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Notes: Abstract Plant competition strongly affects the species composition of managed grassland. To identify relevant processes, Festuca pratensis (Huds.) and Dactylis glomerata (L.) were grown as monocultures or mixtures in boxes placed in the field for two seasons and subjected to two cutting frequencies. Root and shoot competition effects were separated using soil and aerial partitions. Shoot competition was analysed by measuring the vertical distribution of the leaf area and root competition by analysing the absorption of tracers. Values of relative yield indicated that the two grasses fully competed for the same limiting resources under the experimental conditions. The competitive ability of F. pratensis was lower during both years relative to D. glomerata. This was mainly related to its consistently lower shoot competitive ability, which was associated with less leaf area in the upper layers of the canopy and shorter leaves. Root competitive ability of F. pratensis changed with season. It was similar to that of D. glomerata during spring and autumn, but less during summer. The lower root competitive ability in summer might be due to the lower root activity of F. pratensis, measured as rubidium (Rb) and strontium (Sr) absorption in July, 0·1 and 0·2 m below the ground. Cutting frequency did not greatly influence the relative importance of root and shoot competition during the first growing season. However, under infrequent defoliation, the competitive ability of F. pratensis decreased markedly in full competition during the second growing season. These results suggest that distinct differences in the canopy structure and root activity of different plant species can affect the relative importance of root and shoot competition during the season.

Notes: The growth of a timothy sward, a timothy/white clover mixture and a multi-grass/white clover mixture were compared and related to climatic factors at two sites of different altitudes.Temperature and radiation had a greater effect on sward growth than did species composition or management. If the reproductive and vegetative growth periods were studied separately, then more than 90% of the variation in yield could be explained by temperature and radiation. During spring reproductive growth, yields per unit increment of temperature or radiation were 2–3 times higher than during summer vegetative growth. Spring growth was affected more by temperature than by radiation and the lower yields at the higher altitude were also related to lower temperatures.

Notes: Seedlings of Rumex obtusifolius L. were grown in gaps (11 cm diameter) in an established sward at Lolium perenne L., Root competition from the surrounding sward was controlled by PVC tubes and shoot competition was controlled by cutting the sward frequently. All combinations, with and without root competition and with and without shoot competition, were used.Plants received either 40 or 80 kg N ha−1 month−1 and 0 or 300 kg K2O ha−1 and were harvested 39 d after emergence; a second harvest was made 12 d later, after shading to 20% of full sunlight.The shoot dry weight per plant of R. obtusifolius was much more affected by root competition than by shoot competition, especially at low nitrogen applications. Potassium supply had no effect.More dry matter was allocated to the leaf petioles under shaded conditions and the specific leaf area was greater. This morphological plasticity enabled R. obtusifolius to grow well under shading and to be affected only slightly by shoot competition from L. perenne.The results suggest that competition for nitrogen was the main factor limiting the initial grown of R. obtusifolius in a ryegrass sward and that the morphological characteristics of R. obtusifolius make it less susceptible to competition for light in the early stages of its development.

Notes: The aim of this experiment was to compare the growth strategies of Rumex obtusifolius L. and Lolium perenne L. at the seedling stage under different constant photon flux densities. Both species were grown simultaneously in growth chambers (20/15°C) at 50, 150 and 500 μmol m−2 s−1 and sampled on six occasions between the 14th and 34th day after seedling emergence. The relative growth rate of R. obtusifolius always exceeded that of L. perenne. R. obtusifolius allocated more dry matter to the leaves, thus resulting in an increase in specific leaf area and, consequently, leaf area ratio. The decrease in the relative growth rate that resulted from reducing the photon flux density from 500 to 50 μmol m−2 s−1 was lower for R. obtusifolius (−38%) than for L. perenne (−53%). The weed was less sensitive that the grass to the reduction in light intensity, mainly because its specific leaf area increased more and because its net assimilation rate decreased less. The results show that seedlings of R. obtusifolius are able to maximize dry matter production at a low photon flux density. This suggests that the establishment of R. obtusifolius seedlings cannot be prevented by the shade of an established sward. Comparaison de la croissance de R. obtusifolius L. et de L. perenne L. sous différentes densités de flux de photonsLe but de cet essai était de comparer la stratégie de croissance de plantules de R. obtusifolius et de L. perenne sous différentes densités de flux de photons. Les deux espèces ont été cultivées simultanément en chambres de croissance (20/15°C) à 50, 150 et 500 μmol m−2 s−1 et récoltées à six reprises entre le 14e et le 34e jour après la levee. Le taux de croissance relatif de R. obtusifolius a toujours été supérieur à celui de L. perenne. Nous avons observé chez R. obtusifolius un investissement plus important en matière sèche dans les feuilles ainsi qu'une surface foliaire spécifique et, par conséquent, un rapport surface foliaire/poids sec total plus élevés. La diminution du taux de croissance relatif consécutive à la réduction de la densité du flux de photons de 500 à 50 μmol m−2 s−1 fut plus faible chez R. obtusifolius (−38%) que chez L. perenne (−53%). La dicotylédone a été moins sensible à la réduction de l'intensité lumineuse que la graminée principalement parce que sa surface foliaire spécifique a davantage augmenté et également parce que son taux net d'assimilation a moins diminué. Nos résultats montrent que les plantules de R. obtusifolius sont capables de produire un maximum de matière sèche sous une faible densité du flux de photons. Ils suggèrent que l'implantation de plantules de R. obtusifolius ne peut pas être empêchée par l'effet d'ombrage d'un couvert en place. Vergleich des Wachstums von Rumex obtusifolius L. mit Lolium perenne L. bei verschiedenen Photonen-BestrahlungsstärkenZiel dieses Versuches war der Vergleich der Wachstumsstrategie der Keimpflanzen von Rumex obtusifolius L. und Lolium perenne L. bei verschiedenen konstanten Photonen-Bestrahlungsstärken. Beide Arten wurden gleichzeitig in Klimakammern (20/15 °C) bei 50, 150 und 500 pimol m−2 s−1 kultiviert, und zwischen dem 14 und 34. Tag nach dem Auflaufen wurden 6 Teilernten genommen. Die relative Wachstumsrate von R. obtusifolius war jener von L. perenne immer überlegen. R. obtusifolius transportierte mehr Trockensubstanz in die Blätter; die spezifische Blattfläche stieg an, und der Anteil Blattfläche an der gesamten Trockensubstanz nahm zu. Wurde die Photonen-Bestrahlungsstärke von 500 auf 50 μmol m−2 s−1 gesenkt, nahm die relative Wachstumsrate bei R. obtusifolius um 38% und bei L. perenne um 53% ab. Der Ampfer reagierte auf die Reduktion der Lichtintensität weniger empfindlich als das Gras, hauptsächlich weil sie ihre spezifische Blattfläche stärker erhöhte und die Nettoassimüationsrate weniger reduzierte. Die Ergebnisse zeigen, daß Keimpflanzen von R. obtusifolius in der Lage sind, auch bei niedrigen Photonen-Bestrahlungsstärken die Trockensubstanzproduktion zu maximieren. Es wird daher vermutet, daß unter dem Schatten eines Pflanzenbestandes das Etablieren der Keimpflanzen von R. obtusifolius nicht verhindert werden kann.

Notes: The effect of regeneration of Lolium perenne and Festuca rubra from embryogenic suspension cells and protoplasts on fertility and growth was evaluated. Embryogenic suspension cultures were either routinely subcultured or cryopreserved and re-established. Phenology, morphology and fertility of regenerated plants were studied for two growing seasons in a replicated field experiment. Most regenerated L. perenne and F. rubra plants showed a delay in inflorescence emergence, a reduced seed yield and differences in morphological traits when compared with seed-grown plants. For L. perenne, performance of plants regenerated from cryopreserved suspension cultures and protoplasts was similar to that of respective plants regenerated from routinely maintained suspension cultures. However, differences in performance were observed for respective regenerants in F. rubra. The phenotypic deviation observed was partly reflected in the randomly amplified polymorphic DNA (RAPD) analysis performed. However, regenerants of both species showing similar, or even superior performance to the seed-grown plants were also found. Embryogenic suspension cells and corresponding protoplasts of L. perenne and F. rubra have the potential for producing fertile, well-performing plants which can be integrated in breeding programs.

Notes: The broad-leaved dock (Rumex obtusifolius L.) is widely distributed, and is perceived as a major weed problem in intensively managed permanent grassland. The effects of nitrogen fertilization (120; 240 or 480 kg N ha−1), cutting frequency (every 4 or 6 weeks) and competing grasses on development, dry matter production and stem growth of Rumex were studied under field conditions. Young plants of Rumex were planted into established pure swards of Lolium multiflorum Lam., Lolium perenne L., Alopecurus pratensis L. and Poa pratensis L. In all treatments, only Lolium multiflorum hindered the growth of R. obtusifolius substantially. Cutting intervals of 6 weeks were more favourable to Rumex than were 4-week intervals. Its proportion of herbage yield increased from 2% at 120 kg N ha−1, taken as an average of the three grass swards (Lolium multiflorum not included), to 18% at 480 kg N ha−1. The reproductive capacity of Rumex was greatly influenced by the competing grass species: in the P. pratensis sward, Rumex developed up to 12.3 stems with inflorescences per plant per year, but only 0–9 stems developed in the L. multiflorum plots.The regrowth potential of R. obtusifolius after intense shoot competition and low interception of radiation was unexpectedly high. It is concluded that established dock plants cannot be controlled to any extent by strong competition of grasses, by variation in cutting frequency or by nitrogen fertilization. Effet de la fertilisation azotée et de la fréquence de coupe sur la compétitivité et la capacité de repousse de Rumex obtusifolius L. dans plusieurs pelouses de graminées Rumex obtusifolius L. est largement répandu, et il est considéré comme un problème majeur dans les prairies spermanentes intensives. Les effets de la fertilisation azotée (120, 240 ou 480 kg N ha−1), de la fréquence de coupe (Chaque 4 ou 6 semaines) et des graminées en compétition sur le développement, la production de matière sèche et la croissance de la tige de Rumex ont étéétudiées en plein champ. Des jeunes plantes de Rumex ont été plantées dans des gazons purs de Lolium multiflorum Lam., Lolium perenne L., Alopecurus pratensis L., et Poa pratensis L. De tous les traitements, seul Lolium multiflorum entrave subtantiellement la croissance de R. obtusifolius. Les intervalles de coupe de 6 semaines sont plus favorables au Rumex que ceux de 4 semaines. Sa proportion du rendement en herbe passe de 2%à 120 kg N ha−1 (moyenne des 3 gazons sans Lolium multiflorum), à 18%à 480 kg N ha−1. La capacité de reproduction de Rumex est fortement influencée par l'espèce de graminée en compétition: pour le P. pratensis, le Rumex a développé jusqu'à 12,3 tiges avec inflorescence par plante et par an, et seulement 0,9 pour les parcelles L. multiflorum.Le potentiel de reprise de R. obtusifolius après une forte compétitivé aérienne et une faible interception des radiations a étéétonnamment élevé. On conclut que l'extension d'un Rumex installé ne peut être combattue par une forte compétition des graminées, la variation dans la fréquence de coupe ou la fertilisation azotée. Einfluss der Stickstoffdüngung und der Schnitthäufigkeit auf die Konkurrenz von Rumex obtusifolius L. in verschiedenen GrasbeständenDer Stumpfblättrige Ampfer (Rumex obtusifolius L.) stellt auf intensiv bewirtschafteten Dauerwiesen ein wichtiges Unkraut dar. Der Einfluss der Stickstoffdüngung (120, 240 oder 480 kg N ha−1), der Schnitthäufigkeit (alle 4 oder 6 Wochen) und konkurrierender Gräser wurde im Freiland auf die Entwicklung, die Trockensubstanzproduktion und das Sprosswachstum von Ampfer untersucht. Junge Ampferpflanzen wurden in etablierte Reinbestände von Lolium multiflorum Lam., Lolium perenne L., Alopecurus pratensis L. und Poapratensis L. gepflanzt. Nur Lolium multiflorum vermochte das Wachstum von Ampfer wesentlich einzuschränken. Schnittintervalle von 6 Wochen förderten Rumex viel stärker als jene von 4 Wochen. Im Mittel der 3 Grasbestände (ohne L. multiflorum) stieg der Ertragsanteil von Rumex von 2% bei 120 kg N ha−1 auf 18% bei 480 kg N ha−1. Das Reproduktionsvermögen von Rumex wurde durch die konkurrierende Grasart stark beeinflusst, im Poa-pratensis-Bestand bildete Rumex 12,3 blütentragende Stengel pro Pflanze aus, im L. multiflorum-Bestand dagegen nur 0,9.Das Wiederaustrieb-svermögen von R. obtusifolius war nach intensiver Sprosskonkurrenz und niedriger Strahlungsabsorption unerwartet hoch. Aus den Ergebnissen ergibt sich, dass etablierte Ampfer-Pflanzen kaum durch konkurrenzstarke Gräser und durch die Änderung der Schnitthäufigkeit oder Stickstoffdüngung verdrängt werden können.

Notes: Photosynthesis is commonly stimulated in grasslands with experimental increases in atmospheric CO2 concentration ([CO2]), a physiological response that could significantly alter the future carbon cycle if it persists in the long term. Yet an acclimation of photosynthetic capacity suggested by theoretical models and short-term experiments could completely remove this effect of CO2. Perennial ryegrass (Lolium perenne L. cv. Bastion) was grown under an elevated [CO2] of 600 µmol mol−1 for 10 years using Free Air CO2Enrichment (FACE), with two contrasting nitrogen levels and abrupt changes in the source : sink ratio following periodic harvests. More than 3000 measurements characterized the response of leaf photosynthesis and stomatal conductance to elevated [CO2] across each growing season for the duration of the experiment. Over the 10 years as a whole, growth at elevated [CO2] resulted in a 43% higher rate of light-saturated leaf photosynthesis and a 36% increase in daily integral of leaf CO2 uptake. Photosynthetic stimulation was maintained despite a 30% decrease in stomatal conductance and significant decreases in both the apparent, maximum carboxylation velocity (Vc,max) and the maximum rate of electron transport (Jmax). Immediately prior to the periodic (every 4–8 weeks) cuts of the L. perenne stands, Vc,max and Jmax, were significantly lower in elevated than in ambient [CO2] in the low-nitrogen treatment. This difference was smaller after the cut, suggesting a dependence upon the balance between the sources and sinks for carbon. In contrast with theoretical expectations and the results of shorter duration experiments, the present results provide no significant change in photosynthetic stimulation across a 10-year period, nor greater acclimation in Vc,max and Jmax in the later years in either nitrogen treatment.

Notes: Abstract. The influence of leaf age, total leaf area and its dispersion in space on canopy photosynthesis were studied using microswards of red clover (Trifolium pratense L.) which were established in the greenhouse. Two varieties, Renova (flowering) and Molstad (non-flowering), were sown in separate plastic boxes at densities of 225, 400 and 625 plants per m2.Vertical distribution of photosynthetically active radiation (PAR), leaf area, leaf age and 14CO2-fixation were determined periodically. Net photosynthesis and dark respiration of canopies were measured. Maximum photosynthetic capacity of individual leaves was measured on plants taken from the intact canopy or from plants where shading of the growing leaves had been prevented.Net photosynthetic rate of canopies increased linearly with leaf area index (LAI) up to an LAI of 3.5 and then declined at higher LAI, independent of variety and sowing density. Below the optimum LAI, net photosynthesis depended mainly on interception of PAR. Decrease in canopy photosynthesis above the optimum LAI was due to a higher proportion of old leaves with decreased photosynthetic capacity, and not to an increase in respiring plant parts. It is concluded that LAI and position of leaf age categories in the canopy are more important than vertical distribution of leaf area in determining canopy photosynthesis of red clover.

Notes: The effect of an elevated partial pressure of CO2 (pCO2) on carbohydrate concentrations in source leaves and pseudo-stems (stubble) of Lolium perenne L. (perennial ryegrass) during regrowth was studied in a regularly defoliated grass sward in the field. The free air carbon dioxide enrichment (FACE) technology enabled natural environmental conditions to be provided. Two levels of nitrogen (N) supply were used to modulate potential plant growth. Carbohydrate concentrations in source leaves were increased at elevated pCO2, particularly at low N supply. Elevated leaf carbohydrate concentrations were related to an increased structural carbon (C) to N ratio and thus reflected an increased C availability together with a N-dependent sink limitation. Immediately after defoliation, apparent assimilate export rates (differences in the carbohydrate concentrations of young source leaves measured in the evening and on the following morning) showed a greater increase at elevated pCO2 than at ambient pCO2; however, replenishment of carbohydrate reserves was not accelerated. Distinct, treatment-dependent carbohydrate concentrations in pseudo-stems suggested an increasing degree of C-sink limitation from the treatment at ambient pCO2 with high N supply to that at elevated pCO2 with low N supply. During two growing seasons, no evidence of a substantial change in the response of the carbohydrate source in L. perenne to elevated pCO2 was found. Our results support the view that the response of L. perenne to elevated pCO2 is restricted by a C-sink limitation, which is particularly severe at low N supply.

Notes: The objective of this investigation was to examine the effect of an elevated atmospheric CO2 partial pressure (pCO2) on the N-sink strength and performance of symbiotic N2 fixation in Trifolium repens L. cv. Milkanova. After initial growth under ambient pCO2 in a nitrogen-free nutrient solution, T. repens in the exponential growth stage was exposed to ambient and elevated pCO2 (35 and 60 Pa) and two levels of mineral N (N-free and 7·5 mol m–3 N) for 36 d in single pots filled with silica sand in growth chambers. Elevated pCO2 evoked a significant increase in biomass production from day 12 after the start of CO2 enrichment. For plants supplied with 7·5 mol m–3 N, the relative contribution of symbiotically fixed N (%Nsym) as opposed to N assimilated from mineral sources (15N-isotope-dilution method), dropped to 40%. However, in the presence of this high level of mineral N, %Nsym was unaffected by atmospheric pCO2 over the entire experimental period. In plants fully dependent on N2 fixation, the increase in N yield reflects a stimulation of symbiotic N2 fixation that was the result of the formation of more nodules rather than of higher specific N2 fixation. These results are discussed with regard to physiological processes governing symbiotic N2 fixation and to the response of symbiotic N2 fixation to elevated pCO2 in field-grown T. repens.