Distribution and retranslocation of (15)N lodgepole pine over eight growing seasons

We studied the distribution and retranslocation of N in 11-year-old Pinus contorta Dougl. trees following a winter application of N at 100 kg ha(-1) as (15)N-urea, (15)NH(4)NO(3) or NH(4) (15)NO(3). In all treatments, there was little uptake of (15)N after the first growing season although labeled N was still present in the soil. In subsequent years, (15)N in the trees was partly retranslocated, and, at the same time, it was diluted by uptake of unlabeled N from the soil. Between Years 1 and 8 after N fertilization, net retranslocation of (15)N from the lower crown (branches formed before fertilization) was 14%, and 18-25% of the (15)N in the trees was translocated to the upper and mid-crown. Overall, uptake of (15)N from nitrate was less than from urea or ammonium. However, when compared with the urea- and ammonium-N sources, (15)N from the nitrate source initially moved as rapidly into the foliage, but a greater proportion of it was retranslocated from the foliage during the second growing season. Nitrogen in foliage and wood formed in the growing season following fertilization was more highly labeled (measured as % N derived from the fertilizer) than in recently formed tissues. Labeling was substantially higher in foliage formed before fertilization than in wood of a similar age. In contrast, N in foliage formed after fertilization had only slightly higher labeling than wood of a similar age, indicating a relatively stable labeling throughout the trees once (15)N uptake had ceased. The concentrations of total and labeled N were substantially higher in foliage than in either wood or bark. There was evidence of N movement into wood tissues formed before fertilization, presumably along rays, and also of N retranslocation out of xylem cells as they matured. This study of internal N cycles was facilitated by the use of (15)N labeling because there was little uptake of labeled N after the first growing season, whereas interpretation based on total N was obscured by substantial uptake of N from the soil. We conclude that retranslocation studies based on measurements of total N content should be avoided.     

Influence of fertilizer timing on seasonal nutrient uptake and dry-matter production by young Pinus radiata in southern western Australia

Seasonal changes in above ground dry-matter, nitrogen (N), and phosphorus (P) accumulation were measured following application of N and P in autumn or spring to 1-year-old Pinus radiata (D. Don). Dry-matter production and nutrient accumulation were measured eight times over two years following fertilization.

All trees produced dry-matter throughout the year, but during the summer, fertilized trees produced more dry-matter than unfertilized trees. In contrast to dry-matter production, nutrient accumulation showed a distinctly seasonal pattern with maximum accumulation of N and P occurring in winter and spring, when rainfall and soil moisture were highest. Accumulation of N and P either slowed markedly or ceased during summer depending on fertilizer treatment. Continued dry-matter production during summer, when nutrient accumulation was low, resulted in the decline of N and P concentrations in needles, branches and stems of all trees. This indicated that nutrients required for new growth during summer were mobilized from existing foliage and wood. Fertilization increased the concentrations of N and P in foliage and wood, and these higher concentrations persisted through summer. Spring fertilization increased N accumulation to a greater extent than autumn fertilization, this effect lasting two years. The greater dry-matter production by fertilized trees during summer indicated that growth during summer was limited by nutrient supply.     

Vascular tissue anatomy of Norway spruce needles and twigs in relation to magnesium deficiency

Vascular tissues of needles and twigs of Norway spruce with low foliar magnesium concentrations were examined by light microscopy. Observations were made on samples from (1) apparently healthy trees, (2) trees exhibiting progressive symptoms of decline, including the yellowing and fall of needles and the death of twigs and branches in the subapical canopy, and (3) trees recovering from symptoms of decline after fertilization with magnesium sulfate. Abnormalities in cambium and phloem anatomy, which were apparent in 2-year-old needles of declining trees, were seen only in needles at least 4 years old in healthy trees, suggesting that needles of declining trees are susceptible to precocious aging. Abnormalities in xylem anatomy were observed in needles 3 years or more in age in declining trees, but were never seen in needles of healthy trees. Over time, anatomical abnormalities in declining trees were seen in progressively younger twigs, which may explain the acropetal development of decline symptoms. After fertilization with magnesium sulfate, normal phloem and xylem were observed in the newly formed tissues.     

Element concentrations and storage in the aboveground biomass of limed and unlimed Norway spruce trees at Höglwald

Twenty years after the application of dolomitic limestone to a well growing, mature Norway spruce stand at Höglwald, Southern Germany an amount of magnesium equal to the input by liming has left the rooting zone. However, 80% of the applied calcium was retained by the ecosystem. A detailed biomass investigation was carried out in order to elaborate whether the stand shows any nutritional benefits and if the aboveground tree biomass represents a sink for calcium and magnesium after liming. Stem wood, bark, branches, twigs, and needles were analysed for the major nutrients. Needles showed higher calcium and magnesium concentrations and significantly lower potassium concentrations on the limed plot. Surprisingly, limed trees had significantly lower magnesium concentrations in the bark but higher concentrations in twigs and needles than unlimed trees. In addition, significantly lower concentrations for potassium, sulphur, and phosphorus were measured in the bark of limed trees, whereas the element concentrations in the wood were rather similar between the two treatments. The low potassium concentrations in bark and needles indicate an impeded potassium uptake after liming presumably due to a calcium potassium antagonism. Magnesium is probably used to compensate the relative lack of potassium. The small differences in the calculated nutrient storage of limed and unlimed trees, however, suggest that the aboveground biomass can be excluded as a relevant sink for the applied calcium and magnesium at the Höglwald site.     

Importance of soil extractable phosphorus distribution for mature Norway spruce nutrition and productivity

Phosphorus is an essential nutrient for forest growth. In this study, we assessed the impact of soil extractable phosphorus using two simple extraction methods on nutrition and productivity of Norway spruce in sixteen mature forest stands on different bedrocks and soils in Bavaria, Southern Germany. Representative trees were sampled for needles, twigs, branches, stem bark, and stem wood. Total phosphorus content in the tree parts and soil phosphorus stock extractable with citric acid and sodium bicarbonate up to a soil depth of 80 cm were determined. We found that easily soil extractable phosphorus is a suitable indicator for estimating phosphorus uptake and stand productivity in Norway spruce. In contrast, organic layer phosphorus showed no significant correlation with aboveground biomass phosphorus contents. In the biomass, the highest phosphorus contents were measured in young needles and twigs, but the highest correlation with soil phosphorus was detected for phosphorus contents in needles and bark. The stock of phosphorus extracted by citric acid down to 40 cm soil depth revealed the best correlation with phosphorus in needles and bark. Therefore, as a supplemental or alternative method to needle analysis, our study suggests the use of phosphorus contents in stem bark to evaluate tree phosphorus nutrition. These results highlight the suitability of the citric acid soil extraction method to characterize plant available phosphorus in Norway spruce ecosystems.     

Needle and stem wood production in Scots pine (Pinus sylvestris) trees of different age,size and competitive status

We studied effects of tree age, size and competitive status on foliage and stem production of 43 Scots pine (Pinus sylvestris L.) trees in southern Finland. The tree attributes related to competition included foliage density, crown ratio and height/diameter ratio. Needle mass was considered to be the primary cause of growth through photosynthesis. Both stem growth and foliage growth were strongly correlated with foliage mass. Consequently, differences in growth allocation between needles and stem wood in trees of different age, size, or position were small. However, increasing relative height increased the sum of stem growth and foliage growth per unit foliage mass, indicating an effect of available light. Suppressed trees seemed to allocate more growth to stem wood than dominant trees, and their stem growth per unit foliage mass was larger. Similarly, trees in dense stands allocated more growth to stem wood than trees in sparse stands. The results conformed to the pipe model theory but seemed to contradict the priority principle of allocation.     

The influence of fertilizers on the occurrence of Armillaria mellea in Scotch pine plantations

The chemical analysis of needles and wood of roots of trees belonging to four stages of disease caused by A. mellea in fertilized (N, P, K, Ca, Mg) and non-fertilized three Scotch Pine plantations were made. The attack caused gradual interruption of absorption and resulted in the exhaustion of some elements (N, P) and concentration of other ones (K, Ca) in needles and in the decrease of P and transitional growth of Mg-level in wood of roots. Considerable growth of concentration of some chemical substances in wood of roots of dead trees was observed. This referred to the fertilized and nonfertilized trees as well. The rate of these changes, however, was different after treatment: fertilization had no effect on the concentration of K in needles of trees in intermediate stages of disease, but it influenced in the lower rate the exhaustion of Mg in wood of roots. The role of K and Mg in defensive reactions is discussed.     

Seasonal changes in amino acids, protein and total nitrogen in needles of fertilized Scots pine trees

Seasonal changes in amino acids, protein and total nitrogen in needles of 30-year-old, fertilized Scots pine (Pinus sylvestris L.) trees growing in Northern Sweden were investigated over two years in field experiments. The studied plots had been fertilized annually for 17 years with (i) a high level of N, (ii) a medium level of N, or (iii) a medium level of N, P and K. Trees growing on unfertilized plots served as controls. In control trees, glutamine, glutamic acid, gamma-aminobutyric acid, aspartic acid and proline represented 50-70% of the total free amino acids determined. Arginine was present only in low concentrations in control trees throughout the year, but it was usually the most abundant amino acid in fertilized trees. Glutamine concentrations were high during the spring and summer in both years of study, whereas proline concentrations were high in the spring but otherwise low throughout the year. In the first year of study, glutamic acid concentrations were high during the spring and summer, whereas gamma-aminobutyric acid was present in high concentrations during the winter months. This pattern was less pronounced in the second year of investigation. The concentrations of most amino acids, except glutamic acid, increased in response to fertilization. Nitrogen fertilization increased the foliar concentration of arginine from < 1 micromol g(dw) (-1) in control trees to a maximum of 110 micromol g(dw) (-1). Trees fertilized with nitrogen, phosphorus and potassium had significantly lower arginine concentrations than trees fertilized with the same amount of nitrogen only. Protein concentrations were similar in all fertilized trees but higher than those in control trees. For all treatments, protein concentrations were high in winter and at a minimum in early spring. In summer, the protein concentration remained almost constant except for a temporary decrease which coincided with the expansion of new shoots. Apart from arginine, the amino acid composition of proteins was similar in all treatments.     

Fall fertilization enhanced nitrogen storage and translocation in Larix olgensis seedlings

Fall nutrient loading of deciduous forest nursery seedlings is of special interest because of foliage abscission and varied translocation patterns. For non-deciduous seedlings in the nursery, fall fertilization typically can reverse nutrient dilution and possibly increase nutrient reserves; however, this technique has received little attention with deciduous conifer trees that translocate nutrients before abscising foliage. We evaluated how fall nitrogen (N) fertilization affected N storage and translocation in the deciduous conifer Olga Bay larch (Larix olgensis Henry) seedlings during the hardening period. Seedlings were supplied with 25 mg N seedling^?1 for 15 weeks before hardening and fall fertilization treatments began with a three week application period of K¹⁵NO₃ at 0, 5, 10 and 15 mg N seedling^?1. During the hardening period, fall N fertilization had little effect on seedling morphology. The N concentration and content of needles decreased dramatically as needles abscised, while that of stems and roots increased. Six weeks after fall N fertilization ceased, all seedlings translocated similar net N from their needles. For the control seedlings, this accounted for 84 % of the N stored in stems and roots. For fall fertilized seedlings, however, the proportion of N stored in stems and roots translocated from needles accounted for only 41–61 % of the total because of absorption of fall fertilizer that was translocated directly to stems and roots. Six weeks after fall fertilization, the distribution pattern of N concentration and content in seedlings was found in this order: stems > fine roots > coarse roots > needles. Our results suggest that providing deciduous conifer seedlings N during hardening, in this case Olga Bay larch, is a way to promote nutrient loading during nursery production.     

Effect of pruning on nitrogen dynamics within crowns of Pinus radiata

Stem injection of (15)N-labeled ammonium sulfate was used to determine effects of pruning on canopy nitrogen dynamics in open-grown Pinus radiata D. Don in New Zealand. Trees were planted in July 1990 and the isotope introduced in December 1994. Tree crowns were divided into three zones: base section, from which branches of pruned trees were removed; mid section, between the pruned zone and the height of the trees at the start of the year in which they were pruned; and top section, which grew predominantly after the isotope was applied. Pruning removed 32% of the green crown length, representing 75% of foliage biomass. Needles were sampled from each region of the crown until July 1996. Branch growth was used to predict foliage biomass for each sampling occasion. Approximately 45% of the applied isotope was recovered from needles sampled in December 1994 (1 week after application and immediately before pruning), two-thirds of which occurred in needles in the base section. Thereafter, changes in isotope content of needles in the base section of unpruned trees largely reflected foliage biomass fluctuations and dilution of the isotope by continued uptake from the unlabeled soil nitrogen pool. Recovery of isotope in needles from the mid-crown section increased by 58 and 86% from December 1994 to July 1995 in control and pruned trees, respectively. Within this crown section, there was evidence of isotope translocation from old to new needles, with both isotope dilution and efflux observed in the needle cohorts that had been present at the time the isotope was applied. Therefore, isotope dynamics did not reflect the dynamics of the total nitrogen pool in the mid-crown section. By July 1996, a small proportion of the applied isotope was recovered from the new foliage formed in the top section of the crown. Within the top section, isotope dynamics closely matched total nitrogen fluxes. Pruning the lower crown did not affect nitrogen dynamics elsewhere in the crown for the following 18 months.     

Soil Properties Affecting the Frost Sensitivity of Beech Bark in Southern Sweden

Decline in pH and leakage of base cations have been recorded in beech forest soils in southern Sweden. This study investigated the influence of soil properties on frost sensitivity in beech bark and evaluated the effects on frost sensitivity of soil treatment with lime, wood ash or excessive N fertilization. Beech trees on five experimental sites were studied. The results indicated that beech trees subjected to a comparatively low C/N ratio and low concentration of nutrient elements in the mineral soil layer were predisposed to a higher sensitivity to frost. Seven years after treatment, no effects of N fertilization were detectable in the soil, but positive effects of liming were recorded. There were no obvious effects of bark ash amendments, owing to large differences among blocks at the site.     

Cesium absorption through bark of Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis>)

Absorption of radiocesium (¹³⁷Cs and ¹³⁴Cs) through bark, and its subsequent translocation into wood and needles, has been suggested as a potential source of tree contamination, but the process is not well understood. Field experiments were conducted to confirm whether Cs could enter a Japanese cedar tree through the bark and how Cs moves within a tree. Stable Cs (¹³³Cs) was applied to the bark at 1.2-m height on 10- and 26-year-old Japanese cedars. The ¹³³Cs concentrations were determined in the bark, sapwood, and heartwood (for 26-year-old cedar only) of stem disks from several heights, as well as in current-year needles from the canopy. The ¹³³Cs concentrations were considerably higher in the sapwood and heartwood of stem disks from 1.2-m height in treated trees than in untreated trees, suggesting that ¹³³Cs penetrated the bark to enter the wood. The average ¹³³Cs concentrations were higher in the heartwood than the sapwood, indicating ¹³³Cs accumulation in the heartwood. High ¹³³Cs concentrations in the needles of treated trees implied acropetal movement of ¹³³Cs to actively growing organs. Our results demonstrate that Cs can enter Japanese cedar trees through the bark and that Cs is transported radially to the heartwood and vertically to the apex.     

Influence of nitrogen fertilization on minerals, carbohydrates, amino acids and phenolic compounds in beech (Fagus sylvatica L.) leaves

Beginning in 1985, plots in a 120-year-old beech (Fagus sylvatica L.) stand in the province of Scania, southern Sweden were fertilized annually with ammonium nitrate. In August 1989, leaves from both fertilized and unfertilized trees were analyzed for mineral nutrients, soluble carbohydrates, amino acids and phenolic compounds. Leaf concentrations of total nitrogen were increased by N fertilization. Ammonium was not detected in the leaves, but NO(3) (-) was increased by up to 65% as a result of N fertilization. Nitrogen fertilization decreased foliar concentrations of P and Cu, leading to a nutrient imbalance relative to N. Of the free amino acids, glutamic acid and aspartic acid predominated and, together with glutamine and asparagine, were significantly increased by N fertilization. The concentrations of amides were 2-4 times higher in fertilized trees than in unfertilized trees. Soluble carbohydrates did not change in response to N fertilization, but total phenolic compounds decreased markedly. The reduced concentration of phenolics may explain the greater susceptibility to parasite attacks of trees exposed to N deposition and excess N.     

An evaluation of the conical approximation as a generic model for estimating stem volume, biomass and nutrient content in young Eucalyptus plantations

Accurately and non-destructively quantifying the volume, mass or nutrient content of tree components is fundamental for assessing the impact of site, treatment, and climate on biomass, carbon sequestration, and nutrient uptake of a growing plantation. Typically, this has involved the application of allometric equations utilising diameter and height, but for accurate results, these equations are often specific to species, site, and silvicultural treatment. In this study, we assessed the value of incorporating a third piece of information: the height of diameter measurement. We derived a more general volume equation, based on the conical approximation, using a diameter projected to the base of the tree. Common equations were developed which allowed an accurate estimate of stem volume, dry weight and nutrient content across two key plantation grown eucalypt species, Eucalyptus grandis W. Hill ex Maiden and Eucalyptus globulus (Labill.). The conical model was developed with plantation-grown E. grandis trees ranging from 0.28 to 15.85 m in height (1.05 g to 80.3 kg stem wood dry weight), and E. globulus trees ranging from 0.10 to 34.4 m in height (stem wood dry weight from 0.48 g to 652 kg), grown under a range of contrasting cultural treatments, including spacing (E. grandis), site (E. globulus) and fertilization (nitrogen and phosphorus) for both species. With log transformed data the conical function (V_con) was closely related to stem sectional volume over bark and stem weight (R² = 0.996 and 0.990, respectively) for both E. grandis and E. globulus, and the same regressions can be applied to both species. Back transformed data compared with the original data yielded modelling efficiencies of 0.99 and 0.97, respectively. Relationships between V_con and bark dry weight differed for the two species, reflecting differing bark characteristics. Young trees with juvenile foliage had a different form of relationship to older trees with intermediate or adult foliage, the change of slope corresponding to heights about 1.5 m for E. grandis and age 1 year for E. globulus. The V_con model proved to be robust, and unlike conventional models, does not need additional parameters for estimating biomass under different cultural treatments. More than 99% of the statistical variance of the logarithm of biomass was accounted for in the model. V_con captures most of the change in stem taper associated with cultural treatments and some of the change in stem form that occurs after the crown base has lifted appreciably. Fertilization increased N and P concentrations in stem wood and bark, and regressions to estimate N and P contents (the products of biomass and concentration) were dependent on treatment. For instance, there was a large growth response to N fertilization in E. globulus corresponding with a change (P < 0.05) in the intercept of the regression to estimate N content.     

Effects of biotic and abiotic factors on δ15N in young Pinus radiata

In a 12-year-old Pinus radiata plantation, three dominant and three suppressed trees removed by thinning were randomly selected, and needles, annual rings from basal stem disks and bark were collected and analyzed to study the relationships of climate, tree age, dominance and growth with tree δ¹⁵N. The high foliar N concentration (1.35–2.73 % N, dw) suggested that N was not limiting tree growth, therefore allowing plants to fractionate versus δ¹⁵N, leading to differences in δ¹⁵N among trees. Most wood δ¹⁵N_air values were below the δ¹⁵N_air natural abundance in the dominant pines (?2.43 to +1.69 ‰) and above it in the suppressed trees (+0.73 to +3.35 ‰), likely due to the access of dominants to exogenous N sources with lower δ¹⁵N_air than those of suppressed. However, no dominance effect was detected in δ¹⁵N_air of bark and needles that decreased in the order: buds (+1.20 to +2.44 ‰) > needles 1 year (?0.27 to +1.43 ‰) > needles 2 years (?0.97 to +0.41 ‰) > bark (?1.18 to +0.15 ‰). Compared with the soil N in the 0–15 cm layer (δ¹⁵N_air = +4.8 ‰), all plant material was ¹⁵N-depleted. Results suggest that seedlings and foliar buds have a less efficient system for N conservation and recycling, with higher losses. The linear regression models showed that both biotic (dominance and tree age) and abiotic factors (temperature in spring–summer and annual precipitation) are needed to explain the wood δ¹⁵N_air satisfactorily.     

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.     

Crown characteristics of juvenile loblolly pine 6 years after application of thinning and fertilization

Total foliage dry mass and leaf area at the canopy hierarchical level of needle, shoot, branch and crown were measured in 48 trees harvested from a 14-year-old loblolly pine (Pinus taeda L.) plantation, six growing seasons after thinning and fertilization treatments.

In the unthinned treatment, upper crown needles were heavier and had more leaf area than lower crown needles. Branch- and crown-level leaf area of the thinned trees increased 91 and 109%, respectively, and whole-crown foliage biomass doubled. The increased crown leaf area was a result of more live branches and foliated shoots and larger branch sizes in the thinned treatment. Branch leaf area increased with increasing crown depth from the top to the mid-crown and decreased towards the base of the crown. Thinning stimulated foliage growth chiefly in the lower crown. At the same crown depth in the lower crown, branch leaf area was greater in the thinned treatment than in the unthinned treatment. Maximum leaf area per branch was located nearly 3–4 m below the top of the crown in the unthinned treatment and 4–5 m in the thinned treatment. Leaf area of the thinned-treatment trees increased 70% in the upper crown and 130% in the lower crown. Fertilization enhanced needle size and leaf area in the upper crown, but had no effect on leaf area and other variables at the shoot, branch and crown level. We conclude that the thinning-induced increase in light penetration within the canopy leads to increased branch size and crown leaf area. However, the branch and crown attributes have little response to fertilization and its interaction with thinning.     

Calorific values and ash contents of different parts of Masson pine trees in southern China

Calorific value of plants is an important parameter for evalu- ating and indexing material cycles and energy conversion in forest eco- systems. Based on mensuration data of 150 sample sets, we analyzed the calorific value （CV） and ash content （AC） of different parts of Masson pine （Pinus massoniana） trees in southern China using hypothesis testing and regression analysis. CV and AC of different tree parts were almost significantly different （P〈0.05）. In descending order, ash-free calorific value （AFCV） ranked as foliage 〉 branch 〉 stem bark 〉 root 〉 stem wood, and AC ranked as foliage 〉 stem bark 〉 root 〉 branch 〉 stem wood. CV and AC of stem wood from the top, middle and lower sections of trees differed significantly. CV increased from the top to the lower sections of the tnmk while AC decreased. Mean gross calorific value （GCV） and AFCV of aboveground parts were significantly higher than those of belowground parts （roots）. The mean GCV, AFCV and AC of a whole tree of Masson pine were 21.54 kJ/g, 21.74 kJ/g and 0.90%, re- spectively. CV and AC of different tree parts were, to some extent, cor- related with tree diameter, height and origin.     

Storage and internal cycling of nitrogen in relation to seasonal growth of Sitka spruce

Three-year-old clonal cuttings of Picea sitchensis (Bong.) Carr. were grown for two years (1988-1989) in sand irrigated with a nutrient solution containing either 1.0 mol N m(-3) (low N) or 6.0 mol N m(-3) (high N) NH(4)NO(3). In 1988, all the N provided was enriched with (15)N to 4.95 atom % (labeled N). In 1989, N was supplied with (15)N at natural abundance (unlabeled N). The recovery of unlabeled and labeled N in new foliage was used to quantify the internal cycling of N. In the high-N treatment, trees had two flushes of shoot growth and a period of rapid root growth, which coincided with the second flush of shoot growth in August. The timing of root growth and the first flush of shoot growth was similar in the low-N treatment, but there was no second flush of shoot growth and a greater proportion of biomass was recovered in roots. By November 1989, the root/needle dry matter ratio was 1.95 for the low-N trees and 1.36 for the high-N trees. Nitrogen was stored overwinter in roots and current-year needles. During the first six weeks of growth in the spring of 1989, stored N was remobilized for new foliage growth. Subsequent growth depended on root uptake of N. Remobilization of stored N was apparently not affected by the current N supply, because the amount of unlabeled N recovered in foliage produced in 1988 was the same for both N treatments. During 1989, the proportion of (15)N remobilized from roots relative to that from leaves produced in 1988 was greater in low-N trees than in high-N trees. In the autumn of both years, there was rapid uptake of N into roots and current-year needles. The effects of N supply on tree growth and nitrogen use efficiency are discussed in terms of the capacity for both N storage and internal cycling.     

Nutrient dynamics in singleleaf pinyon (Pinus monophylla Torr & Frem.) needles

Nutrient retranslocation from existing foliage of singleleaf pinyon pine (Pinus monophylla Torr. & Frem.) did not meet the nutrient demands of new needles on the same branch. Singleleaf pinyon pines, with an average age of 177 years, located at an elevation of 2300 m on a ridge in The Sweetwater Range in Nevada, USA retained needles for 18 or more years and increased needle number per branch by 17-18% annually. With age, needles gained weight and accumulated Ca, Mg, Fe, Zn, P, and Mn. The amount of N per needle showed no consistent trend with needle age. Only Cu and K declined in concentrations and amounts with needle age and these differences were not statistically significant. Retranslocation of nutrients at needle abscission could provide only 2 to 13% of the N, P, K, Fe, Cu, and Mg required by new needles. We conclude that nutrient capital in existing foliage constitutes a long-term nutrient reserve for the tree rather than a mobile nutrient pool.