁殖4个世代,按分蘖节的繁殖世代划分龄级,现实与潜在无性系构件的年龄谱均以1龄级比重最大,随着龄级的增加明显减少,呈增长型的年龄结构,羊草无性系分株的生产力主要与分株形成及生长的时间长短有关,形成时间越早、生长时间越长的分蘖株对无性系的物质生产和营养繁殖的贡献越大,羊草无性系在空间扩展与物质贮存上具有一定的可调节性。     

Tall fescue EST-SSR markers with transferability across several grass species

Tall fescue (Festuca arundinacea Schreb.) is a major cool season forage and turf grass in the temperate regions of the world. It is also a close relative of other important forage and turf grasses, including meadow fescue and the cultivated ryegrass species. Until now, no SSR markers have been developed from the tall fescue genome. We designed 157 EST-SSR primer pairs from tall fescue ESTs and tested them on 11 genotypes representing seven grass species. Nearly 92% of the primer pairs produced characteristic simple sequence repeat (SSR) bands in at least one species. A large proportion of the primer pairs produced clear reproducible bands in other grass species, with most success in the close taxonomic relatives of tall fescue. A high level of marker polymorphism was observed in the outcrossing species tall fescue and ryegrass (66%). The marker polymorphism in the self-pollinated species rice and wheat was low (43% and 38%, respectively). These SSR markers were useful in the evaluation of genetic relationships among the Festuca and Lolium species. Sequencing of selected PCR bands revealed that the nucleotide sequences of the forage grass genotypes were highly conserved. The two cereal species, particularly rice, had significantly different nucleotide sequences compared to the forage grasses. Our results indicate that the tall fescue EST-SSR markers are valuable genetic markers for the Festuca and Lolium genera. These are also potentially useful markers for comparative genomics among several grass species.Electronic Supplementary Material Supplementary material is available for this article at .     

Persistence and production of perennial grasses under water deficits and extreme temperatures: importance of intraspecific vs. interspecific variability

Extreme climatic events are expected to increase in frequency and magnitude as a consequence of global warming. Grasslands cover a large proportion of the European continent and contribute to both agricultural production and ecosystem services through inter and intraspecific genetic variability. This study analysed the effects of summer droughts and heat waves on the persistence and production of perennial forage grasses. Mediterranean and temperate populations of Dactylis glomerata L. and Festuca arundinacea (Schreb.) were compared at both Mediterranean and temperate sites in France. By manipulating canopy temperatures and water availability, grass swards in the field were subjected to cumulative summer and spring water deficits (CSSWD) ranging from 329 to 707 mm to test different projected climatic conditions and extreme summer events. Under controlled summer heat waves (6–21 days at a mean daily canopy temperature higher than 30–35 °C), there was no increase in membrane damage to surviving aerial tissues. Plant stress was thus mainly generated through greater soil water deficit. Under the greatest CSSWD, annual biomass production was reduced on average by 60% and 30% with temperate and Mediterranean populations, respectively. Thresholds for a significant increase in summer tiller mortality were seen at CSSWD higher than 450 mm for temperate populations and 550 mm for Mediterranean populations. The latter displayed lower predawn leaf water potentials in summer and recovered through intense tillering in the subsequent seasons. Under the most extreme CSSWD, fewer than 20% of tillers of temperate populations survived and their nitrogen uptake ability was drastically altered. The higher potential productivity of Mediterranean populations in winter was associated with greater frost sensitivity. The identification of thresholds for vulnerability and the determination of the role of genetic diversity will improve the management of plant resilience and the design of new plant material to cope with climate change.     

Mealybug, Phenococcus solani, and barley aphid, Sipha maydis, response to endophyte-infected tall and meadow fescues

Mealybugs and aphids are insects which damage grass species. The effects of fungal endophytes on the feeding of the mealybug, Phenococcus solani Ferris (Homoptera: Pseudococcidae), and barley aphid, Sipha maydis Passerini (Homoptera: Aphididae), on tall fescue, Festuca arundinacea Schreb. and meadow fescue, Festuca pratensis Huds., were studied under greenhouse conditions. Mealybugs preferred endophyte‐free (E–) clones over their endophyte‐infected (E+) counterparts. E+ plants had a significantly lower number of mealybugs than E– plants. A mixture of E+ and E– plants supported intermediate mealybug numbers, between pure plantings of E+ and E– grasses. Barley aphids released on to plant materials were deterred from feeding and could not persist on E+ plants. E– plants did not survive because of aphid damage, while E+ plants generally re‐grew, but were damaged to some degree. The results showed that the use of pure stands of endophyte‐infected grasses or a mixed stand of infected and non‐infected plants may increase the persistence and durability of turf and forage grass species in the presence of foliar damaging insects.     

Relationship between Contents of Chlorophyll (a+b) (SPAD values) and Nitrogen of Some Temperate Grasses

In a field experiment the chlorophyll (a+b) (SPAD readings) and nitrogen contents of three grass species (Festuca arundinacea Schreb., Lolium perenne L., and Lolium multiflorum Lam.) and three intergeneric hybrids of Festuca pratensis Huds. × Lolium multiflorum Lam. and Festuca arundinacea Schreb. × Lolium multiflorum Lam. were measured. Close relationships were found between SPAD readings and nitrogen leaf content (r ² = 0.873^** 0.491^** and 0.938^**) for the 1^st, 2^nd, and 3^rd cut, respectively. SPAD readings and N contents were closely correlated (r = 0.836^**) confirming that SPAD measurements could be used in grass selection and/or breeding for high N concentration in herbage. This revised version was published online in August 2006 with corrections to the Cover Date.     

Vernalization,gibberellic acid and photo period are important signals of yield formation in timothy (Phleum pratense)

Timothy (Phleum pratense) is a widely grown perennial forage grass in the Nordic region. The canopy consists of three tiller types, of which the stem forming vegetative elongating (ELONG) tiller and generative (GEN) tillers contribute the most to dry matter yield. In this study, the regulation of tiller formation by vernalization, day length (DL) [12 h, short day length (SD); 16 h, long day length (LD)] and gibberellic acid (GA) was investigated in two timothy cultivars. Vernalization resulted in a shift of ELONG to GEN tillers. No vernalization was required for the development of ELONG tillers but SD strictly arrested stem elongation. Vernalization is an important regulator of tiller development but it seemed to be upstream regulated by DL. LD was essential for floral transition and could not be substituted by GA and/or vernalization treatments. Genotypic variation was found in the development of GEN tillers. The ability to produce GEN tillers was associated with significant upregulation of PpVRN3. PpVRN1 expression peaked at the time of vegetative/generative transition, and PpVRN3 after the transfer to LD, suggesting them to have similar functions with cereal vernalization genes. PpVRN1 alone was not sufficient to activate flowering, and upregulation of PpVRN3 possibly together with PpPpd1 was required. Although vernalization downregulated PpMADS10, this gene did not act as a clear flowering repressor. Our results show that flowering signals alter the tiller composition, so they have important effects on yield formation of timothy.     

Influence of endophyte infection, plant age and harvest interval on Rhopalosiphum padi survival and its relation to quantity of N-formyl and N-acetyl loline in tall fescue

Rhopalosiphum padi L. (Homoptera: Aphididae) is sensitive to loline alkaloids present in tall fescue, Festuca arundinacea Shreb., infected with the endophytic fungus, Acremonium coenophialum Morgan-Jones & Gams. Aphid survival was higher on endophyte-free plants regardless of plant age after germination or age of regrowth tissue after clipping. Survival of aphids on endophyte-infected grass was lower on young tissue but increased as plants aged, although it never reached the same level on endophyte-free plants. Both N-formyl and N-acetyl loline increased as uncut or regrowth tissue aged; however, this was influenced by the age of the plant at the initial cut and the clipping frequency. Although even small amounts of loline cause high aphid mortality, the aphids are able to survive on endophyte-infected plants if the tillers have senescing leaves which contain lower amounts of loline. Preference for senescing leaves may help R. padi avoid plant parts containing high amounts of toxic allelochemicals, thus contributing to higher numbers of aphids on older, endophyte-infected plants.     

Expression of the bacteriophage T4 lysozyme gene in tall fescue confers resistance to gray leaf spot and brown patch diseases

Tall fescue (Festuca arundinacea Schreb.) is an important turf and forage grass species worldwide. Fungal diseases present a major limitation in the maintenance of tall fescue lawns, landscapes, and forage fields. Two severe fungal diseases of tall fescue are brown patch, caused by Rhizoctonia solani, and gray leaf spot, caused by Magnaporthe grisea. These diseases are often major problems of other turfgrass species as well. In efforts to obtain tall fescue plants resistant to these diseases, we introduced the bacteriophage T4 lysozyme gene into tall fescue through Agrobacterium-mediated genetic transformation. In replicated experiments under controlled environments conducive to disease development, 6 of 13 transgenic events showed high resistance to inoculation of a mixture of two M. grisea isolates from tall fescue. Three of these six resistant plants also displayed significant resistance to an R. solani isolate from tall fescue. Thus, we have demonstrated that the bacteriophage T4 lysozyme gene confers resistance to both gray leaf spot and brown patch diseases in transgenic tall fescue plants. The gene may have wide applications in engineered fungal disease resistance in various crops.     

Leaf and Tiller Production of Prairie Grass (Bromus willdenowii Kunth) and Two Ryegrass (Lolium) Species

A detailed morphological study of three prairie grass cultivars(Bromus willdenowii Kunth) was conducted under ‘vegetative’and ‘reproductive’ growth conditions (short andlong photoperiods) and at different temperatures. Perennialryegrass (Lolium perenne L.) and Westerwolds ryegrass (Loliummuhiflorum Lam.) were compared during vegetative growth. Prairie grass had higher leaf appearance rates (leaves per tillerper day) and lower site filling (tillers per tiller per leafappearance interval) than the ryegrass species. Tillering rates(tillers per tiller per day) were also lower, except under vegetativeconditions at 4C. Low tiller number in prairie grass was notdue to lack of tiller sites but a result of poor filling ofthese sites. Lower site filling occurred because of increaseddelays in appearance of the youngest axillary tiller and lackof axillary tillers emerging from basal tiller buds. In prairiegrass, no tillers came from coleoptile buds while only occasionallydid prophyll buds develop tillers. Low tiller number in prairiegrass was compensated for by greater tiller weight. Prairiegrass had more live leaves per tiller, greater area per leafand a high leaf area per plant. Considerable variation between cultivars was found in prairiegrass. The cultivar ‘Bellegarde’ had high leaf appearance,large leaves and rapid reproductive development, but had lowlevels of site filling, tillering rate, final tiller numberand herbage quality during reproductive growth. ‘Primabel’tended to have the opposite levels for these parameters, while‘Grasslands Matua’ was intermediate and possiblyprovided the best balance of all plant parameters. prairie grass, Bromus willdenowii Kunth, perennial ryegrass, Lolium perenne L., Westerwolds ryegrass, Lolium multiflorum Lam., temperature, photoperiod, leaf appearance, leaf area, tillering, site filling, tillering sites, yield     

Enhanced tolerance of transgenic tall fescue plants overexpressing 2-Cys peroxiredoxin against methyl viologen and heat stresses

Plant 2-Cys peroxiredoxins (2-Cys Prx) has both peroxidase and chaperon function. We overexpressed an Arabidopsis 2-Cys Prx in transgenic tall fescue (Festuca arundinacea) plants to confer tolerance against heat and methyl viologen (MV) stress. Transgenic plants were generated by Agrobacterium-mediated genetic transformation, and integration and expression of the transgene was confirmed by Southern, northern and western blot analyses. Compared to control plants, transgenic plants had significantly less electrolyte leakage and thiobarbituric acid-reactive substances (TBARS) when exposed to heat or MV. Under heat stress (42°C), transgenic plants maintained their chlorophyll fluorescence (Fv/Fm) for 24 h while control plants lost chlorophyll fluorescence very quickly. We conclude that the high levels of 2-Cys Prx proteins in transgenic plants protect leaves from oxidative damage probably due to chaperon activity.