Quantitative Analysis of Cold Hardening and Dehardening in Lolium

The change in cold hardiness of three Lolium multiflorum Lam.varieties was followed in plants exposed to hardening or dehardeningconditions at a range of temperatures. Hardening and dehardeningwere analysed as quantitative processes dependent upon temperatureand time. Their time courses changed exponentially to an asymptotewhich was logistically related to temperature. Both componentsof the model were fitted simultaneously. Parameters of biologicalinterest, such as the initial rates of hardening and dehardeningfor a given temperature and the percentage of the process completedin a given time, were derived and compared for varieties ofcontrasting hardiness. The analysis demonstrated the importanceof dehardening in determining hardiness. Similar results wereobtained when the model was applied to hardening and dehardeningdata for Lolium perenne L. The potential of this quantitative analysis for distinguishingbetween possible mechanisms of cold hardiness is discussed andfurther experiments required to characterise the kinetics ofhardening and dehardening more fully are identified. Lolium multiflorum, Lolium perenne, cold hardening, cold dehardening, processes, model, varieties     

Frost tolerance and biochemical changes during hardening and dehardening in contrasting white clover populations

Successful winter survival of perennial plants, like white clover, is dependent on proper timing of both hardening and dehardening. The purpose of this study was to investigate the regulation of these processes in two cultivars (AberCrest and AberHerald) and two Norwegian ecotypes (Særheim collected at 58°46′N lat. and Bodø at 67°20′N lat.) of white clover (Trifolium repens L.). For hardening and dehardening, plants were exposed to controlled temperature conditions and frost hardiness of stolons was tested by programmed freezing at the rate of 3°C per hour. In addition, stolons were analysed for starch, soluble sugars and soluble amino acids. Cultivars AberCrest and AberHerald, selected for growth at low temperature and winter hardiness in the United Kingdom, were significantly less hardy than the Norwegian populations. After six weeks of hardening (2 weeks at 6°C and 4 weeks at 0.5°C), estimated LT₅₀ values were ?13.8, ?13.0, ?17.8 and ?20.3°C for AberCrest, AberHerald, Saerheim and Bodø, respectively. The rate of dehardening increased with increasing temperature. At low temperature (6°C), the northern ecotype from Bodø was more resistant to dehardening than AberHerald. However, at 18°C the absolute rate of dehardening (°C day^?1) was twice as high in Bodø as in AberHerald plants. Stolon elongation during dehardening was initiated at lower temperatures in AberHerald than in plants of the Bodø ecotype. The content of total soluble sugars, sucrose and the amino acids proline and arginine were significantly higher in hardy plants of Bodø than in those of AberHerald. Sucrose levels decreased during dehardening and correlations between sucrose content and LT₅₀ during this process were statistically highly significant for both Bodø and AberHerald. The least hardy populations of white clover were characterized by thick stolons, long internodes and large leaves.     

Regulation of the loss of frost hardiness in Pinus radiata by photoperiod and temperature

Abstract. In controlled environments, the interactive effects of warm (16: 8°C, day: night) and cool (12: 4°C, day: night) temperatures and long (13.5 h) and short (10 h) photoperiods on the dehardening of seedlings of Pinus radiata D. Don were investigated. In another experiment, the effect of four photoperiods from 9 to 14 h was examined. In a third, dehardening at constant temperatures from 5 to 17°C was followed. There was no evidence for an interaction between photoperiod and temperature. Dehardening was temporarily delayed by photoperiods below about 10 h, but there was no other quantitative effect of photoperiod. At constant temperatures, the rate of dehardening was initially constant but declined as the minimum summer frost hardiness was reached. In the initial phase the rate of dehardening was a linear function of temperature, increasing from 0.05°C day⁻¹ at 8°C to 0.30 °C day⁻¹ at 17°C. Temperature controlled the loss of frost hardiness by regulating the rate of dehardening.     

Effect of Photoperiod and Irradiance Changes Upon Development of Freezing Tolerance and Accumulation of Soluble Carbohydrate in Seedlings of Lolium perenne Grown at 2 {degrees}C

Two contrasting cultivars of Lolium perenne were exposed toa range of daily radiation integrals during hardening at 2°Cfor 15 d. The maximum induced freezing tolerance measured asLT₅₀ (temperature for 50 % kill) differed markedly between thecultivars. The observed LT₅₀ values were unaffected by changesin the radiation integral above 10 mol m^–2 d^–1,whereas accumulation of water-soluble carbohydrate showed astrong positive correlation with the radiation integral overthe entire range of the experiment. The correlation betweenLT₅₀ and soluble carbohydrate content at the end of the hardeningperiod was poor and showed no obvious connection with genotype.Fructan polymers and sucrose were the major components of thesoluble carbohydrates in both cultivars. The depression of freezingpoint attributable to the accumulation of soluble, osmoticallyactive carbohydrate was not sufficient to account for the observedchanges in LT₅₀ in the hardy genotype. These results are discussedin relation to the interactions between growth, photosynthesisand assimilate partitioning during hardening. Lolium perenne, hardening, freezing tolerance, irradiance, carbohydrate, fructan     

Application of Impedance Spectroscopy for Selecting Frost Hardy Varieties of English Ryegrass

The frost hardening and frost damage of 12 varieties of Englishryegrass (Lolium perenne) was studied by electrical impedancespectroscopy. For the measurement of the impedance spectrum(80 Hz to 1 MHz) a 10 mm length sample was cut from the stemabove the growing point, but the growing point was included.The impedance spectra were analysed by an asymmetric distributedcircuit model. The impedance spectra were measured at two phasesof hardiness and after freezing, i.e. (a) before hardening,(b) after hardening in controlled conditions, and (c) aftercontrolled frost exposure at -16 °C after hardening. Twomodel parameters, i.e. intra- and extracellular resistance,increased with hardening. The intracellular resistance and theskewness factor before hardening, and the ratio between thosetwo parameters before and after hardening, strongly correlatedwith hardening of different varieties of English ryegrass. Theextracellular resistance and the relaxation time decreased asa result of the frost exposure at -16 °C. Cold acclimation; electrical impedance; English ryegrass; frost hardiness; impedance spectroscopy; Lolium perenne     

The Effect of Low Temperature upon Starch, Sucrose and Fructan Synthesis in Leaves

Measurements on plants in a temperature gradient tunnel showthat diurnal accumulation of starch in illuminated leaves wasgreatly reduced at temperatures below 8 °C, whereas sucrosesynthesis was less affected under similar conditions. High chillingsensitivity for leaf starch accumulation was observed in a numberof chilling resistant temperate species. Enzymes of sucroseand fructan metabolism from mature leaves of Lolium temulentumwere less strongly inhibited at low temperatures than enzymesinvolved in starch synthesis. These results are discussed inrelation to carbon partitioning in species which grow and aremetabolically active at chilling temperatures. Lolium temulentum, starch, sucrose, fructan, temperature, enzyme activity, carbon partitioning     

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     

Changing Environmental Effects on Frost Hardiness of Scots Pine During Dehardening

The effects of raised temperature and extended photoperiod onthe dehardening of quiescent and winter-hardy Scots pine saplingswere examined in an open-top-chamber experiment. The saplingswere exposed during winter to natural, square-curve fluctuating(between 1 and 11 °C with a 14 d interval), and constant(6 °C) temperatures with a natural and an extended (17 h)photoperiod. Frost hardiness of needles was determined by controlledfreezing tests and visual damage scoring. The constant 6 °Ctemperature treatment caused a gradual dehardening of needleswhereas under fluctuating temperatures the level of frost hardinessfluctuated. Trees exposed to extended photoperiods were lesshardy than under natural photoperiods after the initiation ofshoot elongation, but before this there were no clear differencesin frost hardiness between different photoperiodic treatments.The results indicate that the frost hardening competence ofScots pine changes during quiescence. Climate change; frost hardiness; hardening competence; photoperiod; Pinus sylvestris, Scots pine; temperature     

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     

Effects of Edaphic Factors on the Soluble Carbohydrate Content of Roots of Lolium perenneL. and Trifolium repens L.

In a 3³x2² factorial experiment with Lolium perenne L. and Trifoliumrepens L. grown at three soil moisture tensions, three levelsof nitrogen and three levels of phosphorus, there was a widerange of percentage soluble carbohydrate (TSC) in the roots.The significant positive correlations between percentage ofTSC and root weight suggest that root growth-rate was proportionalto the concentration of TSC in the roots. In other experiments with Lolium perenne grown at five constantsoil temperatures and factorial combinations with three soilmoisture treatments with and without manure, there was declinein the percentage of TSC with increasing temperature. This impliedthat the respiration sink strength in the roots did not controlthe partitioning of photosynthate to the roots. No firm conclusions can be drawn from this work, but it is inferredthat there is a dynamic balancing mechanism in the foliage whichcontrols the partitioning of photosynthate between foliage androot growth. This mechanism appears to divert carbohydrate toroots in inverse proportion to root activity.     

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.     

Effect of Temperature on Nitrogenase Activity in White Clover

Single, clonal plants of white clover were grown without inorganicnitrogen in four contrasting day/night temperature regimes,with a 12 h photoperiod, in controlled environments. Root andnodule respiration and acetylene reduction activity were measuredin a flow-through system during both day and night for plantsacclimated to day/night regimes of 23/18, 15/10 and 10/5 ?C.Similar measurements were made on plants acclimated to 20/15?C and stepwise at temperatures from 4 to 33 ?C. Peak rate of ethylene production, nitrogenase-linked respirationand basal root + nodule respiration increased approximatelylinearly from 5 to 23 ?C both in temperature-acclimated plantsand in plants exposed to varying measurement temperatures. Themeasured attributes did not vary significantly between day andnight. Temperatures above 23–25 ?C did not further enhancethe rate of ethylene production, which remained essentiallythe same up to the maximum measured temperature of 33 ?C. The measurements of nitrogenase-linked respiration between 5and 23 ?C, during both day and night, demonstrated a constant‘energetic cost’ of acetylene reduction of 2.9 µmolCO₂ µmol C₂H₄^–1,. Over the same temperature range,the approximate activation energy of acetylene reduction was60 kJ mol^–1. The integrated day plus night nitrogenase-linkedrespiration accounted for 13.4–16% of the plant‘snet shoot photosynthesis in a single diurnal period: there wasno significant effect of temperature between 5 and 23 ?C. Key words: Trifolium repens, white clover, temperature, N₂ fixation, respiration     

Expression of Vernalization Genes in Near-isogenic Wheat Lines: Effects of Night Temperature

Four near-isogenic lines of wheat (Triticum aestivum L.em Thell)were used to compare selected night temperatures for their effectivenessas vernalizing temperatures. All treatments (conducted withina phytotron) had a common day temperature of 20 °C for 12h and night temperatures were 4, 7, 10, 13 and 20 °C. Interpretationof results for reproductive development was confounded by threeinteracting factors, their relative importance varying withgenotype. Firstly, development rate was generally slower atlower night temperatures. Secondly, in contrast, there was atendency for lower night temperatures to hasten developmentrate if vernalization requirements were satisfied. Thirdly,the lower night temperatures provided a more favourable environmentfor leaf production such that for some genotypes, vernalizedplants had higher final leaf numbers than unvernalized plants.Only for the genotype with the strongest vernalization response(vrn1 vrn2) did hastening of development due to vernalizationoverride any delaying effects. For this genotype, 4, 7 and 10°C were vernalizing temperatures. For the other three genotypes,any hastening of development due to vernalization was outweighedby delaying effects of lower night temperatures. Spikelet numberand days to anthesis were positively correlated in three ofthe four genotypes. It appeared that differences in spikeletnumber were a direct result of night temperature influencingthe duration of the spikelet phase and/or rate of spikelet initiation.Plant size at flowering was determined by the differential effectsof night temperature on growth and development rates. Triticum aestivum L., wheat, vernalization, night temperature, isogenic lines     

Response of ovule development and post-pollen production processes in male-sterile tomatoes to chronic, sub-acute high temperature stress

In order to determine the effects of high temperature on ovuledevelopment and reproductive processes subsequent to pollenproduction, nine day/night temperature combinations were imposedover a 9 month period as four separate experiments, each withthree treatments, including one common treatment. In order toeliminate known effects of high temperatures on pollen productionand stylar position, high temperature treatments were appliedonly to male-sterile tomatoes (Lycopersicon esculentum Mill.).Pollen was obtained from male-fertile plants given optimal growthconditions. This allowed comparison of mean daily temperaturesfrom 25–29 C; day/night temperature differentials (DIFs)of 2, 6, and 10C; day temperatures of 28, 30 and 32C at nighttemperatures of 22, 24, and 26C; and night temperatures of22, 24 and 26C at day temperatures of 28, 30 and 32C. Averageweight per fruit and flower number did not demonstrate a consistentpattern of response to high temperature. Other reproductivecharacteristics (% fruitset, total number and weight of fruitper plant, and seediness index) decreased as mean daily temperaturerose from 25C to 26C and from 28C to 29C. The primary parameteraffecting these variables was mean daily temperature, with daytemperature having a secondary role. Thus, in determining reproductiveresponses of tomato to temperatures within this range, day temperature,night temperature and DIFs do not need to be considered independentlyof their effect on mean daily temperature. If this relationshipholds true in other species, and for pre-pollen production pro-cesses as well, modelling the effects of projected climate changeshould be simplified. Key words: Global climate change, therrnotolerance, stress resistance, ovule development, pollination     

Influence of Thermoperiod on Growth and Development in Cucumber

We studied the influence of gradient temperature regimes on various parameters of the formation of shoots and roots of cucumber plants, such as rate of leaf appearance, rate of growth, duration of growth and length of leaves, and the rate of growth shoots organs and roots. The plants were grown under the controlled conditions: at different combinations of day and night temperature, illumination 100 W/m², and 12 h photoperiod. The comparison of constant and fluctuating diurnal temperature regimes has shown that in the optimal area for all studied indices, the highest values were recorded at the constant daily temperature (25°C for all growth indices of shoots and 20°C for growth of roots), while all gradient regimes either did not affect, or exerted inhibitory effects on the plant. Outside the optimum area, the effects of gradient temperatures differed. The main acting fluctuating temperatures, that exerted stimulating effects, combined low hardening (15°C) and optimal temperatures (25°C), which was earlier described for animals. The 15/35 and 35/15°C combinations were unambiguously inhibitory, since both temperatures are hardening for the cucumber. A lesser stimulating effect of gradient temperatures on the developmental rate in a plant, as compared to poikilothermic animals, could be due to a greater autonomy of plant ontogenesis because of autotrophy and, correspondingly, a greater degree of homeostasis. The mechanisms accounting for the responses to temperature gradients are similar in different groups of ectotherms.     

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.     

A Comparison of the Growth of the C4 Grass Spartina anglica with the C3 Grass Lolium perenne at Different Temperatures

Dunn, R., Thomas, S. M., Keys, A. J. and Long, S. P. 1987. Acomparison of the growth of the C₄ grass Spartina anglica withthe C₃ grass Lolium perenne at different temperatures.—J.exp. Bot. 38: 433–441. S. anglica is one of the few C₄ species which occurs naturallyin cool temperate zones. It is known to attain photosyntheticrates which equal or exceed those of C₃ grasses over the temperaturerange typical of the spring and summer in cool temperate climates.This study examines whether S. anglica can also attain comparablegrowth rates at these temperatures. Seedlings of S. anglicaand L. perenne cv. S23 were grown in controlled environmentsat 10,15,20 and 25 °C. Quantitative growth analysis wasconducted by taking frequent harvests to determine the progressionsof leaf area and plant weight of individual plants with time.Quadratic regressions were found to describe these progressionswell. Instantaneous derived growth parameters were calculatedfrom the fitted regressions. Both absolute and relative growthrates of S. anglica were significantly lower than for L. perenne,this being largely attributable to a lower ratio of leaf areaproduction per unit of plant dry weight. Although the amountof dry matter invested into leaves was similar, the leaf areaper unit of leaf dry weight was lower in S. anglica. In comparisonto L. perenne, the rate of dry matter accumulation per unitof leaf area (ULR) was higher in S. anglica at 25 °C andinitially equal at 10 °C. Prolonged exposure to 10 °Csteadily reduced ULR in S. anglica which approached zero at80 d. Although growth in S. anglica is reduced more by low temperaturethan it is in L. perenne, by comparison to other C₄ speciesthe assimilatory capacity of S. anglica is more tolerant oflow temperature exposure. Key words: C4 photosynthesis, temperature, quantitative growth analysis     

The Effect of Air Humidity on Photosynthesis of Ryegrass and White Clover at Three Temperatures

Increasing leaf-air vapour pressure deficit (VPD) decreasedthe stomatal conductance and the photosynthetic rate of leavesof ryegrass (Lolium perenne L.) and white clover (Trifolhimrepens L.) at light saturation and at lower irradiance. In ryegrassboth conductance and photosynthesis, and in clover photosynthesis,decreased less with increasing VPD in low irradiance than theydid at an irradiance which saturated photosynthesis. In ryegrass,relative to their values at 10 mb, photosynthesis and conductancedecreased less with increasing VPD at 25 °C than at 20 or16·5 °C. In white clover, relative conductance (butnot photosynthesis) was less reduced at 25 than at 16·5°C Measurements of VPD of air in the leaf canopy of a field-growncrop are combined with the observed responses of photosynthesisto VPD and temperature in a model. This shows that high VPDis likely to depress photosynthesis significantly and that,during a typical day, the rate of light saturated photosynthesismay remain fairly steady, because the depression of photosynthesisdue to rising VPD is offset by the stimulation due to risingtemperature Perennial ryegrass, Lolium perenne L., White clover, Trifolhim repens L., photosynthesis, leaf conductance, water vapour pressure deficit, temperature     

The Effect of Temperature on Photosynthesis of Ryegrass and White Clover Leaves

The rate of photosynthesis of leaves of perennial ryegrass (Loliumperenne L.) and white clover (Trifollum pratense L.) grown atdifferent temperatures was measured at a range of temperatures.There was a small effect of the temperature at which a leafhad grown on its photosynthetic rate, but a large effect ofmeasurement temperature, especially in bright light, where photosyntheticrates at 15°C were about twice those at 5°C. It appearsthat temperature could affect sward photosynthesis in the field.Ryegrass and clover had similar photosynthetic rates which respondedsimilarly to temperature. Lolium perenne L., ryegrass, Trifolium pratense L., white clover, photosynthesis, temperature, irradiance     

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