Nitrogen Use within the Growing Leaf Blade of Tall Fescue

Leaf elongation rate (LER) of grasses depends on N supply and is expressed mostly through cell production, whereas most N in mature leaf tissues is chloroplastic. Our objective was to evaluate a possible competition for N between cell production and chloroplast development processes, utilizing the gradient of cell development along the leaf growth zone of tall fescue (Festuca arundinacea Schreb.). Under the two contrasting N regimes, total N content was highest in the cell production zone, declined sharply as cells elongated, and remained relatively constant in more distal positions, at values close to those measured in mature tissues. A similar pattern was found for N in proteins and nucleic acids that were not soluble in 80% ethanol. Content of N compounds soluble in 80% ethanol was higher in the cell production and elongation zones than in mature parts of the leaf. NO3- N content was low in the cell production zone and increased in the cell elongation zone for high-N plants. The deposition rate of total N in the growth zone was much higher with plants in high N than in those shifted to no N. For both N regimes, most N was deposited during cell production and early cell elongation. Little N was deposited during cell maturation where ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) was being actively synthesized. This suggests that synthesis of Rubisco, and probably other chloroplastic proteins, occurs largely from recycling of N that was previously incorporated into proteins during cell production. Thus, Rubisco content in mature tissues is more closely associated with N deposited during cell production than with N deposited during its biosynthesis.     

Nitrogen Effects on Leaf Anatomy within the Intercalary Meristems of Tall Fescue Leaf Blades

Longitudinal elongation contributes most to leaf area expansionof grasses and its rate is known to be strongly affected byN. Our objective was to determine the effect of two N regimes(N₀and N+) on the gradient of leaf tissue formation in meristemsof two contrasting tall fescue (Festuca arundinacea Schreb.)genotypes. Proportions of epidermal, mesophyll and vasculartissue as well as intercellular air space were determined throughoutthe base of actively elongating leaves. The area of leaf transversesections nearly doubled between the ligule and the distal endof the growth zone (about 30 mm), and was mainly associatedwith lateral epidermal and mesophyll cell division in the proximal5.0–7.5 mm. Further increase in transverse area was dueto the formation of intercellular airspace and transverse expansionof epidermal cells. Depending on genotype and N treatment themesophyll, epidermis, vascular bundles and air space comprised45–54%, 20–28%, 6–9%, and 17–21%, respectively,of transverse leaf area in the distal part of the growth zone.After a slight increase close to the leaf base, the area ofvascular tissue remained constant throughout the growth zone.The proportion of air space to mesophyll space was higher atN₀than at N+ because mesophyll area was enhanced by N+ to agreater degree than by N₀. In the genotype with slow leaf elongation,the increase in cross-sectional leaf area was due to an increasein both leaf width and leaf thickness. In the genotype whichhad faster leaf elongation and wider leaves, only leaf thicknesswas enhanced by N+. Copyright 2001 Annals of Botany Company Festuca arundinacea(Schreb.), tall fescue, leaf anatomy, growth zone, nitrogen     

Carbohydrate Metabolism in Leaf Meristems of Tall Fescue : II. Relationship to Leaf Elongation Rates Modified by Nitrogen Fertilization

Our objective was to examine alterations in carbohydrate status of leaf meristems that are associated with nitrogen-induced changes in leaf elongation rates of tall fescue (Festuca arundinacea Schreb.). Dark respiration rates, concentrations of nonstructural carbohydrates, and soluble proteins were measured in leaf intercalary meristems and adjacent segments of elongating leaves. The two genotypes used differed by 43% in leaf elongation rate. Application of high nitrogen (336 kilograms per hectare) resulted in 140% higher leaf elongation rate when compared to plants receiving low nitrogen (22 kilograms per hectare). Leaf meristems of plants receiving high and low nitrogen had dark respiration rates of 5.4 and 2.9 microliters O₂ consumed per milligram structural dry weight per hour, respectively. Concentrations of soluble proteins were lower while concentrations of fructan tended to be slightly higher in leaf meristems of low-nitrogen plants when compared to high-nitrogen plants. Concentrations of reducing sugars, nonreducing sugars, and takadiastase-soluble carbohydrate of leaf meristems were not affected by nitrogen treatment. Total nonstructural carbohydrates of leaf meristems averaged 44 and 39% of dry weight for low- and high-nitrogen plants, respectively. Within the leaf meristem, approximately 74 and 34% of the pool of total nonstructural carbohydrate could be consumed per day in high- and low-nitrogen plants, respectively, assuming no carbohydrate import to the meristem occurred. Plants were able to maintain high concentrations of nonstructural carbohydrates in leaf meristems despite a 3-fold range in leaf elongation rates, suggesting that carbohydrate synthesis and transport to leaf intercalary meristems may not limit leaf growth of these genotypes.     

Fructosyltransferase Activities in the Leaf Growth Zone of Tall Fescue

High concentrations of water-soluble carbohydrates, mainly fructan, accumulate in the growth zone of tall fescue (Festuca arundinacea Schreb.) leaf blades. We studied sucrose-hydrolyzing activities in the leaf growth zone because of their importance in carbohydrate partitioning. Sucrose hydrolysis in the basal 1.5 cm was largely due to fructosyltransferases, which had activities up to 10 times higher than in fully developed leaf tissue. Three fructosyltransferases (F1, F2, and F3) were purified from the leaf growth zone. Each synthesized, from either sucrose or 1-kestose, a mixture of trisaccharides and higher-order oligofructans identical with the low-degree of polymerization fructan extracted from similar plant tissue. The highly purified fructosyltransferases retained ability (13%) to transfer fructose from sucrose to water. Time-dependent and substrate-dependent studies, using sucrose as the substrate, showed proportional production of fructose and glucose, indicating that both products are from the same enzyme. Fructosyltransferase was calculated to contribute about half the total transfer of fructose to water in the basal 1.5 cm. Invertase activity increased to near 2.0 cm when fructosyl transfer to sucrose and other oligofructans decreased. Invertase was the major activity for sucrose hydrolysis at positions distal to 3.0 cm.     

Defoliation and Leaf Age Influence on Ergot Alkaloids in Tall Fescue

A controlled environment experiment was conducted to determinethe influence of defoliation on the regrowth and developmentof two tall fescue (Festuca arundinacea Schreb.) host-endophyte(Neotyphodium coenophialum Morgan-Jones and Gams (Glenn, Bacon,Price and Hanlincomb. nov. ) associations (DN2 and DN11), knownto differ in morphology and alkaloid production capacity. Defoliationtreatments included an uncut control, and clipping to a 5- or10-cm residue height. In a separate experiment, leaf age effectswere determined on an uncut plant canopy. Ergot alkaloid concentrationwas greatest in pseudostem and least in harvested leaf. Non-infectedplants were devoid of alkaloid. Alkaloid production (expressedas a function of dry matter) and yield were greater in uncutthan clipped plants, and were greater in DN11 than DN2 plants.Alkaloid production increased with increasing N concentrationin both associations; however, the rate of production was influencedby concentration of non-structural carbohydrate. Leaf age influencedleaf mass and alkaloid concentration and yield. Oldest leaves(>6 weeks after appearance) of DN2 and DN11 had the lowestconcentrations of ergot alkaloid, while leaves that were 2 to4 weeks old had the greatest alkaloid concentration and yield(alkaloid concentrationxleaf mass). Repeated defoliation reducedthe production of alkaloids, but did so as a function of non-structuralcarbohydrate. Our data suggest that alkaloid production canbe modified by defoliation as well as by host-endophyte association. Neotyphodium coenophialum ; Festuca arundinacea ; N; non-structural carbohydrate     

Effect of Ploidy on Quality of Tall Fescue, Italian Ryegrass x Tall Fescue and Tall Fescue x Giant Fescue Hybrids

Selected quality parameters were measured for forage leaf tissuefrom a spaced-plant nursery. The genotypes used were Ky 31 tallfescue and hybrids of Italian ryegrass (Lolium multiflorum Lam.)x tall fescue (Festuca arundinacea Schreb.) and tall fescuex giant fescue [Fescue gigantea (L ) Vill.]. Hybrid ploidy rangedfrom 2n = 28 to 84 chromosomes. Forage quality was characterizedby neutral detergent fibre (NDF), acid detergent fibre (ADF),total soluble carbohydrates (TSC) nutritive value index (NVI),hemicellulose, and in vitro dry matter disappearance (DMD). Quality of tall fescue, as measured by increased DMD, was improvedby hybridization with giant fescue. Improved DMD and NVI correlatedwith lower NDF and ADF in the hybrids. A few hybrids of Italianryegrass x tall fescue (2n = 28) were higher in some qualityparameters than Ky 31. Tall fescue x giant fescue hybrids (2n= 80 to 84), as a group, had significant quality improvementover Ky 31 in higher DMD and NVI and lower NDF and ADF. Whilesome individual hybrids within each group were significantlyhigher in quality, only the 2n = 80 to 84 chromosome group wasconsistently higher than Ky 31. Prediction equations for DMD,NDF, and ADF were established based on solvent extraction withnear-infrared reflectance spectroscopy (NIRS). Linear correlationcoefficients between chemical measurement and NIRS for eachquality parameter were 0–95 or higher. Acid detergent fibre, neutral detergent fibre, dry matter disappearance, hemicellulose, nutritive value index, Festuca arundinacea, Festuca gigantea, Lolium multiflorum     

Analysis of the Nitrogen Response of Leaf Extension in Lolium temulentum Seedlings

Lolium temulentum seedlings were grown on a nutrient mediumcontaining NH₄NO₂ at 0, 0·1, 0·5, 1·0 and4·3 mmoll^–1 as the sole N source. Relative andabsolute extension rates, maximal leaf size, duration of extensiongrowth, rate of leaf appearance and plastochron index were determinedfrom the parameters of Richards functions fitted to lengthsof laminae measured at intervals after sowing. The final lengthof leaf I was relatively insensitive to N whereas mean relativeextension rate was increased and duration of growth decreasedwith increasing NH₄NO₂ concentration. Leaves 2 and 3 enlargedprogressively with N at concentrations up to 1·0 mmoll^–1but were unresponsive thereafter. There was no significant correlationbetween final length and mean relative extension rate for leaves1 to 3. Leaves 4 to 6 continued to show increasing length beyond1·0 mmoll^–1 N and final length was significantlycorrelated with mean relative extension rate. Increasing N increasedthe rate of leaf appearance by decreasing the duration of leafextension and plastochron. These results are discussed in relationto the control of leaf and N turnover. Lolium temulentum, rye grass, leaf extension, nitrogen, Richards function, growth analysis     

Response of Fructan to Water Deficit in Growing Leaves of Tall Fescue

Changes in dry matter and water-soluble carbohydrate components, especially fructan, were examined in the basal 25 mm of expanding leaf blades of tall fescue (Festuca arundinacea Schreb.) to assess their roles in plant response to water deficit. Water was withheld from vegetative plants grown in soil in controlled-environment chambers. As stress progressed, leaf elongation rate decreased sooner in the light period than it did in the dark period. The decrease in growth rate in the dark period was associated with a decrease in local relative elongation rates and a shortening of the elongation zone from about 25 mm (control) to 15 mm. Dry matter content of the leaf base increased 23% during stress, due mainly to increased water-soluble carbohydrate near the ligule and to increased water-soluble, carbohydrate-free dry matter at distal positions. Sucrose content increased 258% in the leaf base, but especially (over 4-fold) within 10 mm of the ligule. Hexose content increased 187% in the leaf base. Content of total fructan decreased to 69% of control, mostly in regions farther from the ligule. Fructan hydrolysis could account for the hexose accumulated. Stress caused the osmotic potential to decrease throughout the leaf base, but more toward the ligule. With stress there was 70% less direct contribution of low-degree-of-polymerization fructan to osmotic potential in the leaf base, but that for sucrose and hexose increased 96 and 67%, respectively. Thus, fructan metabolism is involved but fructan itself contributes only indirectly to osmotic adjustment.     

Specific Leaf Weight in Zones of Cell Division, Elongation and Maturation in Tall Fescue Leaf Blades

Patterns of change in specific leaf weight (SLW), water-solublecarbohydrate (WSC) content and leaf width were used to delineatethe region of secondary cell wall accumulation, and determinethe rate of increase in structural material along a developingleaf blade of tall fescue (Festuca arundinacea Schreb.). Structuralspecific leaf weight (SSLW) was determined by subtracting WSCmass from dry weight to emphasize structural material. Becausemeristematic activity, cell elongation, and cellular maturationare arranged successively in the grass leaf, these patternsrepresent a developmental sequence through which each segmentof the leaf blade passes. Patterns were generally similar fortwo genotypes, one selected for high (HYT) and the other forlow (LYT) yield per tiller, for a single genotype grown at 17or 25 C, and for two field-grown populations which differedin leaf area expansion rate (LAER). In all three studies, the elongation zone of the developingleaf had 31 to 39 per cent WSC on a dry weight basis. The LYTgenotype had a higher SLW at all stages of development whengrown at 17 than at 25 C, due to greater WSC accumulation.At 20 C, the HYT genotype had a higher SLW all along the elongatingleaf blade than the LYT genotype. This difference was due toa difference in SSLW, while WSC content was similar. The LERwas 64 per cent higher in the high population than the low,but elongation zones were similar in WSC. In all cases, SSLWwas high in the meristematic region, lowest near the distalend of the cell elongation zone, then increased linearly astissue matured. Rate of increase in SSLW was 8.5 and 5.2 g m^–2d^–1 for the HYT and LYT genotypes, respectively, and 7.6and 6.7 g m^–2 d^–1 for the high and low LAER populations,respectively. Festuca arundinacea Schreb., tall fescue, specific leaf weight, leaf width, water-soluble carbohydrates, leaf elongation rate     

Growth Rates and Carbohydrate Fluxes within the Elongation Zone of Tall Fescue Leaf Blades

Investigations were performed to better understand the carbon economy in the elongation zone of tall fescue leaf blades. Plants were grown at constant 21°C and continuous 300 micromoles per square meter per second photosynthetic photon flux density where leaf elongation was steady for several days. Elongation occurred in the basal 20 mm of the blade (0-20 millimeters above the ligule) and was maximum at 9 to 12 millimeters. Eight 3-millimeter long segments were sampled along the length of the elongation zone and analyzed for water-soluble carbohydrates. Sucrose concentration was high in the zone of cell division (0-6 millimeters) whereas monosaccharide concentration was high at and distal to the location where cell elongation terminated (20 millimeters). Fructan concentration increased in the basal part, then remained constant at about 85% of the total mass of water-soluble carbohydrates through the remainder of the elongation zone. Data on spatial distribution of growth velocities and substance contents (e.g. microgram fructan per millimeter leaf length) were used to calculate local net rates of substance deposition (i.e. excess rates of substance synthesis and/or import over substance degradation and/or export) and local rates of sucrose import. Rates of sucrose import and net deposition of fructan were positively associated with local elongation rate, whereas net rates of sucrose deposition were high in the zone of cell division and those of monosaccharide were high near the termination of elongation. At the location of most active elongation imported sucrose (29.5 milligrams per square decimeter per hour) was used largely for synthesis of structural components (52%) and fructan (41%).     

Effects of the Fungal Endophyte Acremonium coenophialum on Nitrogen Accumulation and Metabolism in Tall Fescue

Infection by the fungal endophyte Acremonium coenophialum affected the accumulation of inorganic and organic N in leaf blades and leaf sheaths of KY 31 tall fescue (Festuca arundinacea Schreb.) grown under greenhouse conditions. Total soluble amino acid concentrations were increased in either the blade or sheath of the leaf from infected plants. A number of amino acids were significantly increased in the sheath, but only asparagine increased in the blade. Infection resulted in higher sheath NH₄⁺ concentrations, whereas NO₃⁻ concentrations decreased in both leaf parts. The effects on amino acid, NO₃⁻, and NH₄⁺ concentrations were dependent upon the level of N fertilization and were usually apparent only at the high rate (10 millimolar) of application. Administration of ¹⁴CO₂ to the leaf blades increased the accumulation of ¹⁴C in their amino acid fraction but not in the sheaths of infected plants. This may indicate that infection increased amino acid synthesis in the blade but that translocation to the sheath, which is the site of fungal colonization, was not affected. Glutamine synthetase activity was greater in leaf blades of infected plants at high and low N rates of fertilization, but nitrate reductase activity was not affected in either part of the leaf. Increased activities of glutamine synthetase together with the other observed changes in N accumulation and metabolism in endophyte-infected tall fescue suggest that NH₄⁺ reassimilation could also be affected in the leaf blade.     

The Effect of Temperature during Growth on the Subsequent Rate of Photosynthesis in Leaves of Tall Fescue (Festuca arundinacea Schreb)

Two experiments are reported in which young plants of tall fescuewere grown in temperature regimes of 20 °C day/15 °Cnight or 10 °C day/5 °C night until the fourth leafon the main stem was fully expanded. These temperature regimeswere then either changed over for individual plants or continuedunchanged up to the seven-leaf stage. Photosynthesis and respirationrates were determined in the fourth and subsequent leaves andalso in ageing leaves, using an infra-red gas analyser in anopen system and at temperatures of 10 and 20 °C in one and10, 15, 20, and 25 °C in the other experiment. Rates of apparent photosynthesis per unit leaf area in fullyexpanded leaves differed little as a result of previous treatmentand were not greatly affected by temperature during measurement.However, the specific leaf area and the rate of apparent photosynthesisper unit dry weight were higher in plants grown at the hightemperature. Leaves from the high-temperature regime had a higheroptimum temperature for apparent photosynthesis, a shorter life,and a lower respiration rate at any one temperature of measurementthan did leaves from the low-temperature regime. After transfer from one temperature regime to the other, therate of apparent photosynthesis of the next leaf to become fullyexpanded was higher in plants transferred from low to high temperatureand lower in plants transferred from high to low than in plantsremaining in either temperature regime; the leaves which subsequentlyexpanded had rates similar to those of unchanged plants. Inleaves which were fully expanded at the time of transfer, therate of apparent photosynthesis rose after transfer to the high-temperatureregime and fell after transfer to the low-temperature regime. These results are discussed in relation to growth-analysis datafrom other plants grown in the same conditions.     

Leaf Extension Rate in Temperate Pasture Grasses in Relation to Assimilate Pool in the Extension Zone

In two growth cabinet experiments the leaf extension rate (LER)was studied under a 14 h photoperiod followed by prolonged darkness,in tillers of the perennial temperate pasture grasses Phalaristuberosa L. cv. Sirosa and Dactylis glomerata L. cv. Currie.Levels of soluble non-structural carbohydrates and total     

Temperature Effects upon Inflorescence and Seed Development in Tall Fescue (Festuca arundinacea Schreb.)

The effects of three temperatures 15, 20, and 25 °C uponinflorescence and seed development in tall fescue (Festuca arundinaceaSchreb) between inflorescence emergence and seed maturity werestudied. Increasing temperature over this range reduced culmlength and the number of florets per spikelet, hastened theonset of anthesis and pollen release, increased relative growth-rateof the florets 9 days after peak anthesis, reduced the periodof seed development and 1000 seed weight No large effects oftemperature upon the percentage of florets setting seed werefound. The practical implications of these results are discussed.     

Accumulation of Ergopeptide Alkaloids in Symbiotic Tall Fescue Grown under Deficits of Soil Water and Nitrogen Fertilizer

The fungus Acremonium coenophialum is endophytically associated with tall fescue (Festuca arundinacea Schreber). Within this symbiotum the fungus produces ergopeptide alkaloids, which are associated with livestock toxicoses. Environmental effects on the production of ergot alkaloids within the symbiotum are unknown. We conducted a greenhouse study of the effects of flooding, nitrogen rate during fertilization (11, 73, and 220 mg of N per pot weekly), nitrogen form (3.4 and 34 mg of N as NH₄⁺ or NO₃^- per pot), and drought stress (-0.03, -0.05, and -0.50 MPa) on ergopeptide alkaloid concentrations in one genotype of nonsymbiotic and symbiotic tall fescue grown in plastic pots. It was determined that the concentration of ergovaline, the major type of ergopeptide alkaloid, was increased but was not as high as that in nonflooded controls. Total ergopeptide and ergovaline concentrations in plants receiving high (220 mg of N per pot) and low (11 mg of N per pot) levels of NH₄NO₃ fertilization were not affected by flooding. The form of nitrogen was important since all concentrations of NO₃^--N increased ergopeptide alkaloid content, as opposed to the effects of NH₄⁺-N, which was effective only at high concentrations (34 mg of N per pot). Ergopeptide concentrations were highest in drought-stressed plants grown at -0.50 MPa and fertilized at the moderate or high N rate. The results suggest that within this genotype, ergopeptide alkaloid biosynthesis by the fungus is not appreciably affected by flooding but is greatly increased by high rates of N fertilization and moderate water deficit.     

水稻叶色氮素反应的基因型间差异

在大田不施氮素及高氮两个处理下,以叶片的SPAD值作为评价水稻氮素利用能力的参数,对146个不同基因型水稻进行了叶色深浅及对氮素敏感性不同的种质资源鉴定。通过测定抽穗前不同生育时期不同基因型水稻叶片的SPAD值,筛选出对氮素反应迟钝且叶色较浅的基因型19个、对氮素反应迟钝且叶色较深的基因型20个和对氮素敏感且叶色较浅的基因型20个、对氮素敏感且叶色较深的基因型11个。     

Photosynthesis in Tall Fescue : IV. Carbon Assimilation Pattern in two Genotypes of Tall Fescue Differing in Net Photosynthesis Rates

We previously reported that the net photosynthetic rate of a decaploid genotype (I-16-2) of tall fescue (Festuca arundinacea Schreb.) was 32 to 41 versus 22 milligrams CO₂ per square decimeter per hour in a hexaploid genotype (V6-802) (Randall, Nelson, Asay Plant Physiol 59: 38-41). The high rate was later correlated with increases in total ribulose 1,5-bisphosphate carboxylase protein (17%) and activity (27%) (Joseph, Randall, Nelson Plant Physiol 68: 894-898). This report characterizes photosynthesis with respect to light saturation and early products of photosynthesis in an attempt to identify regulatory metabolic site(s) in these two genotypes. Analysis of the early products of photosynthesis indicated that both genotypes fixed CO₂ via the Calvin-Benson cycle with phosphoglyceric acid as the initial primary product. Both genotypes had similar ¹⁴C-labeled intermediates. Sucrose was the primary sink of ¹⁴CO₂ assimilation. After 10 min of ¹⁴CO₂ assimilation with attached leaves, sucrose accounted for 89% (decaploid) and 81% (hexaploid) of the total ¹⁴C incorporated. In 10 min, this amounted to 1.3 (decaploid) and 0.8 (hexaploid) μmol [¹⁴C]sucrose formed g fresh weight⁻¹ and reflected the observed differences in photosynthetic rates. There was limited labeling of starch (1%) and fructan (1%). Results of total nonstructural carbohydrates and P_i analysis also demonstrated sucrose was the predominant carbohydrate in fescue leaves. Quantitative differences in sucrose and P_i between the two genotypes may reflect changes in partitioning and this possibility is discussed.     

Host Status of Endophyte-Infected and Noninfected Tall Fescue Grass to Meloidogyne spp

Tall fescue grass cultivars with or without endophytes were evaluated for their susceptibility to Meloidogyne incognita in the greenhouse. Tall fescue cultivars evaluated included, i) wild-type Jesup (E+, ergot-producing endophyte present), ii) endophyte-free Jesup (E-, no endophyte present), iii) Jesup (Max-Q, non-ergot producing endophyte) and iv) Georgia 5 (E+). Peach was included as the control. Peach supported greater (P ≤ 0.05) reproduction of M. incognita than all tall fescue cultivars. Differences in reproduction were not detected among the tall fescue cultivars and all cultivars were rated as either poor or nonhosts for M. incognita. Suppression of M. incognita reproduction was not influenced by endophyte status. In two other greenhouse experiments, host susceptibility of tall fescue grasses to two M. incognita isolates (BY-peach isolate and GA-peach isolate) did not appear to be related to fungal endophyte strain [i.e., Jesup (Max-Q; nontoxic endophyte strain) vs. Bulldog 51 (toxic endophyte strain)]. Host status of tall fescue varied with species of root-knot nematode. Jesup (Max-Q) was rated as a nonhost for M. incognita (BY-peach isolate and GA-peach isolate) and M. hapla, a poor host for M. javanica and a good host for M. arenaria. Bulldog 51 tall fescue was also a good host for M. arenaria and M. javanica, but not M. incognita. Jesup (Max-Q) tall fescue may have potential as a preplant control strategy for M. incognita and M. hapla in southeastern and northeastern United States, respectively.     

Carbon Balance of Tall Fescue (Festuca arundinacea Schreb.): Effects of Nitrogen Fertilization and the Growing Season

The C balance of a tall fescue sward grown under different ratesof N fertilization in summer, autumn, and spring was calculatedusing models derived from measurements of shoot growth, canopygross photosynthesis, shoot respiration and of C partitioningto the roots. Under the diverse growing conditions associatedwith the seasons and the N fertilization, C utilization forabove- and below-ground biomass accumulation never exceeded39 and 14% of the canopy gross photosynthesis, respectively.Carbon losses attributed to root respiration and exudation,which were estimated by difference between canopy net photosynthesisand total growth, ranged between 3 and 30% of canopy gross photosynthesis.Seasonal differences in shoot growth could be attributed tothe amount of intercepted radiation, the radiation-use efficiencyand the C partitioning to the roots. The effect of N deficiencyon shoot growth can be attributed to its effects on canopy photosynthesis(principally resulting from changes in intercepted photosyntheticallyactive radiation) and C partitioning. In comparison with theeffect on shoot growth, the effect of the N deficiency on thecanopy gross photosynthesis per unit of light intercepted overthe regrowth cycle was limited. It is concluded that most ofthe effect of N fertilization on shoot growth is due to changesin C partitioning which result in faster leaf area developmentand greater light interception.Copyright 1994, 1999 AcademicPress Tall rescue, Festuca arundinacea Schreb., carbon balance, nitrogen, grass, fertilization     

The Effect of Age on the Rate of Apparent Photosynthesis in Leaves of Tall Fescue (Festuca arundinaceaSchreb.)

n apparatus is described in which the rate of apparent photosynthesisof single attached or detached grass leaves was determined bymeasuring with an infra-red gas analyser the uptake of carbondioxide from a stream of air passed over the leaves. In one experiment the apparatus was used to determine the apparentphotosynthesis at one time of the three youngest leaves removedfrom a tiller of tall fescue plants grown in the field. Resultsshowed that each older leaf had a lower rate of apparent photosynthesisthan its successor. To determine the real effect of age of theleaf without any possible interaction with position of the leafor an after-effect of environment, measurements of apparentphotosynthesis were made in two further experiments on leavesattached to the main stem on plants grown in controlled environments.Results were obtained from samples of the fifth leaf of increasingage from full expansion. These showed a progressive declinein apparent photosynthesis with increasing age: in one experimentthis was preceded by a small initial rise after complete leafexpansion. There were no marked changes in the weight of theleaf as it aged and, in consequence, the form of the resultswere similar when expressed on either a leaf area or dry weightbasis.