Growth, allocation and tissue chemistry of Picea abies seedlings affected by nutrient supply during the second growing season

One-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were grown hydroponically in a growth chamber to investigate the effects of low and high nutrient availability (LN; 0.25 mM N and HN; 2.50 mM N) on growth, biomass allocation and chemical composition of needles, stem and roots during the second growing season. Climatic conditions in the growth chamber simulated the mean growing season from May to early October in Flakaliden, northern Sweden. In the latter half of the growing season, biomass allocation changed in response to nutrient availability: increased root growth and decreased shoot growth led to higher root/shoot ratios in LN seedlings than in HN seedlings. At high nutrient availability, total biomass, especially stem biomass, increased, as did total nonstructural carbohydrate and nitrogen contents per seedling. Responses of stem chemistry to nutrient addition differed from those of adult trees of the same provenance. In HN seedlings, concentrations of alpha-cellulose, hemicellulose and lignin decreased in the secondary xylem. Our results illustrate the significance of retranslocation of stored nutrients to support new growth early in the season when root growth and nutrient uptake are still low. We conclude that nutrient availability alters allocation patterns, thereby influencing the success of 2-year-old Norway spruce seedlings at forest planting sites.     

Water relations and morphological development of bare-root jack pine and white spruce seedlings: seedling establishment on a boreal cut-over site

Bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) seedlings were planted on a boreal cut-over site and subsequent growth and seedling water relation patterns were monitored over the first growing season. Comparison of morphological development between white spruce and jack pine showed jack pine seedlings had greater new root development and a lower new shoot/new root ratio, while white spruce seedlings had greater new shoot development. Seasonal water relation patterns showed white spruce seedlings to have a greater decrease in xylem pressure potential (_x) per unit increase in transpirational flux density in comparison to jack pine seedlings. These results suggest that the greater resistance to water flow through the soil-plant-atmosphere continuum in white spruce seedlings compared to jack pine seedlings may be due to the relative lack of new root development in white spruce. Stomatal response of the seedlings showed that as absolute humidity deficit between the needles and air (AHD) increased, needle conductance (gwv) decreased in both species, but at low AHD levels white spruce had gwv approximately 35% higher than jack pine. For white spruce seedlings, gwv decreased as _x became more negative in a predictable curvilinear manner, while gwv of jack pine seedlings responded to _x with a threshold closure phenomenon at approximately - 1.75 MPa. Tissue water potential components for jack pine and white spruce seedlings at the beginning and end of the growing season showed jack pine to reach turgor loss at 76% relative water content while white spruce reached turgor loss at 88% relative water content. White spruce seedlings showed osmotic adjustment over the growing season, with an osmotic potential at turgor loss of - 1.27 MPa and - 1.92 MPa at the beginning and end of the growing season, respectively. Jack pine did not show any osmotic adjustment over the growing season. The implication of morphological development on water relation patterns are discussed with reference to successful seedling establishment.     

Weed control increases nitrogen retranslocation and growth of white spruce seedlings on a reclaimed oil sands soil

Early establishment of seedlings in reclaimed oil sand areas is often limited by low nutrient and water availability due to factors such as strong understory vegetation competition. Management practices such as nursery fertilization and field weed control could help early establishment of planted seedlings and reclamation success. We investigated the effect of nursery nutrient loading and field weed control on the growth, nitrogen (N) retranslocation within seedling components, and plant N uptake from the soil for white spruce (Picea glauca [Moench] Voss) seedlings planted on a highly competitive reclaimed oil sands site for two years. Exponential fertilization during nursery production increased the root biomass but not the nutrient reserve in the seedling. In the field experiment, on average across the treatments, 78 and 49% of the total N demand of new tissue growth in the first and second year were met by N retranslocation, respectively. Though exponential fertilization did not affect N retranslocation, it increased the percent height and root collar diameter growth. Weed control increased not only the growth of seedlings by increasing soil N availability, but also N retranslocation within the seedlings in the second year after outplanting. We conclude that vegetation management by weed control is feasible in improving the early growth of white spruce seedlings planted on reclaimed soils and facilitate tree establishment in the oil sands region. Optimization of the nursery exponential N fertilization regime for white spruce may further help with early revegetation of reclaimed oil sands sites.     

Premature shoot growth termination allows nutrient loading of seedlings with an indeterminate growth strategy

Nutrient loading of nursery seedling stock of species with an indeterminate growth strategy is challenging and poorly understood. Here, we explore the use of two potential techniques for nutrient loading of trembling aspen (Populus tremuloides Michx.) seedlings: (1) exponential fertilization and (2) early shoot growth termination in order to divert assimilated nutrients and carbon to storage rather than to growth. In the first study, aspen seedlings were treated with either exponential or constant fertilization rates, both of which supplied the same amount of nutrients over the growing season. Exponential fertilization resulted in overall poor planting stock form (stunted seedling growth and weak root development) and produced only marginal improvements of nutrient status. As a result, the exponential fertilization regime studied cannot be recommended as a treatment for aspen seedlings. In the second study we treated seedlings with a 2 × 2 factorial combination of fertilization and shoot growth inhibitor (SGI) applications with the fertilizer treatments varying in terms of mid-season fertilizer concentrations. Seedlings with SGI application had much higher tissue nutrient and carbon reserve concentrations than seedlings without a SGI application. In addition, nutrient uptake appeared to be more efficient in SGI treated seedlings, which could potentially result in significant reductions of nutrient application rates during aspen seedling production in nurseries. Overall, early shoot growth termination using a SGI appears to be an effective technique to produce nutrient loaded aspen seedlings.     

Wachstum,Mykorrhiza und Frostresistenz von Fichtenjungpflanzen bei Düngung mit verschiedenen Stickstoffgaben

Growth, mycorrhiza and frost resistance of Picea abies seedlings following fertilization with different levels of nitrogen . NPK fertilization with varying levels of nitrogen causes increased growth of spruce seedlings and faster flushing in spring. During the growing season, the frost resistance of all organs of the plants, especially new needles and shoots is the lower, the more nitrogen the plants have received. Consequently, spruce plantations, especially at high altitudes, may be liable to late frost damage.     

Nutrient enrichment of white spruce seedlings during nursery culture and initial plantation establishment

Growth and nutrient dynamics of bare-root white spruce (Picea glauca (Moench) Voss) seedlings were monitored for three years in a nursery and for one season after planting to assess effects of three nursery fertilization regimes. Nitrogen (totaling 0, 650, and 1300 kg N ha(-1) for 3 years) was applied conventionally in equal amounts during the growing season. The high-N treatment, representing a nutrient loading regime, was also applied seasonally at exponentially increasing rates as a fourth treatment. By the end of the 3-year nursery rotation, the N treatments had stimulated growth by 104-180% and nutrient uptake by as much as 381, 224 and 145% for N, P and K, respectively, inducing large accumulations of N and P in both conventionally and exponentially loaded seedlings. Compared with exponentially loaded seedlings, the concentrations of nutrients were less in conventionally loaded seedlings although their biomass was larger (31%). High nutrient reserves in the seedlings at the end of the nursery rotation resulted in increased biomass production (40-190%) after planting, which was related to the ability of the seedlings to retranslocate internal reserves for new growth, despite little or no net uptake of nutrients during the first season after planting. Compared with conventionally loaded seedlings, retranslocation was greater in exponentially loaded seedlings, which had accumulated larger and more readily available nutrient reserves during the nursery phase. The shortness of this study limits its usefulness for predicting the persistence of the loading response after planting, but we postulate that the high nutrient status of loaded seedlings at the end of the nursery rotation will contribute to future growth through increased nutrient storage and retranslocation, thus prolonging the loading response.     

Fall fertilization of Pinus resinosa seedlings: nutrient uptake, cold hardiness, and morphological development

? Fall fertilization may increase plant nutrient reserves, yet associated impacts on seedling cold hardiness are relatively unexplored.

? Bareroot red pine (Pinus resinosa Ait.) seedlings in north-central Minnesota, USA were fall fertilized at the end of the first growing season with ammonium nitrate (NH₄NO₃) at 0, 11, 22, 44, or 89 kg N ha^?1. Seedling morphology and cold hardiness [assessed by freeze induced electrolyte leakage (FIEL)] were evaluated six weeks after fertilization and following the second growing season.

? Seedling height and number of needle primordia increased with fertilizer rate for both sampling years. Seedlings fertilized with 44 and 89 kg N ha^?1 attained target height (15 cm) after the second growing season. Shoot and root N concentration increased after the first growing season in fall fertilized seedlings compared to controls. Fall fertilized seedlings had lower FIEL (i.e., increased cold hardiness) compared to controls when tested at ?40 °C after the first growing season, but no significant differences in FIEL of control and fertilized seedlings were observed after the second growing season.

? Results suggest that fall fertilization of red pine seedlings can help render desired target height in the nursery, while maintaining or increasing cold hardiness levels.

     

Physiological responses of black spruce layers and planted seedlings to nutrient addition

We investigated effects of nutrient addition on several physiological characteristics of 60-cm-tall black spruce (Picea mariana Mill. B.S.P.) layers (i.e., rooted branches of overstory trees) and 20-cm-tall planted seedlings on a clear-cut, N-limited boreal site. After two growing seasons, current-year and one-year-old needles of fertilized trees (layers and seedlings combined) had higher net photosynthetic rates (A(n)) and maximum capacity of Rubisco for CO(2) fixation (V(max)) than unfertilized trees. One-year-old needles of fertilized trees had higher stomatal conductance (g(s)), higher water-use efficiency, and lower intercellular to ambient CO(2) ratio than unfertilized trees. Additionally, fertilized trees had higher predawn and midday shoot water potentials than unfertilized trees. Stomatal conductance of 1-year-old needles was 23% higher in seedlings than in layers, but there were no significant differences in g(s) of current-year needles between the regeneration types. For both needle age-classes, A(n) and V(max) of layers were 25 and 40% higher, respectively, than the corresponding values for seedlings. The higher values of A(n), V(max) and foliar N concentration of layers compared with seedlings after two growing seasons may be associated with the larger root systems of the layers compared with the transplanted seedlings.     

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.     

The nutrient status of Norway spruce in pure and in mixed-species stands

Atmospheric deposition of N and S appears to have caused nutrient imbalance in Norway spruce stands in southern Sweden. This calls for a change of forest management to procedures that promote nutrient balance. Studies have shown lower soil acidity in Norway spruce/deciduous mixed stands than in spruce monocultures, but the tree nutrient status in such mixtures has not been much investigated so far.

The nutrient status of Norway spruce foliage and top mineral soil chemistry in monocultures and in stands mixed with beech, birch, or oak was investigated through paired comparisons on 30 sites in southern Sweden (27 sites) and eastern Denmark (three sites). In total, 45 mixed stands and 34 pure stands were included in the study.

Spruce needles from mixed stands had higher concentrations and ratios to N of K, P, and Zn than needles from pure spruce stands. Among the mixed stands, the K status appeared to be positively correlated with the percentage of deciduous tree basal area. Soil samples from mixed stands had a higher Mg concentration, base saturation, and BC/Al ratio than soil samples from pure stands. The spruce needle nutrient status was comparable in pure stands on fertile sites and in mixed stands on poor sites. We did not detect any differences in spruce tree growth between pure and mixed stands.

This paper discusses possible reasons for a positive effect on the tree nutrient status in mixed-species stands and the possibility of using mixed-species stands as a forest management procedure to avoid nutrient imbalance.     

Exponential nutrient loading: a new fertilization technique to improve seedling performance on competitive sites

Reforestation efforts in Ontario have become increasingly more reliant on containerized planting stock since these seedlings are less costly to produce and plant than bare-root stock. Container seedlings, however, tend to be more susceptible to competing vegetation when planted on weed-prone sites often requiring release by chemical herbicides. We have developed cultural techniques to improve the competitiveness of containerized seedlings by promoting initial outplanting performance thus reducing the need for early vegetation control. The approach is based on two new preconditioning practices, exponential fertilization and nutrient loading, which induce a steady-state build up of nutrient reserves in seedlings for outplanting. Exponential nutrient loading integrates these practices with high dose fertilization inducing luxury consumption. Steady-state culture corresponds better with the competitive outplanting environment, since stable internal nutrient accumulation in the greenhouse phase conforms with steady-state nutrient uptake of natural exponentially growing vegetation in the field, and exponential nutrient delivery to container-restricted root systems also better simulates nutrient flux reached by expanding roots in a field soil with constant nutrient availability. Combined with nutrient loading, the higher nutrient reserves and improved nutrient balance in seedlings contribute to enhanced stress resistance and increased growth performance. This paper reviews the theory and practice of exponential nutrient loading during seedling culture, presents results of growth and nutritional responses of seedlings planted on competitive sites, and suggests appropriate criteria for quality testing of nutrient loaded stock.     

Seasonal patterns of colonization of Norway spruce needles by Lophodermium piceae

Summary Colonization of living Norway spruce needles by Lophodermium piceae was studied in southern Sweden. One‐month to 3‐year‐old needles were assessed for frequency of infection. Needles appearing in the growing season of 1996 were monitored and sampled at 1 to 4 week intervals over a period of 14 months. Infection frequency of the current‐year needles increased from around 20% in July to approximately 40% in August 1996. This level had then doubled to approximately 80% after another growing season in August 1997. Multiple infections were rare in the needles during the first growing season, but after the second growing season at least 70% of the needles had multiple infections. The average number of infections in 9‐, 21‐ and 33‐month‐old needles were 1.3, 6.3 and 8.4, respectively. The size of the individual mycelia was estimated to be small in all needles, none of the individuals could be isolated from more than two adjacent needle sections, about 1.5 mm long. In conclusion, the number of L. piceae infections per needle increased during ageing of needles but mycelial growth is probably postponed until needle senescence and death.     

Seasonal nutrient dynamics in white pine and white spruce in response to environmental manipulation

Seasonal retranslocation in white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) was examined in response to silvicultural treatments (scarification, annual fertilization application, and annual control of competing vegetation with herbicide) that changed both environmental conditions and the growth rate of the trees. Four years after plantation establishment and initial treatment, nutrient accumulation in current-year needles of white pine and retranslocation from 1-year-old needles were increased following the vegetation control treatment, which increased resource availability (nutrients, water and light) and, hence, growth rate. Nutrient accumulation also increased in current-year white spruce needles following the same treatment, whereas retranslocation decreased in 1-year-old white spruce needles. Correlations of retranslocation (N, P and K) with growth rate (shoot biomass increment) showed a strong positive relationship for white pine and a negative relationship for white spruce. Retranslocation of K was correlated with foliar and soil K concentrations; the availability of this nutrient was also significantly reduced by vegetation control. A general theory for the control of nutrient retranslocation in conifers, which is not based exclusively on either sink strength or soil nutrient availability, is proposed. We conclude that retranslocation response is species specific and related to the potential phenotypic growth response to changing environmental conditions and to short-term imbalance in the supply versus the demand for nutrients.     

Responses of Pinus sylvestris and Picea abies Seedlings to Limited Phosphorus Fertilization and Treatment with Elevated Ozone Concentrations

Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] seedlings were exposed to high phosphorus (HP) or low phosphorus (LP) availability for one growing season in the open field, and to combined P availability and elevated ozone (O 3 ) concentrations (0, 55, 110 and 210 ppb for Scots pine and 0, 40, 75 and 150 ppb for Norway spruce, respectively) for 28 days in controlled laboratory chambers. Compared with HP, the LP treatment reduced Scots pine current-year (C) shoot and root dry masses and Norway spruce total dry mass, whereas the highest O 3 concentrations increased the magnesium concentration of Scots pine C needles and P concentrations of the C needles of both tree species. Chlorophyll a, a+b and carotenoid concentrations of Scots pine C needles were significantly higher in the LP treatment compared with HP under the highest O 3 concentration (210 ppb). In the mesophyll tissue of C needles of both tree species, LP treatment increased the size of mitochondria and elevated O 3 -induced granulation of chloroplast stroma and disintegration of cytoplasm. Exposure to elevated O 3 concentrations increased swelling of chloroplast thylakoids and reduced the amount of vacuolar tannin in the LP Scots pine C needles. The results suggest disturbances in needle photosynthetic machinery due to acute exposure to the combination of elevated O 3 and low P availability. However, clear additive effects were found only in needle P concentrations < 1 mg g -1 in short-term O 3 exposure.     

Temporal variation in nutrient concentrations of Pinus sylvestris needles

Temporal variation in nutrient concentrations of Scots pine (Pinus sylvestris L.) needles was studied during a three‐year period in three stands of differing stages of development. Concentrations of N, P and K varied significantly between years; this variation was related to differences in needle dry weight. Concentrations of all measured nutrients (N, P, K, Ca, Mg, Mn, Cu, Zn, Fe, B) and Al varied between seasons; this variation was related to nutrient mobility and the annual physiological cycle. Concentrations of the mobile nutrients N, P and K decreased in spring and early summer during shoot and needle elongation and increased in late summer and autumn during needle senescence and litterfall. Concentrations of Mg, Cu, Zn and B followed somewhat similar patterns. The poorly mobile nutrients Ca, Mn and Fe accumulated gradually in needles during each growing season. Needle nutrient concentrations were stable during the nonactive period.     

The effect of carbon and nutrient loading during nursery culture on the growth of black spruce seedlings: a six-year field study

We tested the effects of exponential nutrient loading and springtime carbon loading during nursery culture on the field performance of black spruce (Picea mariana (Mill.) B.S.P.). Seedlings were grown from seed with a conventional, fixed dose fertilizer (10 mg N seedling⁻¹) or an exponential nutrient loading regime (75 mg N seedling⁻¹). The following spring, seedlings were exposed for two weeks to either ambient (370 ppm) or elevated levels of CO₂ (800 ppm) and then planted in the field; seedling growth was followed for the next six years. Exponential nutrient loading increased seedling height, stem diameter and leader growth, with the largest increases in height and leader length occurring in the first three years after outplanting. Carbon loading increased seedling height and leader length, but only in seedlings that had been exponentially nutrient loaded. A combination of carbon and nutrient loading increased shoot height 26%, stem diameter 37% and leader length 40% over trees that received neither treatment. These results demonstrate that the growth enhancement seen under exponential nutrient loading is maintained under field conditions for at least six years. Carbon loading just before outplanting was a useful supplement to nutrient loading, but was ineffective in the absence of nutrient loading.     

Effects of low-dose liming and nitrogen fertilization on stemwood growth and needle properties of Picea abies and Pinus sylvestris

Stemwood growth, needle weight and nutrient concentrations in needles after liming and nitrogen fertilization were studied in two randomized block trials. The treatments were a single application of 500 kg lime per hectare, 1000 kg limer per hectare, 150 kg N per hectare and combinations of the two compounds. The trials are situated in the southern part of Sweden on mesic mineral soils. One of the experiments was located in 32–55-year-old highly productive Picea abies stands and the other in a 31-year-old Pinus sylvestris stand.As calculated by growth-ring analysis, liming alone did not significantly affect the growth during a 10-year period following treatment. Nitrogen fertilization increased growth in pine, but not in spruce. Lime in combination with N gave about the same effect in pine as the pure N treatment, whereas lime and N interacted in spruce and resulted in increased stemwood growth.Current-year needles were sampled during the winter, weighed and analysed for nutrient concentrations. In the N-treated plots the N concentration increased in both tree species, whereas the needle weight increased only in pine. In two cases, lower concentrations were indicated. In the first, there was a tendency of decreased N concentrations in spruce needles after liming. In the second, the magnesium concentrations decreased in needles of both species, especially in the treatments that included nitrogen. The concentrations of the measured nutrients were never below levels considered to imply a severe deficiency.     

Nitrogen availability modifies the ozone responses of Scots pine seedlings exposed in an open-field system

Three-year-old Scots pine (Pinus sylvestris L.) seedlings were exposed to either ambient or elevated (1.5-1.6 x ambient) ozone concentration ([O3]) for three growing seasons in an open-field fumigation facility where they were irrigated during the growing season with a nutrient solution providing nitrogen (N) at 70 (LN treatment), 100 (control) or 150% (HN treatment) of the optimum supply rate. Treatment effects were most evident during the third year of exposure, when the ambient [O3] + HN treatment enhanced whole-plant biomass, root/shoot dry weight ratio, needle pigment concentrations and the number of chloroplast plastoglobuli in the mesophyll cells in current-year (C) needles, whereas it reduced starch accumulation in C needles and abscission of 2-year-old (C+2) needles. In the control fertilization, 3 years of exposure to elevated [O3] decreased stem-base diameter and increased K concentration and electron density of chloroplast stroma in C needles. Plants in the HN treatment exposed for 3 years to elevated [O3] had significantly lower heights, current-year main shoot length and root/shoot dry mass ratio than control plants, and increased abscission of C+2 needles. In contrast, O3-induced changes in the ultrastructure of mesophyll cells were most evident in seedlings grown for 3 years in the LN treatment. We conclude that, in Scots pine, a relatively O3-tolerant species, chronic O3 exposure leads to cumulative growth reduction, increased needle abscission and changes in carbon allocation that are strongly influenced by plant N availability.     

The effect of late summer fertilization on the frost hardening of second-year Scots pine seedlings

In this study the effect of summer fertilization on the initiation of frost hardening of containerized second-year Scots pine (Pinus sylvestris L.) seedlings is studied. During the second growing season three different fertilization programs (water soluble NPK with micronutrients) determined by electrical conductivity of peat water extract (0.2, 0.5 and 1.2 mS cm^-1) were initiated. The growth and nutrient concentrations of needles were monitored during the fertilization period. The frost hardiness of seedlings was assessed on four separate occasions at two week intervals from August 7 to September 18. This assessment was based on artificial freezing tests and visual damage scoring of tissue browning on current-year needles. Clear differences in foliar N, P and K concentrations were observed between the fertilization treatments. Fertilization prolonged the growing period of needles and increased root collar diameter. In all the tests, the highest fertilization level resulted in the highest level of frost hardiness. The difference between the fertilization treatments ranged from 1 °C to 2.2 °C. Frost hardiness increased with an increase in foliar nitrogen concentration and slightly less consistently with increases in foliar phosphorus and potassium concentrations.     

Effects of boron fertilization in the nursery or after planting on the performance of Norway spruce seedlings on boron-poor sites

Many forest soils in Finland are poor in boron (B), especially otherwise fertile (nitrogen-rich) soils and peatlands. Boron deficiency causes growth disorders affecting young Norway spruce (Picea abies) stands. We studied the effects of B fertilization on the growth and morphology of Norway spruce seedlings. Boron was applied in a nursery and/or after planting in a nursery-field and on a reforestation site, both known to be low in B. The B applied in the nursery increased foliar B concentration in a linear relationship to the amount of B fertilizer but did not affect seedling growth and morphology. Even the highest foliar B concentration (approx. 400 mg kg^?1) did not have harmful effects on the seedlings. Therefore, B fertilization in the nursery can be used for increasing B storage in needles. Boron fertilization at planting increased foliar B concentration in the first season. Afterward, the concentration decreased but remained above the deficiency limit for at least 4–5 years. In the B-poor nursery-field, nonfertilized seedlings began to produce multiple leaders in the second year. On the reforestation site, fewer multiple leaders were found. At the nursery-field site, height growth was stimulated by B fertilization at planting. In conclusion, 200–400 mg m^?2 (2–4 kg ha^?1) of B applied at planting has longer-lasting effects on seedling B status than does B fertilization applied in the nursery, and can improve seedling quality and growth in the first years after planting on B-poor sites.