Experimental manipulations of water and nutrient input to a Norway spruce plantation at Klosterhede,Denmark

At Klosterhede, Denmark experimental manipulations of water and nutrient inputs to a forest stand were carried out under a 1200 m² roof simulating i) summer drought, ii) removal of N and S input, and iii) optimal nutrition and water availability. In practise, manipulation of element fluxes in a complex system like a forest may cause unintended changes such as climatic effects from the roof which may interfere with the results of the intended manipulation. This paper illustrates the extent and effects of such unintended changes.The most important climatic change under the roof was a 50% reduction of photosynthetic light which caused a decrease in moss cover on the ground floor. Soil temperatures under the roof were decreased 0.3°C during summer and increased 0.2°C during winter. Air humidity was not changed. Throughfall water amount decreased close to the stems whereas element concentrations in throughfall increased close to the stems and with tree size on the ambient plots. This distinct spatial pattern was changed under the roof to a more variable water distribution from the sprinkling system and to constant element concentrations in the sprinkling water. This loss/change of spatial variability in the input was reflected in the soil solution. The concentration patterns found in soil water (increasing close to the stem and with tree size) in the ambient plots were no longer observed under the roof.Exclusion of throughfall by the roof disturbed the internal cycle of nutrients leached from the canopy. The removal of canopy leached Ca and K by the roof caused a decline in soil water concentrations, even on the drought plot where natural throughfall was only cut off during two summer months. The internal cycle of Ca and K had to be restored by addition of Ca and K under the roof. Further, leachable elements and soluble compounds may be washed out from litterfall during the collection period on the roof, and thus lost from the internal cycle by by exclusion of throughfall.It is important to consider these problems in the design of new manipulation experiments and in the interpretation of the results. Some recommendations are given.     

Effects of irrigation and balanced fertilization on nutrient cycling in a Douglas fir stand

The effects on growth and nutrient status of an increased availability of all major nutrients was studied in a 40 year old Douglas fir stand. The nutrient amounts were applied by daily sprinkling and were in fixed, optimal proportions to the estimated annual uptake of N. Irrigation was included to avoid drought stress. The nutrient applications were also done in combination with a lower NH₄ load, realized under a roof that was placed above the ground. According to nutritional standards nitrogen supply was optimal and was related to the moderately high atmospheric N inputs. Nutrition of P, K and in some years Ca seemed critical.In all four studied growing seasons water additions resulted in a growth increase, of 30 to 40%. Irrigation decreased needle fall. Nutrient applications increased the needle concentrations of P and K considerably, but this did not result in a growth enhancement. Fertilization also increased K and P in shedded needles. Only minor leaching losses were calculated. The recovery in trees of fertilizer K and P was maximally 15%. Input-output budgets suggested that P was strongly retained in the soil. The decrease in NH₄ load had slightly decreased N availability and soil acidification, and seemed to enhance tree growth. The use of tree growth as a parameter to evaluate the effects of an improved nutritional balance is discussed.     

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.     

Effects of water and nutrient supply on amount and on nutrient concentration of litterfall and forest floor litter in Eucalyptus globulus plantations

A field experiment was initiated in March 1986 in central Portugal to evaluate the influence of water and nutrient supply on the productivity of E. globulus. The treatments applied were, irrigation plus fertiliser, irrigation and the application of fertilisers in rainfed plots. The control received neither fertilisers nor irrigation. The annual pattern litterfall was measured over a period of 5 years and the litter layer was quantified 6 years after planting. The amount of litterfall varied with the treatments. Simultaneous water and nutrient supply increased significantly litterfall, in respect to control. In rainfed conditions the timing of the maximum of litterfall was anticipated relative to the irrigated plots. However, the time of maximum litterfall did not coincide with the dry season but rather with period of maximum growth in each treatment. The N and P concentration was higher in the litterfall of the two fertilised treatments than in the others. The lowest concentrations of N and P in the leaf litter were coincident with the summer period in all treatments. The withdrawal of N and P ranged between 32 and 65% according to treatment and season. The deliverance of nutrients through litterfall was strongly increased by simultaneous water and nutrient supply. The supply of fertilisers in rainfed conditions promoted higher deliverance of nutrients than in the plots irrigated only. The mass of the litter layer was significantly increased by simultaneous water and nutrient supply. Application of fertilisers induced an increase in N and P concentration and a decrease in C/N ratio of the litter layer. Treatments and C/N values did not influence apparently the proportion of carbon, N and P released through mineralization from the litter layer. The non-fertilized treatments showed a more efficient N cycling than the others.     

The Role of Soil Nutrients in Boundaries between Mangrove and Herbaceous Assemblages in a Tropical Estuary

We aimed to determine the role of soil nutrients in the herbaceous‐tree boundary in a tropical mangrove forest. We assembled plots in this boundary such that each was initially covered by ~50% of each life form. We added NPK to three of the plots monthly and monitored them, along with the control plots, for 18 mo. In plots with added NPK, mangrove branches grew over and shaded the herbaceous plants, resulting in the presence of mostly bare soil. The mangrove trees eventually covered ~80% of the experimental plots, but the proportion of cover in the control plots remained unaltered. We conclude that soil nutrient level modulates the mangrove‐herbaceous plant interaction in steps, and from a given level there is a shift in the dominant life form. This mechanism leads to a sharp transition over a smooth nutritional gradient.     

油茶低产林养分需求和时间配置动态变化研究

以常宁油茶低产林为研究对象,分析了油茶低产林树体各器官及土壤养分时间动态变化。结果表明:油茶低产林树体在不同生长期需求的大量元素均为全N、Ca和全K最多,而需求的Mg和全P最少;需求的微量元素均为Mn和Fe最多,Cu和Cd最少;春梢期供应的土壤养分元素是速效N和Mg,夏梢期主要供应的土壤元素是速效K、全N、全P、Fe和有机质,果实成熟期主要供应的土壤养分元素是全K,开花期主要供应的土壤元素是速效P和Ca;不同时间油茶低产林养分需求为春梢期(28.36%)夏梢期(26.17%)果实成熟期(22.75%)开花期(22.73%);土壤养分供应为夏梢期(2999.83±87.04 mg/kg)果实成熟期(2703.93±292.26 mg/kg)开花期(2554.60±508.84 mg/kg)春梢期(2385.88±199.62 mg/kg);油茶低产林在不同生长期需要的养分和土壤供应的养分并不一一对应。研究结果可为油茶低产林的施肥时间配置和养分时间变化提供科学依据,在春梢期多施肥。     

Size-dependent responses to summer drought in Scots pine, Norway spruce and common oak

In this study, we provide a detailed analysis of tree growth and water status in relation to climate of three major species of forest trees in lower regions of Bavaria, Southern Germany: Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and common oak (Quercus robur). Tree-ring chronologies and latewood δ¹³C were used to derive measures for drought reaction across trees of different dimensions: growth reduction associated with drought years, long-term growth/climate relations and stomatal control on photosynthesis. For Scots pine, growth/climate relations indicated a stronger limitation of radial growth by high summer temperatures and low summer precipitation in smaller trees in contrast to larger trees. This is corroborated by a stronger stomatal control on photosynthesis for smaller pine trees under average conditions. In dry years, however, larger pine trees exhibited stronger growth reductions. For Norway spruce, a significantly stronger correlation of tree-ring width with summer temperatures and summer precipitation was found for larger trees. Additionally, for Norway spruce there is evidence for a change in competition mode from size-asymmetric competition under conditions with sufficient soil water supply to a more size-symmetric competition under dry conditions. Smaller oak trees showed a weaker stomatal control on photosynthesis under both dry and average conditions, which is also reflected by a significantly faster recovery of tree-ring growth after extreme drought events in smaller oak trees. The observed patterns are discussed in the context of the limitation-caused matter partitioning hypothesis and possible species-specific ontogenetic modifications.     

Effects of understory dwarf bamboo on soil water and the growth of overstory trees in a dense secondary <Emphasis Type="Italic">Betula ermanii</Emphasis> forest,northern Japan

The effects of understory dwarf bamboo (Sasa kurilensis) on soil water and the growth of overstory trees were studied in a dense secondary forest of Betula ermanii in northern Japan. Four plots were established in a Betula ermanii forest with Sasa kurilensis in the understory. The Sasa was removed in two of the plots. The annual increment of the trunk diameter for each tree was measured in the first two years from the commencement of the experiment. Soil water potential was similar in the plots following significant rainfall, but was found to be greater in the plot without Sasa between rainfall events. This suggests that the removal of Sasa slows the reduction of soil water after rainfall. The relative growth rate of the trunk diameter of Betula ermanii increased with tree size in all of the plots because taller trees strongly suppressed smaller ones in the dense forest. The growth rates of Betula ermanii were higher in the plots without Sasa. However, the difference in growth rates between all of the plots tended to be smaller in smaller size classes, possibly because smaller trees were strongly suppressed by larger ones, irrespective of the presence/absence of Sasa. Therefore, the removal of Sasa increased soil water and encouraged the growth of larger Betula ermanii in dense forest during the first two years after the Sasa was removed. The present study suggests that Sasa can reduce the growth of larger Betula ermanii in dense forest by limiting available soil water to these trees.     

Effects of water and nutrient applications in a Scots pine stand to tree growth and nutrient cycling

In a Scots pine forest stand, demineralized water and a complete set of nutrients with water were applied to the soil by means of frequent irrigation for four years in order to eliminate water and nutrient shortage of the trees. Apart from this optimization, dissolved (NH₄)₂SO₄ was irrigated at a rate of 120 kg N ha^-1 y^-1 to create a situation of N excess. Effect of treatments on tree growth and chemical composition of soil water and vegetation were monitored. From the first treatment year onwards basal area growth increased by ca. 35% as a result of the increased water supply. Nutrient applications increased K and P concentrations in pine needles immediately, but growth was enhanced only in the fourth treatment year and coincided with an improved K supply. Most of the applied P and K was retained in the soil, and only 6% was recovered in the vegetation. Tree nutrient status did not respond on Ca and Mg applications, whereas Ca and Mg seepage losses were increased with ca. 5 kg ha^-1 y^-1. The applied NH₄ was mostly retained in the 0–20 cm surface soil and caused a drastic increase of Al in soil solution. Tree growth was stimulated initially by extra NH₄, but was hampered after three years obviously because of a decreased P nutrition. The applied base cations were absorped to the soil and the accompanying anions were leached, thus temporarily increasing the acidification of the soil solution.     

红松人工林优势木竞争指数影响因子

在温带湿润气候区东段不同立地条件下的红松人工林内设置面积为600 m~2(20 m×30 m)的70块矩形固定样地。在每个样地内,选取5颗最高且长势较好的优势木作为研究的对象木,用Voronoi图确定优势木相应的竞争木,测定每一块样地内优势木与竞争木之间的距离。采用Hegyi单木竞争指数模型,分析优势木在不同样地水平上的种内竞争强度,探讨林分生长因子、地形因子、土壤养分因子对优势木竞争指数的影响,并对这3类因子与优势木竞争指数进行相应的拟合和相关性分析。结果表明：红松人工林优势木竞争强度随着优势木胸径的增大而变小,并且两者之间的关系服从幂函数;红松优势木的竞争指数与其树高、胸径、冠幅呈极显著的相关关系(P<0.01);坡向、坡位、海拔对竞争指数影响极显著(P<0.01);红松人工林优势木竞争指数的大小与土壤氮磷钾含量均呈极显著的相关关系(P<0.01);pH值对优势木竞争指数的影响不显著。当红松人工林优势木平均胸径达到45cm,优势木平均树高和冠幅大于周围竞争木时,其对周围资源的利用程度增大,林木会发生自然稀疏现象,其所受的竞争压力减小。红松喜光性强,对水分的要求高,...     

Constraint on woody cover in relation to nutrient content of soils in western southern Africa

The influence of soil nutrients on woody plants is poorly understood. Are trees – fire and other disturbance factors being equal – generally promoted by nutrient‐rich or by nutrient‐poor soils? To determine the edaphic parameters controlling woody cover, we sampled soils and summed the extent of the crowns of trees and tall shrubs on 364 plots at 20 sites in Namibia and adjacent South Africa, ranging from desert lichen‐fields to caesalpiniaceous woodland with associated mean annual rainfall of 11 mm to 535 mm. Our analysis included the macro‐nutrients N, P, Mg, K and Ca and the trace elements Mn, Fe, Cu and Zn. A boundary line analysis showed that woody cover was densest, with the greatest large‐scale heterogeneity, at intermediate nutrient contents, but consistently constrained at extreme nutrient richness as well as poverty. If aridity exerted the ultimate constraint at extreme nutrient richness, no such correlation with climate apparently applies at extreme nutrient poverty, where our graphs show an ‘oligotrophic decline’. Notwithstanding the importance of water, we suggest that extreme nutrient richness and poverty both favour grasses over tree seedlings. This is because catabolic dystrophy – a regime in which the supply of catabolic nutrients shortfalls their demand – is unlikely in environments where nutrient richness allows catabolic rates to match anabolic rates or where nutrient poverty constrains anabolic rates. We also reason that surpluses of photosynthate resulting from dystrophy can be allocated to lignin and that the potential for woody growth thus corresponds to soils of intermediate nutrient content. This explains why woody cover is consistently but not homogeneously densest in nutritionally intermediate plots in our dataset. Hence, the abundance of woody plants in various biomes may be determined partly by soil nutrient content, particularly of Cu, Zn, and other elements indispensible for catabolism.     

Alleviation of Summer Drought Boosts Establishment Success of <Emphasis Type="Italic">Pinus sylvestris</Emphasis> in a Mediterranean Mountain: An Experimental Approach

We performed an irrigation experiment to study the impact of summer drought on Pinus sylvestris establishment at its southernmost distribution limit. Watering was done during the first growing season simulating mesic summer conditions, and we monitored the consequences for survival and growth during the first growing season and the delayed consequences on the second growing season. In addition, we considered the heterogeneity created by the microhabitats, where seeds are found after dispersal (bare soil, under shrubs, and under adult pines). Summer drought was the main mortality factor in all the microhabitats. Watering increased emergence and doubled seedling survival compared to non-watered control sites. Differences were even higher when the cumulative effect on emergence and survival was considered, with an overall recruitment of 22.4% in watered plots vs. 7.9% in control. Irrigation increased growth in bare soil and under shrubs, but had scant effect on growth under pines, suggesting that radiation was the limiting factor in this microhabitat. The positive effect of irrigation on growth parameters persisted during the second growing season despite water was not added the second year, showing delayed consequences of drought on seedling performance. Summer drought thus limits Pinus sylvestris establishment in these southernmost forests by reducing both recruitment and growth. This might lead to the development of a remnant dynamic in these relict populations under the current regional increase in dryness and rainfall variability associated with global warming.     

Radial growth response of trees to seasonal soil humidity in a subtropical forest

Tree growth is the most important factor in determining the carbon sequestration processes of forest ecosystems. However, the growth phenology (seasonal growth pattern) and responses of tree growth to climatic variables vary considerably among different species, especially between deciduous and evergreen species. Thus, it is crucial to explore the seasonal growth patterns of different tree species in relation to climate to better understand the responses of tree physiology to climate changes, especially in mixed-species forest stands. In this study, we monitored the daily basal area increments of 220 individuals belonging to 15 common broadleaved tree species, nine deciduous and six evergreen species, in mixed-species experimental stands in subtropical China and analysed the relationships between radial stem growth and seasonal climate at a high-temporal resolution. We fitted daily increments of stem diameters with four frequently used nonlinear models and chose the best model for each species. The results showed that the evergreen trees grew faster than the deciduous trees, both annually and within the growing season. The tested nonlinear models (Korf, Weibull, logistic and Gompertz) produced good fits for the growth patterns of all species. Overall, the evergreen species began stem growth earlier and finished later during the growing season than that of the deciduous species. Within the growing season, the radial growth of trees in mixed stands containing both types of species was strongly positively correlated with humidity. In spring, increases in both temperature and moisture increased the daily relative basal area increment of all species. Maximum growth rates occurred when the soil water content reached its highest level and gradually decreased when the soil water content decreased. In summer, high temperatures combined with low amounts of precipitation led to heat-induced summer drought, to which the evergreen trees appeared to be more tolerant than the deciduous trees, which was reflected in the reduced stem growth of the latter. These results indicate the different climate-dependent seasonal growth strategies of evergreen and deciduous trees related to the trade-off described by the leaf economics spectrum, i.e., short-lived leaves with higher assimilation rates in deciduous and longer-lived leaves with a greater drought tolerance in evergreen species.     

Abundance of trees and grasses in a woodland savanna in relation to environmental factors

The relationships of plant species associations and underlying environmental factors in a woodland savanna in South Africa were investigated. 40 plots were included with 25 tree and 17 grass species dominating the arboreal and ground cover. Correspondence Analysis described the relationships between soil moisture retention, soil nutrients and the abundance of trees and grasses. Dry matter indices represented the accumulated effects of rainfall, fire and grazing of the herbaceous layer. Variations in the abundance of plants corresponded to well-defined gradients of soil nutrients. The distribution of grass and tree species along the ordination axes indicated that soils with high water retention capacity and high nutrient contents provided a suitable substrate for many of the tree species sampled. However, grass species abundance was high in plots with porous soils and poor nutrient availability.     

Dryland management regimes alter forest habitats and understory arthropod communities

Dryland forests, those characterised as having low precipitation and soil nutrients, account for over a quarter of forests globally. Increasing their productivity often relies on irrigation and fertilisation, but the impacts on the wider habitat are largely unknown. Understory invertebrates, in particular, play key roles in forest systems (e.g. nutrient cycling), but their responses to dryland forest management practices are untested. We investigated the impacts of irrigation, fertilisation and a combination of both on soil chemistry, understory vegetation, tree growth and understory arthropod communities in a Eucalyptus plantation to establish linkages between dryland management and ecosystem responses. Fertilisation increased all soil nutrients (N, NO₃N, P and K) with similar effects on the chemical composition of understory grasses. Fertilisation also caused declines in foliar silicon concentrations, an important herbivore defence in grasses. Irrigation increased growth of both understory plants (+90%) and trees (+68%). Irrigation increased the abundance of ground‐dwelling arthropods by over 480% relative to control plots, but depressed higher level taxon arthropod diversity by 15%, declining by a further 7% (?22%) in combined treatment plots. Irrigation also caused a surge in the abundance of Collembola (+1300%) and Isopoda (+323%). Fertilisation drove increases in the abundance of Isopoda (+196%) and Diptera (+63%), whereas fertilisation combined with irrigation increased populations of Thysanoptera (+166%) and Acarina (+328%). Airborne arthropods were less affected, but fertilisation increased the abundance of Apocrita (+95%) and depressed populations of Thysanoptera (?77%). Diptera abundance was positively related to understory vegetation growth, whereas the abundance of other groups (Collembola, Isopoda, Thysanoptera and Acarina) correlated positively with tree growth. We proposed that the large increases in populations of key detritivores, Collembola and Isopoda, were linked to increased leaf litter from enhanced tree growth in irrigated and combined treatment plots. Our findings suggest that dryland management can increase both plant productivity and abundance of arthropods, but cause arthropod diversity at the higher taxon level to decline overall.     

Water relations of citrus exocortis viroid-infected grapefruit trees in the field

Citrus viroids (CVds) were used to limit citrus tree size ('dwarfing'). It is hypothesized that changes also occur in the hydraulic properties of the conducting system, thereby affecting water relations. Grapefruit trees (var. Star Ruby on Troyer Citrange rootstock) in northern Israel were infected with citrus exocortis viroid (CEVd). The orchard was drip irrigated. The infected (I) and apparently healthy (H) control plots were subdivided into wet (W) and dry (D) irrigation treatments receiving 100% and 70% of the recommended irrigation quantity for the region. Soil water content, soil temperature, water uptake, leaf and stem water potential, and leaf conductance were measured in addition to climatic variables. Water uptake response to increasing spring-time soil temperatures was greater in healthy than in infected trees. Leaf conductance in the infected trees was lower than the non-infected trees only in the sixth year after inoculation. Stem water potential was significantly lower in the CEVd-infected trees than in the control trees. Water uptake was lower and hydraulic resistance higher in infected than in healthy trees. It was concluded that CEVd causes a reduction in the water uptake ability of the root and canopy system. Possible mechanisms for this are discussed.     

Stand dynamics modulate water cycling and mortality risk in droughted tropical forest

Transpiration from the Amazon rainforest generates an essential water source at a global and local scale. However, changes in rainforest function with climate change can disrupt this process, causing significant reductions in precipitation across Amazonia, and potentially at a global scale. We report the only study of forest transpiration following a long‐term (>10 year) experimental drought treatment in Amazonian forest. After 15 years of receiving half the normal rainfall, drought‐related tree mortality caused total forest transpiration to decrease by 30%. However, the surviving droughted trees maintained or increased transpiration because of reduced competition for water and increased light availability, which is consistent with increased growth rates. Consequently, the amount of water supplied as rainfall reaching the soil and directly recycled as transpiration increased to 100%. This value was 25% greater than for adjacent nondroughted forest. If these drought conditions were accompanied by a modest increase in temperature (e.g., 1.5°C), water demand would exceed supply, making the forest more prone to increased tree mortality.     

Drought alters timing, quantity, and quality of wood formation in Scots pine

Drought has been frequently discussed as a trigger for forest decline. Today, large-scale Scots pine decline is observed in many dry inner-Alpine valleys, with drought discussed as the main causative factor. This study aimed to analyse the impact of drought on wood formation and wood structure. To study tree growth under contrasting water supply, an irrigation experiment was installed in a mature Scots pine (Pinus sylvestris L.) forest at a xeric site in a dry inner-Alpine valley. Inter- and intra-annual radial increments as well as intra-annual variations in wood structure of pine trees were studied. It was found that non-irrigated trees had a noticeably shorter period of wood formation and showed a significantly lower increment. The water conduction cells were significantly enlarged and had significantly thinner cell walls compared with irrigated trees. It is concluded that pine trees under drought stress build a more effective water-conducting system (larger tracheids) at the cost of a probably higher vulnerability to cavitation (larger tracheids with thinner cell walls) but without losing their capability to recover. The significant shortening of the growth period in control trees indicated that the period where wood formation actually takes place can be much shorter under drought than the 'potential' period, meaning the phenological growth period.     

Foliar demand and resource economy of nutrients in dry tropical forest species

Important phenological activities in seasonally dry tropical forest species occur within the hot‐dry period when soil water is limiting, while the subsequent wet period is utilized for carbon accumulation. Leaf emergence and leaf area expansion in most of these tree species precedes the rainy season when the weather is very dry and hot and the soil cannot support nutrient uptake by the plants. The nutrient requirement for leaf expansion during the dry summer period, however, is substantial in these species. We tested the hypothesis that the nutrients withdrawn from the senescing leaves support the emergence and expansion of leaves in dry tropical woody species to a significant extent. We examined the leaf traits (with parameters such as leaf life span, leaf nutrient content and retranslocation of nutrients during senescence) in eight selected tree species in northern India. The concentrations of N, P and K declined in the senescing foliage while those of Na and Ca increased. Time series observations on foliar nutrients indicated a substantial amount of nutrient resorption before senescence and a ‘tight nutrient budgeting’. The resorbed N‐mass could potentially support 50 to 100% and 46 to 80% of the leaf growth in terms of area and weight, respectively, across the eight species studied. Corresponding values for P were 29 to 100% and 20 to 91%, for K 29 to 100% and 20 to 57%, for Na 3 to 100% and 1 to 54%, and for Ca 0 to 32% and 0 to 30%. The species differed significantly with respect to their efficiency in nutrient resorption. Such interspecific differences in leaf nutrient economy enhance the conservative utilization of soil nutrients by the dry forest community. This reflects an adaptational strategy of the species growing on seasonally dry, nutrient‐poor soils as they tend to depend more or less on efficient internal cycling and, thus, utilize the retranslocated nutrients for the production of new foliage biomass in summer when the availability of soil moisture and nutrients is severely limited.     

Influence of nutrient and water stress on Norway spruce production in south Sweden — the role of air pollutants

A field experiment primarily designed for simulating the indirect effects of air pollutants for a 25-year-old Norway spruce stand in SW Sweden is presented (The Skogaby project). Treatment include irrigation; artificial drought; ammonium sulphate addition; nitrogen-free-fertilization and irrigation with liquid fertilizers including a complete set of nutrients. The experiment has a randomized block design with four replicates per treatment. Growth response on an areal basis of basal area, height and dry mass of stems, branches and needles after up to four years of treatment are presented. Dry mass is estimated using allometric equations based on destructive samplings of trees.The stand suffered from temporary water stress during all four years investigated despite 970– 1160 mm of annual precipitation. Irrigation resulted in improved above-ground dry mass production (stem, bark, branches, needles, litter fall) by 20% during the first 3 years of treatment, whereas 2 years of drought treatment followed by 1 year of recovery led to 10% reduced dry mass growth. During year 2 of recovery, however, basal area growth was only about half of that of the control.Nitrogen, markedly, was a growth limiting nutrient, although the stand got approx. 20 kg N ha^-1 y^-1 from deposition. Ammonium sulphate addition (100 kg N ha^-1 y^-1) resulted in 31% improved dry mass production whereas irrigation with liquid fertilization (100 kg N ha^-1 y^-1) including all important nutrient elements led to 57% increased dry mass growth after 3 years of treatment. Basal area growth of the latter treatment gradually increased and during year 4 of treatment was 123% larger than the control. Nitrogen-free-fertilization resulted in a small improvement of dry mass production (+10%).After 3 years of treatment, the amount of needles had increased markedly for both treatments including irrigation, whereas drought treated trees instead had decreased their needle amount vs control. The increase in needle amount occurred as a result of both larger formation of needles and higher preservence of old needles, the opposite relations being found for the drought treated trees. At the same point larger needle formation in combination with a higher shedding of older needles was found for trees treated with ammonium sulphate and nitrogen-free-fertilizer.It is concluded that there is no stage of N saturation in the Skogaby site as there is no leaching of N from the control plots and N fertilization results in both increased tree growth and N uptake.