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     

The Effect of Light Quality on Shoot Extension Growth in Three Species of Grasses

The effect of light quality on the extension growth of vegetativeshoots and on the final size of their leaves was investigatedin plants of Lolium multiflorum, Sporobolus indicus and Paspalumdilatatum. Three experimental approaches were used, (a) redor far-red end-of-day irradiations of sunlight-grown plants,(b) different red/far-red ratios of white light in a growthroom and (c) sunlight enrichment with radiation of differentred/far-red ratios or with different amounts of far-red lightduring the photoperiod. Plants treated with end-of-day far-redor low red/far-red ratios throughout the photoperiod developedlonger leaves and, as a result, longer shoots. This effect wasmore marked in leaf sheaths than in blades. Tiller extensionand leaf sheath length increased with the amount of far-redadded to sunlight in a simple hyperbolic relationship. Theseresults show that vegetative grass shoots respond to light qualityin a way similar to internodes of dicotyledonous plants. Lolium multiflorum Lam., Sporobolus indicus (L.), Paspalum dilatatum (Poir.), leaf growth, tiller growth, photomorphogenesis     

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     

Effects of Light Quality on Tiller Production in Lolium spp

Variations in phytochrome status in grasses were provoked by end of day red and far-red treatments and above canopy irradiations. Lolium perenne L. and Lolium multiflorum Lam. plants developed more tillers when illuminated with higher red/far-red ratios. These results show that branching of grasses is controlled by phytochrome activity in a way similar to that in dicotyledon plants.     

The Growth and Survival of Severely-shaded Tillers in Lolium perenne L.

The effect of shading a single tiller to below its compensationpoint for a period of 5 weeks in vegetative plants of Loliumperenne L. cv. S23, was studied in two different experimentseach employing two light regimes, one of which was common toboth experiments. In the first experiment tillers in the axils of the first leafwere shaded three weeks from appearance at both 40 and 70 Wm^–2. None of the shaded tillers died and they continuedto produce new leaves and increase in dry weight but at a reducedrate. In the second experiment, tillers with one emerged leafin any leaf axil position were shaded at 70 W m^–2 andin a treatment in which light was reduced to 13 W m^–2after initial growth at 70 W m^–2. As in the first experimentall shaded tillers survived at 70 W m^–2 but in the 70 13 W m^–2 transfer regime all shaded tillers died. In the second experiment shaded tillers in both light regimeswere supplied with ¹⁴C-assimilate by translocation from theremainder of the plant but in the 70 13 W m^–2 the initialsupport was withdrawn within 5 weeks of shading. The results are discussed in terms of the physiological relationshipsbetween the tillers of the grass plant. Lolium perenne L., growth of tillers, survival of tillers, effect of light     

Rates of Leaf and Tiller Production in Young Spaced Perennial Ryegrass Plants in Relation to Soil Temperature and Solar Radiation

In an experiment designed to investigate the rate of leaf appearanceand tiller production in young spaced plants of three clonesof perennial ryegrass grown in the field, it was found thatthe rate of leaf appearance per tiller increased linearly withmean soil temperature up to approx. 14 °C. The rate of productionof tillers in relation to the rate of leaf appearance (sitefilling) appeared to be virtually independent of weather conditions.In plants which were adequately established, but still relativelysmall, site filling was equal to or exceeded the theoreticalsteady state; all the tiller buds which were being formed weredeveloping into visible tillers. Thus the relative rate of tillerproduction was controlled by the rate of leaf appearance. Inlarger plants site filling was less complete, and site redundancieswere probably caused by within-plant competition for light atthe tiller bases. Lolium perenne L., perennial ryegrass, tillering, leaf production, solar radiation, soil temperature     

The Effects of Plant Density on Shoot and Leaf Lamina Angles in Lolium multiflorum and Paspalum dilatatum

The effects of plant density on shoot and leaf lamina zenithangles (i e the angles with respect to the vertical) were investigatedin Lolium multiflorum and Paspalum dilatatum plants grown inpots either outdoors or in a glasshouse Tillers appeared abovethe ligule of their subtending leaf with a small zenith anglebut, in plants grown at low densities, gradually adopted a morehorizontal position In plants grown at high densities this gradualincrease in shoot zenith angle with age was strongly reducedThe average shoot zenith angle at a given time was lower (ie tillers were more erect) at high, compared to low plant densitiesShoot number per plant and average shoot zenith angle per plantwere directly related In P dilatatum plants the leaf lamina remained nearly horizontalat any plant density This observation indicates that the movementof the leaf lamina, pivoting at the point of attachment to theleaf sheath, compensated the movement of the shoot, pivotingat the base of the leaf sheath-tube Shoot angle responses to plant density would place leaf laminaeat different heights within the canopy, affecting their abilityto compete for light and their susceptibility to wind impactand herbivore grazing Foliage architecture, Lolium multiflorum Lam, Paspalum dilatatum Poir, plant density, neighbour effects, shoot angle, leaf angle     

Effect of Flowering on the Photosynthetic Capacity of Ryegrass Leaves Grown With and Without Natural Shading

The photosynthetic capacity of newly expanded leaves of vernalizedor non-vernalized plants of S24 perennial ryegrass (Lolium perenneL.), grown in long or short photoperiods, was measured in twoexperiments. In the first, leaves were protected from shadingduring development, while in the second, the natural shade ofneighbouring tillers in a sward was allowed. In the first experiment there was little effect of vernalization,day length or flowering, and leaves in all treatments had photosyntheticrates at 250 W m^–2 of between 28 and 32 mg CO₂ dm^–2h_–1.In the second experiment the photosynthetic rate of successiveleaves fell as sward leaf area increased. This downward trendwas reversed, however, in flowering tillers in the vernalizedlong-day treatment, while in the other treatments, which didnot flower, photosynthetic capacity continued to fall. It isconcluded that the leaves of reproductive tillers have highphotosynthetic capacities because stem extension carries themto the top of the canopy where they are well illuminated duringexpansion. Lolium perenneL, ryegrass, photosynthetic capacity, flowering, shading, vernalization     

Tillering Responses to the Light Environment and to Defoliation in Populations of Perennial Ryegrass (Lolium perenneL.) Selected for Contrasting Leaf Length

Tillering responses to the light environment and to defoliationwere studied in two populations of perennial ryegrass (LoliumperenneL.) selected for contrasting leaf lengths. The objectiveof this study was to determine whether differences in tilleringresponses between populations, as a result of management practices,affected their competitive ability. Young seedlings were exposed,under controlled conditions, to defoliation, neutral shading(decreased photosynthetic photon flux), low red:far-red ratioand/or decreased blue light. Selection for longer leaves reducedthe tillering rate. After defoliation, this difference betweenshort- and long-leaved populations was magnified. Defoliationdecreased both site filling and phyllochron of the long-leavedpopulation but had no effect on the short-leaved population.Lowering the photosynthetic photon flux reduced the phyllochronin both population. Decreasing the red:far-red ratio reducedtillering rate by reducing site filling, whereas decreasingblue light had no significant effects on tillering. Tilleringresponses to photosynthetic photon flux and to red:far-red ratiowere similar in the two populations selected for contrastingleaf length. The implication of these tillering responses indefining the competitive ability of the grass plants is discussedin relation to their management.Copyright 1999 Annals of BotanyCompany Blue light, defoliation, far-red, irradiance, leaf size,Lolium perenne(L.), perennial ryegrass, photomorphogenesis, phyllochron, shading, site filling, tiller production.     

Tillering Responses of Lolium multiflorum Plants to Changes of Red/Far-Red Ratio Typical of Sparse Canopies

Casal, J. J., Sáchez, R. A. and Deregibus, V. A. 1987.Tillering responses of Lolium multiflorum plants to changesof red/far-red ratio typical of sparse canopies.—J. exp.Bot. 38: 1432–1439. Plants of Lolium multiflorum were grown either in growth roomsunder radiation sources providing equivalent PAR but differentR/FR ratios between 0·46 and 2·36, or under sunlightsupplemented with a series of small fluence rates of FR throughoutthe photoperiod. In both kinds of experiments a reduction oftillering was found along with small depressions of R/FR ratiobelow the values typical of sunlight. In this range both thesquare-root-transformed rate of tillering and the site-fillingrate were linearly related to the estimated phytochrome photo-equilibriaand the slope was very steep under conditions that allowed hightillering rates. The addition of low fluence rates of red lightat the base of the shoots raised tillering of plants exposedto low R/FR ratios to the level of plants grown under high R/FRratios in spite of the ratios received by the rest of the plant.Moreover, irradiation of plant bases with intermediate-low R/FRratios reduced tillering indicating that this would be one siteof perception. These results show a high sensitivity to changesof R/FR ratio typical of different degrees of shade in sparsecanopies. This suggests that by monitoring these changes Loliummultiflorum plants would be able to perceive an anticipatedsignal of impending competition in growing canopies.     

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     

A Method for in Vitro Storage of Lolium multiflorum Lam

A means of storing Lolium multiflorum Lam. stock plants in vitrohas been developed over a period of three years. Variousexplantsand cultureconditions were examined. Shoot tips 0.3–0.9mm long were found to be best for establishing storage culturesbecause they gave a high yield of plantlets in culture and importantpathogens are eliminated. For sub-culturing, after each annualcycle of storage at 2–4 °C, shoot tips, tiller buds,tiller bases and nodes can be used. Tiller buds were most abundantand best for increasing the number of stored plantlets whennecessary. There was no advantage of including an auxin in theculture medium for shoot tips and Murashige and Skoog's mediumwith 0.2 mg l^–1 kinetin has been adopted for initiating,sub-culturing and storing cultures. The optimum temperaturerange for regenerating plantlets was 20–25 °C. Lightwas necessary for a high rate of plantlet regeneration and lightquality and intensity affected their growth. Lolium multiflorum Lam., ryegrass, storage in vitro, tissue culture     

The Influence of Defoliation on the Distribution of Assimilates in Lolium multiflorum Lam

Autoradiographs were made of plants of Lolium multiflorum Lam.after ¹⁴CO₂ had been fixed by selected leaves. The results showedthat labelled compounds were not translocated to other tillersbut were moved to the whole root system. This pattern of distributionwas changed when all or some of the tillers on the plant weredefoliated. Where a single undefoliated tiller remained, itinitially supplied the cut tillers with ¹⁴C-containing products,thus reintegrating a system of apparently independent tillers.When all the tillers were partially defoliated, labelled compoundswere no longer translocated to the root system. A further experimentsuggested that root reserves were not mobilized for regrowthfollowing defoliation. These results are discussed in termsof the integration of a grass plant in the vegetative state.     

Growth response of threeFestuca rubra clones to light quality and arbuscular mycorrhiza

The influence of the arbuscular mycorrhizal fungus—Glomus etunicatum and changes in light quality (decrease of red/far-red ratio) on the growth of threeFestuca rubra clones, ecotypes originating from a mountain grassland region, was studied in a growth chamber experiment. Inoculation with the arbuscular mycorrhizal fungus and low red/far-red ratio decreased both the number of tillers and the biomass of treated plants. Significant interactions between the treatments were found and most of the growth characteristics were reduced further when both treatments were applied simultaneously. Inoculation with the arbuscular mycorrhizal fungus also resulted in reduced maximum height of tillers, whereas low red/far-red ratio caused the maximum height of tillers to increase. Differences in plasticity were found for the threeFestuca rubra clones. Response to one treatment was strongly modified by the other treatment. This indicates that the arbuscular mycorrhizal fungus and red/far-red ratio can differentially influence the growth ofFestuca rubra clones and thus modify their relative competitive abilities which can consequently have implications for the coexistence mechanisms within plant population, thereby potentially influencing plant canopy and community structure.     

Effects of Blue Light on the Vertical Colonization of Space by White Clover and their Consequences for Dry Matter Distribution

The morphology of white clover is very sensitive to the lightenvironment, especially to the ratio of red:far-red light andto photon irradiance. However, less is known about the effectsof blue light on clover morphogenesis. Cuttings of white cloverwere grown for 56 d in two controlled chambers receiving lightwith similar photosynthetic efficiency and phytochrome photoequilibriumstate but different levels of blue light: some plants were grownunder orange light (very low blue light, 0.02 µmol m^-2s^-1)or under white light containing blue light (83 µmol m^-2s^-1).Other plants were switched from white light to orange lightorvice versa,after 30 d. The absence of blue light modifiedthe growth habit of clover and raised the laminae in the upperlayer of the canopy by increasing petiole length, and petioleangle from the horizontal, and by raising stolons above theground surface. Moreover, the absence of blue light had no effecton total leaf area and total dry weight per plant, but increasedthe leaf area and biomass of petioles of the main axis. Largerpetioles and laminae were associated with the allocation ofmore dry weight to the petiole at the same petiole thicknessbut with thinner laminae. These results indicate that a decreasein blue light is involved in the perception of, and adaptationto, shading by the plant.Copyright 1997 Annals of Botany Company Biomass allocation; blue light; growth habit; leaf area; light quality; photomorphogenesis; Trifolium repensL.; white clover     

Effect of Light Quality (Red:Far-red Ratio) at the Apical Bud of the Main Stolon on Morphogenesis of Trifolium repens L.

A controlled environment experiment investigated whether thered:far-red (R:FR) ratio of light at the apical bud of the mainstolon could alter plant morphogenesis in clonal cuttings ofwhite clover (Trifolium repens L.) The apical bud included theapical meristem, five to six developing leaf primordia withassociated axillary bud primordia and stipules and the firstemerged folded leaf until development was greater than 0·3on the Carlson scale. Three light regimes were imposed on theapical bud by collimating light from R or FR light-emittingdiodes so that the R:FR ratio of light incident at the apicalbud was set at 0·25, 1·6 or 2·1, withoutsignificantly altering photosynthetically active radiation.The effect of these light regimes on white clover seedling growthwas also tested. At a low R:FR ratio seedling hypocotyl and cotyledon lengthswere significantly longer. However, with the cuttings, the lighttreatments did not alter node appearance rate or internode lengthof the main stolon, petiole length, area of leaves or totalshoot dry matter. There was one significant photomorphogeneticresponse in the cuttings, a delay of 0·5 of a phyllochronin the appearance of branches from axillary buds in the lowR:FR ratio treatment relative to the other treatments. Wherebranch appearance was delayed plants had fewer branches. Thisdifference could be ascribed solely to a delay in branch appearanceas there were no significant treatment effects on either theinitiation of axillary bud primordia within the apical bud,the probability of branching or on the rate of growth of branchesafter appearance. Because treatment of the apical bud inducedonly one of the many previously observed responses of whiteclover to a decrease in the R:FR ratio of light, we concludethat other plant organs must also sense the quality of incidentlight.Copyright 1994, 1999 Academic Press White clover, Trifolium repens, apical bud, light quality, red:far-red ratio, light-emitting diode, branching, axillary buds, photomorphogenesis     

Tillering responses to red:far-red light ratio during different phenological stages in Eragrostis curvula

Tillering responses to light quality in different phenological stages of a perennial warm-season grass Eragrostis curvula were investigated in controlled environments. In vegetative plants, the tillering rate was greater (P<0.01) in the high (1.1–1.3) than in the low (0.59–0.70) red:far-red ratio (R/FR) light regime. Tillering rates were higher in the low R/FR treatment when the plants in the high R/FR regime reached the reproductive stage, while the plants in the low R/FR regime remained vegetative. After the reproductive tillers were removed by defoliation, more tillers were produced in the defoliated plants grown in the high R/FR regime. When the plants in both light treatments entered the reproductive stage, the tillering rate under the two light regimes became similar, suggesting a significant interaction between tillering and inflorescence development. The more advanced inflorescence development in the high R/FR regime may have reduced assimilate availability to tiller growth and overshadowed the effect of high R/FR on tillering. Both tillering and inflorescence development appeared to be controlled by R/FR ratio. The higher rate of aerial tiller production on the reproductive culms during the post-anthesis period in the high R/FR regime suggested that high R/FR ratio stimulated not only basal tillering, but also aerial tillering.     

Cessation of tillering in spring wheat in relation to light interception and red : far-red ratio

BACKGROUND AND AIMS: The production of axillary shoots (tillering) in spring wheat (Triticum aestivum) depends on intraspecific competition. The mechanisms that underlie this competition are complex, but light within the wheat canopy plays a key role. The main objectives of this paper are to analyse the effects of plant population density and shade on tillering dynamics of spring wheat, to assess the canopy conditions quantitatively at the time of tillering cessation, and to analyse the relationship between the tiller bud and the leaf on the same phytomer. METHODS: Spring wheat plants were grown at three plant population densities and under two light regimes (25 % and 100 % light). Tiller appearance, fraction of the light intercepted, and red : far-red ratio at soil level were recorded. On six sampling dates the growth status of axillary buds was analysed. KEY RESULTS: Tillering ceased earlier at high population densities and ceased earlier in the shade than in full sunlight. At cessation of tillering, both the fraction of light intercepted and the red : far-red ratio at soil level were similar in all treatments. Leaves on the same phytomer of buds that grew out showed more leaf mass per unit area than those on the same phytomer of buds that remained dormant. CONCLUSIONS: Tillering ceases at specific light conditions within the wheat canopy, independent of population density, and to a lesser extent independent of light intensity. It is suggested that cessation of tillering is induced when the fraction of PAR intercepted by the canopy exceeds a specific threshold (0.40-0.45) and red : far-red ratio drops below 0.35-0.40.     

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     

Light-mediated bioregulation of tillering and photosynthate partitioning in wheat

The influence of plant population density on spectral distribution of light received by wheat ( Triticum aestivum L. cv. Coker 797) seedlings was measured under field conditions, and effects of red and far-red light on tillering and photosynthate partitioning were studied in controlled environments. Spectral distribution of light was measured in sunflecks at soil level in close-, intermediate-, and wide-spaced field populations during the tillering stage. Close-spaced seedlings received higher far-red/red light ratios than wide-spaced plants because of the larger amount of far-red reflected from green leaves of the more numerous nearby plants. The far-red/red light ratios in all population densities were higher in late afternoon than at noon. Close-spaced plants developed fewer tillers, less roots and longer leaves than wide-spaced seedlings under field conditions. In controlled environments, a higher far-red/red ratio during photosynthetic periods resulted in fewer tillers and longer leaves; whereas, brief red or far-red exposures at the end of each day had a more pronounced effect. Wheat seedlings that received 5-min exposures to far-red light at the end of the photosynthetic period each day for 20 consecutive days developed fewer tillers, longer leaves, less roots, and a higher shoot/root biomass ratio. The effects of far-red light were reversed by red light. The light spectral shifts associated with field plant population densities and the responses to red and far-red treatments under controlled environments suggest that phytochrome serves as a sensing mechanism that detects the amount of competition from other plants, and regulates the development of tillers and the partitioning of photosynthate between shoots and roots.