The Growth and Carbon Economy of Selection Lines of Lolium perenne cv. S23 with Differing Rates of Dark Respiration: 2. Grown as Young Plants from Seed

Young plants of two selection lines of Lolium perenne cv. S23with ‘fast’ and ‘slow’ rates of ‘maturetissue’ respiration were individually grown from seed,together with plants of S23, their common parent, in 9.2 cmpots in a controlled environment at 20/15 °C day/night temperatures. No significant differences were found between the genotypesin leaf extension and tiller production during this early stageof their growth. They did differ however, by an average of 26%,in the rate of dark respiration of fully expanded leaf laminae.The use of a simple model demonstrated that such a differencein respiration could alone account for the different rates ofdry matter production shown by the selection lines when grownas young crops from seed. Possible penalties of ‘slow’respiration are also considered. Lolium perenne L., ryegrass, respiration, maintenance respiration, stimulated swards, leaf growth, tiller production, carbon economy     

The Growth and Carbon Economy of Selection Lines of Lolium perenne cv. S23 with Differing Rates of Dark Respiration: 1. Grown as Simulated Swards During a Regrowth Period

Simulated swards of each of two selection lines of Lolium perennecv. S23 with ‘fast’ and ‘slow’ ratesof ‘mature tissue’ respiration were establishedin growth rooms at 20/15 °C day/night temperatures and studiedover four successive regrowth periods of 46, 30, 26 and 53 daysduration. The ‘slow’ line outyielded the ‘fast’,both in harvestable shoot (above a 5 cm cut) and in root andstubble. Its advantage increased over successive regrowth periodsto 23 per cent (total biomass). Gas analysis measurements onthe entire communities (including roots), during the final regrowthperiod, showed that the ‘slow’ line had a 22–34per cent lower rate of dark respiration per unit dry weight.This enabled it to maintain its greater mass of tissue for thesame cost in terms of CO₂ efflux per unit ground area. Halfthe extra dry weight produced by the ‘slow’ line,relative to the ‘fast’, could be attributed to itsmore economic use of carbon. The rest could be traced to a 25per cent greater tiller number which enabled the ‘slow’line to expand leaf area faster (though not at a greater rateper tiller), intercept more light and fix more carbon, earlyin the regrowth period. Lolium perenne L., ryegrass, respiration, maintenance respiration, tiller production, simulated swards, canopy photosynthesis, carbon economy     

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     

Effect of Cutting on Production and Tillering in Prairie Grass (Bromus willdenowii Kunth) Compared With Two Ryegrass (Lolium) species. 2. Reproductive Plants

Effects of time of initial cut and subsequent cutting interval(1, 2 or 4 weeks) were examined in a glasshouse during summer1988 for reproductive plants of three prairie grass cultivars(Bromus willdenowii Kunth), Westerwolds ryegrass (Lolium multiflorumLam.) and perennial ryegrass (Lolium perenne L.). Measurementswere made of tiller and leaf numbers, sites of tillering, reproductivedevelopment, and herbage quality and yields. Effect of timeof initial cutting on regrowth appeared to be independent ofstage of reproductive development and unrelated to any of themeasured plant parameters. Characteristics for each cultivaras identified during undisturbed growth prior to the initialcuts, were modified by the subsequent cutting frequencies. Perennial ryegrass had the highest yields under frequent cuttingwith high herbage quality. Westenwolds ryegrass and the prairiegrass cultivars ‘Grasslands Matua’ and ‘Primabel’had the highest yields with infrequent cutting, but lower herbagequality than in perennial ryegrass. At each cut, tiller deathin prairie grass was determined by the number of reproductivetillers, and in the ryegrasses also by the numbers of elongatedvegetative tillers. Recovery of tiller numbers was rapid andprimarily from inhibited tiller buds at the base of reproductivetillers. In prairie grass, tiller numbers were relatively unaffectedby cutting frequency, but at frequent cutting, many axillarytillers originated from vegetative tillers rather than frominhibited tiller buds at the base of reproductive tillers. Prairie grass, Bromus willdenowii Kunth, Westerwolds ryegrass, Lolium multiflorum Lam., perennial ryegrass, Lolium perenne L., first (initial) cut, cutting frequency, regrowth, leaf appearance, tillering, yield, nitrogen, water-soluble carbohydrates, herbage quality     

Primary and Secondary Induction Requirements for Flowering of Contrasting European Varieties of Lolium perenne

The flowering requirements of six European varieties of Loliumperenne L. were studied in controlled environments. In experimentson primary induction, flowering was recorded after transferto long days (LD) in a greenhouse at 12–24°C. In experimentson secondary induction, primary induction was first accomplishedat 6°C/10 h daylength for 12 weeks. When evaluated by the50% heading criterion, the requirement for duration of primaryinduction at 6°C/8 h daylength was <3 weeks in Mediterranean,5–6 weeks in Central European and 7–8 weeks in Scandinavianvarieties. While ‘Veyo’ (Italy) flowered profuselyregardless of temperature or daylength during primary induction,critical temperatures for primary induction in SD and LD were15 and 11°C in ‘Baca’ (Czech Republic) and 11and 7°C in ‘Falster’ (Denmark). The criticalphotoperiod for secondary induction at 15°C ranged from12 h in ‘Veyo’ and 14 h in ‘Baca’ to16.5 h in ‘Falster’ and 17.5 in ‘Kleppe’(Norway). The critical number of LD cycles varied correspondingly.While the Central and North European varieties required fewerLD cycles for 50% heading at 18 than at 12°C, ‘Veyo’showed the opposite response. It is concluded that the requirementsfor both primary and secondary induction of Lolium perenne increasewith increasing latitude of origin of the germplasm. In oneexperiment, 39–87% of the inflorescences came from tillersthat were not visible on transfer from primary to secondaryinduction, thus it is also concluded that there is no juvenilestage in tillers of Lolium perenne. Copyright 2000 Annals ofBotany Company Daylength, flowering, juvenility, perennial ryegrass (Lolium perenne L.), primary induction, secondary induction, temperature, varieties, vernalization     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.): 3. Partition of Assimilates between Root and Shoot during the Transition from Vegetative to Reproductive Growth

The partition of ¹⁴C labelled current assimilates to root insimulated swards of Lolium perenne cv. S24 was measured duringthe transition from vegetative growth in autumn to reproductivegrowth in spring under close to natural conditions of lightand temperature. Assimilate partitioning was also measured in‘established’ swards cut three times during thegrowing season and in vegetative ‘seedling’ swardsgrowing in autumn and in spring. All measurements were madewhen the swards had achieved more than 90 per cent light interception,and all swards were abundantly supplied with water and mineralnutrients. During autumn there was a gradual decrease in the proportionof assimilates partitioned to the roots in both the ‘established’and the ‘seedling’ swards. In the established swards,partition to roots was low over winter, increased during earlyspring, but decreased dramatically, later in the spring, whenstem elongation began. In contrast, in the unvernalized vegatativeseedling swards in spring, partition to roots remained high. The seasonal pattern of assimilate partitioning is consideredin relation to changes in the natural environment and the rateat which the crop fixed carbon in photosynthesis. A decreasein the proportion of assimilates partitioned to roots duringlate spring was significant in increasing the production ofshoot at that time but seasonal differences in partition contributedvery little to the marked differences in shoot growth betweenthe spring and autumn crop. Lolium perenne L., perennial ryegrass, partition of assimilates, flowering     

Response to Selection for Dark Respiration Rate of Mature Leaves in Lolium perenne and its Effects on Growth of Young Plants and Simulated Swards

Selections for slow and for fast rate of dark respiration ofmature leaves were made from within Lolium perenne cv. S23.Selected parents were pair-crossed to provide 15 F₁ familieswith slow respiration and 15 with fast. Dark respiration was inherited and families with contrastingrates were subjected to sequentially harvested growth analysesfrom the third leaf stage to that of 95 per cent light interceptionin a growth room. Seven periods of regrowth of simulated swardsof the families were then recorded. During development of theprimary canopy, growth of the selections did not differ untilthe final harvest interval. At this stage slow respiration familieshad faster (P < 0.05) net assimilation rate and greater plantdry weight (P < 0.05) and leaf area index (P < 0.05) thanthe fast respiration group. Relative growth rate followed thesame trend. In the swards after each regrowth dry matter yieldof the slow respiration group was greater than that of the fast. In another experiment, simulated swards of six slow respirationfamilies yielded more than swards of six fast respiration familiesover sequential regrowth periods in a glasshouse from May toNovember: S23 was intermediate. Differences were most duringAugust and September. Crop growth rate at each harvest correlated(P < 0.05 or P < 0.01) with previously determined leafrespiration at 25 °C. Leaf protein levels in August weaklycorrelated (r = +0.57, P < 0.05) with respiration rate perunit dry weight but there was a significant residual negativecorrelation (r = –0.67, P < 0.05) between the rateper unit protein and growth at that time. Results are discussedin relation to the concept of ‘maintenance-relàted’respiration. Lolium perenne L., perennial ryegrass, respiration, maintenance respiration, relative growth rate, leaf area ratio     

Associations between enzyme genotypes and dark respiration in perennial ryegrass,Lolium perenne L.

Summary In this study, we determined whether relationships existed between dark respiration and genotype at five enzyme polymorphisms in perennial ryegrass, Lolium perenne L. Positive correlations were found between Q ₁₀ of dark respiration and genotype at the phosphoglucomutase (PGM) and 6-phosphogluconate dehydrogenase (6PGD) loci. Plants doubly homozygous for the common allele at these loci were found to have Q ₁₀ values 20% higher than those for double heterozygotes. In plants that were heat stressed for five consecutive days, Q ₁₀ was found to be negatively correlated with apparent vigor after stressing. Individuals homozygous for PGM and 6PGD (with higher Q ₁₀ values) exhibited more apparent damage following the stress than heterozygous individuals. Both PGM and 6PGD occupy positions in metabolism with regulatory potential. Although caution must be used in assigning causal relationships, the results suggest that specific forms of these enzymes are directly related to, or are correlated with, the determinants of respiratory efficiency in L. perenne.     

The Growth of Perennial Ryegrass: A Model

The annual pattern of production of the perennial ryegrass cropis largely affected by the seasonal weather pattern and thechange from the vegetative to the reproductive phase of growth. Analysis of the relative importance of the factors governingthe growth of the reproductive crop and the regrowth of thevegetative crop is complex. Often physiological variables arecorrelated as well as climatic variables, which influence thephysiology and morphology of the crop. Much experimental data has been collected, in the field andunder controlled environments, concerning the growth of thegrass crop. This information has been used to construct a mathematicalmodel of the crop. In order to evaluate the problems of modellingthe grass crop the model has been used to predict the net growthof the crop in terms of leaf, ‘stem’, root and deadmatter using daily irradiance data. The influence of some cropcharacteristics on vegetative regrowth have been considered.The model has also been used to assess the relative importance,for yield, of some physiological characteristics of the reproductivegrass crop. Lolium perenne, perennial ryegrass, growth, mathematical model, yield     

Effect of Selection for Dark Respiration Rate of Mature Leaves on Crop Yields of Lolium perenne cv. S23

Growth and nutritive quality of two polycross progeny (GL 83and GL 112) from F₁ plants of Lolium perenne ex cv. S23 showingslow dark respiration rates of mature leaves were compared inthe field with S23 in 3 years. Plots were cut either nine (simulatedgrazing) or five (conservation) times annually. In the firsttrial, sown in 1977, dry matter yields from plots of GL 83 werecompared with S23 during 1978 and 1979. A similar trial sownin 1979 compared GL 83, S23 and GL 112 during 1980. The slowrespiration populations displayed consistently greater (6–13per cent) annual dry matter yields than S23, mostly during midto late summer. In the first trial the difference in yield wasgreater in 1979 than 1978 and more under conservation than simulatedgrazing. In the second trial GL 83 and GL 112 yielded 11 and13 per cent more dry matter than S23 under simulated grazingand 5–6 per cent under conservation. There were no consistentdifferences in dry matter digestibility, water soluble carbohydratesor protein between the populations. Cellulose content of GL83 and to a much lesser extent GL 112, tended to be greaterthan that of S23 from July to September. Analysis of one harvestfrom another trial revealed no significant difference in energyvalue, as measured by adiabatic bomb calorimetry, between GL83 and S23. Thus, it appears that the greater dry matter productionof the progeny with slower mature leaf dark respiration ratesreflected a true increase in total energy yield not apparentlyassociated with any agronomically undesirable characteristics.The general significance of the results for crop improvementare discussed. Lolium perenne L., perennial ryegrass, yield of dry matter, energy value, dry matter digestibility     

Response of Populations of Lolium perenne cv. S 23 with Contrasting Rates of Dark Respiration to Nitrogen Supply and Defoliation Regime 2. Grown as Mixtures

Two populations of perennial ryegrass cv. S 23, selected forcontrasting rates of mature leaf tissue respiration, were grownin 50: 50 replacement mixtures to investigate their relativecompetitive abilities in a range of conditions. The 18 mixedsimulated swards were subject to three levels of nitrogen supply(14.5, 32 and 173.5 ppm) from establishment, and to three cuttingfrequencies (at 1-, 3- and 6-week intervals), 7 weeks aftersowing. The slow-respiring population yielded more than the fast. Thissuperiority was greater in mixtures (50%) than in monocultures(10%). Its superiority was as great in weekly-cut swards (51%), where there was little or no aerial competition, as in 6-weekly-cutswards (50%), where shoot interference was maximal, indicatingthat its advantage lay in below-ground competition. The limitingresource for which competition occurred could not be positivelyidentified. Lolium perenne, perennial ryegrass cv. S 23, population, simulated sward, defoliation, nitrogen, respiration, dry-matter production, 50: 50 replacement mixture     

Response of Populations of Lolium perenne cv. S 23 with Contrasting Rates of Dark Respiration to Nitrogen Supply and Defoliation Regime: 1. Grown as Monocultures

Simulated swards of two populations of perennial ryegrass cv.S 23 selected for contrasting rates of mature leaf tissue respirationwere grown in a glasshouse. From establishment, the swards weresubject to three levels of nitrogen supply (14.5, 32 and 173.5ppm N) and from the first harvest 7 weeks after sowing, to threecutting frequencies (at intervals of 1, 3, and 6 weeks). Throughout the 18-week experiment, the herbage yield and themean tiller weight of the slow-respiring population were 10%greater on average than those of the fast-respiring population.Increasing nitrogen supply (from 14.5 to 173.5 ppm N) enhancedthe yield advantage of the slower-respiring population—moreso under infrequent cutting (from nothing to 22%) than underfrequent cutting (from 6 to 13%). Both maximum absolute yields,and the greatest yield advantage of the slow-respiring populationover the fast, were achieved when high nitrogen was combinedwith infrequent defoliation. Lolium perenne, perennial ryegrass cv. S23, respiration, nitrogen defoliation, simulated sward, dry-matter production, monoculture, population     

Short-day and Low-temperature Induction in Lolium

The requirements for floral induction of several outbreedingpopulations of Lolium perenne, L. multiflorum, and L. rigidumhave been studied. Floral induction can be brought about byeither low temperature (0–30° C.) or short day (8hr.) independently, but short day is ineffective at low temperatures. The exact inductive requirement varies with the population.The summerannual Westerwolds ryegrass needs neither cold norshort day, the winter-annual Wimmera ryegrass shows a quantitativeresponse, while the perennial varieties from north-west Europehave an obligatory requirement for either cold or short daybefore floral induction. This relation between the annual orperennial habit and the inductive requirement was confirmedin a number of Mediterranean collections of L. perenne and L.rigidum. Considerable genetic variation in inductive requirement couldbe detected within each outbreeding population, and rapid responseto selection proved possible in Wimmera, Irish, and Kent ryegrass.     

Development of a genomic microsatellite library in perennial ryegrass (Lolium perenne) and its use in trait mapping

Background and Aims: Perennial ryegrass (Lolium perenne) is one of the key forageand amenity grasses throughout the world. In the UK it accountsfor 70 % of all agricultural land use with an estimated farmgate value of £6 billion per annum. However, in termsof the genetic resources available, L. perenne has lagged behindother major crops in Poaceae. The aim of this project was thereforethe construction of a microsatellite-enriched genomic libraryfor L. perenne to increase the number of genetic markers availablefor both marker-assisted selection in breeding programmes andgene isolation. Methods: Primers for 229 non-redundant microsatellite markers were designedand used to screen two L. perenne genotypes, one amenity andone forage. Of the 229 microsatellites, 95 were found to showpolymorphism between amenity and forage genotypes. A selectionof microsatellite primers was selected from these 95 and usedto screen two mapping populations derived from intercrossingand backcrossing the two forage and amenity grass genotypes. Key Results and Conclusions: The utility of the resulting genetic maps for analysis of thegenetic control of target traits was demonstrated by the mappingof genes associated with heading date to linkage groups 4 and7.     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.). 1. Response of Leaf Extension to Temperature during the Transition from Vegetative to Reproductive Growth

The potential for leaf extension of plants of Lolium perennecv S24 growing in small artificial communities under naturalconditions was measured as the plants progressed from the vegetativeto the reproductive state In two consecutive years, 1975 and 1976, ‘simulated swards’were sown in autumn and overwintered in an open, unheated glasshouseIndividual swards from the batch sown in 1975 were brought into a growth room on two occasions in spring 1976 to measuretheir potential for leaf extension at a range of temperatures(5–20 °C) Individual swards from the batch sown inautumn 1976 were brought in to the growth room on 15 occasionsbetween November 1976 and May 1977 and their potential for leafextension was measured at a single temperature of 15 °CFrequent dissections were made in both years to describe changesin the developing apex. The potential for leaf extension at 15 °C decreased fromc 17 mm day^–1 in November to c 10 mm day^–1 in mid-winter.In January, the potential rapidly increased threefold to reach30mm day^–1 by mid February The increase began coincidentwith the earliest stages of floral initiation and was completedby the time of double-ridge formation Spring-grown vegetativeplants, however, showed potential rates of < 20 mm day^–1at 15 °C The results are discussed in relation to reproductive developmentand to changes in the carbohydrate strategy of the plants inearly spring Lolium perenne L perennial ryegrass, leaf extension, temperature response     

Glucose utilization by 6pgd genotypes in Lolium perenne

To gain a deeper understanding of the mechanisms underlying associations between allozyme genotypes and rates of respiration in Lolium perenne, V_max K_m and rates of glucose flux through glycolysis and the pentose phosphate pathway were estimated for the three genotypes of the 6pgd locus. K_m V_max and V_max/K_m differed significantly among genotypes. Values of K_m for the 11, 12, and 22 genotypes were 0.29, 0.25, 0.13, while the values of V_max/K_m for die 11, 12, and 22 genotypes were 3.79, 3.85, 6.70. Flux through the pentose shunt did not differ among genotypes at 20 °C, but at 35 °C the rates of flux for the 11, 12, and 22 genotypes were 0.15, 0.25 and 0.42, respectively. Thus, the 6PGD allozyme genotypes differ markedly in both enzyme kinetic characteristics and in flux through a metabolic pathway. These associations reveal potentially causal relationships between allozyme genotypes and rates of respiration.     

Effect of Cutting on Production and Tillering in Prairie Grass (Bromus willdenowii Kunth) Compared With Two Ryegrass (Loliunt) Species. 1. Vegetative Plants

Effects of cutting to 3 or 6 cm stubble height at frequenciesof 1, 2 or 4 weeks were investigated in young, vegetative, spacedplants of prairie grass (Bromus willdenowii Kunth), a tetraploidWesterwolds ryegrass (Lolium multiflorum Lam.) and perennialryegrass (Lolium perenne L.). Plants were grown for 8 weekson nutrient solution in a growth cabinet at day/night temperaturesof 15/10°C and in a short photoperiod. Leaf appearance,site filling and tiller numbers were all reduced by both increasedcutting frequency and a lower cutting height. The extent ofthese reductions was least in prairie grass and greatest inWesterwolds ryegrass. Relative growth rates in all species respondedin a similar manner with more frequent cutting and a lower cuttingheight. Cutting frequency had the greatest effect on growthrates, although the effect of cutting height increased withtime. Results confirm general recommendations of long intervalsbetween defoliations to achieve high yields for vegetative prairiegrass, but the relative amounts of dry matter harvested underthe different cutting regimes may be no different to that obtainedin ryegrass. Stubble height was of lesser importance in determiningyields. A typical U-shaped curve for depletion and recovery in water-solublecarbohydrates occurred after defoliation in all species, butlevels remained low at frequent cutting. Water-soluble carbohydratelevels in stubble and roots were higher in prairie grass. Regrowthat the end of the experiment was highly correlated with totalstubble and root weights (r = 0.84), while regrowth per tillershowed a good correlation with water-soluble carbohydrate content,although the response varied between species. Prairie grass, Bromus willdenowii Kunth, perennial ryegrass, Lolium perenne L., Westerwolds ryegrass, Lolium multiflorum Lam., cutting frequency, cutting height, leaf appearance, tillering, site filling, yield, regrowth, nitrogen, water-soluble carbohydrates     

Water Use Characteristics of Dactylis glomerata L., Lolium perenne L. and L. multiflorum Lam. Plants

Cocksfoot (Dactylis glomerata L.). perennial ryegrass (Loliumperenne L.) and Italian ryegrass (L. multiflorum Lam.) plantswere grown on deep (75–95 cm) columns of soil in glasshousesand growth rooms with and without irrigation. The species inwhich growth declined least rapidly after water had been withheldwere those which transpired most slowly. During early establishmentin the glasshouse cocksfoot transpired least because of slowroot growth. In the growth room, when root systems were deeperand denser, perennial ryegrass transpired least because of lowleaf water conductance. Results are discussed in relation to(a) drought resistance in the three species; (b) breeding forincreased drought resistance through modifying root distributionand leaf water conductance; and (c) the use of isolated soilcolumns in water relations studies. Dactylis glomerata L., Lolium perenne L., Lolium multiflorum Lam., cocksfoot, perennial ryegrass, Italian ryegrass, transpiration, roots, leaf water conductance     

Flowering of S. 24 Perennial Ryegrass (Lolium perenne L.) in Short Days at Low Temperature

The flowering of the quantitative long-day plant perennial ryegrass(Lolium perenne L.) in short days (8 h), when grown at low temperature(9/4 °C) and under natural summer daylight, is presentedas evidence for the replacement of specific environmental requirementsfor flowering by alternative stimuli in a grass.