Fleshy fungi in grass fields. Dependence on fertilization, grass species, and age of field

An experimental field with grass had 48 squares which were treated with combinations of NPK fertilizers. The fungus flora was studied 1978–1980 by numerous yearly observations. A total of 40 species were noted: 36 agarics and 4 gasteromycetes. 24 species were confined to squares where nitrate was applied. 5 of these species were recorded as fairy ring formers. The fairy rings developed exclusively in the +N squares. Some common species grew in both +N and –N squares, while a few species were confined to –N squares. None of the latter group was found in abundance. The squares were divided in parcels sown with 4 species of lawn grasses. Several fungi showed preference for one or two grass species. The field consists of two parts, sown in 1977 and 1972 respectively. The immigration pattern into the new field is described.     

Fleshy fungi in grass fields. II. Precipitation and fructification

An experimental field with grass had 192 parcels treated with combinations of NKP fertilizers and sown with 4 species of grass. Species of fleshy fungi in the parcels were counted over 4 years with 17–24 observations per year. Precipitation was measured at the field. The occurrence of the species varied with the distribution of the precipitation through the season. In early summer and after prolonged desiccation the fungus flora developed 25–30 days after heavy rain, while after mowing and after shorter desiccation periods, a new crop of fruit bodies had developed after 5 days. These intervals correspond to the maturation period of the mycelia, and to the period of the development of primordia already formed. Differences in the yearly periodicity of the individual species are discussed.     

The effect of a nitrification inhibitor on the concentration of nitrate in grass during growth

Summary In an experiment on clay-loam soil ammonium sulphate with or without the nitrification inhibitor 2-chloro-6-(trichloromethyl) pyridine (at 2% of the weight of N in fertilizer) or sodium nitrate were applied at 100 or 200 lb N/acre to the seedbed before sowing S22 Italian ryegrass and again after the first cut. Eighty-four days after the first dressing, all the grass given 100 lb N/acre contained similar amounts of nitrate-N; with 200 lb N/acre, grass given ammonium sulphate alone contained most nitrate-N and grass given ammonium sulphate plus inhibitor least. Forty-two days after the second dressing, all the grass given 100 lb N/acre again contained similar amounts of nitrate-N and with 200 lb N/acre, grass given sodium nitrate contained most nitrate-N and that given ammonium sulphate plus inhibitor least.     

Feeding behaviour of yearling and older hybrid grass carp

Hybrid grass carp resulting from the cross of a female grass carp, Ctenopharyngodon idella (Val.), and a male bighead carp, Hypophthalmichthys ( Aristichthys ) nobilis Rich., 12–18 months old ( c . 300 mm T.L.) were studied in a two-part experiment to determine feeding preference and total daily consumption fish^-1 on selected species of aquatic plants. Fish were maintained in circular pools with 6840·8 1 of water inside a temperature-controlled greenhouse. Preference tests were conducted at three temperature ranges; 25–28° C, 17–20° C and 12–15° C. Based on the time to complete consumption or the relative quantity consumed, the most preferred plant was Lemna gibba when in combination with six other species. Chara sp., Najas guadalupensis and Potamogeton peciinatus were readily consumed and considered to be of about equal preference. Ceratophyllum demersum and Myriophyllum brasiliense were least preferred. Hybrid grass carp generally consumed as much plant material species^-1 and in the same order of preference at the 12–15°C range as they did at 25–28° C. In the second part, mean daily consumption (g) fish^-1 at 25·7–31·0° C for five plant species tested separately was as follows: Chara sp. 369·8; Lemna gibba 178·2; Najas guadalupensis 172·6; Hydrilla verticillata 106·4 and Ceratophyllum demersum 8·8.     

Influence of ammonium and nitrate supply on growth,nitrate reductase activity and N-use efficiency in a natural hybrid pine and its parents

Aims Selection of tree species with a high capacity to assimilate N and efficiently utilize N resources would facilitate the success of initial tree seedling establishment in infertile soils. The preference for N forms was tested using three pine species (Pinus densata, Pinus tabuliformis and Pinus yunnanensis). Pinus densata is a natural diploid hybrid between P. tabuliformis and P. yunnanensis .Methods Seedlings of three pine species were supplied with nitrate-N, ammonium-N (at two different pH regimes) or combined ammonium and nitrate as a nitrogen source in perlite culture in a controlled environment.Important findings Seedlings of P. densata had higher total biomass and net photosynthesis when supplied with nitrate-N and ammonium nitrate than with ammonium-N. In parental species, total biomass and net photosynthesis for P. yunnanensis seedlings was higher in ammonium-N than in nitrate-N, whereas the other parental species P. tabuliformis had the highest total biomass among species for all treatments except ammonium with CaCO 3. Most morphological traits in P. densata seedlings were intermediate between its two parental species. However, N-use efficiency and photosynthetic N-use efficiency of P. densata significantly exceeded both parents when supplied with nitrate-N and ammonium nitrate. The results suggested that the diploid hybrid tree species P. densata has a preference for nitrate and is not well adapted to ammonium-N as a sole nitrogen source regardless of the growth medium pH. Based on changes in environmental conditions, such as predicted future temperature increases in high altitude areas associated with climate change, P. densata is likely to be increasingly competitive and have wide adaptation in high altitude regions.     

Seasonal distribution of topsoil ammonium and nitrate under legume-grass and grass swards

Losses of soil N through leaching and N₂ fixation by legumes often are related to soil nitrate concentration. The seasonal distribution of soil ammonium and nitrate concentrations under ungrazed legume-grass and grass swards were evaluated on two experiments that were established in 1983 (Exp. 1) and in 1984 (Exp. 2). Treatments were white clover (Trifolium repens L.) (WC), red clover (Trifolium pratense L.) (RC), and birdsfoot trefoil (Lotus corniculatus L.) (BT), each grown with tall fescue (Festuca arundicacea Schreb.) (TF) at two legume proportions, and a pure stand of TF. The concentrations of both forms of N were measured in the top 20-cm layer during 2 years in Exp. 1 and for 1 year in Exp. 2. The concentrations of nitrate and ammonium were least in winter and spring, and greatest in summer. The concentration of nitrate for the mixtures decreased in the order WC-TF, RC-TF, and BT-TF in both summers of Exp. 1 but there were no mixture differences in Exp. 2. The concentration of soil ammonium was not affected by the treatments applied. We conclude that the concentration of soil nitrate usually was small for these swards but became greater and often dependent on species and legume proportion during summer. The concentration of soil ammonium also was greater in summer but was not affected by species or legume proportion. Journal of Paper no. J.-13359 of the Iowa Agric. and Home Econ. Exp Stn., Ames. Project 2281. Supported in part by the Facultad de Agronomía, Montevideo, Uruguay. Journal of Paper no. J.-13359 of the Iowa Agric. and Home Econ. Exp Stn., Ames. Project 2281. Supported in part by the Facultad de Agronomía, Montevideo, Uruguay.     

Positive feedbacks to growth of an invasive grass through alteration of nitrogen cycling

Understanding the mechanisms by which invasive plants maintain dominance is essential to achieving long-term restoration goals. While many reports have suggested invasive plants alter resource availability, experimental tests of feedbacks between invasive plants and soil resources are lacking. We used field observations and experimental manipulations to test if the invasive grass Microstegium vimineum both causes and benefits from altered soil nitrogen (N) cycling. To quantify M. vimineum effects on N dynamics, we compared inorganic N pools and nitrification rates in 20 naturally invaded and uninvaded plots across a range of mixed hardwood forests, and in experimentally invaded and uninvaded common garden plots. Potential nitrification rates were 142 and 63?% greater in invaded than uninvaded plots in forest and common garden soils, respectively. As a result, soil nitrate was the dominant form of inorganic N during peak M. vimineum productivity in both studies. To determine the response of M. vimineum to altered nitrogen availability, we manipulated the dominant N form (nitrate or ammonium) in greenhouse pots containing M. vimineum alone, M. vimineum with native species, and native species alone. M. vimineum productivity was highest in monocultures receiving nitrate; in contrast, uninvaded native communities showed no response to N form. Notably, the positive response of M. vimineum to nitrate was not apparent when grown in competition with natives, suggesting an invader density threshold is required before positive feedbacks occur. Collectively, our results demonstrate that persistence of invasive plants can be promoted by positive feedbacks with soil resources but that the magnitude of feedbacks may depend on interspecific interactions.     

禾草内生真菌对其它微生物的影响研究进展

禾草内生真菌在宿主植物的茎叶等地上组织中普遍存在,不仅能够提高禾草对生物与非生物逆境的抗性,而且能够对周围环境中的不同微生物类群产生影响。主要总结了禾草Neotyphodium/Epichlo内生真菌对病原真菌、丛枝菌根真菌和土壤微生物的影响及其作用机理。发现禾草内生真菌普遍存在对病原真菌的抑制作用,而对丛枝菌根真菌存在不对称的竞争作用,且因种类而异。禾草内生真菌对土壤微生物群落的作用则会随着土壤类型和时间等外界因素发生变化。禾草内生真菌对不同类群微生物的影响机制主要包括:通过生态位竞争、抑菌物质分泌、诱导抗病性等对病原真菌造成影响;通过根系化学物质释放、营养元素调节、侵染条件差异等对丛枝菌根真菌造成影响;通过根际沉积物和凋落物等对土壤微生物群落造成影响。禾草内生真菌产生的生物碱能提高宿主植物对包括昆虫在内草食动物采食的抗性,影响病原菌的侵入、定殖和扩展;根组织分泌物中包含次生代谢产物能够抑制菌根真菌、土传病原真菌及其它土壤微生物的侵染与群落组成;也可能通过次生代谢物影响禾草的其它抗性。因此,禾草内生真菌在植物-微生物系统中的作用应该给予更多的关注和深入研究。     

Nitrogen uptake by size-fractionated plankton in permanently well-mixed temperate coastal waters

Nitrogen uptake by net- (15–200 µm), nano- (1–15µm) and picoplankton (<1 µm) was measured overseasonal cycles at two stations with different patterns of biologicaland chemical cycles in the Morlaix Bay (western English Channel).Though assimilable dissolved N nutrient pool at both stationswas nitrate-dominated, characteristics of biomass and N uptakeby netplankton differed from conventional patterns in two respects.In the first, biomass (26–30%) and N uptake (36–43%)were less important than those of nanoplankton. In the second,the netplankton did not show any marked preference for nitrateover ammonium (nitrate to ammonium uptake ratios of 0.98 and1.08). In contrast, nanoplankton had a preference for ammoniumover nitrate (ammonium to nitrate uptake ratios of 2 and 1.2).N uptake by picoplankton was only 8% of total N uptake at bothstations and was supported mainly by regenerated N (66% ammoniumand 17% urea), with nitrate uptake detectable in only one instanceand nitrite uptake in none. Substrate-dependent uptake of ammoniumin all fractions and a higher ammonium uptake in the nanoplanktonfraction in summer at both stations when ambient ammonium concentrationswere high indicated that while nitrate may satisfy a part ofN requirements, availability of ammonium and its flux throughnanoplankton determine the magnitude of total N uptake in thesewaters. Most of the N uptake in picoplankton appears to be autotrophic,suggesting that a substantial part of heterotrophic uptake,if any, could be localized in the fractions >1 µm,and mediated by free-living and particle-bound bacteria.     

Inertia in an ombrotrophic bog ecosystem in response to 9 years' realistic perturbation by wet deposition of nitrogen,separated by form

Wet deposition of nitrogen (N) occurs in oxidized (nitrate) and reduced (ammonium) forms. Whether one form drives vegetation change more than the other is widely debated, as field evidence has been lacking. We are manipulating N form in wet deposition to an ombrotrophic bog, Whim (Scottish Borders), and here report nine years of results. Ammonium and nitrate were provided in rainwater spray as NH₄Cl or NaNO₃ at 8, 24 or 56 kg N ha^?1 yr^?1, plus a rainwater only control, via an automated system coupled to site meteorology. Detrimental N effects were observed in sensitive nonvascular plant species, with higher cumulative N loads leading to more damage at lower annual doses. Cover responses to N addition, both in relation to form and dose, were species specific and mostly dependent on N dose. Some species were generally indifferent to N form and dose, while others were dose sensitive. Calluna vulgaris showed a preference for higher N doses as ammonium N and Hypnum jutlandicum for nitrate N. However, after 9 years, the magnitude of change from wet deposited N on overall species cover is small, indicating only a slow decline in key species. Nitrogen treatment effects on soil N availability were likewise small and rarely correlated with species cover. Ammonium caused most N accumulation and damage to sensitive species at lower N loads, but toxic effects also occurred with nitrate. However, because different species respond differently to N form, setting of ecosystem level critical loads by N form is challenging. We recommend implementing the lowest value of the critical load range where communities include sensitive nonvascular plants and where ammonium dominates wet deposition chemistry. In the context of parallel assessment at the same site, N treatments for wet deposition showed overall much smaller effects than corresponding inputs of dry deposition as ammonia.     

Nitrogen nutrition inAtriplex hastata L.

Summary The investigation was carried out in order to ascertain whyAtriplex hastata, previously recorded as a nitrophilous plant, can colonise fly ash which has a low nitrogen content.Sinapis arvensis, a weed which does not grow on fly ash, was selected as a control plant. Solution-culture experiments showed that the two species differed in their responses to high and low levels of nitrogen, the growth rate ofS. arvensis being much more severely affected by low levels of 0 to 50 ppm.A. hastata was found to make very efficient use of its nitrogen supply. Both species grew better with nitrate than with ammonium nitrogen. InA. hastata, uptake of nitrogen was not influenced by pH when only nitrate was present. Where both nitrate and ammonium were available, nitrate was absorbed preferentially at low pH and ammonium at high pH. Some properties of fly ash which render it a suitable medium for growth ofA. hastata are discussed.     

Tall fescue EST-SSR markers with transferability across several grass species

Tall fescue (Festuca arundinacea Schreb.) is a major cool season forage and turf grass in the temperate regions of the world. It is also a close relative of other important forage and turf grasses, including meadow fescue and the cultivated ryegrass species. Until now, no SSR markers have been developed from the tall fescue genome. We designed 157 EST-SSR primer pairs from tall fescue ESTs and tested them on 11 genotypes representing seven grass species. Nearly 92% of the primer pairs produced characteristic simple sequence repeat (SSR) bands in at least one species. A large proportion of the primer pairs produced clear reproducible bands in other grass species, with most success in the close taxonomic relatives of tall fescue. A high level of marker polymorphism was observed in the outcrossing species tall fescue and ryegrass (66%). The marker polymorphism in the self-pollinated species rice and wheat was low (43% and 38%, respectively). These SSR markers were useful in the evaluation of genetic relationships among the Festuca and Lolium species. Sequencing of selected PCR bands revealed that the nucleotide sequences of the forage grass genotypes were highly conserved. The two cereal species, particularly rice, had significantly different nucleotide sequences compared to the forage grasses. Our results indicate that the tall fescue EST-SSR markers are valuable genetic markers for the Festuca and Lolium genera. These are also potentially useful markers for comparative genomics among several grass species.Electronic Supplementary Material Supplementary material is available for this article at .     

Relationships between soil seed bank,vegetation and soil fertility along an urbanisation gradient

Questions: How does urbanisation influence soil mineral nitrogen stocks (nitrate and ammonium stocks) and what are the consequences of these modifications on the functional diversity of the herbaceous vegetation (vascular plants) and the seed bank? Location: Nine study sites were located on an urbanisation gradient in the city of Rennes, France. Methods: Three urbanisation levels were defined: urban areas (high grey/green ratio), suburban (medium grey/green ratio) and periurban (low grey/green ratio). For each urbanisation level, nitrates and ammonium stocks were quantified; the herbaceous vegetation was surveyed as well as the soil seed bank (using the seedling emergence method). Results: Nitrate concentration increased with urbanisation (the nitrate level in urban plots was twice the concentration in periurban ones) whereas the ammonium level was higher in periurban areas than in urban areas. In urban plots, the vegetation and the seed bank were more nitrophilous, whereas the nitrogen requirement was lower for periurban species. The relationship between the seed bank and the above‐ground vegetation was not significant. Conclusions: The higher nitrate concentration in the urban area appeared to be related to higher concentrations of atmospheric pollutants found in this area and lower ammonium levels may be related to the higher temperature in urban areas (leading to higher nitrification rates). The shift in the composition of the seed bank and vegetation appeared to be a consequence of higher nitrate stocks. The dissimilarity between the seed bank and vegetation may be caused by enhanced emergence of nitrophilous species in urban areas.     

Microsatellite diversity of the agriculturally important alpine grass Poa alpina in relation to land use and natural environment

BACKGROUND AND AIMS: The Alpine Meadow Grass Poa alpina is common in subalpine and alpine natural sites and agriculturally used land, where it is an important fodder grass. Natural factors and human land use are supposed to have been shaping its genetic diversity for hundreds of years. The species comprises sexually and vegetatively reproducing plants. The aim of this study was to investigate the effects of agricultural land use, environmental factors and the mode of reproduction on the distribution of its microsatellite diversity within and among populations and to analyse whether its genetic diversity is correlated with plant species diversity in grassland parcels. METHODS: Genetic diversity of P. alpina was assessed with five microsatellite markers for 569 plants originating from 20 natural sites and from 54 grassland parcels of different cultural tradition, land use and altitude in the Swiss Alps. Due to polyploidy and frequent aneuploidy of the species, data analyses were based on the presence of microsatellite bands. KEY RESULTS: A low but significant differentiation was found in microsatellite bands among natural sites and agriculturally used parcels, while their microsatellite band diversity within populations did not differ. An increased differentiation was found in microsatellite bands with increasing geographic distance among parcels, and a differentiation among grazed and mown parcels, and among sexually and vegetatively reproducing populations. Band richness of sampled plants per village was higher for villages where parcels represented more different land-use types. Within populations, microsatellite band diversity was higher in grazed than in mown parcels. CONCLUSIONS: The diversity of human land use in the Alps was associated with genetic diversity of P. alpina. Therefore, the ongoing socio-economically motivated land-use changes, which reduce the number of different land-use types, will affect the genetic diversity of P. alpina negatively.     

内生真菌对草坪植物病原真菌抑制作用的比较

分别从野生牧草羽茅(Achnatherum sibiricum (L.) Keng)、栽培种高羊茅(Festuca arundinacea Schreb.)(品种Millennium)、栽培种黑麦草(Lolium perenne L.) (品种Justus)中分离出内生真菌Neotyphodium sp.、N. coenophialum和N. lolli,通过体外培养法比较了这3种内生真菌对草坪植物病原真菌的抑制作用.结果表明,从羽茅中分离的内生真菌Neotyphodium sp.在两菌相交前对所有供试的病原真菌都有一定的抑制作用,其中对枝孢霉属(Cladosporium sp.)、弯孢霉属(Curvularia sp.)和拟茎点属(Phomopsis sp.)病原真菌的抑制效果尤为显著,对峙培养3d后的抑菌率分别达70.1%、52.3%和30.9%,营养竞争作用、重寄生作用是其主要的拮抗机制;从高羊茅中分离的内生真菌N. coenophialum对枝孢霉属病原真菌存在一定的抑制作用;而从黑麦草中分离的内生真菌N. lolli与病原真菌对峙培养时,病原真菌菌落慢慢侵占整个营养空间,内生真菌停止生长并逐渐褐变死亡.体外培养结果说明Neotyphodium sp.对供试病原真菌的拮抗效果优于N. coenophialum和N. lolli,由此推测Neotyphodium sp.与宿主植物羽茅的共生可能有利于宿主植物抵抗病原真菌的侵扰.     

Note on high-nitrate para grass (Brachiaria mutica) silage given to rabbits

Silage prepared from high-nitrate (2.76% NO^?₃) wilted para grass (Brachiaria mutica) had high ammonia content (25.3% of total N as NH₃-N) and pH (7.6), and low Flieg index (7) and nitrate content (0.093% NO^?₃). Aeration for 2 h after unloading improved the Flieg index (50), and reduced ammonia and nitrate contents. However, on longer aeration nitrate content increased.One month of feeding rabbits on fresh para grass of low nitrate (Diet A) and high nitrate (Diet B) content, and silage prepared from high-nitrate para grass (Diet C) caused toxic symptoms in Groups B and C (rabbits on Diets B and C, respectively), resembling those of nitrate and ammonia toxicity, respectively. Erythrocyte count, haemoglobin and packed cell volume were high and leucocyte count low in Group B, while a reverse trend was observed in Group C. Nitrate was detected in different body fluids of rabbits of Group B by qualitative test. Blood and urinary ammonia nitrogen were high in rabbits of Group C.     

Co-existing grass species have distinctive arbuscular mycorrhizal communities

Arbuscular mycorrhizal (AM) fungi are biotrophic symbionts colonizing the majority of land plants, and are of major importance in plant nutrient supply. Their diversity is suggested to be an important determinant of plant community structure, but the influence of host-plant and environmental factors on AM fungal community in plant roots is poorly documented. Using the terminal restriction fragment length polymorphism (T-RFLP) strategy, the diversity of AM fungi was assessed in 89 roots of three grass species (Agrostis capillaris, Festuca rubra, Poa pratensis) that co-occurred in the same plots of a field experiment. The impact of different soil amendments (nitrogen, lime, nitrogen and lime) and insecticide application on AM fungal community was also studied. The level of diversity found in AM fungal communities using the T-RFLP strategy was consistent with previous studies based on clone libraries. Our results clearly confirm that an AM fungal host-plant preference exists, even between different grass species. AM communities colonizing A. capillaris were statistically different from the others (P < 0.05). Although grass species evenness changed in amended soils, AM fungal community composition in roots of a given grass species remained stable. Conversely, in plots where insecticide was applied, we found higher AM fungal diversity and, in F. rubra roots, a statistically different AM fungal community.     

Response of conifer seedlings to nitrate and ammonium sources of nitrogen

Summary Differences in growth responses of Douglas fir, western hemlock, Sitka spruce, and white spruce to nitrate and ammonium N sources were examined in sand culture and artificial soil culture. Effects of the two forms of N on growth, needle area, and N uptake of three Douglas fir halb-sib progenies were examined in a second sand culture. Response of Douglas fir to the two forms of N was followed over two years in nursery soil of different pH levels. In sand culture 1 mean seedling dry weight of all species, except hemlock, was greatest when ammonium N and nitrate N were provided in equal amounts. In all species, except Sitka spruce, ammonium alone resulted in greater growth than nitrate alone. Use of ammonium N resulted in greater growth of all species, than was obtained with nitrate N, at pH values in the region 5.4 and 7.5 in artificial soil culture. Only Douglas fir showed substantial differences due to N source below pH 5. Growth of all species was greater at pH 5.4 than at 7.5 in each N source treatment. Growth of Douglas fir seedlings was greatest with ammonium N and least with nitrate N in sand culture 2. Supply of nitrate and ammonium in equal proportions resulted in intermediate growth. Leaf area/plant weight ratio was unaffected by N source. Analysis of nutrient solutions showed appreciable nitrification of ammonium N during the 7 days between solution changes. In the three greenhouse experiments, with little exception, increase in proportion of ammonium in N supply resulted in increase of seedling tissue N concentration. This effect was more pronounced in roots than shoots. Total N uptake by ammonium fed seedlings was about double the N uptake of nitrate fed seedlings in sand culture 2. Nursery grown Douglas fir seedlings showed greater growth response to ammonium sulphate than to calcium nitrate, and this appeared due entirely to form of N supply in the first year. A similar response in the second year was partly due to greater soil acidification by ammonium sulphate. Compared with calcium nitrate, ammonium sulphate increased N concentration of one-year old shoots, but this difference was not detected by foliar analysis of two-year old seedlings.     

The use of mean pool abundances to interpret 15N tracer experiments

A pulse dilution ¹⁵N technique was used in the field to determine the effect of the ammonium to nitrate ratio in a fertilizer application on the uptake of ammonium and nitrate by ryegrass and on gross rates of mineralization and nitrification. Two experiments were performed, corresponding approximately to the first and second cuts of grass. Where no substantial recent immobilization of inorganic nitrogen had occurred, mineralization was insensitive to the form of nitrogen applied, ranging from 2.1–2.6 kg N ha^-1 d^-1. The immobilization of ammonium increased as the proportion of ammonium in the application increased. In the second experiment there was evidence that high rates of immobilization in the first experiment were associated with high rates of mineralization in the second. The implication was that some nitrogen immobilized in the first experiment was re-mineralized during the second. Whether this was nitrogen taken up, stored in roots and released following defoliation was not clear. Nitrification rates in this soil were low (0.1–0.63 kg N ha^-1 d^-1), and as a result, varying the ratio of ammonium to nitrate applied markedly altered the relative uptake of ammonium and nitrate. In the first experiment, where temperatures were low, preferential uptake of ammonium occurred, but where >90% of the uptake was as ammonium, a reduction in yield and nitrogen uptake was observed. In the second experiment, where temperatures and growth rates were higher, the proportion of ammonium to nitrate taken up had no effect on yield or nitrogen uptake.