Competition for nitrogen sources between European beech (Fagus sylvatica) and sycamore maple (Acer pseudoplatanus) seedlings

To investigate the short‐term consequences of direct competition between beech and sycamore maple on root N uptake and N composition, mycorrhizal seedlings of both tree species were incubated for 4 days (i.e. beech only, sycamore maple only or both together) in an artificial nutrient solution with low N availability. On the fourth day, N uptake experiments were conducted to study the effects of competition on inorganic and organic N uptake. For this purpose, multiple N sources were applied with a single label. Furthermore, fine roots were sampled and analysed for total amino acids, soluble protein, total nitrogen, nitrate and ammonium content. Our results clearly show that both tree species were able to use inorganic and organic N sources. Uptake of inorganic and organic N by beech roots was negatively affected in the presence of the competing tree species. In contrast, the presence of beech stimulated inorganic N uptake by sycamore maple roots. Both the negative effect of sycamore maple on N uptake of beech and the positive effect of beech on N uptake of sycamore maple led to an increase in root soluble protein in beech, despite an overall decrease in total N concentration. Thus, beech compensated for the negative effects of the tree competitor on N uptake by incorporating less N into structural N components, but otherwise exhibited the same strategy as the competitor, namely, enhancing soluble protein levels in roots when grown under competition. It is speculated that enhanced enzyme activities of so far unknown nature are required in beech as a defence response to inter‐specific competition.     

Foliar free polyamine and inorganic ion content in relation to soil and soil solution chemistry in two fertilized forest stands at the Harvard Forest,Massachusetts

Polyamines (putrescine, spermidine, and spermine) are low molecular weight, open-chained, organic polycations which are found in all organisms and have been linked with stress responses in plants. The objectives of our study were to investigate the effects of chronic N additions to pine and hardwood stands at Harvard Forest, Petersham, MA on foliar polyamine and inorganic ion contents as well as soil and soil solution chemistry. Four treatment plots were established within each stand in 1988: control, low N (50 kg N ha^-1 yr^-1 as NH₄NO₃), low N + sulfur (74 kg S ha^-1 yr^-1 as Na₂SO₄), and high N (150 kg N ha^-1 yr^-1 as NH₄NO₃). All samples were analyzed for inorganic elements; foliage samples were also analyzed for polyamines and total N. In the pine stand putrescine and total N levels in the foliage were significantly higher for all N treatments as compared to the control plot. Total N content was positively correlated with polyamines in the needles (P 0.05). Both putrescine and N contents were also negatively correlated with most exchangeable cations and total elements in organic soil horizons and positively correlated with Ca and Mg in the soil solution (P 0.05). In the hardwood stand, putrescine and total N levels in the foliage were significantly higher for the high N treatment only as compared to the control plot. Here also, total foliar N content was positively correlated with polyamines (P 0.05). Unlike the case with the pine stand, in the hardwood stand foliar polyamines and N were significantly and negatively correlated with foliar total Ca, Mg, and Mn (P 0.05). Additional significant (P 0.05) relationships in hardwoods included: negative correlations between foliar polyamines and N content to exchangeable K and P and total P in the organic soil horizon; and positive correlations between foliar polyamines and N content to Mg in soil solution. With few exceptions, low N + S treatment had effects similar to the ones observed with low N alone for both stands. The changes observed in the pine stand for polyamine metabolism, N uptake, and element leaching from the soil into the soil solution in all treatment plots provide additional evidence that the pine stand is more nitrogen saturated than the hardwood stand. These results also indicate that the long-term addition of N to these stands has species specific and/or site specific effects that may in part be explained by the different land use histories of the two stands.     

Nitrogen uptake and the importance of internal nitrogen loading in Lake Balaton

1. The importance of various forms of nitrogen to the nitrogen supply of phytoplankton has been investigated in the mesotrophic eastern and eutrophic western basin of Lake Balaton.
2. Uptake rates of ammonium, urea, nitrate and carbon were measured simultaneously. The uptake rates were determined using N and C methodologies, and N2‐fixation was measured using the acetylene‐reduction method. The light dependence of uptake was described with an exponential saturation equation and used to calculate surface‐related (areal) daily uptake.
3. The contribution of ammonium, urea and nitrate to the daily nitrogen supply of phytoplankton varied between 11 and 80%, 17 and 73% and 1 and 15%, respectively. N2‐fixation was negligible in the eastern basin and varied between 5 and 30% in the western region of the lake. The annual external nitrogen load was only 10% of that utilized by algae.
4. The predominant process supplying nitrogen to the phytoplankton in the lake is the rapid recycling of ammonium and urea in the water column. The importance of the internal nutrient loading is emphasized.     

In situ ion exchange resin bags to estimate forest site quality

Cation and anion exchange resin bags were placed just under the humus layer at five adjacent forest sample sites with differing site quality classes in order to assess the available nutrient supply. For comparison, humus samples were collected from the same sites. Nutrients were extracted from humus samples by conventional extraction methods and by shaking together with ion exchange resin bags. Ca and Mg corresponded best to differences in site quality class, of all analysed ions in thein situ resin bag eluates. Thein situ resin bag adsorption of NH₄−N, Na and Mn also showed a positive correlation with site quality. The adsorption of PO₄−P was negatively correlated to site quality class. Inadequate amounts of exchange resin, or leaving resin bagsin situ for too long a time result in the replacement of already adsorbed ions by ions with higher ion exchange constants.     

硅钙钾镁肥对南方稻田土壤酸性和盐基离子动态变化的影响

南方稻田土壤大面积酸化是水稻生产的主要限制因子.尽管石灰作为酸化土壤调理剂已广泛应用,但大量或长期施用石灰不仅会引起土壤板结,而且会导致土壤钙、钾、镁等元素的平衡失调.硅钙钾镁肥由于溶解度更低、养分全面是良好的替代材料.为了明确硅钙钾镁肥阻控土壤酸化的效果和作用,本研究采用连续4年的硅钙钾镁肥田间定位试验,以农民习惯施肥为对照,分析在农民习惯施肥基础上增施750、1125、1500和1875 kg·hm^-2硅钙钾镁肥下稻田土壤pH、交换性酸、交换性盐基离子和有效硅的动态变化.结果表明: 农民习惯施肥导致土壤pH、土壤交换性盐基和盐基饱和度逐年下降,土壤交换性酸逐年增加.与之相反,硅钙钾镁肥处理显著提高了土壤pH值,提高幅度随硅钙钾镁肥施用次数或用量的增加而增大.连续多次施用硅钙钾镁肥有效促进了盐基离子在土壤中的累积和土壤交换酸的消耗,特别是土壤交换性Ca²⁺、Mg²⁺的累积和土壤交换性Al³⁺的消耗,硅钙钾镁肥用量越大,积累或消耗的量越多,但速率相对越慢.土壤交换性酸消耗量中,硅钙钾镁肥释放的交换性盐基离子和相应碱贡献了108.8%,是交换性酸减少的主要途径.硅钙钾镁肥在改良稻田土壤酸性的同时,土壤有效硅含量逐年增加,增幅随硅钙钾镁肥施用量的增加而显著增大.总之,农民习惯施肥导致土壤持续酸化,酸化率为2.86 kmol H⁺·hm^-2·a^-1,硅钙钾镁肥能有效阻控酸化过程,产生了大量碱(9.69~18.44 kmol OH^-·hm^-2·a^-1),释放的Ca²⁺、Mg²⁺盐基离子和相应碱是土壤酸化阻控的主要作用因子.     

Temporal variation in nutrient uptake capacity by intact roots of mature loblolly pine

Nutrient uptake is generally thought to exhibit a simple seasonal pattern, but few studies have measured temporal variation of nutrient uptake capacity in mature trees. We measured net uptake capacity of K, NH⁺₄, NO₃^–, Mg and Ca across a range of solution concentrations by roots of mature loblolly pine at Calhoun Experimental Forest in October 2001, July 2001, and April 2002. Uptake capacity was generally lowest in July; rates in October were similar to those in April. Across a range of concentrations, antecedent nutrient solution concentrations affected the temporal patterns in uptake in July but not in October or April. In July, uptake of NH⁺₄, Mg and Ca was positively correlated with concentration when roots were exposed to successively lower concentrations, but negatively correlated with concentration when exposed to successively higher concentrations. In contrast, uptake in October was constant across the range of concentrations, while uptake increased with concentration in April. As in studies of other species, we found greater uptake of NH⁺₄ than NO₃^–. Temporal patterns of uptake capacity are difficult to predict, and our results indicate that experimental conditions, such as experiment duration, antecedent root conditions and nutrient solution concentration, affect measured rates of nutrient uptake.     

Differences in Earthworm Densities and Nitrogen Dynamics in Soils Under Exotic and Native Plant Species

Previous studies of the invasion of two exotic plants – Berberis thunbergii and Microstegium vimineum – in hardwood forests of New Jersey have shown a significant increase of pH in soils under the invasive plants as compared with soils from under native shrubs (Vaccinium spp). We present a further investigation of soil properties under the exotic plants in question. We measured the densities of earthworms in the soil under the two exotics and the native shrubs in three parks in New Jersey. In the same populations we also measured the extractable ammonium and nitrate in the top 5 cm of the soil, as well as the respiration of the soils and the potential rates of mineralization (aerobic lab incubation). In addition, we measured the nitrate reductase activity in leaves of the two exotic plants and several native shrubs and trees. Although there were differences between parks, we observed significantly higher earthworm densities in the soil under the exotic species. The worms were all European species. Soil pH, available nitrate and net potential nitrification were significantly higher in soils under the two exotic species. In contrast, total soil C and N and net ammonification were significantly higher under native vegetation. Nitrate reductase activities were much higher in the leaves of exotic plants than in the leaves of native shrubs and trees. Changes in soil properties, especially the change in nitrogen cycling, associated with the invasion of these two plant species may permit the invasion of other weedy or exotic species. Our results also suggest that even if the two exotic species were removed, the restoration of the native flora might be inhibited by the high nitrate concentrations in the soil.     

Alpine plants show species-level differences in the uptake of organic and inorganic nitrogen

As an estimate of species-level differences in the capacity to take up different forms of N, we measured plant uptake of ¹⁵N-NH₄ ⁺, ¹⁵N-NO₃ ^– and ¹⁵N, [1]-¹³C glycine within a set of herbaceous species collected from three alpine community types. Plants grown from cuttings in the greenhouse showed similar growth responses to the three forms of N but varied in the capacity to take up NH₄ ⁺, NO₃ ^– and glycine. Glycine uptake ranged from approximately 42% to greater than 100% of NH₄ ⁺ uptake; however, four out of nine species showed significantly greater uptake of either NH₄ ⁺ or NO₃ ^– than of glycine. Relative concentrations of exchangeable N at the sites of plant collection did not correspond with patterns of N uptake among species; instead, species from the same community varied widely in the capacity to take up NH₄ ⁺, NO₃ ^–, and glycine, suggesting the potential for differentiation among species in resource (N) use.     

Content of Oxalate in Actinidia Deliciosa Plants Grown in Nutrient Solutions with Different Nitrogen Forms

Kiwifruit plants (Actinidia deliciosa cv. Hayward) were grown in Hoagland nutrient solution with calcium nitrate, potassium nitrate, ammonium nitrate or ammonium chloride as the nitrogen source. Plants grown in the solution with nitrate nitrogen displayed a higher oxalate content, greater shoot length and leaf area, and higher content of ascorbic acid and NO₃ ^– ions in the leaves. Plants grown in the solution with ammonium nitrate, and particularly with ammonium chloride, showed low oxalate content, low content of ascorbic acid and NO₃ ^–, high content of Cl^– and Na⁺, low shoot length and leaf area. Oxalate formation appeared to be connected with the assimulation of nitrate, more precisely with nitrate reduction, while ammonium nitrogen assimilation did not induce the synthesis of oxalic acid.     

Varietal differences in uptake and utilization of nitrogen and other macro-elements in seedlings of barley, Hordeum vulgare

Growth and uptake of N, P, S, K, Ca and Mg in barley ( Hordeum vulgare L.) were studied in water culture using young plants of 17 cultivars. Large varietal differences were obtained in dry weight production and mineral accumulation. The differences were not the same for plants grown in high- and low-salt media. For plants grown under both conditions there was a good correlation between dry weight production and total N content. Total shoot contents of K and Ca were closely correlated with shoot dry weight. Utilization of P and S in high- and low-salt plants and Mg in low-salt plants was variable in relation to dry weight production in both types of nutrient conditions. The correlation between dry weight and total content of Mg in high-salt plants was good. These differences in mineral economy between young barley plants were partly caused by varietal differences in relative growth rate, and in high-salt seedlings also by differences in seed content of N. The significance of root size, and of uptake, root-shoot partitioning and use-efficiency of specific elements differed; all four factors were important for P and S, but had varying impact on K, Mg and Ca. For N, differences in root size and ion accumulation were the most important factors causing varietal variation in mineral nutrition. – In a special experiment seedlings of barley were transferred to N-free nutrient solution after six days of adequate N supply. There was no significant varietal differences in use-efficiency ratio of N. Root/shoot partitioning of N was unaffected.     

Measuring Nitrogen and Phosphorus Uptake by Intact Roots of Mature Acer saccharum Marsh., Pinus resinosa Ait., and Picea abies (L.) Karst

A common method for measuring uptake by intact roots in situ is the depletion method, wherein intact fine roots are separated from soil and placed in nutrient solution. The difference between initial and final amounts of nutrient in solution is attributed to root uptake. Variations on this method include applying pretreatment solutions, training roots to grow into bags or trays, and varying concentrations of nutrient solution. We tested whether variations in methods affected measured net uptake rates of NH ₄ ⁺ , NO ₃ ⁻ , and PO ₄ ³⁻ . Intact roots of 60 year-old sugar maple (Acer saccharum Marsh.), red pine (Pinus resinosa Ait.), and Norway spruce (Picea abies (L.) Karst.) were given one of four treatments prior to measuring net uptake. “Trained” roots were grown in a sand-soil mixture. “Recovered” roots were excavated and allowed to recover in nutrient solution for two or four days (“two-day recovery” and “four-day recovery”, respectively). “No recovery” roots were excavated and used immediately in experiments. We exposed roots to three concentrations of nutrient solutions to observe the effects of initial nutrient solution concentration. Initial nutrient solution concentration was an important source of variation in measured uptake rates, and N uptake was stimulated by low antecedent concentrations. We found no significant differences in net uptake rates between pretreatments for any of the species studied, indicating that our pretreatments were not effective in improving measurement of uptake. Such pretreatments may not be necessary for measuring net uptake and may not hinder the comparison of rates measured using variations of the depletion method.     

Differences in the success of sugar maple and red maple seedlings on acid soils are influenced by nutrient dynamics and light environment

Variation in tolerance to nutrient limitations may contribute to the differential success of sugar maple ( Acer saccharum Marsh.) and red maple ( Acer rubrum L.) on acid soils. The objectives of this study were to examine these relationships as influenced by light environment and test whether sensitivity to nutrient stress is mediated by oxidative stress. First-year sugar maple and red seedlings were grown on forest soil cores contrasting in nutrient availability under high or low light intensity. Foliar nutrition, photosynthesis, growth and antioxidant enzyme activity were assessed. Photosynthesis and growth of sugar maple were significantly lower on nutrient-poor soils and were correlated with leaf nutrient status with Ca and P having the strongest influence. For red maple, only chlorophyll content showed sensitivity to the nutrient-poor soils. High light exacerbated the negative effects of nutrient imbalances on photosynthesis and growth in sugar maple. Antioxidant enzyme activity in sugar maple was highest in seedlings growing on nutrient-poor soils and was inversely correlated with photosynthesis, Ca, P, and Mg concentrations. These results suggest that: (1) sugar maple is more sensitive to nutrient stresses associated with low pH soils than red maple; (2) high light increases sugar maple sensitivity to nutrient stress; (3) the negative effects of nutrient imbalances on sugar maple may be mediated by oxidative stress.     

Membrane phospholipid composition during maturation of seeds of Acer platanoides and Acer pseudoplatanus in relation to desiccation tolerance

Membrane phospholipid composition was investigated in seeds of two species from the genus Acer: Norway maple (Acer platanoides L.) — tolerant to desiccation, and sycamore (Acer pseudoplatanus L.) — intolerant to desiccation, during their maturation, from 1 August to 25 September 1995, at weekly intervals. Seeds of Norway maple acquire tolerance to desiccation at the end of August ie. about 125 days after flowering (DAF). Phospholipid composition during development revealed marked differences between studied seeds. Seeds of Norway maple after acquiring tolerance to desiccation contained much more phosphatidylcholine (PC) and phosphatidylethanolamine (PE), compared to sycamore. The ratio of PC/PE in mature Norway maple seeds was evidently higher than those in sycamore. The level of unsaturated fatty acids in the phospholipid fraction substantially increased in Norway maple seeds during development and the saturation of PC and PE was less than in sycamore. The results suggest that phospholipid composition may be involved in desiccation tolerance of Norway maple seeds.     

Changes in nutrient distribution in forests and soils of Walker Branch watershed,Tennessee, over an eleven-year period

Changes in vegetation, litter, and soil nutrient content were measured in selected plots on Walker Branch watershed, Tennessee, from 1972–73 to 1982. The watershed has been allowed to revert to forest since 1942, before which it consisted of small subsistence farms and woodland pastures. Changes in Ca status were of particular interest because initial nutrient cycling characterizations indicated that net Ca accumulation in vegetation could have caused large decreases in soil exchangeable Ca²⁺ within 20 years.Decreases in forest floor and subsoil (45–60 cm) N, exchangeable Ca²⁺, and Mg²⁺ content were noted in several plots from 1972 to 1982. Surface soils (0–15 cm) showed either no change or, in some cases (e.g., N and exchangeable K⁺ in certain plots), increases over the 11-year period. Reductions in forest floor and subsoil exchangeable Ca²⁺ and exchangeable Mg²⁺ on cherty, upper slope oak-hickory and chestnut oak forests were most striking. The changes in Ca²⁺ are thought to be due primarily to high rates of Ca²⁺ incorporation into woody tissues of oak and hickory species. Reductions in forest floor and subsoil exchangeable Mg²⁺ could not be accounted for by woody increment; leaching may have played a major role in causing these decreases. Changes in P and exchangeable K⁺ were variable, with both increases and decreases.There were significant increases in exchangeable Al³⁺ in both subsoils and surface soils of certain plots, but these were not accompanied by decreases in exchangeable base cations or consistent decreases in pH. Dissolution of interlayer Al from 2:1 clays may be the cause of the exchangeable Al³⁺ increases.These results suggest a general decline in fertility, especially with regard to Ca and Mg in those forests with low soil Ca and Mg supplies. Monitoring of further changes (if any) in these ecosystems will continue as the currently aggrading forests approach steady state.     

稳定性铵态氮肥在黑土和褐土中的氮素转化特征

以稳定性氯化铵为氮源,采用室内培养的方法,研究0.20、0.50、1.00 g N·kg^-1干土3种浓度的稳定性铵态氮肥在黑土、褐土中的氮素转化特征.结果表明: 在褐土中,随着氯化铵添加量的增加,土壤中发生硝化作用的时间逐渐推迟,添加0.20、0.50 g N·kg^-1干土处理开始发生明显硝化反应的时间分别为第3、7天,在高浓度氮量(1.00 g N·kg^-1干土)添加下硝化作用受到明显抑制;在黑土中,各浓度氮量添加处理开始发生硝化反应的时间相同,均为第3天,且随着添加量的增加,硝化作用潜势逐渐减弱.只加铵态氮肥的处理中,添加0.20 g N·kg^-1干土的氯化铵氮肥在褐土和黑土中的硝化反应时间分别可维持3周和2周左右;添加0.50 g N·kg^-1干土的氯化铵氮肥在褐土和黑土中的硝化反应时间分别可维持4周和3周左右.与单施氯化铵相比,黑土和褐土在0.20、0.50 g N·kg^-1干土添加浓度下,按纯氮量的1.0%添加3,4-二甲基吡唑磷酸盐(DMPP)、4.0%添加二氰二胺(DCD)均能显著抑制硝化作用,降低硝态氮的含量,抑制硝化作用潜势.综上,在褐土中,随着氯化铵添加浓度增加,土壤硝化作用受到抑制效果大于黑土.在0.20、0.50 g N·kg^-1干土外源铵态氮时,添加抑制剂可以显著抑制铵态氮的硝化作用.因此室内硝化抑制剂培养试验时,建议铵态氮添加量不超过1.00 g N·kg^-1干土,以0.50 g N·kg^-1干土效果最好.     

Culture of mycorrhizal tree seedlings under controlled conditions: Effects of nitrogen and aluminium

A sand culture system was developed for growth of mycorrhizal seedlings under monoxenic conditions, with frequently renewed nutrient solution The composition of the nutrient solution resembled that of a forest soil solution, based on long-term measurements from forest sites at Soiling, northern Germany. Seedlings of Picea abies (L.) Karst. inoculated with Lactarius rufus (Scop.) Fr. were grown in this culture system. Plants developed rapidly, having almost totally mycorrhizal root systems. Nitrate at 2.7 mM in the nutrient solution and applied over a 13 week period had no negative effect on mycorrhizal development. Ammonium at 2.7 mM reduced the degree of mycorrhizal infection slightly, in such a way that the degree of mycorrhizal infection was reduced to a much less extent than the total number of root lips. Hence, the impact of NH⁺₄ may be primarily on root development and not on mycorrhizal fungal colonization. When the concentrations of NO-₃ and NH⁺₄ used in the present study are compared to those found in forest soil solutions, NO-₃ and NH₄⁺ would not appear to influence mycorrhizal development negatively under natural conditions. Aluminium at 0.8 mM and applied over a 13 week period reduced Mg uptake into roots and needles by 52 and 64%, respectively, resulting in needle chlorosis and strongly reduced photosynthetic activity. From a comparison of this study with others, no major difference in physiological response to aluminium exposure between non-mycorrhizal seedlings and seedlings colonized with Lactarius rufus was found.     

Nitrogen limitation in mycorrhizal Norway spruce (Picea abies) seedlings induced mycelial foraging for ammonium: implications for Ca and Mg uptake

Although many studies support the importance of the external mycelium for nutrient acquisition of ectomycorrhizal plants, direct evidence for a significant contribution to host nitrogen nutrition is still scarce. We grew nonmycorrhizal seedlings and seedlings mycorrhizal with Paxillus involutus (Batsch) Fr. in a sand culture system with two compartments separated by a 45-m Nylon mesh. Hyphae, but not roots, can penetrate this net. Nutrient solutions were designed to limit seedling growth by nitrogen. Hyphal density in the hyphal compartment, host N status and shoot growth of mycorrhizal seedlings significantly increased in response to NH₄ ⁺ addition to the hyphal compartment. Labeling the compartment only accessible to hyphae with ¹⁵NH₄ ⁺ showed that the increase in N uptake in the mycorrhizal seedlings was a result of hyphal N acquisition from the hyphal compartment. These results indicate that hyphae of P. involutus may actively forage into N-rich patches and improve host N status and growth. In the mycorrhizal seedlings, which received additional NH₄ ⁺ via their external mycelium, the increase in NH₄ ⁺ supply less negatively affected Ca and Mg uptake than in nonmycorrhizal seedlings, where the additional NH₄ ⁺ was directly supplied to the roots. This was most likely due to the close link of NH₄ ⁺ uptake and H⁺ extrusion, which, in the nonmycorrhizal seedlings, lead to a strong acidification in the root compartment, and subsequently reduced Ca and Mg uptake, whereas in the mycorrhizal seedlings the site of intensive NH₄ ⁺ uptake and acidification was in the hyphal and not in the root compartment. Our data support the idea that the ectomycorrhizal mycelium connected to an N-deficient host may actively forage for N. The mycelium may also be important as a biological buffer system ameliorating negative influence of high NH₄ ⁺ supply on cation uptake.