Seasonal changes in foliar macronutrients (N,P, K,Ca and Mg) inEucalyptus saligna Sm. andE. wandoo Blakely growing in rehabilitated bauxite mine soils of the Darling Range,Western Australia

Summary Seasonal changes in the foliar concentration of macronutrients (N, P, K, Ca and Mg) in sapling trees ofEucalyptus saligna Sm. andE. wandoo Blakely growing in rehabilitated bauxite mined areas in the Darling Range of Western Australia are described. Foliar N concentration decreased with age of the fully expanded leaf tissue. Leaf N concentrations were also high when rates of litter decomposition were expected to be high during the period of early spring. The greatest foliar N difference between trees growing in good soil conditions and those from poorer soil conditions also occurred during this period. Levels of P in leaves were highest in young developing leaves but once the leaves reached full size, no seasonal trend in P concentration was observed. Foliar K was lower during the winter and probably related to the period of maximum leaching by precipitation. High foliar K during summer, however, could be related to the role of K in lowering cellular water potential. Leaf Ca was highest during early sping. Low mobility of cellular Ca during the cool portion of the year was indicated. Foliar Mg showed a weak pattern of decreasing concentration with leaf age. The best season for sampling for these broadleafed evergreen species to provide information on plant nutrient status appears to be in spring.     

Nutritional imbalance caused by nitrogen excess is correlated with the occurrence of watermark disease in white willow

Willows in the Flemish region are threatened by the watermark disease, a wood disease caused by the bacterium Brenneria salicis (Hauben et al. Syst Appl Microbiol 21:384–397, 1998). Affected trees are observed to grow at specific sites which seem more sensitive to periodical watermark symptom development than other sites. In this study we aim at revealing abiotic site factors that could play a role in triggering the watermark disease. During three consecutive years we analyzed stands of young isogenic willows (Salix alba L. cv. Lichtenvoorde), which were planted on four different sites in an agricultural environment. These test sites were chosen, based on the occurrence/absence of watermark disease in the old willows that grow on these sites. Two sites had old willow pollards with clear watermark symptoms, and on the two other sites the old willows were unaffected. Differences in abiotic characteristics between the sites were analysed and their effects on the nutrient status in willow were measured in the isogenic willows that were newly planted. Brenneria salicis-specific multiplex PCR revealed the presence of this pathogenic bacterium in both the old and the freshly planted young willows. At each site, the physical and chemical characteristics of soil and groundwater were investigated; the young tree growth was monitored and nutritional status of the willow plantings was assessed by foliar analysis. Physical soil properties including soil type and profile water availability, and chemical soil analysis were unable to explain differences between healthy and affected sites. Diseased sites, however, had significantly higher N concentrations and traces of herbicides in the groundwater. In young trees, foliar levels of N and S were always greater at the affected sites, while P contents were lower. Dry mass based N concentration in the leaves of the newly planted trees increased systematically over the years, but more at affected than at healthy sites. Also, chlorophyll content was significantly greater at the affected sites. Growth rate of the young isogenic trees was twice as fast at the diseased sites. The presented data suggest that application of fertilisers in agriculture causes an imbalance in the nutritional status of Salix alba L. Excess nitrogen makes this species fast-growing, reflected by an average foliar N:P ratio >10, and more susceptible for watermark disease, indicated by an average foliar N:Ca ratio exceeding 1.5.     

兰州市南山和北山3种乔木叶片生态化学计量特征的对比研究

叶片生态化学计量是了解生物地球化学循环的窗口,也是认识植物的生长状况及生态适应性的关键,研究人工林叶片生态化学计量特征对进一步优化生态工程建设具有重要意义。为揭示兰州市南北两山主要绿化树种侧柏、圆柏和云杉环境适应性的异同,对其叶片生态化学计量特征进行了对比研究。结果显示:云杉、圆柏叶片C含量(496.02、497.64 g/kg)显著高于侧柏叶片(484.12 g/kg),侧柏叶片N、P含量(17.70、1.66 g/kg)均显著高于圆柏(14.04、1.42 g/kg)和云杉(15.62、1.40 g/kg)。C、N、P的变异系数均为圆柏<侧柏<云杉;云杉、圆柏叶片C∶N (33.32、35.42)、C∶P (389.57、360.52)显著高于侧柏叶片(28.17、306.07),云杉叶片N∶P (11.88)显著高于圆柏叶片(10.20)和侧柏叶片(10.98)。C∶N、C∶P、N∶P的变异系数均为圆柏<侧柏<云杉;侧柏和云杉叶片N含量与P含量正相关、与C∶N、C∶P负相关,而3种乔木叶片P含量与C∶P、N∶P负相关(且C∶P与N∶P正相关)。圆柏叶片C含量与N含量正相关,侧柏叶片C含量与N含量负相关但与叶片C∶N、C∶P正相关,侧柏和云杉叶片C∶N与C∶P正相关。冗余分析表明,SOC∶STN是影响3种乔木叶片化学计量的主导因子,圆柏叶片C、侧柏叶片P与SOC正相关,3种乔木叶片N都与SOC、SOC∶STN和SOC∶STP正相关;云杉叶片N,侧柏、云杉叶片P和STP负相关,而STP与侧柏叶片C∶P、N∶P及云杉叶片C∶N正相关;3种乔木叶片C∶N与SOC∶STN、SOC∶STP负相关。研究结果表明:面对养分贫瘠的土壤环境,3种乔木叶片生态化学计量的适应策略有所不同,其中云杉和圆柏采取防御策略,而侧柏采取竞争倾向策略。虽然云杉是兰州市较适宜的造林树种,但其生长与其他两种乔木一样,都受N元素的限制。     

Phosphorus limitation of forest leaf area and net primary production on a highly weathered soil

We tested the hypothesis that P was the nutrient limiting net primary production of a nativeMetrosideros polymorpha forest on a highly weathered montane tropical soil in Hawaii. A factorial experiment used all combinations of three fertilizer treatments: nitrogen (N), phosphorus (P) and a mix of other essential nutrients (OE), consisting primarily of mineral derived cations and excluding N and P. P addition, but not N or OE, increased leaf area index within 12 months, foliar P concentration measured at 18 months, and stem diameter increment within 18 months. Stem growth at 18 months was even greater when trees fertilized with P also received the OE treatment. N and P additions increased leaf litterfall and N and P in combination further increased litterfall. The sequence of responses suggests that increased available P promoted an increase in photosynthetic area which led to increased wood production. P was the essential element most limiting to primary production on old volcanic soil in contrast to the N limitation found on young volcanic soils.     

Phenology and nutrition of miombo woodland trees in Zambia

Seasonality in nitrogen (N) and phosphorus (P) concentration in soil and shoots of five Brachystegia-Julbernardia (miombo) woodland trees was studied from September 1991 to March 1993 at two regrowth miombo sites in central Zambia. Shoot growth started in the dry season (September–November) and lasted until April during the 1991/92 season but had virtually ceased by January 1993 during the 1992/93 season. The shoot growing season was associated with low foliar N/P ratios. These ratios were much lower (<5) during the 1991/92 season than in the 1992/93 season (12–15). The increase in foliar N/P ratios after the shoot growing season was caused by a sharp drop in foliar P concentration, apparently due to reabsorption before leaf fall. There were no annual variations in biomass N concentration in contrast to P. During the 1992/93 growing season P concentrations in foliage and wood were a quarter and a third, respectively, of the 1991/ 92 levels. Since the short shoot growing season observed during the 1992/93 season is typical of savanna woodland trees in southern Africa, the high biomass P concentration and longer growing season in 1991/92 season were exceptional and may have been related to reduced competition by shallow rooting herbaceous plants caused by the severe drought of that season.     

Nutritional disturbances of young Scots pines caused by pine bark bugs in a dry heath forest

This study deals with the effect of pine bark bugs (Aradus cinnamomeus Panzer) on the nutrition of young Scots pines (Pinus sylvestris L.). Soil and needle samples for analytical purposes were collected from a young pine stand growing on a dry barren mineral soil afflicted by pine bark bugs.The damage to vascular tissues caused by pine bark bugs disturbed the nutrition of the trees, especially in the top part of the crown. The foliar calcium, magnesium, manganese and sulphur concentrations were highly reduced. Scots pines suffered from a lack of calcium, magnesium, nitrogen and phosphorus. These deficiencies were secondary by nature, because no differences were observed between the nutrient concentrations of the underlying soil of the healthy and affected trees. The symptoms of trees damaged by pine bark bugs resembled most of all calcium deficiency symptoms.     

云冷杉林林隙内倒木腐烂等级及其形成的微立地类型对土壤理化性质的影响

为揭示倒木腐烂等级及其形成的微立地类型对土壤理化性质的影响,分析了小兴安岭凉水国家级自然保护区谷地云冷杉林由倒木形成的林隙内不同腐烂等级倒木形成的3种微立地(丘顶、坑底和倒木下)的土壤理化性质(容重、毛管孔隙度、总孔隙度、毛管持水量、饱和持水量、土壤有机碳、全氮、全磷、有效磷、速效钾和pH值)和化学计量特征(C/N、N/P和C/P)的差异.结果表明: 倒木腐烂等级对土壤物理性质的影响不显著,而倒木形成的微立地类型对其影响显著.除土壤饱和持水量外,倒木下的其他土壤物理性质均最好,坑底最差;随着倒木腐烂等级的上升,除有效磷外的其他土壤养分含量呈显著上升趋势;3种微立地中,坑底土壤养分含量最低,pH最高;随倒木腐烂等级的上升,3种微立地浅层土壤C/N、N/P和C/P下降,而丘顶和坑底深层土壤C/N上升,N/P和C/P下降,C/P的变化趋势与土壤有效磷的一致.综上,随倒木腐烂等级的提高,土壤养分含量显著增加,且不同类型微立地之间差异明显,其中坑底始终最低.     

N and P in New Zealand Soil Chronosequences and Relationships with Foliar N and P

The growth of forest species in soil development chronosequences becomes increasingly phosphorus (P)-limited with time, as P is weathered, eroded and leached from soil. Foliar nitrogen (N) concentrations also tend to decrease with soil age when vegetation may be limited in both N and P. Here we report on soil development in temperate rain forests along three New Zealand chronosequences that have minimal pollution and disturbance from human activities, at Franz Josef, Waitutu and Central Volcanic Plateau, and on factors influencing soil net N mineralization (aerobic; 56 days) and foliar N and P concentrations. Except in very young soils (<500 years), at least 85% of total-P in mineral soil (0–10 cm) was transformed to organic-P. In each chronosequence, total-P declined with time, and foliar N:P ratios (mass) generally increased from 8 to 15–18, suggesting P was more limiting than N in the oldest soils of the chronosequence. There was a negative relationship between net N mineralization and C:N ratio for mineral soil. For the FH (organic) layer, net N mineralization had the strongest relationships with total-N concentration (positively) and C:organic-P ratio (negatively); however, relationships varied with forest group, suggesting that other factors were also important. Foliar P of kamahi (Weinmannia racemosa Linn. f.), a dominant canopy species, was related to soil organic-P, suggesting mineralization was an important process for tree nutrition.Foliar N was positively related to N concentration in the FH layer, but was not significantly related to any measured property in mineral soil, possibly because of the wide range of soils. The consistent declines in both soil and foliar P across the contrasting chronosequences strongly suggest that vegetation becomes progressively P-limited during long-term ecosystem development.     

The impact of available water content in previous agricultural soils on tree growth and nutritional status in young hybrid aspen plantations in Estonia

The impact of soil available water content (AWC) on the growth and foliar nutrient concentrations of trees in 7-yr-old hybrid aspen (Populus tremula L. x P. tremuloides Michx.) plantations established on abandoned agricultural lands was studied. AWC in the topmost 75 cm soil layer was significantly related to height growth and foliar N concentration of hybrid aspens. The correlations were stronger on well-drained automorphic soils and in the case of a dry growing season with insufficient precipitation. AWC over 150–160 mm can be considered sufficient, and below 120 mm insufficient for fast growth of hybrid aspen. The differences in AWC were less pronounced in the top 25 cm soil layer but were more noticeable in the 25–50 and 50–75 cm soil layers. AWC estimated as a function of soil specific surface area and bulk density was shown here to be a significant indicator for site selection for establishing hybrid aspen plantations. Foliar N concentration over 2.7% can be considered optimal and below 2.4%, insufficient for hybrid aspen on the studied soils. Foliar concentrations of P (on average 0.24%) and K (on average 0.79%) varied little and, thus, did not correlate with tree growth. The most suitable previous agricultural soils for afforestation with hybrid aspen would be moderately drained Albeluvisols, Luvisols and Planosols.     

秦岭地区华北落叶松人工林地土壤养分和酶活性变化

以秦岭地区不同林龄(5年生、10年生、20年生、30年生和40年生)华北落叶松人工林为研究对象,采用野外调查采样和室内分析相结合的方法,研究了不同林龄华北落叶松人工林地土壤剖面p H值、有机质养分和酶活性的变化。结果表明:土壤p H值随着林龄有降低趋势,随着土层深度的增加有升高趋势。土壤有机质和土壤速效氮、速效磷和速效钾在近熟龄期显著高于幼龄期。土壤中的磷酸酶、脲酶、蔗糖酶和过氧化氢酶活性随着林龄都呈"高—低—高"的趋势,幼龄期的土壤蔗糖酶活性显著高于近熟龄的,而磷酸酶、脲酶和过氧化氢酶活性是近熟龄期的显著高于中幼龄。土壤速效养分和酶活性都随着土层深度的增加有显著的降低。相关分析表明磷酸酶活性与有机质、速效氮、速效磷、速效钾和脲酶活性呈极显著的正相关性(P0.01),脲酶与有机质、速效氮和速效钾呈极显著正相关。蔗糖酶与过氧化氢酶活性显著负相关(P0.05),与p H值有一定的正相关性。秦岭地区华北落叶松人工林进入近熟林之后土壤肥力有一定的恢复,而在中幼龄阶段土壤养分比较缺乏,尤其是氮磷。     

Nutrient requirements of exotic tree species in Zimbabwe

Pot and field experiments were conducted in the greenhouse and at three field sites (Marondera, Domboshawa and Makoholi) in Zimbabwe to examine the effects of soils and fertilizers on nutrient uptake and growth of 6 exotic tree species (Eucalyptus camaldulensis, E. grandis, E. tereticornis, Leucaena leucocephala, Casuarina cunninghamiana, and Acacia holosericea). Plant growth, N and P contents of all species were increased by the application of N, P, K and micronutrient fertilizers. The effect of individual nutrients (N, P, K and micronutrients) and their combination on Eucalyptus species was further investigated in a pot experiment using soil from Domboshawa. Eucalyptus species responded only to N application and no significant interactions were found between N and the other elements. Nutrient uptake results showed that E. camaldulensis and E. tereticornis removed more cations than the N-fixing trees but only in the fertilized treatments. L. leucocephala and C. cunninghamiana were higher in P, but no clear trends were observed for N. Plant growth and nutrient uptake by E. camaldulensis. C. cunninghamiana and A. holosericea were assessed in the field at the three sites. Plant species grown in the Marondera site had greater height and diameter at breast height (DBH) than those in the two other sites. These results followed trends in soil nutrient contents. The analysis of foliage revealed differences in the nutrient concentration of leaves from different trees, but no effect of site was found. The interrelationships between plant characteristics, soil and foliage nutrients were examined. In a pot experiment, mineral N was the only variable correlated with growth response and nutrient uptake, while in the field the only significant correlation was obtained with soil pH.     

Foliar N application reduces soil NO<Stack><Subscript>3</Subscript><Superscript>−</Superscript></Stack>-N leaching loss in apple orchards

A comparison of the effects of foliar and soil N application was made in field-grown mature fruiting Gala/M9 apple trees (Malus domestica Borkh) in 2001 and 2002 growing seasons under Pacific Northwest growing conditions in southern British Columbia, Canada. The trees, six years old at the start of the experiment, were treated: (1) with 5 g/l urea sprays supplied every two weeks (7 times) from mid May to mid August (total about 50 g N/tree/year), (2) with the same amount of N applied to the soil with the same timing and quantity as for the foliar treatment, and (3) with no N (control). Leaf color (as SPAD readings) and N concentrations (mg/g), and soil NH₄⁺-N and NO₃^–-N were measured periodically throughout the two seasons. Leached NO₃^–-N was monitored monthly via an anion exchange probe from June to October in 2001 and from May to November in 2002. Shoot length was measured in October and N concentration of one-year-old wood and roots was determined in December of each growing s eason. Soil N application significantly increased shoot length relative to control or foliar N application. Leaf color, leaf N, and N concentration of one-year-old wood and roots were similarly increased relative to control by both soil and foliar N application. These treatments also increased fruit yield relative to control. There was no significant difference in yield and fruit quality between soil and foliar N applications. Soil N application increased soil NH₄⁺-N and NO₃^–-N content in the root zone, and also increased the NO₃^– leaching loss below the root zone especially late in the growing season. Our results suggested that tree N status and yield and fruit quality could be maintained by multiple urea sprays during the growing season in apple orchards, and foliar N application will reduce the risk of soil NO₃^–-N leaching.     

Regulation of nitrogen partitioning in field-grown almond trees: Effects of fruit load and foliar nitrogen applications

Two treatments were employed to influence the amount of amino nitrogen (N) transport in phloem. In walnut trees (Juglans regia L.), developing fruit significantly reduced the efflux of foliar-applied ¹⁵N-enriched urea from treated spurs over a 33-day period in comparison with similarly-treated defruited spurs. Those data suggest that local aboveground demand for N influences vascular transport of amino N. In another experiment, a 1% urea solution was applied foliarly to 5-year old `Mission' almond trees [Prunus dulcis (Mill.) D. A. Webb] to increase the concentration of amino N in the phloem. The effect of foliar N treatments on a) the transport and distribution of labelled urea N within the trees over the experimental period and b) the uptake of soil-applied labelled N were determined by replicated whole tree excavation, fractionation into various tree components and mass spectrometric analyses of the ¹⁴N/¹⁵N ratios. Concentrations and composition of amino acids in the phloem and xylem saps of control trees and trees receiving foliar-applied urea were also determined. In foliar urea-treated trees, the amino acid concentrations increased significantly in leaf and bark phloem exudate, within 24 and 96 h, respectively. Foliar-applied urea N was translocated to the roots of almond trees over the experimental period and decreased soil N uptake. The results of these experiments are consistent with the hypothesis that aboveground N demand affects the amount of amino N cycling between shoots and roots, and may be involved in the regulation of soil N uptake. This revised version was published online in June 2006 with corrections to the Cover Date.     

Foliar and twig macronutrients (N,P, K,Ca and Mg) in selected species ofEucalyptus used in rehabilitation: sources of variation

Summary The macronutrient variation within four 6 year oldEucalyptus saligna and four 5 year oldE. wandoo growing on rehabilitated bauxite pits was determined. Significant differences in mean nutrient concentrations were generally recorded between good soil condition sites and poor soil sites, between tree individuals, branch height, and plant organ type; but mean nutrient values were not different among canopy aspects. Fully expanded leaves of the current year provided the most uniform nutrient levels among the plant organs and showed major differences between sites with good soil nutrient conditions and those with poor conditions. Differences in foliar and branch levels of N, P, K, Ca and Mg, the variation between sites, canopy heights and plant organ types, and the use of foliar nutrient levels to indicate deficiencies are discussed.     

秦岭中幼林龄华北落叶松针叶与土壤的碳氮磷生态化学计量特征

为探究华北落叶松人工林物质循环规律与养分元素的分配格局,将华北落叶松针叶和土壤结合起来,以秦岭地区7年(7a)、12年(12a)、22年(22a)生华北落叶松人工林为研究对象,综合探究中幼林龄华北落叶松针叶和土壤的C、N、P含量及C∶N∶P化学计量特征。结果表明,不同林龄间土壤C和P含量的差异显著,7a和12a的C和P含量显著高于22a的,N含量差异不显著,随土层深度的增加,土壤C、N、P均呈下降趋势;22a土壤的C∶P和N∶P显著高于7a和12a,说明随着林龄增加,P限制了人工林正常的生长发育,C∶N在林龄之间的差异不显著;不同林龄针叶C差异不显著,N和P差异显著,N和P含量在中龄林中最高;22a针叶的C∶N和C∶P均表现为显著下降,3种林龄华北落叶松针叶的N∶P在6.8—9.3之间,说明幼、中林龄华北落叶松主要受N的限制;华北落叶松针叶和土壤的C、C∶N之间呈显著的正相关,P、C∶P之间呈显著的负相关,N、N∶P之间相关性不显著。中林龄华北落叶松人工林土壤P元素含量低,主要是通过养分的再吸收来满足植株的生长需求,而非通过吸收土壤养分。     

哈巴湖国家级自然保护区油蒿群落生态化学计量特征对群落生物量和物种多样性的影响

以宁夏哈巴湖国家级自然保护区内的油蒿群落为研究对象,测定了油蒿群落不同发育阶段植物叶C、N、P含量,分析了植物叶C∶N∶P比化学计量特征,探讨了油蒿群落生态化学计量特征对物种多样性和生物量的影响。结果表明,在哈巴湖国家级自然保护区,油蒿群落发育早期,植物生长受N限制,发育中后期,植物生长受P限制;油蒿群落发育早期阶段的植物具有较高的叶C∶N比和较低的叶N∶P比,而油蒿群落发育后期阶段的植物具有较低的叶C∶N比和较高的叶N∶P比。随着叶C∶N比的增大,油蒿群落生物量呈指数函数显著降低,物种多样性呈对数函数降低的趋势,但不显著;而随着叶N∶P比的增大,油蒿群落生物量和物种多样性均呈幂函数显著增加,表明生态化学计量特征对油蒿群落的物种多样性和生物量有重要影响。     

Inhibition of growth,and effects on nutrient uptake of arctic graminoids by leaf extracts — allelopathy or resource competition between plants and microbes?

Previous research has shown that plant extracts, e.g. from boreal dwarf shrubs and trees, can cause reduced growth of neighbouring plants: an effect known as allelopathy. To examine whether arctic and subarctic plants could also be affected by leaching of phytochemicals, we added extracts from the commonly occurring arctic dwarf shrubs Cassiope tetragona and Empetrum hermaphroditum, and from mountain birch, Betula pubescens ssp. tortuosa to three graminoid species, Carex bigelowii, Festuca vivipara and Luzula arcuata, grown in previously sterilized or non-sterilized arctic soils. The graminoids in non-sterilized soil grew more slowly than those in sterilized soil. Excised roots of the plants in non-sterilized soil had higher uptake rate of labelled P than those in sterilized soil, demonstrating larger nutrient deficiency. The difference in growth rate was probably caused by higher nutrient availability for plants in soils in which the microbial biomass was killed after soil sterilization. The dwarf shrub extracts contained low amounts of inorganic N and P and medium high amounts of carbohydrates. Betula extracts contained somewhat higher levels of N and much higher levels of P and carbohydrates. Addition of leaf extracts to the strongly nutrient limited graminoids in non-sterilized soil tended to reduce growth, whereas in the less nutrient limited sterilized soil it caused strong growth decline. Furthermore, the N and P uptake by excised roots of plants grown in both types of soil was high if extracts from the dwarf shrubs (with low P and N concentrations) had been added, whereas the P uptake declined but the N uptake increased after addition of the P-rich Betula extract. In contrast to the adverse extract effects on plants, soil microbial respiration and soil fungal biomass (ergosterol) was generally stimulated, most strongly after addition of the Betula extract. Although we cannot exclude the possibility that the reduced plant growth and the concomitant stimulation of microbial activity were caused by phytochemicals, we believe that this was more likely due to labile carbon in the extracts which stimulated microbial biomass and activity. As a result microbial uptake increased, thereby depleting the plant available pool of N and P, or, for the P-rich Betula extract, depleting soil inorganic N alone, to the extent of reducing plant growth. This chain of events is supported by the negative correlation between plant growth and sugar content in the three added extracts, and the positive correlation between microbial activity, fungal biomass production and sugar content, and are known reactions when labile carbon is added to nutrient deficient soils.     

Nitrogen availability patterns in white-sand vegetations of Central Brazilian Amazon

Addressing spatial variability in nitrogen (N) availability in the Central Brazilian Amazon, we hypothesized that N availability varies among white-sand vegetation types (campina and campinarana) and lowland tropical forests (dense terra-firme forests) in the Central Brazilian Amazon, under the same climate conditions. Accordingly, we measured soil and foliar N concentration and N isotope ratios (δ¹⁵N) throughout the campina-campinarana transect and compared to published dense terra-firme forest results. There were no differences between white-sand vegetation types in regard to soil N concentration, C:N ratio and δ¹⁵N across the transect. Both white-sand vegetation types showed very low foliar N concentrations and elevated foliar C:N ratios, and no significant difference between site types was observed. Foliar δ¹⁵N was depleted, varying from −9.6 to 1.6‰ in the white-sand vegetations. The legume Aldina heterophylla had the highest average δ¹⁵N values (−1.5‰) as well as the highest foliar N concentration (2.1%) while the non-legume species had more depleted δ¹⁵N values and the average foliar N concentrations varied from 0.9 to 1.5% among them. Despite the high variation in foliar δ¹⁵N among plants, a significant and gradual ¹⁵N-enrichment in foliar isotopic signatures throughout the campina–campinarana transect was observed. Individual plants growing in the campinarana were significantly enriched in ¹⁵N compared to those in campina. In the white-sand N-limited ecosystems, the differentiation of N use seems to be a major cause of variations observed in foliar δ¹⁵N values throughout the campina–campinarana transect.     

Tree mineral nutrition is deteriorating in Europe

The response of forest ecosystems to increased atmospheric CO₂ is constrained by nutrient availability. It is thus crucial to account for nutrient limitation when studying the forest response to climate change. The objectives of this study were to describe the nutritional status of the main European tree species, to identify growth‐limiting nutrients and to assess changes in tree nutrition during the past two decades. We analysed the foliar nutrition data collected during 1992–2009 on the intensive forest monitoring plots of the ICP Forests programme. Of the 22 significant temporal trends that were observed in foliar nutrient concentrations, 20 were decreasing and two were increasing. Some of these trends were alarming, among which the foliar P concentration in F. sylvatica, Q. Petraea and P. sylvestris that significantly deteriorated during 1992–2009. In Q. Petraea and P. sylvestris, the decrease in foliar P concentration was more pronounced on plots with low foliar P status, meaning that trees with latent P deficiency could become deficient in the near future. Increased tree productivity, possibly resulting from high N deposition and from the global increase in atmospheric CO₂, has led to higher nutrient demand by trees. As the soil nutrient supply was not always sufficient to meet the demands of faster growing trees, this could partly explain the deterioration of tree mineral nutrition. The results suggest that when evaluating forest carbon storage capacity and when planning to reduce CO₂ emissions by increasing use of wood biomass for bioenergy, it is crucial that nutrient limitations for forest growth are considered.     

Early growth and nutritional response to resource competition in cocoa-shade intercropped systems

Intercropping is often promoted for effective mutualism between species, thus compensating for external inputs. However, for optimal farm design resulting in superior production and nutrition, an accurate assessment of plant inter- and intra-specific competition is required. In predominant shade tree-cocoa (Theobroma cacao) systems, inconclusive evidence remains on species interactions, limitations to resource availability and subsequent growth and nutritional response, particularly in early growth. We examined cocoa biomass and foliar nutrition as well as nutrient supply through rates of decomposition and N mineralization after 1-year growth. Our approach employed fertilization and mixed planting treatments in an additive design of cocoa in monoculture (control), under artificial shade, and intercropped under two separate shade species (Terminalia superba and Newbouldia laevis). Intercropping had no effect on cocoa biomass production in comparison to monoculture cocoa. However, artificial shading stimulated foliage and root production both with and without fertilization, suggesting strong effects of light regulation on growth in the absence of belowground competition. Nutritionally, intercropping suppressed K uptake in cocoa foliage as K concentration was reduced by 20–25%, signifying dilution of this nutrient, presumably due to interspecific competition for mobile elements. Foliar N content under N. laevis was raised, where N concentration kept up with growth under this intercropped species. Intercropping also delayed decomposition rates, suggesting slower but sustained release of available nutrients into the topsoil. Cocoa under artificial shade, both with and without fertilization, exhibited the greatest nutrient responses as compared to unfertilized monoculture cocoa, where P uptake was stimulated most (175 and 112%), followed by K (69 and 71%), and then N (54 and 42%). Intercropping with shade trees failed to increase cocoa biomass, however, nutrient uptake was sustained for N and P, suggesting low interspecific competition. When fertilizers are undesirable or unavailable, intercropping of appropriately selected shade trees will not competitively suppress early growth of cocoa but will improve light regulation and nutritional status of cocoa saplings.