Modified exponential nitrogen loading to promote morphological quality and nutrient storage of bareroot-cultured Quercus rubra and Quercus alba seedlings

Abstract

Exponential nutrient loading has been used to improve nursery fertilizer uptake efficiency of conifer seedlings, but the technique has received little attention in the culture of temperate deciduous hardwoods. This study examined responses of northern red oak (Quercus rubra L.) and white oak (Q. alba L.) seedlings to modified exponential nitrogen loading during bareroot nursery culture using a broad range of nutrient supply from 0 to 3.35 g nitrogen (N) per plant per season for 18 weeks in Indiana, USA. Seedling growth and nutritional parameters followed a curvilinear pattern that ranged from deficiency to toxicity with increased fertilization consistent with trends depicted in the proposed model for nutrient loading. Fertilization increased plant dry mass by 113–260% for red oak and 49–144% for white oak. Severe nutrient deficiency occurred under indigenous soil fertility, and limited phosphorus and potassium uptake were found to inhibit seedling growth at higher N supply. The sufficiency and optimum rates were determined to be 0.84 and 1.68 g N per seedling per season, respectively, under the current cultural conditions. Fertilization at 1.68 g N per plant increased N content by 40% in red oak and 35% in white oak. This approach may be used to help refine nursery fertilization practices in hardwood culture to produce high-quality seedlings for field planting.     

Stand development and production dynamics of loblolly pine under a range of cultural treatments in north-central Florida USA

The objectives of this study were to examine the effects of stand development and soil nutrient supply on processes affecting the productivity of loblolly pine (Pinus taeda L.) over a period approximately equal to a pulpwood rotation (18 years). The experiment consisted of a 2×2 factorial combination of complete and sustained weed control and annual fertilization treatments (C: control treatment, F: fertilization, W: weed control, FW: combined fertilization and weed control), located on a Spodosol in north-central Florida, USA. The reduction of soil nutrient limitations through fertilization or control of competing vegetation resulted in dramatic increases in almost every measure of productivity investigated, including height (19.7 m in the FW treatment versus 12.5 m in the C treatment at age 18 years), basal area (FW=44.2 m² ha⁻¹, F=39.6 m² ha⁻¹, W=36.6 m² ha⁻¹, C=19.9 m² ha⁻¹ at age 16 years), stemwood biomass accumulation (114 Mg ha⁻¹ in FW versus 42.8 Mg ha⁻¹ in C at age 18 years), foliar nitrogen concentration (1.53% in plots receiving fertilization versus 1.06% in unfertilized plots at age 17 years) and leaf area index (age 16-year peak projected of approximately 3.3 at age 9–10 years in F and FW plots, 2.5 in the W treatment and 1.5 in the C plots). Cultural treatments also decreased the growth ring earlywood/latewood ratio, and accelerated the juvenile wood to mature wood transition. While soil nutrient supply was a major determinant of productivity, production changes that occurred within treatments over the course of stand development were equally dramatic. For example, between age 8 and 15 years, stemwood PAI in the FW treatment declined by 275%; similarly large reductions occurred in the F and W treatments over the same time period. The reductions in PAI in the treated plots were linearly related to stand BA, suggesting the decline in productivity was associated with the onset of inter-tree competition. Responses of stemwood PAI to re-fertilization treatments at age 15 years suggests that the declines in growth and growth efficiency with time were partially attributable to nutrient limitations.     

Interactive Effects of Mycorrhization and Fertilization on Growth,Nutrition, and Water Relations of Sweet Birch

Abstract

Pisolithus tinctorius(Pers.) Coker & Couch was evaluated as an ectomycorrhizal associate of sweet birch (Betula lentaL.). Sand culture and mine soil growth media were employed, and three fertilization treatments were imposed by application of differing Hoagland's nutrient solution #2 concentrations. The high fertilization treatment suppressed mycorrhizal formation by P. tinctoriusin both growth media, while the medium rather than the low treatment induced the most extensive colonization by this mycobiont. Seedlings with induced P. tinctoriusmycorrhization exhibited enhanced above— and below-ground growth according to shoot dimensions and dry weight and root weight and length. However, these responses were frequently absent with high fertilization, especially in the mine soil. Inoculated seedlings had higher foliar N, P, and K concentrations overall but lower Ca, Mg, and S concentrations than uninoculated seedlings. Those with P. tinctoriusalso had lower concentrations of several potentially phytotoxic trace elements, specifically Zn, B, Mo, and Al. Furthermore, xylem pressure and soil water potentials measured during a simulated drought episode indicated that water uptake was enhanced by these mycorrhizae. Again, however, the above physiological responses were often compromised by high fertilization. Collectively, these results indicate that P. tinctoriusis a viable mycobiont for the inoculation of sweet birch, and when substrate fertility is not excessive, can avail this host of physiological benefits that result in substantial growth enhancement.     

Variation in Stemwood Nutrient Concentrations in Scots Pine Growing on Peatland

Radial and longitudinal variation in the nutrient concentrations of Scots pine (Pinus sylvestris L.) stemwood and the effect of fertilization (PK and NPK) were studied on three peatland sites in eastern Finland. The results showed distinctive patterns in longitudinal and radial nutrient concentrations in stemwood. The N, P and K concentrations were highest in the youngest tissues both in the radial and longitudinal directions independent of the site type or fertilization. Ca and Mn accumulated in the older parts of stemwood. Increased N, P, K and Ca concentrations were found 9 yrs after fertilization in the annual rings formed during the first-5-yr period after fertilization. Decreased Mn concentrations were found in the annual rings formed during the 5-and-9-yr periods after fertilization. The results indicate that in the intensive utilization of Scots pine stemwood, e.g. whole stem harvesting instead of stem harvesting to a fixed diameter (7-8 cm in Finland), nutrient loss increases proportionally more than the amount of harvested biomass. The results also imply that this wood contains more undesirable elements, such as K, which may cause problems in wood combustion in power plants, and that fertilization raises these concentrations in wood. However, the concentrations in unfertilized trees were generally at about the same level as in Scots pine stemwood grown on infertile mineral soil sites.     

Heterobasidion annosum root rot in Picea abies: Modelling economic outcomes of stump treatment in Scandinavian coniferous forests

Abstract

The economic outcomes of stump treatment against spore infections of the root rot pathogen Heterobasiodion annosum s. l. were analysed based on simulations in four stands typical of Swedish forestry and forest management: (A) Norway spruce [Picea abies (L.) Karst.] stand on former agricultural soil (SI?=?32), (B) Norway spruce stand (SI?=?26) on forest land; (C) Mixed stand of Norway spruce and Scots pine (Pinus sylvestris L.) (SI?=?24) with only H. parviporum present, i.e. no infection of Scots pine and no possibility of interspecies spread of disease between hosts; and (D) same as C, but H. annosum s. str. (Fr.) Bref and H. parviporum Niemelä & Korhonen present, i.e. interspecies spread of disease possible. Models for disease development, growth and yield and cross-cutting were used in the simulations. The simulated decay frequency in Norway spruce trees ranged between 2 and 90%. Stump treatment at the previous final felling and in all thinning operations was profitable at interest rates 1 and 3% in stands A, B and D, but not in stand C. In stand C, no stump treatment at all or treatment in thinnings only gave the highest net future value. Implications for stump treatment in practical forestry are discussed.     

Trends of tree growth in Swedish forests 1953–1992: An analysis based on sample trees from the national forest inventory

Average tree height and basal area growth for Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) in Sweden were studied as functions of species, age, stand density, location and year of inventory in the period 1953–1992, on the basis of sample tree data from the National Forest Inventory. A highly significant annual increase of both height and basal area growth was found, of the magnitude 0.5–0.8%, during the 40 yr period. Possible reasons for the trend are discussed. The altered way of cutting in the early 1950s, from selective cutting to clear felling and thinning from below, has had a large impact. Also, improved regeneration methods, nitrogen fertilization and ditching have increased growth. The increasing atmospheric deposition of nitrogen is another possible factor.     

Lime,sewage sludge and mineral fertilization in a silvopastoral system developed in very acid soils

In recent years, in the European Union, sewage sludge production has been increased as a result of EU policy (European directive 91/271/EEC). Organic matter and nutrient sewage sludge contents, principally nitrogen, indicate it can be used as fertilizer. The objective of the experiment was to compare the effect of no fertilization, three doses of sewage sludge, with or without liming, and the fertilization usually used in the region applied over a period of 3 years on pasture production and tree growth in a silvopastoral system. The experiment was conducted in the northwest of Spain. The soil was very acid (soil pH = 4.5) and had very low nutrient levels, especially P, that is related to site index. It was sown with a grass mixture (25 kg ha⁻¹ of Lolium perenne L. 10 kg ha⁻¹ of Dactylis glomerata L. and 4 kg ha⁻¹ of Trifolium repens L.) in Autumn 1997 under a plantation of 5-year-old Pinus radiata D. Don at a density of 1,667 trees ha⁻¹. Liming and sewage sludge fertilization increased soil pH and reduced saturated aluminium percentage in the interchange complex (Al/IC) in the soil, coming up the effect before with liming. Medium and high sewage sludge doses increased pasture production in the two first years. In a silvopastoral system, positive tree growth response to different fertilization treatments depended on tree age, initial soil fertility, soil pH, the relationship of competition with pasture production and previous liming application.     

Depth of water table prior to ditch network maintenance is a key factor for tree growth response

Abstract

In boreal-drained peatland forests, tree growth is retarded by the gradual deterioration of drainage ditch networks. In order to avoid the development of suboptimal growth conditions, ditch network maintenance (DNM) operations (ditch cleaning and/or complementary ditching) are annually conducted on an area of about 70,000 ha in Finland. The previous studies indicate that the depth of the water table prior to DNM may influence the magnitude of the growth response to DNM. Tree growth does not necessarily increase after DNM at sites with large stand volume and subsequent low water levels due to tree stand evapotranspiration. We investigated how the pre-treatment water table depth (pre-WTD) in late summer relates to the growth of Scots pine (Pinus sylvestris L.) stands after the DNM operation in 12 field experiments. The increase in mean annual volume growth caused by DNM was negatively related to the pre-WTD, with the highest growth response in stands where the pre-WTD was less than 25–30 cm. DNM did not clearly increase stand growth in sites where the pre-WTD was more than 35–40 cm below the soil surface. There was a high variation in growth response to DNM between the water levels from 20 to 35 cm below the soil surface, indicating that factors other than water table depth are also needed as decision criteria for assessing the appropriate timing of ditch network maintenance.     

Autumn Fertilization in the Nursery Affects Growth of Picea abies Container Seedlings After Transplanting

Second-year Norway spruce seedlings [Picea abies (L.) Karst.] grown in containers were divided into three fertilization levels in August [5, 15 and 25 mg nitrogen (N) seedling^?1]. The resulting foliar concentrations of N were 11.0, 13.1 and 15.8 g kg^?1, respectively. Seedling height (mean 26.0 cm) did not differ among treatments. The next spring, the seedlings were tested in two experiments. (1) The seedlings were transplanted into pots containing sandy soil in the greenhouse, where they were fertilized with either pure water or nutrient solution (22 mg N l^?1). (2) The outplanting performance of the autumn-fertilized seedlings was tested on a sandy field. In the greenhouse experiment, the autumn fertilization level affected height growth and root egress of the seedlings significantly, but less than fertilization with a nutrient solution after planting. In the field experiment, during the first season after transplanting shoot growth of the seedlings increased with the level of autumn fertilization. After the second and third seasons, the seedling stem volume was highest with the highest level of autumn fertilization. These results suggest that, by improving the preplanting nutrient status of seedlings, the growth of shoot, stem diameter and root biomass can be enhanced after planting, especially on nutrient-poor soils. However, heavier autumn fertilization than that used here may yield a greater and more persistent increment in growth.     

Main Findings and Future Challenges in Forest Nutritional Research and Management in the Nordic Countries

Previous research has documented that nitrogen fertilization can lead to a significant tree growth increase in large parts of Scandinavia. Nutrient management of Nordic forests is currently under debate, mainly because environmental values are believed to be unnecessarily endangered by these actions. Nutrient management including fertilization, liming, recycling of wood ash or application of alternative nutrient sources can counteract nutrient deficiencies and imbalances, and be used for compensating the nutrients removed through harvesting. For successful implementation of nutritional management measures, several questions should still be studied. These questions mainly concern diagnosis of the need for applications, nutritional sustainability and treatment effects on the forest ecosystem and the surrounding environment.     

Long-term effects of stump harvesting on soil properties and tree growth in Scots pine and Norway spruce stands

Abstract

Effects of stump harvesting on the properties of surface soil and on the density, structure and growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) stands were estimated in a field trial in western Finland. The experiment was established in 1977 and measured in 2010. Stems and logging residues were harvested after clear-cutting, and stumps were lifted and removed from half of the experimental area. Sixteen plots were planted with pine seedlings and 16 with spruce. The main effects of stump harvesting were improved survival of planted trees and an increase in natural regeneration. No clearly negative effects were noted in the stand development. Stump harvesting had no or minimal effects on the properties of the organic layer and those of the 0- to 10-cm mineral-soil layer. Soil properties did not differ between tree species. Pine production was higher on plots with stump removal compared to plots without soil treatment.     

The impact of climate,drainage and fertilization on the survival and growth of pinus sylvestris L. in afforestation of open,low‐production peatlands

The objective of this study was to analyse the conditions for forest production on open, low‐production peatlands in Sweden with respect to climate, and water and nutrient regimes. The study focused on survival and growth of Scots pine (Pinus sylvestris L.) seedlings, planted 18 yrs ago in five experimental areas evenly distributed between south and north Sweden. Different ditch spacing and NPK fertilizer treatments were combined systematically in all experiments. Survival was positively correlated with temperature sum, fertilization and drainage intensity. Tree growth was not influenced by variations in temperature sum after merely draining, but in combination with fertilization, growth was strongly correlated with climate. In the southern experiments, fertilization increased stand growth eight to nine times, whereas stands on the northern sites did not respond to fertilization. The most important fertilizer element was P. The application of N had no effect on growth. More intensive drainage increased stand growth by 60%. In the southern areas, height development in the most intensive drainage and fertilizer treatments indicates a mean annual increment of 6–7 m³ ha^?1, and no sign of decline in growth was seen. Turf‐planting had positive effects on both survival and growth, especially in less intensively drained plots. A large proportion of damaged trees was observed in the experiment. The frequency of damaged trees was positively correlated with treatment intensity but negatively correlated with temperature sum. The results show clearly that merely draining is not sufficient for successful afforestation of low‐production peatland sites. Fertilization by P and K is a necessary prerequisite, but the effect of fertilization varies with climatic conditions, probably owing to the amount of plant‐available N.     

Fertilization in a Fourth Rotation Eucalyptus grandis Plantation with Minimal Management

ABSTRACT

Successive rotation and wood harvest of fast growing trees, like Eucalyptus grandis, can deplete soil nutrient pools. In these intensively used soils, productivity can decrease, and fertilization can be necessary to recover soil fertility. The aim of this work is to evaluate if fertilization with N and P increases tree growth and foliar nutrients in an Eucalyptus grandis plantation growing over a soil used for three rotations of this species. In the same experiment, both fertilization on seedlings and coppiced trees were evaluated. One and two years after planting, any fertilization treatment increased tree growth, even though coppiced trees had higher total height and DBH than replanted trees. Coppiced trees had lower specific leaf area, as well as lower N and P leaf concentration. The lack of promotion in growth due to fertilization can be explained by the inputs of nutrients from the harvest residues of the previous rotation. As residues were never burned in this site, its decomposition released a high part of nutrients necessary to sustain growth during the first two years. We conclude that in sites where residues were left on site and were not burned, nutrient availability is enough and so fertilization at establishment is not necessary. It remains to be determined if fertilization is necessary at mid or end rotation, in order to balance nutrients export with harvest wood.     

Effect of Nutrient Optimization on Branch Characteristics in Picea abies (L.) Karst

The number and size of branches are major determinants of log grading. To study the effect of fertilization on branch characteristics in Norway spruce, trees were sampled in a nutrient optimization experiment in northern Sweden. Data were collected from 24 40-yr-old trees from fertilized and control plots after 12 yrs of annual nutrient application. On the fertilized plots, all essential macronutrients and micronutrients were supplied in irrigation water every second day during the growing season. Fertilization increased the number of new branches, but its effect on branch angle and branch mortality was small. Fertilization increased the diameter of the branches that were formed after the initiation of the treatment, but also the diameter of the branches that were formed before the fertilization commenced. Fertilization enhanced the general growth of a tree, but there were no significant changes in the relationships between stem and branch characteristics. Improved growth rate of trees, as an effect of fertilization, may impair branching quality of logs. Such negative effects may, however, to some extent be counteracted by planting density at stand establishment and later thinning regimens.     

Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

Long-term Growth Effects Following Forest Nitrogen Fertilization in Pinus sylvestris and Picea abies Stands in Sweden

Forest nitrogen (N) fertilization induces changes in the soil and soil microorganisms that could hypothetically affect the long-term productivity of the soil. The tree growth response following a normal (i.e. 150 kg N ha^?1) single shoot N fertilization has a duration of 7–10 yrs. The aim of this study was to investigate whether any residual effects on tree growth persist, which could be attributed to previous N fertilization. The study included six Pinus sylvestris L. and three Picea abies (L.) Karst. experimental sites, sampled for growth parameters 14–28 yrs after the last fertilization. Residual growth effects were on average small, and not statistically significant. Negative residual growth effects of varying duration could be discerned, especially at low-fertility P. sylvestris sites. However, there was an overall tendency for growth to increase in the long term after N fertilization. The main conclusion is that operational forest N fertilization with a normal N dose should not be regarded as a threat to long-term forest production.     

Estimates of above-ground biomass and nutrient accumulation in Mimosa scabrella fallows in southern Brazil

Naturally regenerated stands of bracatinga (Mimosa scabrella Bentham) are harvested for firewood after six to eight years of unregulated growth, debris burnt and the area planted to one cycle of intercropped maize (Zea mays L.) and beans (Phaseoulus vulgaris L.). Burning breaks dormancy of bracatinga seed (> 80% germination) marking the onset of a new fallow-crop cycle. This production system has been practiced for nearly 100 years in Southern Brazil, covering some 60,000 ha in 3,000 small farms. An estimation of above-ground biomass and nutrient accumulation was made using literature data on stand age, population numbers, tree sizes, tree biomass partitioning and concentration of major nutrients in tree tissues. A simple simulation model, used to quantify above-ground nutrient pathways and their temporal dynamics, confirmed that six to eight years is the optimal rotation length. Biomass and nutrients deposited onto the soil, peak at stand age six years, which may result in significant soil fertility improvement prior to crop planting. At year six, estimated total above-ground biomass amounts to 83 Mg ha^–1; 44 Mg ha^–1 available as firewood and 39 Mg ha^–1 to be returned to the soil. Roughly half the amount of nutrients fixed in the above-ground bracatinga biomass would be exported in firewood and subsequent grain crops.     

Relationships between local stand density and local species composition and nutrient content in the topsoil of pure and mixed stands of silver fir (Abies alba Mill.)

This study examined topsoil samples from the humic mineral horizon in four pure silver fir (Abies alba Mill.) and five mixed stands with beech (Fagus sylvatica L.), spruce (Picea abies L.; Karst.) and pine (Pinus sylvestris L.). In every stand, 60–70 sampling locations were chosen using a stratified random scheme that represented a gradient in local stand density and local species composition. One working hypothesis was that stand density and species composition affect the content of nutrients in the mineral humic soil horizon. Specifically, the analysis statistically tested for relationships between general and species-specific local stand density measures (basal area, sum of tree diameters weighted by distance, canopy openness) and the contents of total C, total N (in four stands), base cations (Ca²⁺, Mg²⁺, K⁺, Na⁺) and Mn²⁺ (in nine stands). A considerable variation in nutrient concentrations was observed; the coefficients of variation ranged from 0.21 to 0.38 for N, from 0.20 to 0.94 for the sum of base cations and from 0.56 to 0.95 for Mn. However, no clear relationship emerged between local stand density and the concentration of N and base cations; in five stands, manganese showed a negative correlation with local stand density. In the mixed fir-beech stand, significantly higher concentrations of Ca, K and Mn were found in the vicinity of beeches, and species-specific stand density explained up to 15, 29 and 34% of total variation in the content of these elements, respectively. Similar tendencies also appeared in the mixed stands with pine, although the relationships were weak. The results obtained for two fir-spruce stands were ambiguous. In sum, in the stands studied, small canopy gaps and inter-crown openings might not be considered ‘‘nutrient hot spots’’, and the admixed species, except for beech, had an insignificant effect on nutrient content in the humic mineral horizon.     

The effects of fertilization on the health status,nutrition and growth of Norway spruce forests with yellowing symptoms

We studied the effects of magnesium (Mg) NPK fertilizer applied in 2000 to maturing Norway spruce (Picea abies /L./ Karst.) stands with yellowing symptoms and situated on poor acidified soils in the ?umava mountains (the Czech Republic). Long-term defoliation, nutrient content in the foliage and diameter increment were evaluated in three specific variants: yellowing trees, trees without yellowing symptoms and fertilized trees. The results indicate differences in soil characteristics, nutrient content and radial growth with response to climate. Fertilization increased Mg content and base saturation in the soil and decreased the amount of exchangeable aluminum and hydrolytic acidity. Differences were significant especially in magnesium, calcium and phosphorus contents in the foliage of yellowing trees. There was also a significant difference in manganese content; yellowing trees showed deficiency in comparison with both fertilized and unfertilized healthy trees. Two years after the initial fertilization (to present), defoliation and yellowing symptoms in the unfertilized yellowing trees were more pronounced compared to other variants. These symptoms were more influenced by fertilization than foliation. The duration of significant positive effects of fertilization on diameter increment lasted for a shorter time than the effect on foliage. Simultaneously, higher sensitivity to climatic factors was observed in the fertilized trees.     

Changes of soil chemistry,stand nutrition,and stand growth at two Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) sites in Central Europe during 40 years after fertilization,liming, and lupine introduction

Long-term (40 years) effects of two soil amelioration techniques [NPKMgCa fertilization + liming; combination of PKMgCa fertilization, liming, tillage, and introduction of lupine (Lupinus polyphyllus L.)] on chemical topsoil properties, stand nutrition, and stand growth at two sites in Germany (Pfaffenwinkel, Pustert) with mature Scots pine (Pinus sylvestris L.) forest were investigated. Both sites are characterized by base-poor parent material, historic N and P depletion by intense litter-raking, and recent high atmospheric N input. Such sites contribute significantly to the forested area in Central Europe. Amelioration resulted in a long-term increase of pH, base saturation, and exchangeable Ca and Mg stocks in the topsoil. Moreover, significant losses of the forest floor in organic carbon (OC) and nitrogen stocks, and a decrease of the C/N ratio in the topsoil were noticed. The concentrations and stocks of OC and N in the mineral topsoil increased; however, the increases compensated only the N, but not the OC losses of the forest floor. During the recent 40 years, the N nutrition of the stands at the control plots improved considerably, whereas the foliar P, K, and Ca concentrations decreased. The 100-fascicle weights and foliar concentrations of N, P, Mg, and Ca were increased after both amelioration procedures throughout the entire 40-year period of investigation. For both stands, considerable growth acceleration during the recent 40 years was noticed on the control plots; the amelioration resulted in an additional significant long-term growth enhancement, with the NPKMgCa fertilization liming + being more effective than the combination of PKMgCa fertilization, liming, tillage, and introduction of lupine. The comprehensive evaluation of soil, foliage, and growth data revealed a key relevance of the N and P nutrition of the stands for their growth, and a change from initial N limitation to a limitation of other growth factors (P, Mg, Ca, and water).