The Growth and Development of Simulated Swards of Perennial Ryegrass: I. Leaf Growth and Dry Weight Change as Related to the Ceiling Yield of a Seedling Sward

The leaf growth, tiller production, light interception, anddry weight increase of a simulated sward of S24 perennial ryegrass(Lolium perenne) were followed during the development of thesward from a collection of two-leaved seedlings to a closedcanopy with an LAI of 23, of which 15 consisted of green leaflaminae. The dry weight of live shoots increased exponentiallyat first, but then entered a long linear phase of increase.This was equivalent to a crop growth rate of 200 Kg ha^–1day^–1 and a conversion efficiency of radiant energy (400–700nm) of 7.2 per cent. Towards the end of the growth period therate of increase of live shoots declined rapidly to zero anda ceiling yield was reached equivalent to 10 metric tons ha^–1.Leaf growth continued at a high rate, but was equalled by therate of leaf death, so that the weight of live leaf tissue remainedconstant. By this time the swards had achieved a stable tillerpopulation (about 1 cm^–1), each tiller bore a constantnumber of live leaves (about three), and the length of eachnewly expanded leaf equalled the length of the old leaf it replaced(about 70 cm). The swards were grown in Perlite so that in theabsence of soil fauna dead leaves accumulated at the base ofthe sward where, after 12 weeks, they accounted for 19 per centof the total weight of dry matter produced.     

Single Leaf and Canopy Photosynthesis in a Ryegrass Sward

Swards of S24 perennial ryegrass (Lolium perenne L.) were grownin the field and harvested by repeated cutting. The rate ofphotosynthesis of the swards and of single leaves of plantstaken from them was measured throughout the growing season.Leaf area and light interception of the sward were also measured. In each growth period, sward photosynthesis increased with increasingleaf area and then declined. In the first growth period, duringwhich the crop flowered, the maximum rate of sward photosynthesiswas higher and was reached later than in succeeding vegetativegrowth periods. The photosynthetic capacity of successive youngest expandedleaves was high throughout the first growth period, and onlydeclined due to leaf age when no further leaves were producedafter flowering. However, in the succeeding vegetative growthperiods, although the first leaves to expand had high photosyntheticcapacities, later leaves had progressively lower capacities. The extent to which changes in the photosynthetic capacity ofsingle leaves accounts for the changes in sward photosynthesisobserved in the field is examined using a mathematical model.     

The photosynthesis of ryegrass leaves grown in a simulated sward

Plants were taken from simulated swards of perennial ryegrass (Lolium perenne) grown in a controlled environment and the rates of photosynthesis of the youngest fully expanded leaves, and the second and third youngest leaves on the same tillers were measured. The youngest leaves had the highest rates and the third the lowest, with the second leaves intermediate. The rate of photosynthesis in bright light of successive youngest expanded leaves decreased as the swards increased in leaf area, but did not when plants were grown so that the main stem was not shaded. When plants were grown at different densities and the photosynthetic rates of leaves of a particular ontogenetic rank were measured, it was found that leaves on plants from higher densities had lower rates of photosynthesis. Also leaves on plants grown in bright light had higher photosynthetic rates than those on plants grown in dim light. It is concluded that the decline in the photosynthetic capacity of successive leaves in a rapidly growing simulated sward is due to the intense shading to which they are subjected during their expansion.     

The Effect of Shading During Vegetative and Reproductive Growth on the Photosynthetic Capacity of Leaves in a Grass Sward

The decline in photosynthetic capacity of successive newly expandedleaves, which occurs as a vegetative grass sward increases inleaf area, was prevented in a field sward of S24 perennial ryegrass(Lolium perenne L.) by protecting tillers from shading by theirneighbours. This adds support for the view that the declineis caused by the shading of leaves during their expansion. However, in a flowering sward where there is no such decline,the photosynthetic capacity of leaves was not reduced by shadingthem with a plastic mesh during expansion, although it was bypegging down tillers so that they developed at the bottom ofthe sward. Lolium perenne, perennial rye-grass, photosynthetic capacity, growth, shading     

Microclimate, Canopy Structure and Photosynthesis in Canopies of Three Contrasting Temperate Forage Grasses: I. Canopy Structure and Growth

The crop growth rates and structures of three temperate foragegrasses Lolium perenne cv. S24, L. perenne cv. Reveille andFestuca arundinacea cv. S170, were examined in the field duringa summer growth period. The growth rates of the varieties wereremarkably similar at 7 g DM m^–2 day^–1. The angularstructures of the varieties were different and they varied duringthe experiment. However, these differences did not seem to affectcrop growth rates. Nevertheless, a decrease in the efficiencyof light energy conversion of approximately 24 per cent wasobserved after a change to a more prostrate form of canopy dueto lodging. There appeared to be an inverse relationship betweenthe number of tillers per unit ground area and the weight ofan individual stem. There were large numbers of relatively lighttillers in S24 whereas S1 70 had fewer but heavier tillers.Furthermore, S24 had many small leaves per unit ground areacompared with SI70 which had fewer longer leaves per groundarea and a slower rate of leaf appearance. There were diurnalchanges in the rates of leaf extension for all the varieties.The mean daily extension rates declined as the canopies developed.     

Competition between grass and clover in spring as affected by nitrogen fertiliser

Detailed measurements of irrigated ryegrass/white clover swards growing without interruption, with or without nitrogen fertiliser in spring, showed that the relative growth rate of clover was as great as that of grass, in the + N sward, and considerably greater than grass, in the – N sward. Clover leaves were not overtopped by grass leaves. Indeed, in both swards, clover had a greater proportion of its leaf lamina area in the upper, well lit, layers of the canopy than grass did. Consequently, clover had a greater mean rate of leaf photosynthesis in situ in the sward than grass. Clover's advantage in photosynthetic rate per leaf area was offset to some extent by its smaller ratio of leaf area to total above-ground dry weight than grass. The consequences of these results for our understanding of competition between grass and clover in mixed swards are discussed.     

The Effects of Shading and Cutting Treatments on the Photosynthetic Rate of Ryegrass Leaves

As the leaf area of miniature simulated swards of S24 perennialreygrass (Lolium perenne L.) grown in controlled environmentincreased, successive leaves which expanded on the main stemof a sample plant within the sward had progressively lower photosyntheticcapacities. When the plants surrounding the sample plant were cut, the photosyntheticcapacity of the next leaf to expand was increased, whether ornot other leaves on the sample plant had been removed. Whenall fully expanded leaves on the sample plant were removed butthe surrounding plants left uncut, the photosynthetic capacityof successive leaves on the sample plant continued to decreaseas in uncut swards. These results support the view that the changes in photosyntheticcapacity in grass swards are consequences of changes in thelight climate of the developing leaf rather than of changesin demand for assimilates or growth substance supply. The declinein photosynthetic capactiy was contributed to by both stomataland residual diffusion resistances.     

Photosynthesis by White Clover Leaves in Mixed Clover/Ryegrass Swards

Measurements of rates of net photosynthesis were made on singleBlanca white clover leaves on plants taken from a field-grown,mixed clover/perennial ryegrass sward during two regrowth periods. Net photosynthesis fell by 20 per cent in the first measurementperiod as leaf area index increased and the grass componentof the crop flowered, but did not change significantly in thesecond measurement period during which the grass remained vegetative. Leaves which had been artificially protected from shading inthe sward did not have significantly different photosyntheticcapacities from leaves in the undisturbed sward, even in thefirst measurement period. As leaf area index and sward height increased, successive cloverpetioles were longer, keeping the newly expanded leaves nearthe top of the sward where they received full light. It is suggestedthat it is this which allows successive clover leaves, unlikethose of vegetative grasses, to attain a high photosyntheticcapacity throughout a growth period. Trifolium repens, Lolium perenne, Photosynthetic capacity, shading, growth     

The Effect of New Tiller Growth on Carbohydrates, Nitrogen and Seed Yield per Ear in Lolium perenne L.

To clarify whether new vegetative tillers that develop aroundanthesis in aLolium perenne seed crop can depress seed yields,the possible competition for carbohydrates or nitrogen betweenthe seeds and new tillers that develop after the onset of anthesiswas investigated. In two greenhouse experiments the number oftillers per plant was varied by a combination of cutting, nitrogensupply, light quality and light intensity treatments. Two genotypeswith different tillering rates were used. Seed yield per earwas largely independent of the number of tillers and regrowthof cut tillers after the onset of anthesis. It increased inone genotype, but only under low light and a reduced nutrientavailability, and no new tillers were produced. The amount ofwater-soluble carbohydrates in the reproductive tillers increasedin both clones under these conditions. Under more favourableconditions the increased tillering rate and regrowth of tillersafter cutting did not adversely affect seed yield per ear ineither clone, although carbohydrate reserves in the floweringtillers were sharply reduced. Tiller removal increased the concentrationand amount of nitrogen in the remaining flowering tillers, irrespectiveof the amount of regrowth. It is concluded that competitionfor carbohydrates or nitrogen between the seeds and new vegetativetillers that develop after the onset of anthesis, is not a majorcause of the low and variable seed yields inL. perenne seedcrops. Processes within the ear itself are probably limiting. Lolium perenne L.; perennial ryegrass; seed yield; seed set; tillering; carbohydrates; nitrogen; competition.     

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     

The Kinetics of Extracellular Glycollate Production by Chlorella pyrenoidosa

Extracellular glycollate is liberated by Chlorella pyrenoidosaduring growth in medium bubbled with air or 3 per cent carbondioxide in air. With air the rate of release of glycollate percell decreases, with 3 per cent carbon dioxide it increases,with increase in cell number. Glycollate is released duringshort-term experiments when C. pyrenoidasa, grown under lowlight and high carbon dioxide, is transferred suddenly to highlight and low carbon dioxide. No other combination of thesefactors produces a comparable release of glycollate. The quantityof glycollate released in short-term experiments increases exponentiallywith the relative growth-rate of the culture from which thecells are derived. A crucial condition for maximum glycollaterelease is that growth of the culture prior to the experimentshould not be limited by carbon-dixoide concentration. The effectof pH is related to its effect on growth-rate; i.e. C. pyrenoidosahas a lower relative growth-rate at pH 8.3 and produces correspondinglyless glycollate than faster growing cultures at pH 6.4. Duringshort-term experiments under high light and low carbon dioxidethe rate of glycollate release drops after 50–100 minutessuggesting exhaustion of the glycollate precursor.     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions I. Carbon Assimilation and Utilization

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and their growth and carbon economyexamined. The swards received a nutrient solution daily, whichcontained either High (220 mg l^–1) or Low (10 mg l^–1)nitrate N. Rates of canopy photosynthesis and respiration, and final drymatter yields were similar in the two treatments although theproportions of grass and clover differed greatly. The Low-Nswards were made up largely of clover. The grass plants in theseswards had high root: shoot ratios and low relative photosyntheticrates – both signs of N deficiency – and were clearlyunable to compete with the vigorously growing Low-N clover plants.These had higher relative growth rates and dry matter yieldsthan their High-N counterparts. In the High-N swards clovercontributed around 50 per cent to the sward dry weight throughoutthe measurement period despite having a smaller proportion ofits dry weight in photosynthetic tissue (laminae) than grassover much of it. The latter was compensated for, initially bya higher specific leaf area than grass, and later by a higherphotosynthetic rate per unit leaf weight. The results are discussedin relation to observed declines in the clover content of swardsafter the addition of nitrogen fertilizer in the field. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, nitrogen, photosynthesis, carbon balance     

Effects of Tiller Separation and Root Pruning on the Growth of Lolium perenne L.

Experiments have been carried out to assess the role of inter-tillerinteractions within plants of Lolium perenne L. cv. S24 as factorswhich could play a part in determining the growth-rate of thewhole plant. In order to do this, the effect of tiller separationon plant growth was studied as well as the influence of theremoval of different parts of the root and shoot system. Wheneverplants were supplied with nutrients by means of culture solutions,separation of tillers brought about a marked increase in theamount of dry matter produced. Root pruning and in particularthe removal of root apices brought about similar increases ingrowth. Separation of tillers in soil-grown plants did not bringabout increased growth. These results are interpreted as indicatingthat separation of tillers or the initial removal of root apicesin plants supplied with nutrients by means of culture solution,promotes root branching which brings about an increased netassimilation rate arising from an increase supply of a growth-promotingsubstance, probably a cytokinin, from the root apices. The resultsare not taken as indicative of an intertiller interaction whichrestricts the growth of intact plants.     

Leaf photosynthesis in pure swards of two grasses (Lolium perenne and Lolium multiflorum) subjected to contrasting intensities of defoliation

Photosynthetic rates were measured on light saturated, fully-expanded leaves in pure swards of Lolium perenne and Lolium multiflorum during late summer using 14-carbon dioxide. These swards were defoliated by cutting at three heights of 3, 6 and 9 cm above the ground. The photosynthetic rates of leaves on tillers in swards cut constantly 3 cm above ground level were higher than those of leaves in swards cut constantly 9 cm above ground level. Additional treatments with various sequences of cuts 3, 6 and 9 cm above ground level were designed to reduce damage to the growing points of tillers whilst ensuring satisfactory harvesting of the shoots. The photosynthetic rates of leaves on tillers cut to various heights above ground level at successive harvests were intermediate between those of similar leaves in the constantly low and high cut swards. The rates of photosynthesis of Italian ryegrass leaves were higher than those of perennial ryegrass leaves for a short time after defoliation of the constantly high cut swards. However, these perennial ryegrass leaves quickly adapted their rates of photosynthesis to the higher irradiances they received after cutting. Thus grass species differing in morphology adjust to management practices by the use of different photosynthetic strategies.     

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 Temperature and Nitrogen Supply on the Growth of Perennial Ryegrass and White Clover. 1. Carbon and Nitrogen Economies of Mixed Swards at Low Temperature

Simulated mixed swards of perennial ryegrass (Lolium perenneL. cv. S23) and white clover (Trifolium repens L. cv. S100)were grown from seed under a constant 10°C day/8°C nighttemperature regime and their growth, and carbon and nitrogeneconomies examined. The swards received a nutrient solution,every second day, which contained either high (220 µgg^–1) or low (40 µg g^–1) nitrate N. The High-N swards had rates of canopy photosynthesis and drymatter production (over the linear phase of growth) similarto those previously shown by mixed swards at high temperature.The Low-N swards grew more slowly; canopy photosynthesis, ata given LAI, was similar to that at High-N but lower LAI's weresustained. Clover increased its contribution to total carbonuptake and total dry weight throughout the period in the Low-Ntreatment and, despite the fact that grass took up most of theavailable nitrate, clover maintained a consistently higher Ncontent by virtue of N₂-fixation. At High-N, grass dominated throughout the measurement period.Earlier, when plants grew as spaced individuals, clover grewless well than grass, but once the canopy was closed it hada similar relative growth rate and thus maintained a steadyproportion of total sward dry weight. It is proposed that earlyin the development of the crop, leaf area production is thelimiting factor for growth, and that in this respect cloveris adversely affected by low temperature relative to grass.Later, as the LAI of the crop builds up, and the canopy becomesfully light intercepting, net canopy photosynthesis plays amore dominant role and here the higher photosynthetic rate perunit leaf area of the clover is crucial. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, low temperature, nitrogen, photosynthesis     

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     

Studies in the Physiological Action of 2,2-Dichloropropionic Acid: II. THE EFFECTS OF LIGHT AND TEMPERATURE ON THE FACTORS RESPONSIBLE FOR THE INHIBITION OF GROWTH

When Lemna minor and Salvinia natans, grown in a constant environment,are subjected to sub-lethal concentrations of 2,2-dichioropropionicacid (DCPA), the relative growth-rates are progressively reduced.These cumulative reductions, which are greater for S. natans,are correlated with decreases in (1) the rate of leaf or frondformation, (2) the mean area per leaf or frond, and (3) thenet assimilation rate. Of these components, the first is themost important and the third is the least. The effects of light intensity (300, 600, 900 f.c.), temperature(20, 25, and 30°C), and concentration of DCPA on both therelative growth-rate and rate of leaf or frond multiplicationhave been examined in multi-factorial experiments. Over theconcentration range selected (100, 200, and 400 mg/l for S.natans and 100, 300, and 6oo mg/l for L. minor) there are positiveeffects of light intensity, temperature, and concentration.For each concentration the order of the depression is maximalunder a combination of the highest temperature and the greatestintensity. Using radioactive DCPA it has been established that uptake isalso a cumulative process, and that S. natans has a greatercapacity to absorb DCPA. The rate of uptake is independent ofthe light intensity but increases with temperature and concentration. DCPA brings about morphological and structural changes. In S.natans, many of the leaves become submerged and the proportionis positively dependent on light, temperature, and concentration.This failure to float is associated with a reduction in thedensity of epiderrnal hairs. It is concluded that the inhibitory effects of DCPA are maximalunder conditions which are optimal for both meristematic activityand accumulation.     

Influence of cocksfoot streak virus on the growth of cocksfoot swards

Small experimental swards of three types-healthy, 50% and 100% streak-virus-infected-were established from single tillers of randomly selected cocksfoot genotypes. In these swards 44% of infected plants died within 2 years compared with 21% of healthy plants. Mortality of infected plants was not increased by the presence of healthy plants in the same sward. In swards containing both healthy and infected plants, increased growth of healthy plants compensated for the low yield of infected ones only when the swards were frequently defoliated. Infrequent cutting apparently allowed infected plants to check the growth of healthy ones, and 50% infected swards yielded little more than 100% infected swards. This difference in response was attributed to differences in the growth habits of healthy and infected plants. Few initially healthy plants became infected during 2 years of the experiment.     

冠层部位对金叶女贞叶片结构的影响

金叶女贞是一种在园林绿化上广泛应用的彩叶植物,上层叶金黄色,下层叶深绿色。本文利用常规植物制片技术,研究了叶绿素缺少的金叶女贞上层金叶和下层绿叶的解剖结构。结果表明:(1)金叶女贞上层金叶和下层绿叶的上表皮无显著差异,但是下层绿叶的下表皮细胞较小,气孔密度较低;(2)下层绿叶的叶肉组织较薄,细胞较大,排列疏松;(3)在下层绿叶中叶绿体类囊体片层排列密集,叶肉组织内单位体积叶绿体数量较少;(4)冠层部位对光合色素的积累有明显影响。叶绿素缺少的金叶女贞叶片结构显著受到冠层部位的影响,光合色素的变化幅度较一般植物更大,不同冠层部位叶片叶色有明显差异,可以作为研究植物冠层部位对叶形态、结构和生理生化影响的好材料。